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My dear Sir James, I I cannot more approj^riately inscribe this Treatise, having for its object the general diffusion of sound Physio- logical knowledge, than to one whose Professional eminence ' is foiuided on his enlightened application of it to the pre- vention and cure of Disease, and who has ever been the consistent advocate of Liberal Education. The grateful sense I entertain of many acts of personal kindness, makes me feel additional pleasure in paying this ' humble tribute. Believe me to remain. My dear Sir James, Your obliged Friend and Servant, WILLIAM L. CAPvPENTEK. University Hall, London, January, 1859. Ik PREFACE. The issue of the present Volume may be considered as an attempt to supply what the Author has long considered to be a deficiency in the literature of this countrj'', — that, namely, of an Educational Treatise on Animal Physiology, which should at the same tmie communicate to its readers the facts of gi-eatest importance as regards theii' practical bearing, and present these in such a form as to place the learner in possession of the essential j^rinciples of Physiological Science. The Author has followed the general plan of the Treatise on Animal Physiology contributed by Professor Milne-Edwards, one of the most eminent Naturalists in France (in which country it is not thought beneath the dignity of men of the highest scientific reputation to write elementary books for the instruction of the beginner), to the " Coui^s Elementaire d'Histoii^e Xaturelle" adopted by the French Government as the text-book of instruction in the Colleges connected with the University of Paris, which requires from every Candidate for its Degree of " Bachelor of Sciences " a competent know- ledge both of Animal and of Vegetable Physiology. He has also had at his disposal the admirable series of Illustrations prepared for that work, which, as a whole, are unsurpassed either in beauty or in exactness. In carrying-out this plan, however, the Author has entirely followed his own judgment ; and has made so much more use b VI PREFACE. of his own materials than of those siippUed by the treatise of Professor Milne-Edwards, that the work may be regarded as almost entirely original. The present Edition, too, has undergone very considerable modifications ; the first chapter, which now contains a complete outline of the Elementary Tissues of the Animal Body, and the last, in which a com- prehensive sketch is given of the principal ])lienomena of Reproduction and Development throughout the Animal Kingdom, having been entirely re-written and illustrated with numerous additional figures. In order to make room for the large amount of new matter now introduced (not less than one-fifth of the entire volume), the second chapter, contain- ing a General View of the Animal Kingdom, has been much abridged ; — a change the Author has the less regretted ])eing obliged to make, since there are now before the public several excellent Elementary Treatises on Zoology, which had no existence at the time this volume originally appeared. Everyone Avho desires to see the study of Physiology duly appreciated as a branch of General Education, must feel gratified at the progress which has been made of late years in the public recognition of its value. The University of London led the way, by the introduction of Animal Physiology into the programme of study to which all Candidates for its Degree of Bachelor of Arts are required to conform. The Universi- ties of Oxford and Cambridge have since admitted it as one of the subjects which Candidates may select for their Bachelor of Arts Examination, and in which they may obtain Honours. And in many of the large Public Educational Institutions with Mdiicli this country is now so abundantly furnished, it forms a 2:>art of the regular course of instruction. It has been the Author's steady aim, not merely to adapt his treatise to the Avants of those who wish to acquire a general knowdedge of the principal facts and doctrines of Physiological Science, but also to render it suitable to that PREFACE. Vll which he considers a far more important purpose of the study, — namely, the culture and discipline of the Mind itself. Having been satisfied, by no inconsiderable experience of ditterent modes of Education, that Natural Science, if judiciously taught, is second in value to no other subject as an educational means, and that it may be made to call forth a more varied and wholesome exercise of the mental powers than almost any other taken singly, — he has kept this purpose constantly in view ; and he trusts that the experience of intelligent In- structors will be found so far to concur with his own, that the study of Physiology may be still more generally introduced into Popular Education. It can only be by the general diffu- sion of sound information on this subject, that the Public Mind can be led to understand the difference between Rational Medicine, and that Empiricism which now presents itself under so many different forms ; that it can appreciate the true value of measures of Sanitary Eeform, the efficiency of which must depend upon the amount of support they receive from an intelligent public opinion ; and that it can be preserved from those Epidemic Delusions, whose preva- lence, from time to time, is not less injurious to the minds of which they lay hold, than is that of Epidemic Diseases to the bodies of those who suffer from them. He has only further to add that, whilst keeping in view the most important practical applications of the Science of Physiology, he has not thought it desirable to pursue these too far; since they constitute the details of the Art of pre- serving Health, which is founded upon it, and which may be much better studied in a distinct form, when this outline of the Science has been mastered. And, for the same reason, he has adverted but slightly to those inferences respecting the Infinite Power, Wisdom, and Goodness, of the Great First Cause, which are more obvious, although, perhaps, not really more clear and valid^ in this Science, than in any other. Vlll PREFACE. Believing, as he does, that such inferences are more satisfac- torily based upon the general manifestations of Law and Order, than upon individual instances of Design, he has thought it the legitimate object of this treatise to lay the foundation for them, by developing, so far as might be, the Principles of Physiology, — leaving it to special treatises on Natural Theology, to build-up the applications. University Hall, London, Jan. 1859. CONTENTS. TAGE INTRODUCTION 1 CHAPTER I. On the Vital Operations of Animals, and the Instruments BY which they are PERFORMED 17 CHEMICAL CONSTITUTION OF THE ANIMAL BODY ... 31 STRUCTURE OF THE PRIMARY TISSUES 36 CHAPTER II. General View of the Animal Kingdom 84 Vertebrata 84 mammals 89 BIRDS 02 reptiles 93 batrachdv 97 fishes 100 Abticulata 102 insects 104 arachnida 105 crustacea 106 cirrhifeda 109 myriapoda 110 annelida ?v>. ENTOZOA Ill MOLLUSCA 112 CEPHALOPODA 116 PTEROPODA 117 GASTEROPODA 118 CONCHIFERA ib. TUNICATA 121 POLYZOA 122 CONTENTS. CHAPTER 11.— Continued. PAGE Radiata 123 echinodepmata 125 ACALEPH^ 128 polypifera 129 Protozoa 185 rhizopoda 136 infusoria 139 porifera 140 CHAPTER III. Nature and Sources of Animal Food 142 CHAPTER IV. Digestion and Absorption 162 prehension of food 163 mastication 166 insaliyation 176 deglutition 178 digestive apparatus 181 gastric digestion : — CHyMIFICATION 188 intestinal digestion :— chylification 193 defecation 195 absorption of nutritive MATERIAL 196 SANGUIFICATION 199 CHAPTER V. Of the Blood, and its Circulation 202 properties of the blood 203 circulation of the blood 216 circulating apparatus of THE HIGHER ANIMALS . . 222 FORCES THAT MOVE THE BLOOD 232 COURSE OF THE BLOOD IN THE DIFFERENT CLASSES OF ANIMALS 240 CONTEXTS. CHAPTER VI PAGE Of Respiration 258 nature of the changes essentially constituting respiration 259 structure and actions of the respiratory apparatus 265 CHAPTER VII. Of Excretion and Secretion 292 general purposes op the excreting processes . . ib. nature of the secreting process.— structure of the secreting organs 298 characters of particular secretions 304 CHAPTER VIII. General Review of the Nutritive Operations. — Formation OF the Tissues 316 GENERAL REVIEW OF THE NUTRITIVE OPERATIONS , . ih. FORMATION OF THE TISSUES 317 REPAIR OF INJURIES 323 CHAPTER IX. On THE Evolution of Light, Heat, and Electricity by Animals 327 animal luminousness ih. ANIMAL heat 332 animal electricity 340 CHAPTER X. Functions of the Nervous System 345 structure AND ACTIONS OF THE NERVOUS SY'STEM IN THE PRINCIPAL CLASSES OF ANIMALS 350 FUNCTIONS OF THE SPINAL CORD. — REFLEX ACTION . . 374 FUNCTKJNS OF THE GANGLIA OF SPECIAL SKNSK. — CON- SENSUAL ACTIONS 380 FUNCTION OF THE CEREBELLUM. — COMBINATION OF MUS- CULAR ACTIONS 384 FUNCTION OP THE CEREBRUM. INTELLIGEXCI': AND WILL . 385 Xll CONTENTS. CHAPTER XI. PAGE On Sensation, and the Organs of the Senses 387 sense of touch 390 sense op taste 395 sense of smell 398 sense of hearing 401 sense of sight 413 CHAPTER XII. Op Animal Motion, and its Instruments 443 contractile tissues.— muscular contractility . . 444 applications of muscular power. — bones and joints . 453 motor apparatus op man. — skeleton and muscles , 464 of the attitudes of the body, and the various kinds of locomotion . 489 CHAPTER XIII. Of THE Production of Sounds : Voice and Speech .... 513 CHAPTER XIV. Of Instinct and Intelligence 525 manifestations of intelligence 546 CHAPTER XV. Of Reproduction 552 gemmiparous or non-sexual reproduction .... 553 sexual reproduction, or generation 557 ANIMAL PHYSIOLOGY. inteoductiojn-. The importance of tho study of Animal Physiology, as a branch of General Education, can scarcely be over-estimated ; and it is remarkable that it is not more generally appreciated. It might have been supposed that curiosity alone would have led the mind of Man to the eager study of those wonderful actions by which his body is constructed and maintained ; and that a knowledge of those laws, the observance of which is necessary for the due performance of these actions, — in other words, for the maintenance of his health, — would have been an object of universal pursuit. That it has not hitherto been so, may be attributed to several caasea The very familiarity of the occurrences is one of these. We are much more apt to seek for explanations of phenomena that rarely present themselves, than of those which we daily witness. The Comet excites the world's curiosity, whilst the movements of the sun, moon, and planets are regarded as things of course. AYe almost daily see vast numbers of animals of different tribes, in active life around us ; their origin, growth, movements, decline, death, and reproduction, are continually taking place under our eyes ; and there seems to common apprehension nothing to explain, where everything is so apparent. And of Man too, the ordinary vital actions are so famiUar, that the study of their conditions appears superfluous. To be born, to grow, to be subject to occasional disease, to decline, to die, is his lot in common with other animals ; and what know- ledge can avail (it may be asked) to avert the doom imposed on him by his Creator ] B 2 INTRODUCTION. In reply to this it is sufficient to state, tliat millions annually perish from a neglect of the conditions which Divine wisdom has appointed as requisite for the preservation of the hody from fatal disease ; and that millions more are constantly suffering various degrees of pain and weakness, that might have been prevented by a simple attention to those principles which it is the province of Physiology to unfold. From the moment of his birth, the infant is so completely subjected to tlie in- fluence of the circumstances in which he is placed, that the future development of his frame may be said to be governed by them ; and thus it depends, in great part, npon the care with wdiich he is tended, and the knowledge by which that care is guided, whether he shall grow up in health and vigour of body and mind ; or shall become weakly, fretful, and self- willed, a source of constant discomfort to himself and to others ; or shall form one of that vast proportion, whose lot it is to be removed from this world before infancy has ex- panded into childhood. The due supply of warmth, food, and air are the principal points then to be attended to ; and on every one of these the greatest errors of management prevail. Thousands and tens of thousands of infants annually perish durmg the few first days of infanc}'', from exposure to cold, which their feeble frames are not yet able to resist ; and at a later period, when the infant has greater power of sustain- ing its own temperature, and is consequently not so liable to sutler from this canse, the seeds of future disease are sown, b}'- inattention to the simple physiological principles, which should regulate its clothing in accordance with the cold or lieat of the atmosphere around. ISTor is less injury done by inattention to the due regulation of the diet, as to the quan- tity and quality of the food, and the times at which it should be given ; the rules for which, simple and easy as they are, are continually transgressed through ignorance or carelessness. And, lastly, one of the most fertile sources of infantile dis- ease, is the want of a due supply of pure and wholesome air ; the effects of which are sure to manifest themselves in some way or other, though often obscurely and at a remote period. It is physiologically impossible for human beings to grow np in a sound and healthy state of body and mind, in the midst of a close, ill-ventilated atmosphere. Those that arc least able to resist its baneful influence, are carried ofl" by the dis- INTRODUCTION. 6 eases of infancy and cliildliood ; and those whose native vigour of constitution enables them to struggle through these, become the victims, in later years, of diseases which cut short their term of hfe, or deprive them of a large part of that enjoyment which health alone can bring. Kor is the effect of these injurious causes confined to infancy, though most strikingly manifested at that period, " The child is father to the man," in body as well as in mind; but the vigorous health of the adult is too often wasted and destro3'ed by excesses, whether in sensual indulgence, in boddy labour, or in mental exertion, to which the very feeling of buoyancy and energy often acts as the incentive ; and the strength which, carefully husbanded and sustained, might have kept the body and mind in activity and enjoyment to the full amount of its allotted period of "threescore years and ten," is too frequently dissipated in early manhood. Or, again, the want of the necessary conditions for the support of life, — the warmth, food, and air, on wliich the body depends for its continued sustenance, no less than for its early deve- lopment, — may cause its early dissolution, even where the individual is gmiltless of having impaired its vigour by his own transgressions. These statements are not theoretical merely : they are based upon facts drawn from observations carried oh upon the most extensive scale. Wherever we find those conditions, which the Physiologist asserts to be most favourable to the preservation of the health of the body, most completely fulfilled, there do sickness and mortality least prevail. A few facts will place this subject in a striking light. " The average mortality of infants among rich and poor in this country (and with little variation throughout Europe) is about one in every four and a-half before the end of the first year of existence. So directly, however, is infant life influenced by good or bad management, that, about a century ago, the workhouses of London presented the astounding result of twenty-three deaths in every twenty- four infants under the age of one year. For a long time this frightful devastation was allowed to go on, iis beyond the reach of human remedy. But when at last an improved system of management was adopted in consequence of a parliamentary inquiry having taken place, the proportion of deaths was speedily reduced from 2, GOO to 450 in a year. Here, then,. B 2 4 INTRODUCTIOX. was a total of 2,150 instances of loss of life, occnrring yearly in a single institution, cliargeable, not against any unalterable decrees of Providence, as some are disposed to contend as an excuse for tlieii' own negligence ; but against tlie ignorance, indiiference, or cruelty of man. And what a lesson of vigi- lance and inquiry' ought not such occurrences to convey, when, even now, with all our boasted improvements, every tenth infant still perishes within a month of its birth /" ^ The effect of attention to cleanliness and ventilation in the reduction of an excessive infantile mortality, has been equally shown in the experience of the Dublin I.ying-in Hospital. At the conclusion of 1782, it was found that out of 17,6.50 inflmts born alive, no fewer than 2,944, or one in every six, had died within the lirst fortnight. By the more cihcient ventilation of the wards, the proportion of deaths during the lirst fortnight was at once reduced to 419 out of 8,033, or but little more than one in twenty; and it has subsequently been still further dimmished. In the island of St. Ivilda, the most northern of the Heb- rides, according to the statement of a gentleman who visited it in 1838, as many as eiyht ont of tvery ten children die between the eighth and twelfth day of their existence ; in consequence of which terrible mortality, the population of the island is diminishing rather than increasing. This is due, not to anything injurious in the position or atmosphere of the island ; for its ** au' is good, and the water excellent : " but to the " tilth in which the inhabitants hve, and the noxious effluvia which pervade their houses." The huts are small, low- roofed, and without windows ; and are used during the winter as stores for the collection of manure, which is carefully laid out upon the floor, and trodden under foot, till it accumulates to the depth of several feet. The clerg}-man, who lives exactly as those around him do^ in every respect, except as regards the condition of his house, has reared a famil}-of four children, all of whom are well and healthy ; whereas, accord- ing to the average mortality arouud him, at least three out of the four would have been dead within the lirst fortnight. It is not a little remarkable that a recent sanitary inquiry carried ov.t by order of the Danish govermnent, into the con- ^ Dr. A. Combe on the Physiological and Moral Management of luiauej. INTRODUCTION. dition of the Icelandic population, should have disclosed the existence of almost precisely similar hahits of life among them, with almost precisely the same results. The dwellings of the great bulk of the peasantry seem as if constructed for the express purpose of poisoning the air which they contain. They are small and low, without any direct provision for ventilation, the door serving alike as window and chimney ; the walls and roof let in the rain, which the floor, chiefly composed of hardened sheep's-dung, sucks up ; the same room generally serves for all the uses of the whole 'family, and not only for the human part of it, but frequently also for the sheep, which are thus housed during the severest part of the winter. The fuel employed in this country chiefly con- sists of cow-dung and sheep's-dung, caked and dried ; and near the sea-coast, of the bones and refuse of fish and sea- fowl ; producing a stench, which to those unaccustomed to it is completely insupportable. In addition to this, the people are noted for their extreme want of personal cleanliness ; the same garments (chiefly of black flannel) being worn for months without having even been taken oil at night. Although the Icelanders enjoy an ahnost complete exemption from many diseases (such as consumption) which are very fatal elsewhere, and the number of births is fully equal to the usual average, the population of the island does not increase, and in some parts actually diminishes. This result is in great measure due, as at St. ELilda, to the very high rate of infiantile mortality ; a large proportion of . all the infants born being carried olf before they are a fortnight old. It is in the httle island of AVestmannoe, and the opposite parts of the coast of Iceland, v/here the bird-fuel is used all the year round, instead of (as elsewhere) during a few months only, that the rate is the highest; the average mortality for many years having been sixty-four out of every hundred, or nearly two out of three, of all the infants born in these locaUties, But it is yet more remarkable that the immediate cause of the high rate of infantile mortality should have been pre- cisely the same in the Workhouses of London, the Lying-in Hospital of Dublin, and the close filthy huts of the peasantry of Iceland and St. Kilda ; for it w^as almost entirely referrible to one single disease, " Trismus nascentium," or, " Lock-jaw of the ISTew-born ; " and this disease has diminished in exact 6 INTRODUCTION. proportion to the improvement of the places it previously infested, in respect to ventilation and cleanliness. Thus, it is 80 rare for a case of it now to occur in London, that many practitioners of large experience have never seen the disease. In the Dublin Lying-in Hospital, the number of deaths from it has been reduced to three or four yearly. And there can- not be a reasonable' doubt, that, by due attention to the same conditions, it might be exterminated from Iceland and from St. Kilda. There is scarcely, in fact, a disease incident to humanity, which is more completely preventible than tliis ; and yet the annual sacrifice of life which it formerly caused in our own country alone, might have been reckoned by tens of thousands. Although the peculiar susceptibiltty of the constitution of children, gives to foul air and other causes of disease a much more destructive influence over them, than the like causes have over persons more advanced in life, yet it is now well ascertained that the rate of mortality among different classes of the community varies in a degree which bears a very close relation to the nature of the conditions under which they live. Thus, whilst the annual average number of deaths in the whole of England and Wales is about. 22 out of every thousand persons living, there are localities in which the annual average exceeds 50 in a thousand, and others in wliich it falls as lov/ as 11 in a thousand. And it is not a little remarkable, that the difference is almost entirely referrible to the mortality produced by Fevers and allied diseases, which, as experience has now fully demonstrated, are absolutely preventible by due attention to the ordmary conditions of health. As the population of England and Wales may at present be estimated at about twenty millions, and its actual mortality at about 440,000, what maybe termed its inevitable mortality — arising from diseases that would not be directly affected by sanitary improvements — would be only one half, or 220,000 ; so that the same number of lives may be considered to be annually sacrificed by the public neglect of the means of pre- serving them, — the deaths from typhus alone being no fewer than 50,000. But as it is scarcely to be supposed that every part of our population could be placed in conditions as favour- able as those which prevail where the rate of morttility is the lowest, v/e may take 13 per thousand as the average to INTRODUCTION. < whicli it may bo safely affirmed, on the basis of actual expe- rience, that the annual mortality may be reduced, by such efficient sanitary measures as render the dwellings of the mass of the population fit for human habitation ; this would give an annual mortality for England and Wales of 260,000, sho^\ing a saving of 180,000 lives annually in that one por- tion of the British empire. And it must be remembered that this amount of mortahty represents a vastly greater amount of sickness, since, for every death, there are numerous cases of severe illness ; so that it would be scarcely too much to affirm that at least a million out of the whole number of such cases annually occurring, are preventible, hke the 180,000 deaths, by adequate provisions for the supply of pure air and water, and by efficient sewerage for the removal of decomposing matters. It cannot be doubted that, even in a mere pecuniary point of view, the expense of such arrange- ments would be amply compensated by the prevention of a vast amount of .that loss of productive labour of various kinds, wjiich is at present due to disease ; and, considered on the large scale, as a question of social economy, the import- ance of sanitary legislation can scarcely be over-rated. But much cannot be expected to be done in this direction, until such an inteUlgeut puhlic opinion shall have been created, by the general dihusion of sound physiological information, as shall be sufficiently forcible to bear down the self-interested opposition of those, who do not see that the value of their pro^Derty will be permanently increased at least in proportion to the amount of money judiciously expended upon it. A more remarkable illustration of what is to be effected by sanitary improvements can scarcely be adduced, than that which is presented by the comparison between the locality termed "the Potteries," in the immediate vicinity of Ken- sington, and the " Model Lodging-houses," which have been erected in various parts of the Metropolis. The site of the group of dwellings constituting the former is far from being insalubrious in itself, and rows of handsome houses are rising up in its unmediate neighbourhood ; but the condition of these dwellings is most hltliy. A few years ago, as many as 3,000 pigs were kept in this locality (the number has since been somewhat dimmished) ; and the boiling of fat and other offal, which is carried on by some of the pig-feeders, some- 8 INTRODUCTION. times taints the air for a mile round. Very few of tlie tene- ments have any water-supply ; the wells are useless, or worse than useless, through the contamination of their water with putrescent liquid which filters down into them; and the drainage of the dwellmgs both for men and pigs is almost entirely superficial, being chiefly discharged into a stagnant piece of water called the " Ocean," which is covered with a filthy slime and bubbles with poisonous gases, and very commonly has dead dogs or cats floating on its surface. It is difficult to conceive anything more horribly offensive than the rears of some of the houses, whose yards are filled with ordure and other filth collected for manure, which is here stored for weeks, or even months, until an opportunity occurs for selling it. And even the public ways are generally covered with black putrescent mire. ISTow, during ten months of the year 1852, when no epidemic prevailed, as many as forty deaths occurred in the Potteries, out of a population of about one thousand, — the mortality being thus at the rate of 48 per thousand annually ; and no fewer i\iMi four-fifths of these deaths occurred at, or beneath, five years of age. In the first ten months of 1849, when cholera was prevalent, the number of deaths was fifty, or about one in twenty of the whole population, twenty-one of these being due to cholera and diarrhoea, and twenty-nine to typhus and other diseases. — On the other hand, in the whole population of the " Model Lodging-houses," amounting to 1,343, only seven deaths took place in the whole twelve months of 1852, or at the rate of scarcely more than 5 per thousand; and although they contain a large proportion of children, yet only half the number of deaths occurred below ten years old. Duruig the prevalence of the cholera- ej)idemic, no cases of that disease occurred among them, although it was raging in their various neighbourhoods ; and from the time that their drainage has been rendered thoroughl}^ efficient, no case of fever has presented itself among their inmates. The experience of Cholera-epidemics is peculiarly valuable, on account of the marked tendency of this disease to search out and expose defects, which have continued to produce other diseases year after year, without having been suspected as the causes of them. The greatest severity in each visita- tion has shown itseK in identical localities, provided those INTRODUCTION. 9 remained in the same foul state as at first ; whilst new loca- lities have been affected, jnst in proportion to the degree in which they have participated in the same conditions ; and those originally attacked have escaped, wherever they had adopted the requisite means of purification. Thus, at JSTewcastle-on- Tyne and Gateshead, the first outbreak occurred in the very same streets, and even in the same houses, in the three visi- tations of 1831, 1848, and 1853. An outbreak which occurred in 1853, at Luton, in Bedfordshire, — ^\;rhere, out of a population of 126 persons, inhabiting twenty-five houses, no fewer than fifty-four attacks of choleraic disease, fifteen of them fatal, took place within three weeks,' — was most dis- tinctly traceable to defect of sewerage, which had been pre- viously manifesting its malign influence on the general health of the town. And the fearful pestilence which devastated the neighbourhood of Golden Square (London) in the autumn of 1854, was no less distinctly traceable to the contamination of the pump-water by the bursting of a sewer into the well. On the other hand, Exeter and ]N"ottingham, which suffered severely in the first epidemic, escaped comparatively un- harmecl in the subsequent visitations ; and this result is plainly clue to the sanitary improvements which had been made in the interval. In 1832 there perished of the epide- mic in Exeter, as many as 402, out of a population of 28,000, or no fewer than one in seventy ; and a vast amount of suffering, with a heavy expense, was entailed upon the town. In 1848-9, on the other hand, out of a population of about 32,600, there were but 44 deaths, or less than one in seven hundred ; and upwards of one-half of these occurred in a single parish, that lies very low, and in the midst of putrid exhalations from the city drains. In Nottingham, with a population of 50,000, there were 296 fatal cases of cholera in 1832, nearly all of these being in the lower part of the town, which was ill-drained, extremely filthy, and densely popu- lated ; but in 1848-9, though the population had increased to 58,000, the number of deaths from cholera was no more than 18, all of these occurring in locaHties, which, in spite of ^hat had been done, retained much of their previous filth. The foregoing are only samples of a vast number of c^eses which might be adduced, in proof of the absolute preventi- bihty of Cholera, and of other diseases of the same class. It 10 INTRODUCTION. may be well to siil^join a few additional facts, derived from the cliolera-exiDerience of 1848-9, which, from its general diffusion, tested, in a very remarkable degree, the relative healthfulness of different provincial towns, and of different metropolitan districts. Thus, among the whole population of the ten towns of Exeter, Derby, Cheltenham, Leicester, ^Nottingham, Eochdale, jN'orwich, Preston, Halifax, and Eir- mingham, amounting to 057,000, there were no more than 238 deaths from cholera ; whilst, in an equal jitopulation inhabiting the towns of iN'ewcastle-nnder-Lyne, Plymouth, Brighton, Merthyr Tydvil, Portsea, Tynemouth, Wigan, Hull, Wolverhampton, and Leeds, the number of deaths was no fewer than 10,4:15, ov fo7'ti/-three times as great. So again, in twenty-five Metropolitan districts, chiefly on the north side of the Thames, having a total population of about 310,000, the number of deaths from cholera was only 389 ; whilst in twenty- two districts, almost entirely on the south side of the river, the number of deaths, out of a population of almost exactly the same amount, was 5,932, or more than twelve times as great. In no instance is there the least difiieulty in accounting for these contrasts. They all point to the same general conclusion; that, namely, of the immense influence which is exercised over human health by the purity of the air that is breathed, and of the water that is drunk ; and it is because these two conditions are in a great degree capable of public regulation, that legislative interference has so much in its power, and is so imperatively called for by the interests of humanity, which S23eak solemnly and distinctly to all who claim the rights of property in the foul " plague-spots " which deface our country, of their bounden diet?/ to render them not unfit for human occupation. But although the magnitude of the evils resulting from the neglect of the conditions of Public Health, gives to this sub- ject the first claim on our consideration, yet it is not the less important that every individual should acquire as much knowledge of the constitution of his body, and of the right means of keeping it in working order, as will save him from seriously damaging either himself or other peoj)le by his ignorance of such matters. It is less than ten years since a fearful sacrifice of life occurred among the deck-j)assengers on board the Irish steamer " Londonderry," who were ordered INTRODUCTION. 11 below by tlie Captain on account of the stormy cliaractcr of the weather, and on whom the hatches Avere closed down, although the cabin which was crowded l)y tlieni had scarcely any other means of ventilation. Out of 150 of these unfor- tunates, no fewer than 70 died of suffocation before the morning, — a catastrophe only second to that which occurred in the "Black Hole of Calcutta," in which 123 out of 14:6 died during one night's coniinement in a room eighteen feet square, provided with only two small windows. Yet the Captain of the " Londonderry " was acquitted of all blame ; since he had done what seemed to him best for the welfare of his passengers, the result being due simply to his astound- ing ignorance of the fact that men cannot live without having air to breathe. Not a year passes without the occurrence of numerous deaths from the like cause ; and yet these are really insignificant, when compared with the vast amount of disease which is constantly attributable to inattention, on the part of individuals, to those simple ineans of securing an adequate supply of air which are within the reach of every one. And when we bear in mind that the respiratory func- tion is only one of the processes whose due performance has to be provided for, and that the regulation of the food and drink, of the excretions, of clothing and tempera,ture, of exercise (bodily and mental) and repose, and of the repro- ductive functions, all fall within rules which it is the pro- vince of Physiology to prescribe, we see how vain it is to expect that the body can be maintained in health, without some acquaintance with that science, or at least with the rules which it lays down. For, although it is quite true that man has within himself certain instincts which afford him a considerable measure of guidance in all these particulars, — hunger and thirst, for example, leading him to take the sustenance v/hich his body requires, weariness tem^^ting hun to needed repose, and so on, — yet it is no less certain that in a state of artificial civilisation these instincts are so often over- borne by acquhed tastes, or by the pressure of other circum- stances, that they cannot alone be safely relied on. Hence it is all the more important that the rules for j)reserving health should be based on an intelligent knowledge of Physiological principles ; otherAvise, like the natural instmcts, they are likely to be put aside as occasion prompts ; Avhereas, in proportion as 12 INTRODUCTIOK the individual is possessed of tlieir rationale, will he be likely to shape his conduct in accordance with them. The general principles of Physiological science, again, will be likely to be thoroughly apprehended, in proportion as they are based on an extended recognition of the phenomena which they comprehend. Every physiologist is now satisfied that the life or vital actions of no one species of animal can be correctly understood, unless compared with those of other tribes of diiferent conformation. Hence, for the student of 13hysiology to confine himself to the observation of what takes place in Man alone, would be as absurd as for the astro- nomer to restrict liimself to the observation of a single planet, or for the chemist to endeavour to determine the properties of a metal by the study of those of that one only. There is not a single species of animal, that does not present us with a set of facts which we should never learn but by observing it ; and many of the facts ascertained by the observation of the simplest and most common animals, throw great light upon the great object of all our inquiries, the Physiolog}^ of Man. For though in liim are combined, in a most wonderful and "unequalled manner, the various faculties which separately exhibit themselves in various other animals, he is not the most favourable subject for observing their action ; for the obvious reason that his machinery (so to speak) is rendered too complex, on account of the multitude of operations it has to perform : so that we often have to look to tjie lowest and simplest animals for the explanation of what is obscure in man, their actions being less numerous, and the conditions which they require being more easily ascertained. The diffusion of Animal life is only one degree less exten- sive than that of vegetable existence. As animals cannot, like plants, obtain their support directly from the elements around, they cannot maintain hfe, where life of some kind has not preceded them. But vegetation of the humblest character is often sufficient to maintain animals of the highest class. Thus the lichen that grows beneath the snows of Lapland, is, for many months in the year, the only food of the rein-deer ; and thus contributes to the support of human races, which depend almost solely upon this useful animal for their existence. No extremes of temj^erature in our atmo- sphere seem inconsistent with animal life. In the little pools INTRODUCTION. 13 formed by the temporary influence of the sun upon the sur- face of the arctic snows, animalcules have been found in a state of activity ; and the ocean of those inhosj^itable regions is tenanted, not only by the whales and other monsters which w^e think of as their chief inhabitants, whose massive forms are only to be encountered " few and far between," but by the shoals of smaller fishes and inferior anunals of various kinds upon Avhicli they feed, and through vast fleets of which the mariner sails for many miles together. On the other hand, even the hottest and most arid portions of the sandy deserts of Africa and Asia are inhabited by animals of various kinds, provided that vegetables can find sustenance there. The humble and toilsome ants make these their food, and become in turn the prey of the cunning ant- lion and of the agile lizard ; and these tyrants are in their turn kept under by the voracity of the birds which are adapted to prey upon them. The waters of the tropical ocean never acquire any high temperature, owing to the constant interchange which is taking place between them and those of colder regions ; but in the hot springs of various parts of the world, we have examjoles of the compatibility of even the heat of almost boiling water v/ith the preservation of animal life. Thus in a hot spring at Manilla which raises the ther- mometer to 187°, and in another in Barbary whose usual tem- perature is 172°, fishes have been seen to flourish. Fishes have been thrown up in very hot water from the crater of a volcano, which, from their lively condition, was apparently their natural residence. Small caterpillars have been found in hot springs of the temperature of 205 ° ; and small black beetles, which died when placed in cold water, in the hot sulphur baths of Albano. Intestinal worms within the body of a carp have been seen alive after the boihng of the fish for eating ; and the inhabitants of some little snail-shells, which seemed to have been dried up within them, have been caused to revive by placing the shells in hot water for the purpose of cleaning them. The lofty heights of the atmosphere, and the dark and rayless depths of the ocean, are tenanted by anunals of beautiful organisation and wonderful powers. Vast flights of butterflies, the emblems of summer and sunshine, may some- times be seen above the highest peaks of the Alps, almost 14 INTRODUCTION. touching witli their fragile wings the hard surface of the never-melting snow. The gigantic condor or vulture of the Andes has been seen to soar on its widely-expanded wings far above the highest peak of Chimborazo, where the baro- meter would have sunk below ten inches. The existence of marine fishes has been ascertained at a depth of from 500 to 600 fathoms ; and in the deep recesses of those caverns in Styi'ia and Carniola, which are inhabited by the curious Proteus (ZooL. § 532), numerous species of insects are found, all of which, hov^ever, like the Proteus, are blind. Having thus glanced at some of those facts which demon- strate the practical importance of the study of Physiology, and having indicated ' the mode in which that study should be pursued, it remains to offer a few observations upon its value with reference to the culture and discipline of the mind itself One of its great advantages is, that it not only calls forth, in a degree second to no otlier, both the ohserving and the reasoning powers ; but that it oifers so much that is attractive by its novelty to those who enter upon it seriously, and make it an object of regular pursuit. For it affords abundant opportunities, even to the beginner, of adding to the common stock of information respecting the structure and habits of the vast number of living beings that peoj^le our globe. The immense variety of the objects which come under the investigation of the physiologist, so far from discouraging the learner, should have the effect of stimulating his exertions, by opening to him new fields for productive cultivation. Of by far the larger part of the organised crea- tion, little is certainly known. Of no single species, — of none of our commonest native animals, — not even of Man himself, — can our knowledge be regarded as anything but im- perfect. Of the meanest and simplest forms of animal life, we know perhaps even less than we do of the more elevated and complex ; and it cannot be doubted that phenomena of the most surprising nature yet remain to be discovered by patient observation of their actions. It was not until very recently, that the existence of a most extraordinary series of metamor- phoses, more wonderful than those of the insect, has been discovered in the jelly-fish of our seas, in the barnacles that INTRODUCTION. 15 attacli tliemsolves to floating pieces of timber, and in the crabs, lobsters, and shrimps of our shores. The very best accounts we have, of the structure, habits, and economy of the loAver tribes of animals, have been furnished to us by individuals who did not think it beneath them to devote many years to the study of a single species ; and as there are very few which have been thus fully investigated, there is ample opportunity for every one to suit his own taste in the choice of an object. And none but those who have tried the experiment, can form an estimate of the pleasure Avliich the study of Nature is capable of affording to its votaries. There is a simple pleasure in the acquisition of knowledge, worth to many far more than the acquisition of wealth. There is a pleasure in looking in upon its growing stores, and watching the expan- sion of the mind which embraces it, far above that which the miser feels in the grovelling contemplation of his hard-sought pelf. There is a pleasure in making it useful to others, com- parable at least to that which the man of generous benevo- lence feels in ministering to their relief with his purse or his sympathy. There is a pleasure in the contemplation of beauty and harmony, wherever presented to us. And are not all these pleasures increased, when we are made aware, — as in the study of ]^ature Ave soon become, — that the sources of them are never-ending, and that our enjoyment of them becomes more intense in proportion to the comprehensiveness of our knowledge ? And does not the feeling that we are not look- ing upon the inventions or contriA'ances of a skilful human artificer, but studying the wonders of a Creative Design infinitely more skilful, immeasurably heighten all these sources of gratification? If it is not every one Avho can feel all these motives, cannot every one feel the force of some ? •There is certainly no science Avhich more constantly and forcibly brings before the mind the poAver, the Avisdom, and the goodness of the Creator. For whilst the Astronomer has to seek for the proofs of these attributes in the motions and adjustments of a universe, Avhose nearest member is at a distance which imagination can scarcely realize, the Physio- logist finds them in the meanest Avorm that aa^c tread beneath our feet, or in the humblest zoophyte dashed by the waves 16 INTRODUCTION. upon our shores, no less than in the gigantic whale, or massive elephant. And the wonderful diversity which exists amongst the several tribes of animals, presents us with a continual variety in the mode in which these adjustments are made, that prevents us from ever growing weary in the search. But it is not only in affording us such interesting objects of regular study, that the bounty of Nature is exiubited. Perhaps it is even more keenly felt by the mind which, harassed by the cares of the world, or vexed by its disap- pointments, or fatigued by severer studies, seeks refuge in her calm retirement, and allows her sober gladness to exert it? cheering and tranquilhzing influence on the spirit. " With tender ministrations, tliou, Nature, Healest thy wandering and distracted child; Thon ]3onrest on him thy soft influences, Thy sunny hues, fair forms, and breathing sweets, The melody of woods, and winds, and waters.- — Till he relent, and can no more endure To he a jarring and dissonant thing Amidst the general voice and minstrelsy, — But bursting into tears wins back his way, Hia angry spix'it healed and harmonized By the benignant touch of love and mercy." COLEBIDGB, DISTINCTIVE CHARACTERS OF ORGANIZED BODIES. l7 CHAPTER I. OP THE VITAL OPERATIONS OP ANIMALS, AND THE INSTRUMENTS BY WHICH THEY ARE PERFORMED. 1. Living beings, wlietlier belonging to tlie Animal or to the Vegetable kingdom, are distinguished from the masses of inert matter of which the Mineral kingdom is made np, by peculiarities of form and size, of structure, of elementary- composition, and of actions. — Wherever a definite form is exhibited by Mineral substances, it is bounded by plane surfaces, straight lines, and angles, and is the effect of the process of crystallization, in which particles of like nature arrange themselves on a determinate plan, so as to produce a regular aggregation; and there is, probably, no Inorganic element or combination which is not capable of assuming such a form, if placed in circumstances adapted to the manifestation of its tendency to do so. The number of different crystalline forms is by no means large ;' and as many substances crystal- lize in several dissimilar forms, whilst crystals resembhng one another in form often have a great diversity of composition, there is no constant correspondence between the crystalline forms and the essential nature of the greater number of mineral substances. If that peculiar arrangement of the molecules which constitutes crystallization should be wanting, so that simple cohesive attraction is exercised in bringing them together, without any general control over their direc- tion, an indefinite or shapeless figure is the result. With this indefiniteness of form, there is an absence of any limit whatever in regard to size : a crystal may go on increasing continuously, so long as there is new material supplied ; but this new material is deposited upon its surface merely, and its addition involves no interstitial change ; the older particles, which were first deposited, and which continue to form the nucleus of the crystal, remaining just as they were. In Or- ganized bodies, on the other hand, we meet with convex surfaces and rounded outlines, and with a general absence of angularity ; and the simplest grades, both of Animal and of c 18 DISTINCTIVE CHARACTEES OF ORGANIZED BODIES. Vegetable life, present themselves under a shape which ap- proaches more or less closely to the globular. From the highest to the lowest, each species has a certain characteristic form, by which it is distinguished ; this form, however, often presents marked diversities at different periods of life, and it is also liable to vary within certain limits among the individuals of which the species is composed. The size of Organized structures, like their form, is restramed within tolerably definite limits, which may nevertheless vary to a certain extent among the individuals of the same species. These limits are most obvious in the higher animals, whilst they seem almost to disappear among certain members both of the Animal and the Vegetable kingdoms, which tend to increase themselves almost indefinitely by a process of gemmation or budding, so as to produce aggregations of enormous size. Such aggregations, however, being formed by the repetition of similar parts, wliich can maintain their existence when detached from one another, may, in some sense, be regarded as clusters of distinct organisms, rather than as smgle individuals. Such is the case, for example, with the wide-spreading forest-tree, and with those enormous masses of coral of which reefs and islands are composed in the Polynesian Archipelago. For every separate leaf-bud of the tree, Uke every single polype of the coral, if detached from its stock, can, under favourable circumstances, perform all the functions of life, and can develop itself into a new fabric resembling that from which it was separated. 2. The differences between Organized and Inorganic bodies, in regard to their structure, are much more important than those which relate to their external configuration. . Every particle of a mineral substance, in which there has not been a mere mixture of components, exhibits the same properties as those possessed by the whole ; the minutest atom of car- bonate of lime, for instance, has all the properties of a crystal of calc-spar, were it as large as a mountain. Hence it is the essential nature of an Inorganic body that each of its particles possesses a separate individuality, and has no relation but that of juxtaposition to the other particles associated with itself in one mass. — The Organized structure, on the other hand, receives its designation from being made up of a greater or less number of dissimilar parts or ory what may be termed the vegetative organs. But in the lower of tribes of Animals, we do not find the animal lunctions to possess this predominance. In fact, among the 30 DISTINCTIVE CHARACTERS OF ANIMALS. many wliicli are fixed to one spot during nearly their whole lives, and which grow and extend themselves like plants, the movements of the body are but few in number, and trifling as to their variety; these movements are only destined to assist in the performance of the organic functions, as by bringing food to the mouth, and water to the respii'atory organs ; and the nervo-muscular apparatus by which they are effected, bears so small a proportion to the organs of nutrition, as to seem hke a mere appendage to them, and is sometimes altogether undiscoverable. Tliis is the case, for example, in the lowest kinds of shell-fish, such as the Oyster, and in the Coral-polypes. 12. Hence we perceive, as we descend the Animal scale, a nearer and nearer approach to the character of Plants ; and this we shall find to be the case, not only in the general arrangement of the organs, but also in the nature of the elementary tissues of which these are composed. For in the higher animals, the whole organism is constructed in such a manner as to admit a free motion in its individual parts. The different portions of the skeleton or hard framework are connected with each other by flexible ligaments, which are adapted to resist a very powerful strain; the muscles are attached to these by fibrous cords or tendons, which, also, can support a vast weight ; and the several muscles and other ]Darts, which need to be mutually connected, but also require a certam power of moving independently of one another, are bound together by a very elastic loosely-arranged tissue, consisting of fibres crossing and interlacing in every direction, the interstices between which are filled with fluid. J^ow to these fibrous tissues, there is nothing analogous in plants, because no freedom of motion is required, or even permitted, among their parts ; and we find them bearing a less and less proportion to the whole, as Ave descend the animal scale. On the other hand, we find the various forms of true cellular tissue, such as predominate in plants (Yeget. Phys. Chap. III.), becoming more and more abundant, as we pass from the highest to the lowest animals, and having more and more important duties to fulfil. But even in the highest Animals, as will hereafter appear, they are the im- mediate instruments of the most important among the organic functions, just as they are in Plants. CIIEMICxVL CONSTITUENTS : ALBUMEN. 31 Chemical Constitution of the Animal Body. 13. By far tlic larger proportion of the Animal fabric is formed at tlie expense of the substance termed A Ibumen ; the composition and properties of which, tlierefore, claim our iirst attention. The fundamental importance of albumen in the animal economy, is shown by the fact that it constitutes, with fat, and a small proportion of certain mineral ingredients, the whole of that mass of nutrient material stored up in the eggs of oviparous animals, which, being appropriated by the germ to the building up of its fabric, is converted by it into the bones, muscles, nerves, tendons, ligaments, glands, mem- branes, &c. of the embryo. We find it also constituting a large proportion of the solid matter of the blood and other nutrient lluids of the adult animal ; and it is the fundamental form to which the various azotized substances employed as food (§ 153) — such as animal flesh, or the gluten of bread — are first reduced by the act of digestion. It is composed of 49 carbon, 36 hydrogen, 14 oxygen, 6 nitrogen, with a minute proportion of sulphur ; it is generally blended, also, with more or less of fatty matter, and with saline and earthy substances. 14. Albumen may exist in two states, — the soluble and insoluble. In the animal fluids it exists in its soluble form; and is united (as an acid to its base) with about 1^ per cent, of soda, forming an albuminate of soda. It is not altered by being cbied at a low temperature, but still retains its power of being completely dissolved in water. When a considerable quantity of it exists in a fluid (as in the white of the G-gg), it gives to it a glairy tenacious character ; but it is nearly tasteless. When such a fluid is exposed to a tempe- rature of about 150°, a coagulation or 'setting' takes place, as in the famihar process of boiling an egg. But if the albumen be present in smaller quantity, the fluid does not form a consistent mass, but only becomes turbid ; and this only after being boiled. Albumen which has been dried at a low temperature, however, may be heated to the boiling point of water, without passing into the insoluble condition ; a fact which is of peculiar interest in relation to the power which the Tardigrada (ZoOL. § 841) possess, of sustaining a very high temperature without the loss of their vitality, when 32 CHEMICAL CONSTITUENTS : ALBUMEN, CASEIN. their bodies have been completely dried up in , the first instance. ISTo trace of organization can be detected in coagulated albumen, which seems to be composed only of a mass of granules ; and in this respect it differs in an im- portant degree from fibrin — as we shall presently see. Albumen may also be made to coagulate readily by the action of acids, especially the nitric (aqua-fortis) ; so that a very small quantity of it may be detected in water, by the tur- bidity produced by adding to it a di'op or two of nitric acid, and then heating it. Now, when thus coagulated, albumen cannot be dissolved again by any ordinary process ; but its solution may be accomplished by rubbing it in a mortar with a caustic alkali, potass or soda. From this solution it may be precipitated again on the addition of an acid in sufficient ({uantity to neutralise the alkali. Albumen is distinguished, then, by its peculiar property of coagulating on the applica- tion of heat, or on being treated with certain acids. 15. JSTearly allied to albumen is the substance termed Casein, which replaces it in milk ; and this is specially worthy of notice here, because it is the sole form in which the young Mammal receives albuminous nourishment during the period of suckling, in which it draws its sustenance from its parent. Like albumen, this substance may exist in two forms, the soluble, and the insoluble or coagulated ; and the jDresence of a small quantity of free alkali seems essential to its continuance in the soluble form. Casein differs from albumen, however, in this, that it does not coagulate by heat, and that it is precipitated from its solution by organic acids, such as the acetic and lactic, which have no coagulating action on albumen. It is further remarkable for the facihty with which its coagulation is effected by the contact of certain animal membranes ; as we see when a small piece of ■rennet (which is the dried stomach of the calf) is put into a large pan of milk in the process of cheese-making, the ' curd' which then separates being composed of casein entangling the oily particles of the milk. In the coagulated state, casein differs but very little from albumen, and is readily converted into it by the gastric fluid. It is remarkable for its power of dissolving the earthy phosphates, as much as 6 per cent, of phosphate of lime being usually obtainable from it ; and it is in this combinatioji, that the large quantity of bone-earth required for the consolidation of the skeleton of the young animal, is introduced into its system. A substance resembling casein is obtainable from the serum of the blood, especially in pregnant females ; and also from the serous fluid which occupies the interstices of the tissues. It is found, also, mingled with albumen, in the yolk of the egg, forming a compound which (before its true character was known) has been distinguished as vitellin. Now as all the liquids con- taining casein have it for their special function to supply formative materials to rapidly-growing tissues, we may with much probability regard it as still more closely related to them than is albumen itself. It differs from albumen but little, if at all, in the ultimate proportions of its elements (§ 1 3). 16. The substance of which muscles are composed, has been commonly considered to be Fibrin (§ 17) ; but it diflers essentially from hbrin in its properties, and is now dis- tinguished as Syntonin. Its chief peculiarity is its solubility in very dilute muriatic acid (1 part to 100 of water), and its precipitation in the form of a jelly when the acid is neutra- lised ; this jelly treated with dilute alkalies forms a solution which coagulates by heat ; and thus it seems to be reduced nearly to the condition of albumen. This is, in fact, very much what takes place in the act of digestion of flesh-meat ; the muscle-substance being first dissolved by the muriatic or other acid of the gastric fluid, and the solution being then rendered alkaline by the mixture of bile and other secretions in the small intestine. 17. In the blood and other nutrient fluids of the animal body, there is found a substance which is so closely related to albumen in its ultimate chemical composition, as not to be dis- tinguishable from it with any certainty ; but which, though fluid whilst circulating in the living vessels, coagulates spon- taneously after having been for a short time withdrawn from them, the coagulum or clot bemg distinguished from that of albumen or fifei^H by the fibrillar arrangement of its particles, which indicates an incipient organization. This substance, termed Fibr'm, may be obtained in a separate form, by stirring fresh-drawn blood with a stick, to which it adheres in threads. In this condition it possesses the softness and elasticity which characterise the flesh of animals, and con- tains about three-fourths of its weight- of water. It may be o 34: CHEMICAL CONSTITUENTS : — FIBRIN. deprived of this water by drying, and then iDecomes a hard and brittle substance ; but, like dried flesh, it imbibes water again when moistened, and recovers its original softness and elasticity. From the recent experiments of Dr. Eichardson, it appears that the coagulation of blood-fibrin depends upon the escape of ammonia, being accelerated by such conditions as favour the liberation of this gas, and retarded or prevented by such as cause its retention in the liquid; whilst, even after the clot has been formed, it may be dissolved by ammonia, forming again when that gas is set free. Pibrin differs from syntonin or muscle-substance in not being dis- solved by very dilute muriatic acid, but being merely caused to swell up into a gelatinous mass, which contracts again when more acid is added. It combines with the earthy phosphates, of which as much as 2^ per cent, is sometimes found in the ash left by its combustion. 18. There can be no doubt that fibrin is formed in the blood and in the other fluids in which it presents itself, at the expense of albumen. What is its precise destination, cannot as yet be clearly specified ; but there are several circumstances which point to the conclusion that it is to be regarded as a transitional stage in the metamorphosis of albumen into the simple fibrous tissues (§ 23.) Thus, when the ordinary clot of blood is examined microscopically, it is found to consist, not, like an albuminous coagulum, of a homogeneous mass of granules, but of a network of im- perfectly-formed fibres, enclosing the red corpuscles in its interstices. A much more distinct network of the same kind may be seen in the colourless coagulum formed by the liquid which may be skimmed off the surface of the blood drawn from persons sufiering under any severe inflammation*; such blood coagulates slowly, and its red corpuscles and the fluid in which they float have an unusual tendency to separate from each other ; and the fibrin previously dissolved in the latter sets into definite fibres, which continue for some days to increase in firmness. It is a liquid of the same kind, charged with fibrin in a peculiarly " plastic " condition, that is poured forth for the formation of new tissue when the repa- rative processes are at work for the healing of a wound or the reunion of divided parts ; and it is by a plug of coagulated fibrin, which gradually comes to present a more and jnore CHEMICAL CONSTITUENTS : — FIBRIN, GELATIN. 35 distinctly fibrous structure, tliat the mouths of divided blood- vessels are closed up, when the flow of blood from them spontaneously stops. In all such cases, tlie fibrous network, if formed out of connexion with a living body, passes after a time into decay ; but if it be formed in a])positioii with living- parts, blood-vessels gradually extend into it from these, its nutrition is maintained and improved, and it progressively comes to present the ordinary characters of the simple fibrous tissues (§ 22). 19. Although the tissues most actively concerned in carrying on the vital operations, retain for the most part the composition of albumen, yet that very large proportion of the fabric of the higher animals whose offices are essentially mechanical, has a very different chemical constitution. • If we boil down either their bones, their skin, or their internal membranes, we shall get a considerable quantity of the sub- stance scientifically termed Gelatin, familiarly glue. Though consisting of the same elements as albimien, its composition is simpler, because these elements are united in smaller propor- tions ; the atom or combining equivalent of gelatin being made up of 13 Carbon, 10 Hydrogen, 5 Oxygen, 2 Nitrogen. The distinctive character of gelatin consists in its solubility in warm water, its coagulation on cooling into a uniform jelly which can be liquefied again by warmth, and its formation of a peculiar insoluble compound v/ith tannin. Gelatin is very sparingly soluble in cold water, though made to swell up and soften by prolonged contact with it. A solution of only one part of gelatin in 100 of hot water is sufiiciently strong for the whole to form a consistent jelly on cooling. The re- action of gelatin with tannin is so decided, that the presence of only one 2:)art in 5000 of water is at once detected by infusion of galls ; and it is in this action that the process of tanning consists, — the gelatinous fibre of the skin, which would speedily pass into decay, being converted into a com- paratively unchangeable substance. The different tissues which have gelatin for their base, yield it to boiling water with different degrees of facihty ; this diversity ap2:)arently depending in some degree upon the definiteness of their organization. Thus the " sound " or air-bladder of the cod, sturgeon, and other fish, which, when dried and cut into strips, is knov/n as isinglass, is very readily acted on ; the d2 36 CHEMICAL CONSTITUENTS : — GELATIN, CHONDRIN. same is the case with, the animal substance of bones from which the earthy matter has been removed ; and in each case the fibrous texture of the hving tissue is but very imperfectly developed. For the extraction of gelatin from the skin, the ligaments, the tendons, and various internal membranes, whose fibrous texture is more pronounced (§ 29), a much longer action of boiling water is required. 20. A peculiar modification of gelatin, which presents itself in Cartilage (or gristle), is distinguished as Chondrin. This requires longer boiling than gelatin for its solution in water; as is seen when a knuckle of veal or of mutton is cooked, the tendons and ligaments about the joint being almost reduced to pulp, whilst the cartilages are scarcely at all softened. The essential properties of chondrin are nearly the same as those of gelatin, and its composition seems nearly identical ; but it is thrown down from its solution by muriatic and acetic acids and some other reagents, which do not disturb a solution of gelatin. 21. It is not yet f Lilly known how the material of the gelatinous tissues is produced in the animal body. There can be no doubt of its being producible from albumen ; since we find it in large proportion in the tissues of animals that have never received gelatin into their bodies in any shape. And although carnivorous animals will receive it as part of their ahment, yet there is strong reason to believe that the gelatin which is thus supplied to them does not really serve to nourish their bodies, but that it is speedily decomposed and got rid of (§ 159). It may be considered as quite certain that the albuminous tissues cannot be formed by the meta- morphosis of gelatin ; whilst conversely, looking to the fact that in the egg and in milk no gelatin is provided for the young animal, although the gelatinous tissues form a yet larger proportion of its body than they do in the adult, we seem entitled to question whether it is possible that these tissues can be formed in any other way than at the expense of the albuminous constituents of the blood. Structure of the Primary Tissues. 22. In considering the structure of the " primary tissues," of which the various organs of animals are composed, it will be convenient first to treat of those which are subservient PRIMARY TISSUES : SIMPLE FIBROUS TISSUES. 37 merely to the physical actions of the framework ; as, for ex- ample, by holding its parts together, by communicating motion, or by giving them mechanical support and protection. — Tlie several parts of the body, even to the very minute divisions of its organs, are held together by what may be termed, in contradistinction to Muscular and Nervous fibre, the simple Jlhrous tissues ; and these are merely endowed, like ordinary cords, with the power of resisting tension or strain, either without themselves yielding to it at all, or with a certain amount of elasticity, which enables them first to yield to a certain degree, and then to recover their previous state. These two quahties are characteristic of two distinct forms of simple fibrous tissue, the v)hite and the yellow. 23. The White fibrous tissue presents itself under various forms, being sometimes composed of fibres so minute as to be scarcely distinguishable, but more commonly presenting itself under the aspect of flattened bands, which are but imperfectly divided into fibres, and have more or less of a wavy aspect (fig. 1). This tissue is resolved, by long boiling, into gelatine ; and when treated with acetic acid, it swells up and becomes transparent, by w^hich peculiarity it can White Fibrous Tissue. dis- The be readily tinguished from the other kind, to be next described. Yellow fibrous tissue presents itself in the form of long, separate, clearly defined fibres, which sometimes branch, and which break short off when overstrained, their extremi- ties bemg disposed to curl up (fig. 2). They are, for the most part, between 1 -5,000th and l-10,000th of an inch in diameter ; but they are often met with both larger and smaller. This kind of tissue un- dergoes but very little change from long boiling, and it is Yellow Fibrous Tissue. 38 PRIMARY TISSUES I AREOLAR TISSUE. not acted on by acetic acid. It is but little prone to decom- position, and will exliibit its peculiar elasticity long after it lias been separated from the body, provided it be kept moist. — These two forms of tissue exist separately in certain parts of the fabric, but they are much more frec[uently combined ; and the proportion of the yellow elastic tissue which exists in any such combination, may be readily determined under the microscope by the use of acetic acid, which renders all the white fibrous structure so transparent, that the yellow fibres are seen completely isolated in the midst of it. 24, One of the tissues which is composed of such an admixture of white and yellow (or non-elastic and elastic) fibres, is the one which was formerly called "cellular," but which is now more correctly designated as Areolar.^ This is composed of a mesh-work of fibres, and of bands of fibrous membrane, which are interwoven in such a manner as to leave very numerous interstices and cavities amongst them, having a tolerably free communication with each other (fig. 3). These Fig. 3. — Portion of Areolar Tissue. cavities are filled during life with a serous fluid ; " and it is a necessary result of the communication between them, that if an accumulation of this fluid takes place to an undue extent, ^ From the Latin areola, a small open space. ^ A fluid resembling the serum of the blood, diluted with water (§ 236). PRIMARY TISSUES : AREOLAR TISSUE. 39 as in dropsy, it descends by gravity to the lowest situation. Hence, tlie legs swell more frequently than any other parts. In its natural state, this tissue possesses considerable elas- ticity ; hence, when we press upon any soft part, and force out the fluid beneath into the tissue around, the original state returns as soon as the pressure is removed. Eut in dropsy, it appears as if the elasticity of the hbres were impaired or destroyed by their being over-stretched ; for when we press with the finger upon a dropsical part, a j^it remains for some time after the finger has been removed. 25. This Areolar tissue is diffused through almost the whole fabric of the adult animal, and enters into the compo- sition of almost every organ. It binds together the minute parts of which the muscles are composed ; it lies amongst the muscles themselves, connecting them together, but yet per- mitting them sufficient freedom of motion ; it exists in large amount between the muscles and the skin ; it forms sheaths to the blood-vessels and nerves, and so connects them with the muscles that they shall not be strained or suddenly bent by the movements of the latter ; and it enters mto the struc- ture of almost every one of the organs which are contained in the cavity of the trunk, uniting its parts to each other, and keeping the whole in its place. But it is a great mistake to assert, as it was formerly common to do, that it penetrates the harder organs, such as bone, teeth, and cartilage. Its ]3urpose obviously is to allow a certain amount of motion among the jiarts it unites ; and we find that the more free this motion is required to be, the larger is the proportion borne by the yellow or elastic fibres, to the white or non-elastic. 2Q. Although the Areolar tissue contains a very large number of blood-vessels and nerves, yet it does so merely because it furnishes the bed or channel in which they are conducted to the parts where they are really wanted. Its own vitality is low, and its sensibility very slight. It is quickly reproduced after injury ; and it is by its means that losses of substance are repaired in tissues of a more elaborate kind, which are not so easily regenerated. 27. The continuity or connectedness of this tissue over the whole siu'face of the body, admits air to pass readily from one part to another ; and the inflation or blowing-up of its cavities with air, which has sometimes happened accidentally, 40 PRIMARY TISSUES : SEROUS MEMBRANES. and lias sometimes been purposely effected, does not produce any disorder in the general functions of the body. In blow- ing the nose violently, some j^art of the membrane lining its cavity has occasionally given way, so as to allow air to pass into the areolar tissue of the face, and especially into that contained in the eyehds, which is particularly loose ; an enor- mous swelling of these parts then takes place, presenting a very frightful appearance, but not attended mth the least danger, and subsiding of itself in a few days. This swelling presents a character to the touch quite different from that which would be occasioned by a similar distension with liquid; for it gives somewhat of the crackling feel that is occasioned by pressing on a blown bladder. A similar inflation of the areolar tissue of the body has sometimes occurred from the formation of an aperture, by disease or injury, in the walls of the lungs or air-passages, and the consequent escape of air during the act of breathing : in one remarkable case of this kind, the skin of the whole body was so tightly distended with air as to resemble a drum. It is intentionally practised by butchers, who " blow up " the areolar tissue of their veal, in order to increase its plumpness of aspect ; and the in- flation of the areolar tissue of the head, in the living state, has been sometimes practised by impostors, in order to excite commiseration. 28. Fibres and shreds of fibro-membrane, resembling those of which areolar tissue is composed, may be so interwoven as to form a continuous sheet of membrane, having a smooth and glistening surface j and in this manner are produced the Serous Membranes that line the different cavities in which the viscera (or organs contained within the skull, the chest, and the abdomen) are lodged. The peculiar manner in which these membranes are arranged, will be explamed hereafter (§ 43). One of their surfaces is always free or. unattached, whilst the other is in contact with the outer wall of the cavity ; and from the free surface, which is covered with a layer of flattened epithelium-cells (fig. 10), a serous fluid is exhaled, which adds to its smoothness. It is by an accumula- tion of this fluid, that di^opsies of the cavities are produced, — such as water on the brain, or in the chest. 29. By the union of fibres of a stronger kind, those firmer tissues are produced, which are employed wherever a greater FIBROUS MEMBRANES AND LIGAMENTS. 41 strain has to be borne. This is the case with the Ligaments, which bind together the bones at the joints, the Tendons, by which the muscles are usually attached to the bones, and the tough Fibrous Membranes that envelope and protect many of the most important viscera. In these any considerable amount of elasticity would be misplaced ; and we conse- c^uently find that they are chielly or entirely composed of the wJiite fibrous tissue. "Whenever an elastic ligament is re- quired, however, we find the white replaced by yellow. One of the best examples of this is seen in the ligament of the neck of many quadrupeds, commonly known as the paxy- waxy ; which is given to the large herbivorous quadrupeds, such as the ox, to assist them in supporting their heavy heads with as Httle exertion as possible ; whilst carnivorous quadrupeds, such as the Hon and tiger, are endowed with it to give them additional power of carrying away heavy bur- dens in their mouths. In Man we scarcely find a trace of it. This yellow fibrous tissue is found, moreover, in the walls of the arteries (§ 248), to which it gives their peculiar elas- ticity; and it also forms the vocal cords of the larynx (§ 681), It is by the same kind of elastic ligament that the claws of the Feline tribe are ch-awn back into their sheaths when not in use, being projected (when required) by muscular action ; and that the two j)ieces of the shell of Bivalve MoUusks are united at the hinge, and are at the same time kept apart for the admission of water between them, except when the animal forcibly draws them together by its adductor muscle (§ 113). 30. All these fibrous tissues, then, are concerned in actions purely mechanical; and there is nothing in their properties which is so distinct from those of inorganic substances, as to require to be considered as vital. We may consider them, therefore, as among the lowest forms of animal tissue ; and accordingly we find that, when the higher forms degenerate or waste away, these appear in their place. Such a degene- ration may take place simply from want of use. Thus if, from palsy or want of power of the nerves, the muscles of the legs are disused for several years, they will lose their peculiar property of contractility (§ 5) ; and it will be found that scarcely any true muscular structure remains, but that it is replaced by some form of fibrous tissue. Or again, if the 42 BASEMENT MEMBRANE : CEIiLS. front of the eye be so injured by accident or disease, that light cannot pass through it to make its impression on the nerve, that nerve, being thrown into disuse, will gradually degenerate into fibrous tissue. Moreover, this change may take place as a part of the regular actions of life ; for there are certain organs in the young animal previous to birth, which are not required afterwards ; and these degenerate in like manner, gradually wasting away, and leaving only traces behind them, — tubes shrivelling into fibrous ligaments, and glandular structures remaining only as areolar tissue. 31. Along every free surface of the body, both external and internal, is spread out a delicate structureless layer, which is termed the Basement or Primary Membrane. This forms the outer layer of the True Skin, lying between it and the Epidermis or scarf-skin (§ 37) ; in the same manner it underhes the Epithelial layer of the Mucous membjanes which line the open cavities of the body (§ 39), and of the Serous membranes which line its closed cavities (§ 43) ; and it occupies the same position in the walls of the blood- vessels, gland-ducts, and other tubes. It is difficult to sepa- rate it, in any of these parts, from the tissues with which it is in contact ; and its characters may be well studied by dis- solving the calcareous part of an oyster or mussel-shell in dilute acid, when it mil be found that layers of a thin trans- parent membrane are left, which have been thrown off at each act of shell-formation, from the surface of the mantle. This elementary membrane, like that which forms the walls of cells (§ 32), is remarkable for the readiness with which it is permeated by fluid, al- though no visible pores can be seen in it. 32. A considerable part of the fabric of even the highest Animal is formed, like the entire organism of the Plant, of Cells, either unchanged or in some way metamorphosed. A cell is a minute bag or Fig. 4.-NUC.EATED Cells; a a, nuclei. ^^^.^^^^ ^^^^^^^ ^^ ^ structure- less membrane, and having its cavity filled with fluid of some kind. In some part of its interior, most commonly adhering JL // '""^C^ K ' \ \ "•:.^ 4 i li > u cells; their mode of multiplication. 43 to its wall, there is usually to be observed a solid collection of granular matter, wbich is termed tlie nucleus (fig. 4, a a). The typical form of tlie cell is globular or oval (fig. 5) ; but when a number of cells are in contact with each other, and are pressed together, their sides become flattened; so that when they are cut across no intervals are seen between them, but their walls are everywhere in contact (fig. 6), just as in »:p^^ '1 Fig. 5. Fig. 6. Rounded Cells in Cartilage Polygonal Cells from Car- OF Bat's Ear. tilage in Mouse's Ear. the section of a vegetable pith. The chemical composition of the nucleus differs from that of the cell- wall ; for whilst tlie latter is dissolved by acetic acid, the former (like the yellow elastic tissue, with which its substance appears to have some relationship) is unchanged by it. When the formation of a cell is complete, and it is not destined to reproduce its kind, the nucleus frequently disappears ; tliis is the case, for example, with the red corpuscles of the blood of Mammaha (§ 229), and also with Fat-cells (§ 46). 33. ]^ew cells may originate in one of two very distinct modes ; either from a pre-existing cell, or by an entirely new production in the midst of an organizable fluid or blastema. The most remarkable example of the first process is presented in the early development of the germ, which entirely consists of an aggregation of cells, every one of which undergoes successive subdivisions into two, so that the total number in the germ-mass is repeatedly doubled (Chap. xv.). The same method of multiplication by hinary suhdivisio7i may be seen to continue throughout life in Cartilage-cells (§ 47), the growth of which almost exactly repeats the history of the growth of the lowest forms of Sea-weeds. The process of sub- division seems to commence in the nucleus, which begins to separate itself into two equal parts, and each of these draws 44 MULTIPLICATION AND NEW PRODUCTION OF CELLS. around it a portion of tlie contents of the cell ; so tliat the cell-wall, which is at first merely doubled inwards by a sort of hour-glass contraction, at last forms a complete partition between the two halves of the original cavity. The process may be repeated either in the same or in a transverse direc- tion, so as to produce four cells, which may be either arranged in a single line O O O O or may form a cluster gg ; and another subdivision of each cell will, of course, again double the entire number. In other cases, however, the nucleus appears to break up at once into several fragments, each of which may draw around it a portion of the contents of the parent-cell, which becomes invested by a cell-wall of its own ; and thus the cavity of the parent-cell may at once become filled with a whole brood of young cells, without any successive subdivision. Generally spealdng, the former method seems to prevail in structures which, lilie Cartilage, have a com- paratively joermri??e?i^ destination ; whilst the latter is followed in cases in which the cells thus formed are destined only for a transitory existence. This is the case especially in Can- cerous structures, which are particularly distinguished by their proneness to the rapid production of cells within cells. 34. The production of new cells in the midst of an or- ganizable blastema or formative fluid, such as is poured out from the blood for the reparation of an injury, is a very different process. This blastema, when first effused, is an apparently homogeneous semi-fluid substance ; as it solidifies, however, it becomes dimly shaded by minute dots, and as it is acquiring further consistence, some of these dots seem to aggregate, so as to form little round or oval clusters, bearing a strong resemblance to cell-nuclei. These bodies appear to be the centres of the further changes which take place in the blastema ; for if it be about to undergo development into a fibrous tissue (§ 18), they seem to be the centres from which the fibrillation spreads ; whilst, if a cellular structure is to be generated, it is from them that the cells take their origin. The first stage of the latter process appears to consist in the accumulation of the substance which the cell is to include, about each nucleus, and around this the cell-membrane is subsequently developed. It is in this mode that the de- velopment of new structures, for the filling up of losses of substance, is provided forj and it appears, from recent ISOLATED CELLS OF ANIMAL FLUIDS. 45 inquiries, tliat the blastema will resolve itself into fibres or into cells, according as the wound is completely secluded from the air, or is exj30sed to it. It is under the former condition that losses of substance are most rapidly and most completely repaired ; whilst it is under the latter that inilanmiation is most Hkely to arise, in consequence of the bad effect pro- duced by the contact of air with the raw surface ; the process of healing, when thus interfered with, going on less favourably as w^eU as more slowly. 35. The very simplest and most independent condition of the animal Cell, is probably to be found in the nutritive fluids of the body ; in which we meet with floating cells that are completely isolated from each other, and which are conse- quently just as self-sustaining as are the separate vesicles of the Yeast-plant, of the Eed Snow, or of other simple cellular Plants. These cells are of two classes. In the blood of animals generally, and in the chyle and lymph of Yertebrata, we find a larger or smaller proportion of colourless corpuscles, which are usually nearly spherical in form, and which exhibit various stages of development into cells, being sometimes little else than collections of granules, without any distinct enveloping membrane, whilst, in other instances, there is a distinct cell-wall, cell-cavity, and nucleus. These bodies, if watched under a sufficiently pov.^erful microscope, may often be seen to undergo very curious changes of form, resembhng those of the Amoeba (§ 129). Besides the foregoing, however, the blood of Yertebrated animals contains a far larger pro- portion of reel corpuscles, which are flattened disks, sometimes circular but more commonly oval, having pellucid and colour- less walls, but having their cavities filled with a pecuhar coloured fluid. As these will be more fully described here- after (§ 229), it is not requisite to do more than notice them here as constituting a most important part of the animal organism, probably not less than a twelfth part of the entire weight of Man and the higher animals, being thus composed of nothing else than these isolated cells. ^ 36. ISText in independence to the cells or corpuscles float- ing in the animal fluids, are those which cover the free ^ The entire weight of the blood of Man seems to be about one-sixth part of that of the body ; and the moist corpuscles constitute about half the entire weight of the blood. 46 SKIN AND MUCOUS MEMBRANES. membranous surfaces of tlie body, and which form the Epidermis, or superficial layer of the skin, and the Epithelium of the internal membranes. And it will be convenient here to consider the entire structure of the Skin, the Mucous Membranes, and the Serous Membranes, which are complex fabrics, chiefly made up of the elementary tissues already described. — These membranes may each be considered as composed of three principal parts, namely, the superficial layer or layers of cells, the basement-membrane whereon the cells lie, and the subjacent texture covered by this, which consists of fibrous tissue compactly interwoven and traversed by blood-vessels, nerves, absorbents, and also containing glands of various kinds. The Skin and Mucous Membrane may, in fact, be regarded as belonging to one and the same type ; for they are continuous with each other wherever one of the open cavities of the body conmiunicates with the surface, as at the mouth, nostrils, and anus ; and in the Hydra (§ 121) it has been experimentally found that the membranous layer covering the body may be made to change places with that which lines the stomach, without any sensible disturbance in the functions of either. The difl'erence between the two essentially consists in this ; that the Skin, being destined especially for the reception of sensations, and for the protection of the soft parts beneath, is more copiously furnished with nerves than with blood-vessels, and has its surface covered by a firm, dry cuticle ; whilst the Mucous Membrane, ministering especially to the organic functions, is comparatively little supplied with nerves, but is abundantly furnished with blood-vessels, and in certain parts with absorbents, whilst its cellular layer is soft and easily permeable by liquids. Both in the skin and in mucous membrane we find a multitude of minute glands, for the separation of particular fluids from the blood ; the nature of these differs with the locality. 37. The fibrous mesh-work of the Cutis or True-Shin is con- tinuous with that of the Ai-eolar tissue which lies immediately beneath it ; so that the two textures are not separated one from the other by any definite boundary (as the examination of a vertical section (fig. 7) clearly proves), but are dis- tinguishable only by the compactness of the one, as contrasted with the looseness of the other. The outer surface of the Cutis usually presents numerous minute elevations or papillce STRUCTURE OF THE SKIN. 47 (fig. 7, i i), wliicli are commonly arranged in rows ; of these, some are organs of touch, being furnished with sensory nerves that end upon a peculiar cushion-like organ in their interior (§ 490) ; but into others no nerves can be traced, so that, as these are copiously supplied with blood-vessels, it is pro- bable that they minister to the nutrition of the epidermis. / Fig. 7. — Vertical Section of the Skin, Showing the diflferent structures which it contains. A, Epidermis ; a a, its outer surface ; a — b, its horny layer ; h — c its inner soft layer, dipping down into the hollow between the papillae ; B, Cutis ; d, arterial twig supplying its vascular papillae; e e, perspiratory glandulas ; /, cluster of fat-cells ; g ff, perspiratory duct, traversing the true skin; h, its continuation through the epidermis; j i, tactile papillae, with their nerves. This is the more probable from the fact that we find these vascular papillse very large and full of blood-vessels in the interior of corns,' warts, and other such productions, formed by a "hypertrophy" or over-nutrition of the epidermis in particular spots ; and also in situations in which the ordinary epidermis is very thick, as it is on the black pads of the foot of the dog or cat. And a highly vascular structure of the 48 STKUCTURE OF THE SKIN. same kind is foinid in tlie matrix or receiDtacle of the growing roots of nails, hoofs, horns, &c, which are only modified forms of epidermis. Imbedded in the substance of the cutis we find, in most situations, the perspiratoyy glands (fig. 7, ee), by which the watery fluid that is continually being exhaled from the skin, is separated from the blood (§ 371) ; these send forth their secretion by canals {g h), which traverse the epidermis in a corkscrew-like manner, and then open upon its surface by oblique valvular orifices. In the Cutis, also, are lodged the hair-follicles (§ 38), which are really pits or depressions of its surface, with a vascular papilla at the bottom of each, supplying nutriment for the abundant development of the cells in which the hair originates, as will be presently described. "Wherever the hair-fol- licles occur, there do we also find sebaceous follicles (fig. 8, a a) ; these are peculiar glandulas, secreting fatty matter, which is poured into the hair- canal, so as to come through it to the surface of the epidermis ; and the use of this secretion, which is particularly abundant in the dark skins of the natives of warm climates, is to pre- vent the cuticle and the hair from being too much dried up by exposure to air. — The surface of the Cutis is covered by a layer of basement-mem- brane (§ 31), which is not traversed either by blood-vessels, nerves, or absorbents ; so that none of these pass into the epidermis which lies on its outer side. 38. The Epidermis, otherwise j,.^ termed the cuticle, or " scarf-skin," is Tkin section'ok the Human composcd of numerous layers of nu- ScALP;— a a, sebaceous glands; cleatecl Cclls j of wllich We find those b, a hair, with its follicle c. -^^ immediate contact with the base- ment-membrane to be nearly spherical ; those a little removed from it to be rendered polygonal by the mutual pressure of their sides ; those nearer the outer surface to be flattened, and this in an increased degree, as we pass from within STRUCTURE OF THE SKIN : EPIDERMIS. 49 outwards, until we arrive at layers composed entirely of dry flat scales, which show but little indication of ever having been cells. There is no doubt, however, that all these forms are but different stages of the existence of one and the same set of epidermic cells ; these taking their origin in the formative fluid exuded on the surface of the basement-mem- brane, and being progressively carried towards the surface by the . successive development of new layers beneath them, whilst the layers above them are thro'wn off", or are worn away ; and at the same time undergoing a change of form, in the first instance from mutual pressure, and afterwards from the loss of their contained fluid. At the same time they are rendered more firm in texture, by the formation of a horny secretion in their interior ; so that the outer layers of epi- dermis form a consistent membrane, which is raised from the surface of the Cutis when fluid infiltrates between them (as when the hand has been long soaked in water), or is poured out by the vessels of the latter (as when a blister is applied) ; whilst the soft internal layers remain in contact with the basement-membrane. — The number of layers varies greatly in different parts, being usually found to be greatest where there is most pressure or friction, as if the irritation deter- mined an increased supply of blood to the spot, and thus favoured an augmented development of epidermic cells. Thus, on the soles of the feet, particularly at the heel and the ball of the great toe, the Epidermis is extremely thick ; and the palms of the hands of the labouring man are distinguished by the horny hardness of their thick cuticle. — It was formerly supposed that a special layer of a soft spongy tissue, termed the rete mucosum, intervenes between the Cutis and the Epidermis ; and that this was the special seat of the colour of the skin in the dark races. It is now well ascertained, however, that this supposed rete con- sists of notliing else than the newly-forming soft layers of the true epidermis ; and that the colouring matter is diffused through the epidermic cells, so as to tinge the entire thickness of the cuticle, although its presence is particularly obvious in the deeper layers. — The Nails may be considered as nothing more than an altered form of Epidermis ; when examined near their origin, they are found to consist of cells which gradually dry into scales that remain E 50 EPIDERMIC APPENDAGES : — NAILS, HAIR, &C. coherent; and when thin sections are treated by a dilute solution of soda, these scales swell out again (as do also those of the cuticle) into globular cells. A new production is continually taking place in the groove of the skin in which the root of the nail is imbedded, and also from the whole of the surface beneath it ; the former adds to the length of the nail • the latter to its thickness. — The structure of Hairs is essentially the same. The base of each is formed of a " bulb," which consists of a mass of epidermic cells developed from the vascular papilla at the bottom of the hair follicle (fig. 8, c) ; and as this narrows into the " shaft " of the hair, a difference shows itself between the cortical or outer layer, and the medullary or pith-like substance of the interior. The former, which is continuous with the outer layers of the epi- dermis, is composed of flattened scales, arranged in an imbri- cated (tile-like) manner, so that the surface of the hair is usually marked by transverse jagged lines ; the latter consists of cells which frequently retain their spheroidal form, hke the inner layers of the epidermis ; but in the human hair these cells are elongated into fibres. It is very seldom that there is any canal in the interior of the Hair, although irregular spaces are not unfrequently left by the drying-up of the fluid con- tents of the cells. The structure of Quills is essentially the same as that of hairs on a large scale ; and we there see the difference very distinctly marked between the cortical portion which forms the "barrel" of the quill, and the medullary portion which forms the white pith-like substance of the stem of the feather. The Scales, where they are really epi- dermic appendages, as is the case in serpents and lizards, are formed upon the same pattern ; and we liave a good example of tJie detachment of the entire epidermis at once (reminding us of the casting of the shell of the crab and lobster) in tlie " sloughing " of the snake. 39. The Mucous Membranes form a sort of internal skin, lining those cavities of the body which open on its surface ; mid the elements of which they are composed are essentially the same, though combined and arranged in a different manner, in accordance with their difference of function. The principal part of the thickness of every ordinar}^ mucous membrane is made up, as in the skin, by the consolidation of areolar tissue, the fibres of which arc continuous with those MUCOUS MEMBRANES : EPITHELIUM. 61 of the ordinary areolar tissue on wliicli the membrane rests ; this layer is copiously furnished with blood-vessels, but it is seldom supplied with many nerves. Thus the mucous mem- brane lining the stomach possesses in health so little sensi- bility, that we are not aware of the contact of the substances taken in as food, unless they are of an acrid character, or of a temperature very dilferent from . that of the body ; and though the mucous membrane linmg the air-passages is very susceptible of certain kinds of irritation, jet it has but little ordinary sensibility in the state of health, except near the entrance to the windpipe. The large supply of blood which these membranes receive, has reference to their active j)artici- pation in the functions of secretion and absorption. One secretion is common to all, that of the mucus by wliicli they are covered ; tliis serves to j^rotect them from the irritation that would otherwise be produced by the contact of solid or liquid substances, or even of aii', with thek free surfaces ; and we see the results of its deficiency, in the inflammation which attacks the membrane, sometunes j^roceeding to its entire destruction, when from any cause the secretion is checked, as it sometimes is by injuries of the nerves sup- plying the part. 40. In every mucous membrane, as in the skin, the fibrous texture is bounded on the free surface by basement-mem- brane, beyond which no blood-vessels pass. And the surface of the basement-membrane is covered by cells, arranged either in a single layer or in multiple layers, conscituting the Epithelium. This, although answering to the Epidermis in structure and position, has a very different character ; for its cells neither dry up nor become horny ; nor do they adliere in such a manner as to form a contmuous membrane, except in the interior of the mouth and 02Sophagus (gullet), where the epithehum is endowed mth somewhat of the firmness of cuticle, in order to resist the abrading contact of hard substances. The epithelium cells of mucous membranes are commonly somewhat flattened ; but in some situations, as on the villi of the intestinal canal (fig. 9, d), they have more of a cylindrical, or rather conical shape, their smaller extremities being in contact with the basement-membrane. The epi- thelial cells are frequently cast off, like the epidermic, espe- cially from the parts that are most concerned in secretion ; E 2 52 MUCOUS MEMBRANES -EPITnELIUM. and tlicy arc as continually replaced by newly-formed cells, ■which are produced on the surface of the basement-mem- brane, at tlie expense of the fluid that transudes through it from the blood-vessels copiously distributed to its under surface. 41. Mucous membrane may either exist in the condition of a simple expanded surface, or may have a much more complex arrangement, by which its surface is greatly increased. The simple mucous membrane, such as that which lines the nose and air-passages, is found, for the most part, where no ab- sorj)tion has to be performed, and where only a moderate amount of secretion is necessary. But where it is to absorb as well as to secrete, it is usually involuted or folded upon itself, in such a manner as to form a series of little projec- tions, and also a number of minute pits (fig. 0). These pro- Fig. 9. — DlAGKAM UEPUF.SKNTING THE MuCOUS MeMBRANE OF THE Intestinal Canal. a a, absorbent vessels ; b b, basement membrane ; c c, epithelium-cells of level surface of membrane ; d d, cylindrical epithelium-cells of villus ; e e, secreting cells of follicle. jections sometimes have the form of long folds ; in other instances they are narrow filaments, crowded together so as almost to resemble the pile of velvet. In either case, the absorbent surface is vastly increased ; but chiclly so by these filaments, which are termed mlli, and act as so many little rootlets. On the other hand, it is in the pits or follicles, that the production of the fluid which is to be separated or secreted from the blood, chiefly takes place. — 'Not only are the flat expanded surfaces of the mucous membrane covered with epithelium cells, but the villi also arc sheathed by them ; and the secreting follicles are lined by the same. STRUCTURE OF GLANDS. SEROUS MEMBRANES. 53 The cells covering the villi (fig. 9, d) perform the important function of selecting and absorbing certain nutritious ele- ments of the food, which they communicate to the absorbent vessels in the interior of the villi. On the otlier hand, the epithelium-cells of the follicles (e) seem to be the real agents in the secreting process ; drawing from the blood, as materials for their own growth, certain elements contained in it ; and falling oflj, when mature, so as to discharge these substances as the product of secretion, giving place to a fresh crop or generation of cells, which go through a series of changes precisely similar to the preceding. 42, Now these follicles arc the simplest types or examples of all the Glandular structures, by which certain products arc separated from the blood, some to be cast forth from the body as unfit to be retained in it, and some to answer particular purposes in the system. In all of them the structure ulti- mately consists of such follicles, sometimes swollen into rounded vesicles, and sometimes extended into 'long and narrow tubes. Each follicle, vesicle, or tube, is composed of a layer of basement-membrane, lined with epithelium-cells, and surrounded on the outside with miimtely distributed blood-vessels ; and it seems to be by the peculiar powers of these cells, that the products of the secreting action, whether bde, saliva, fatty matter, or gastric fluid, are formed (see Chap. VII.). — Hence we see that the act of Secretion is, in animals as in plants, really performed by cells. It is neces- sary to bear in mind, however, that a simple transudation of the watery parts of the blood may take place without any proper secreting action, in the dead as in the living body ; it is in this manner that the serous fluid of areolar tissue and serous membrane is poured out, and that the watery portion of the urine is separated. 43. The Serous Membra7ies which line the closed cavities of the body, though composed of the same elements as the skin and mucous membranes, hfive a much simpler structure, and can scarcely be said to minister directly to any important vital function. The tissue of which Serous membrane is principally composed, scarcely differs, except in its greater density, from the laxer areolar tissue whereby the membrane is attached to the walls which it covers like i^laster ; it is but sparingly supplied either with blood-vessels or absorbents; and 54 ARRANGEMENT OF SEROUS MEMBRANES. it contains very few nerves. The smooth surface of the mem- brane forms one unbroken plane, being neither raised into villi, nor depressed into follicles ; and its basement-membrane is covered with a single layer of flat epithehum-cells, which are closely applied to it and to each other, like the pieces of a pavement (fig. 10). It is with such a membrane that every one of those great cavities is lined, which contains important viscera ; and it is also continued on to the outer surface of these viscera, so as to afford them an external coating* over every part save that by which they are attached. Thus the heart is suspended Pavemen?' ^^epxthe- freely, by the large vessels proceeding from LiuM Cells OF Serous its summit, witliin a bag or sac of fibrous Membrane. membrane peculiar to itself, which is termed the pericardium. The cavity of this bag is completely Imed by the serous membrane (fig. 11, p' ), which closely embraces the vessels, and which then bends down over the surface of the heart, so as to enclose it in the envelope p. Hence it will be seen that tliis membrane, whilst including the heart, and allowing it to ?\ "^m-'' I \ communicate with its vessels, lf/^,,J"""^P forms a completely shut sac; and it may be likened to a / common double cotton or woollen night-cap, which has a similar cavity between its Fig.ll.-DlAGRAMOFTHEPERICARmUM. ^WO ^CrS, tllO hCad bcing o a, auricles; t; «, ventricles ;/;, pulmonary really On the OUtside of artery, c, aorta; py. pericardium. ^j^-^^ ^j^-j^^ SCemiug tO be within the envelope. The two layers of the pericardium, though separated in the diagram for the sake of distinctness, are really in mutual contact, save when separated by the in- terposition of fluid poured out in disease. Each of the lungs, in like manner, is suspended in a closed sac of its own, termed the pleura; and the surface of the lung is covered by a serous membrane, which is reflected over the wall of the pleural cavity. SEROUS AND SYNOVIAL MEMBRANES. 55 A similar arrangement exists in the great cavity of the ab- domen ; hut the number and the complex relations of the viscera which this contains, give to the disposition of its serous membrane, termed the peritoneu7n, a peculiar complica- tion. The cavity of the skull also is lined by a serous mem- brane, termed the arachnoid, and this is prolonged over the surface of the brain, and enters its lateral ventricles (§ 458). The chief purpose of these membranes appears to be to faci- litate the movements of the included organs, by forming smooth surfaces which shall freely glide over each other ; this , is evidently of great importance, where such constantly- moving organs as the heart and lungs are concerned. Their surfaces are kept constantly moist with a serous fluid which exudes from the blood ; but in the state of health this fluid does not accumulate in their cavities, being al^sorbed as fast as it is poured out. Various forms of dropsy, however, — such as "water on the brain," "water on the chest," and " ascites," or dropsy of the abdomen — are the result of the increased outpouring of fluid into the serous cavities of the arachnoid, the pericardium, the pleura, and the peritonemn respectively. 44. ISTearly allied to the Serous mem- branes are the Synovial, which form closed sacs in the interior of joints, covering the \ ends of the cartilages, and then hning the t\ fibrous capsule which passes from one bone ' to the other. The mode of their arrange- ment will be understood from the accom- panying diagram ; in which a a represent the extremities of the two bones which are jointed together, h b the layers of car- tilage with which they are severally covered, and the dotted hne c c the synovial mem- brane, which is seen to form the sac or bag cf c', whilst at the points c c c c it is reflected upon the cartilages of the joints. In point of fact, however, the Synovial membrane is not ordinarily traceable as a distinct layer over the surface of these cartilages, but seems to have become incorporated with them ; for though in the embryo its presence may be distinctly proved Diagram OF THE STRUC- TURE OF A Joint. a a, extremities of the bones, covered with cartilage ; b b, layer of cartilage closely co- vered with synovial membrane : c c' c, re- flected layer of syno- vial membrane form- ing synovial capsule. 5Q SYNOVIAL MEMBRANES. — CILIATED EPITHELIUM*. hj the continuity of its blood-vessels over tlie entire car- tilage, yet these are found to retreat gradually as the joint is brought into use, until at last they only form a circle round the border of the cartilage. Some of the Synovial mem- branes, as that of the knee-joint, are furnished with little fringe -like projections, somewhat resembling the villi of mucous membranes (§ 41) ; these are extremely vascular, and are furnished with an epithelium which very readily falls off; and there is a strong probability that they are concerned in the secretion of the synovial fluid, which is much denser than the ordinary serous transudation, having from 6 to 8 per cent, of additional albumen, and presenting a glairy appearance like that of white of egg. It is interesting to see that the same purpose may thus be served by the extension of the membrane in either direction, either out- wards into a villous filament, or imvards into a follicle ; the function being determined in each case rather by the attributes of the cells, and by the supply of blood, than by the form which the secreting surface may happen to present. 45. The cells of Epithelium, whether flattened or cylindrical, are observed to be furnished in particular situations with a fringe of delicate filaments, which are termed cilia. These, although of extreme minuteness, are organs of great importance in the animal economy, on account of the extra- ordinary motor powers Avith which they are endowed. The form of the cilia is usually a little flattened, and tapering gradually from the base to the point. Their size is extremely variable ; the largest that and in transverse section at B; j ^ obscrVcd being about their cilia are seen at b, their i • i i nuclei at c; at a is shown one of l-500th of an incli ]n length, and these cells unusually elongated, the Smallest 1-1 3,000th. "When in motion, each filament appears to bend from its root to its point, returning again to its original state, like the stalks of Fig. 13. — Ciliated Epithelium Cells; as seen sideways at A, CILIARY MOVEMENT. 57 corn when depressed by the wind; and if a number be affected in succession with this motion, the appearance of progressive waves following one another is produced, as when a corn-field is agitated by repeated gusts. When the ciliary motion is taking place in full activity, however, nothing can be distinguished save the whirl of particles in the surround- ing liquid; and it is only when the rate of movement slackens, that the shape and size of the individual filaments, and the manner in which their stroke is made, can be made out. The motion of the cilia is not only quite independent (in all the liigher animals at least) of the will of the animal, but is also independent even of the life of the rest of the body ; being seen to continue after the death of the animal, and even going on with j)erfect regularity in parts separated from the body. Thus, isolated epithelium-cells have been seen to swim about actively in water, by the agency of their cilia, for some hours after their detachment from the mucous membrane of the nose ; and the regular movement of cilia has been noticed fifteen days after death, in the body of a tortoise in which putrefaction was already far advanced. In the gills of the Eiver Mussel, which are amongst the best objects for the study of this most curious phenomenon, the movement endures with similar pertinacity. — The purpose of this remarkable agency is obviously to propel fluids over the surfaces which are furnished with cilia. We find it taking the most important share in the functions of life among the lowest classes of animals. Thus, in Animalcules of various kinds, the cilia are the sole instruments, not merely for the production of those currents in the water which may bring them the requisite sujDplies of air and food, but also for pro- pelling their own bodies through the hc|uid. In most Zoophytes, and in the inferior Mollusks, which pass their lives with little or no change from one spot to another, the motion of the ciha lining the alimentary canal and clothing the gills (where such have a special existence), draws into the mouth the minute currents wliich serve as food, and also renews the layer of water in contact with the respiratory surface. The gills of Fishes are not furnished with ciha, another provision being .made by muscular action for conti- nually driving fresh streams of water over them ; but the motion may be very well seen upon the gills of the young 58 CILIA. — FAT CELLS. Tadpole or larva of the Water N"ewt, wliicli liang down as fringes on either side of the neck. In the higher air-breathing animals, the function of the cilia is much more limited. They clothe the mucous membrane which hnes the air-passages ; and their function appears to be, in that and other cases, to prevent the accumulation of the secretion with which the membrane is kept moist, by keeping up a continual onward movement of it towards the outlet of the passage. In some other cases, however, we find the ducts of secreting organs furnished with cilia, whose action is obviously to assist in carrying the products of secretion towards their outlet. 46. Passing on, now, to those tissues of animals of which cells constitute the permanent components, instead of being successively thrown .off and replaced as they are in the Epidermis and Epithelium, we may first notice the Adipose tissue, or Fat, in which the oily and fatty matters of the body are for the most part contained. This tissue is composed of minute cells or vesicles (fig. 14), having no communication with each other, but lying side by side in the meshes of the areolar tissue, which serves to hold them together, and through which also the blood- vessels find their way to them. Erom the fluid in these vessels, the fatty matter is separated in the first place by the secreting action of the ff? ■ >jf cells j and it is prevented ^ 14.-F.T CELLS, HIGHLY ^om malvlug its way through MAGNIFIED. tlie vcry thin walls of the cells, by the simple expedient of keeping these constantly moist with a watery fluid, the blood. ^ The blood-vessels have also the power of taking back the fatty matter again into the circulation, when it is wanted for other purposes in the economy. These deposits of fatty matter answer several important objects. They often assist the action of moving parts, by giving them support without interfering with their free motions ; thus the eye rests on a sort of cushion of fat, on which it can freely turn, and through which the muscles ^ Thus oil will nob pass into blotting-paper, if this have been previously moistened with water. FAT. CARTILAGE. 59 pass that keep it in play. It also affords, by its power of re- sisting the passage of heat, a warm covering to animals that are destined to live in cold climates ; and it is in these that we find it accumulated' to the largest amount. Further, being deposited when nourishment is abundant, it serves as a store of combustive material, which may be taken back into the system, and made use of in time of need. The causes which peculiarly contribute to the production of fat, will be considered hereafter (§ 1 G2). 47. Another tissue of which cells form the principal part, is that termed Cartilage or gristle. Its simplest state is that of a mass of firm substance, composed of chondiin (§20), through which are scat- tered a number of cells, at a greater or less distance from one another. In the sunple cellular cartilages, such as those wliich cover the ends of' the bones where they glide over one another so as to form Ni^>^-^ moveable joints, no trace of structure can be seen in the intervening substance. Fig. i5, — Section op But in cartilages which have to resist not ^^ . cartilage, •^ - . . . Showing its cells imbea- oniy pressure but also extension or strain, ded in intercellular sub- we find the space between the cells partly stance. occupied by fibres, Avhich resemble those of ligaments ; and such are tevmed Jlbro-cartilages. . They are found in Man be- tween the vertebrae of which the spinal column is made up (§ 71); and also uniting the bones of the pelvis (§ 645). Sometimes, where elasticity is required, the fibres are those of the yellow fibrous tissue (§ 23) ; this is the case with the cartilage which forms the external ear. Cartilage is not penetrated by blood-vessels, at least in its natural state. The blood is brought to its surface by a set of vessels which bulge out into dilatations or swellings upon it, so that a large quan- tity of fluid comes into the immediate neighbourhood of the cartilage, being only separated from it by the thin walls of the vessels ; and it appears that this fluid, or so much of it as is required, is absorbed by the nearest cells, and trans- mitted by them to the cells in the interior, so that the whole substance is nourished. This is precisely the mode in which the interior of the large sea-weeds (whose tissue consists of cells imbedded in a gelatmous substance, and therefore bears GO CARTILAGE. BONE. a close resemblance to animal cartilage) obtains its nourish-, ment from the surrounding fluid. 48. The permanent Cartilages seem to undergo very little change from time to time. Their wear is slow; and, being purely mechanical, it is confined to the surface. It is replaced by the materials absorbed from the blood, which are employed in the development of new cells, — sometimes within the old ones, sometunes in the space between them. When a portion of cartilage has been destroyed, however, by disease or injury, it is not renewed by true cartilaginous structure, but by what seems a condensed areolar tissue. Although cartilage does not usually contain vessels, yet these may be rapidly deve- loped in its substance, by a process which will be described hereafter (§ 393), when it becomes inflamed. This may be often seen to take place. The front of the eye is formed by a transparent lamina of a substance somewhat resembUng cartilage, which bulges like a watch-glass : this, which is termed the cornea (§ 533), is properly nourished only by vessels that bring blood to its edge, where it is connected with the tough membrane that forms the white of the eye. But when the cornea becomes inflamed, minute vessels may be seen to spread over it, proceeding from its circular edge towards its centre ; and at last some of these often become of considerable size. Under proper treatment, however, these vessels gradually shrink and disappear ; and the cornea becomes nearly as transparent as before. 49. Many parts exist in the state of Cartilage in the young animal, which are afterwards to become Bone; and it has been commonly believed that all bone has its origin in a cartilaginous structure. This, however, is not the fact, as will be presently shown. Before attempting to explain the formation of Bone, it will be desirable to describe its structure. When we cut through a fully formed bone, such as that of the thigh, we find that the shaft or elongated portion is a hollow c^dinder ; of which the walls are formed by what appears to be solid bone ; whilst the interior is filled, in the living state, by an oily substance laid up in cells, and termed marrow. Towards the extremities, however, the struc- ture of bone is very different. The outside wall becomes thinner ; and the interior, instead of forming one large cavity, is divided into a vast number of small chambers, like those STRUCTURE OP BONE. 61 of areolar tissue, by thin bony partitions, whicli cross each other in every direction, forming what is called the " cancel- lated " structure. These chambers or cancelli are filled with marrow, like the central cavity, with which they communi- cate. In the flat bones, moreover, — such as those of the head — we find that the two surfaces are composed of dense plates of bone, Hke that which forms the shaft of the long bones ; but that between them there is a layer of cancellated structure, filled in like manner with marrow. But when we examine with the microscope a thin section of even the densest bony matter, we find it traversed by a network of minute canals, continuous with the central cavity. These canals usually run, in the shafts of long bones, in the direction *'''- —^ of their length ; and are con- i f nected, every here and there, by j i liiiiF r^ cross branches (fig. 16). They i. ,|i||| are termed the Haversian canals, \ : ' "! after the name of their disco- i^ : j verer, Havers. — The hning mem- »' brane of the large central cavity ■; . is copiously supplied with blood- i jljljll vessels; and this sends off pro- 5^/,. . : . , ., .lilllll longations into the cancelli at ^ the extremities of the bone, and ^^^- ^6.— diagram reprksenting . , ,, T-T • 1 rm ^"^ STRUCTURE OF A PORTION OF into the Haversian canals, ihus the shaft of a long bone. blood is conveyed into the in- %L''3vie''ecUo";%%yc/ruriace terior of the bone : but no vessels seen m longitudinal section ; i, Ha- 1, 1111.1-i-j. veisian systems cut across, each can be traced absolutely into its having an Haversian canal in its texture, so that all the spaces f^ntre; 9,, Haversian systems cut ,.,'.,. ., __ -'• . lon^tudinally ; /, lamellae near the which lie between the Haversian surface of bone, destitute of Haver- canals are as destitute of vessels as ''^" systems. is healthy cartilage. These spaces are provided with nutriment by the following very remarkable arrangement. 50. When we cut across the shaft of a long bone, and examine a thin section with a microscope, we of course see the open extremities of the Haversian canals (fig. 17, a); just as we 'see the cut ends of the ducts and vessels of wood, when we make a transverse section of a stem. Around each of these apertures, the bony matter is arranged in concentric rings, which are marked out and divided 62 STRUCTURE OF BONE. by circles of little dark spots; and when these spots are examined with a higher magnifying power, it is seen that they ai-e small flattened cavities, from which proceed a number of extremely minute tubules (A). These tubules pass out Fig. 17.— Transverse Section of Bone. Showing the concentric rings round a a, the Haversian canals. At A are seen some of tlie cavities with^their radiating tubes, more highly magnified. from the two flat sides of each cavity; one set passes invv'ards, towards the centre of the rmg, and the other outwards, to- wards the rmg that next surrounds them. These minute tubuh, which are far smaller than the smallest blood-vessels, may thus be traced into every part of the substance of the bone ; and those proceeding from dilTcrent rings are so con- nected with each other, that a communication is- established between the innermost and the outermost circles. The tubuli which open upon the sides of the Haversian canals, are thus enabled to take up the nourishment with which they arc COMPOSITION OF BONE. 63 supplied by the blood-vessels, and to transmit it to tlie outer circles, or those furthest removed from those vessels ; and in this manner, a much more active nutrition takes place in bone than that vi^hich is performed in cartilage. It has been proved by various experiments, that the substance of bone is undergoing continual change ; and it is owing to the comparative activity of its nutritive processes, that bone is so readily and perfectly repaired, when it has been broken by violence or has been injured by disease. 51. But the peculiarity of Bone consists, not so much in this remarkable arrangement of its organic structure, as in its solidity and firmness. This is given to it by the union of a large quantity of mineral matter with the organic substance of its tissue. The mineral matter of bones consists almost entirely of two compounds of Lime ; the carbonate, with which we are familiar in the form of limestone and chalk ; and the phosphate, which is seldom found as an ingredient of rocks or soils, except where it has been derived from animal remains. The latter greatly predominates, at least in the bones of the higher animals. We may easily separate the animal and the mineral portions of the bony tissue. If we soak a small bone for some time in muriatic acid much diluted with water, the compounds of lime are entirel}'- removed from it, and the organic substance remains ; the latter is now quite flexible, and almost trans2:)arent, so that the distribution of its vessels (if they have been pre- viously injected with colouring matter) may be distinctly seen. On the other hand, if we subject a bone to strong- heat, the animal portion will be burnt out, and the earthy matter will remain. The form of the bone will be still retained ; but the cohesion between the earthy particles is so slight, that the least touch will break them asunder. Thus we see that the hardness of bone, or power of* resisting pres- sure, is given by the earthy matter; whilst its tenacity, or power of holding together, depends upon the animal portion. Although the animal substance which remains after the solu- tion of the mineral matter, has been commonly described as Cartilage, yet it is not so in reahty ; for it consists not of chondrin, but of gelatin ; and instead of being made up of an aggregation of cells united by an intervening substance, it may be torn into layers of an indistinctly-fibrous matting. In fact, 64 COMPOSITION AND DEVELOPMENT OP BONE. it corresponds closely with the wliite fibrous tissue (§ 23), both in structure and composition ; and so far from this view of its nature being inconsistent with the history of the formation of bone, it will be found to be in entire harmony with it. The proportion which the mineral bears to the annual substance of bone is very constant, when the 'proper 'osseous tissue alone is taken into account ; being almost exactly two of the former to one of the latter, or 66f per cent, to 33^ per cent. But when the composition of entire hones, including the contents of the Haversian canals and cancelli, is compared, the proj)ortion of mineral to animal matter is found to vary greatly in different classes of anunals, in the same animal at different ages, and even in different bones of the same individual ; the mineral matter predominating in bones of a comj^act texture, and the animal in those Avhose substance is more sjoongy. 52. In the first development of the embryo, a sort of mould of cartilage is laid down for the greater part of the bones ; though, in the case of the fiat bones, this mould is generally limited to the central portion, the place of their marginal part being occupied by a fibrous membrane only. The process of ossification, or bone-formation, commences with the deposit of calcareous matter in the intercellular substance of the cartilage, so as to form a sort of network, in the interspaces of which are seen the remains of the cartilage-cells. The tissue thus formed can scarcely be considered as true bone, for it contains neither lacunae nor cancdicidi. Before long, however, it undergoes very important changes ; for many of the partitions are removed, so that the minute chambers which they separated coalesce into larger ones ; and thus are formed the cancelli of the spongy substance, and the Haversian canals of the more compact. These are at first much larger than they are subsequently to become ; for they are gradually narrowed by deposits of true bony tissue, which successively take place upon their interior walls, at the expense of the materials supphcd by the blood brought thither by their contained vessels ; and it is by this forma- tion of concentric layers around the cavities of the Haversian canals, that the appearance of concentric rings is produced, wliich we have just seen to be presented by transverse sec- tions of long bones. In old bones the Haversian canals are 80 nearly filled by these deposits, that there is barely room DEVELOPMENT OF BONE : OSSIFICATION. G5 for the blood-vessels to pass along them. And it is through their complete blocking up, by a continuance of the same gi'owth, that the supply of blood is cut oif from the interior of the bone which forms the antlers of the deer, so that they die and fall off ; their shedding and rcne^ral being an annual process.^ — Whilst the formation of the Haversian canals and cancelli is being effected by the partial removal of the first formed partitions, a complete cavity is formed in the centre of the shaft of every lo7ig bone (at least in Mammals and Birds), by the entire removal of the solid tissue. This cavity is at first not much larger than one of the Haversian canals ; but as the bone grows in diameter by additions to the exterior of its shaft, so is the cavity in its interior augmented by the removal (by absorption) of the first-formed bone ; and this double process continues until the bone has attained its full diameter. The formation of new bone on the exterior of the shaft seems to be the result of the consolidation of the fibrous tissue of the periostetn^i (or membrane covering the bone) by calcareous deposit ; the lacunse being probably the cavities of cells which were entangled in the fibres, and the canalicuii being outgrowths from these ; and new fibrous tissue being formed on the outside of the periosteum, to replace that which has been taken into the bone. Thus it comes to pass, that after a time none of the bone first formed in its cartilaginous mould any longer remains, the whole of it having been removed by absorption ; since the central cavity of the perfect bone is much larger than the entire carti- laginous shaft in which it originated. And thus it also comes to pass, that (as gelatin is the basis of fibrous tissue) bones yield gelatin, not chondrin, upon being long boiled, — The increase of the shaft in length, however, is the result of a different process. In all bones of any considerable dimen- sions, the process of ossification commences in more than one l^oint at a time. In the long bones, there are usually three such points; one for the shaft, and the others for the two ^ It is commonly stated that the death of the antlers is due to the formation of a bony I'ing at their base, which cuts off the supply ot blood from the " velvet" vv'hich covers them ; but though this may con- tribute to produce the effect, it is by no means the sole cause, as the interior of the antlers is supplied with blood from the vessels of the bone from which they sprout, and not from those of the "velvet" only. QQ DEVELOPMENT OP BONE : OSSIFICATION. extremities. Long after the ossification of the shaft and of the extremities has been completed, these parts remain sepa- rated from each other by tlie interposition of a thin layer of unconsolidated cartilage ; so that, although the bone appears firm and complete, its three portions fall apart, if it be macerated sufficiently long in water for the cartilage to decay. Now it is by the progressive consolidation of the cartilage at these two junctions, and by the continual forma- tion of new cartilage as the old is taken into the bone, that the length of the shaft continues to increase up to adult age ; and then, its full size having been attained, the whole thickness of the intervening layer of cartilage is replaced by bone, so that the shaft and extremities become firmly con- solidated. — The general history of the formation of the Jiat bones is nearly the same. In these, when they are large, or have projecting out-growths, there are several centres of ossi- fication ; and although the first ossification takes place in the substance of cartilage, yet the subsequent growth seems to be effected mainly by the consolidation of fibrous mem- brane. 53. The foregoing description applies chiefly to those higher and more complete forms of Bone, which are found in Birds and ]\Iammals. In lieptiles and Fishes, the process of ossification is stopped short, as it were, at an early period ; and thus the texture of their bones resembles that which we find the skeleton to present in the earlier life of the higher animals. — The long bones of Eeptiles (with one remarkable exce})tion in the Fterodactylus, § 6G9, which is adapted to the life of a Bird) have no one central cavity, but are pene- trated by numerous large Haversian canals, like those of very young bone ; and various pieces remain separate in them throughout life, which, originating in distinct centres of ossi- fication, subsequently coalesce in Birds and Mammals. Tliis permanent separation is still more remarkable in the bones of Fishes ; and it is consequently in them that we can best .study the real composition of the skeleton, — every piece which originates in a distinct centre of ossification, being, in the eye of the philosophical anatomist, a separate bone. Further, there is a large group of Fishes in which the skeleton retains the cartilaginous character through life ; a certain quantity of mineral matter being deposited in the BONES OF FISHES : TEETH. 67 cartilage, but its conversion into true bony structure never taking place. In a few, not even a firm cartilage is produced; and all the trace of a skeleton is a cylinder formed of hex- agonal cells, resembling those of the pith of plants, which takes the place that is generally occupied by the " boclies " of the vertebrae (§ 71). Such a cylinder, "vvhich is termed the chorda dorsalis, precedes the formation of the vertebral column in other vertebratecl animals (§757). In the curious Arnphioxzis (ZooL. § 642), even this is wanting ; and the only rudiment of the bony skeleton is to be found in the fibrous sheath that surrounds the nervous centres, and sends off prolongations between the successive transverse bands of muscles, which are attached to these, as they are in other fishes to the ribs and the spines of the vertebrge. 54. In connexion with the structure of Bone, it will be convenient to describe that of Teeth, although the general description of the form and development of these organs will be more appropriately given in connexion with the account of their instrumental uses (§§ 181 — 183). The principal part of the substance of all teetli is made up of a solid tissue, which has been appropriately called Dentine. Of tliis sub- stance, one variety, which is peculiarly close in texture, and susceptible of a high polish, is familiarly known as 'diiori/. The more perfect forms of dentine, such as present them- selves in Man and the Mammalia generally, consist of a- hard transparent substance formed by the union of animal matter and calcareous salts (chiefly phos- phate of lime), in the proportion of about 28 of the former to 72 of the latter ; the mineral matter thus bearing a somewhat larger ratio to the organic, than it' does. in bone. This dentinal substance is traversed by minute tubuli of about 1-1 0,000th of an inch in diameter, Avliich appear as dark Lines, generally very close to- Fig. is. gether ; these pass in a radiating poetiok of df.ntink (highly magni- manner from the central cavity fied), showing its tubular structure, of the tooth, diverging from each other as they approach its exterior; but when seen in only a small part of their p 2 68 STRUCTURE OF TEETU. course, they appear to be nearly parallel (fig. 18), tliougli visually more or less wavy. They occasionally divide into two branches, which continue to run, at a little distance from one another, in the same parallel direction ; and they also frequently give off small lateral branches, which again send off smaller ones. In some animals the tubuli may be traced at their extremities into minute cavities analogous to the lacunas of bone ; and the lateral branchlets also occasionally terminate in similar cavities. Thus the whole tooth may be likened, in some degree, to a single Haversian system in bone ; the central cavity, which is lined by a vascular mem- brane, representing the Haversian canal, while the radiating tubuli of the former correspond with the radiating canaliculi of the latter ; the chief difference lying in the absence of lacunae along the course of the radiating tubes. In a large proportion of Fishes, however, there is no single central cavity, but the whole tooth is traversed by a system of medullary canals, not only resembling the Haversian, but actually con- tinuous with those of the bone on which the tooth is im- planted; and as each of these is the centre of a distinct system of radiating tubuli, the resemblance of their dentine to bone is very close. A somewhat similar condition of the dentine (obviously a lower or less specialized form of this substance) presents itself in certain Eeptiles and Mammals. — In the Teeth of Man and most other Mammals, and in those of many Reptiles and some Fishes, we find two other sub- __„_ stances, one of them harder and /~r^p |tr~7 " ''' !'a.^ ' the other softer than dentine. ^^~£^ 'ri ' ^'-^-^^ former, which is called ^P-^^^^ 'W] Ena7nel^ consists of long pris- C- — --~-w '|il>s j matic cells, which pass from one v~--~--J^--'^--jv4 surface to the other of the thin \3J^pifr:£Ez£?|/|i' layer formed by this substance w-^isf— Hs^/# jj I over the crown, or sometimes in I C^g^?tS=f^f|| the interior of the tooth (§ 182). I — ^Si^^&MiEi These prisms are usually hex- Fig. 19- agonal in form, as is seen in Portion OF Enamkl (highly niap;ni-|;j,^^j^S^ej,Sg gCCtion (fig. 19); and ned), showing its component prisms. . . \ r> J } ^ their course is usually more or less wavy. In teeth which have to sustain an extraordinary amount of compression (as is especially the case with those of TEETH. MUSCLE AND NERVE. 69 the Eodentia), the enamel-prisms cross and interlace with one another, in such a manner as to prevent that separation which would readily occur if the direction of all of them were the same. Of all the tissues of the animal body, the Enamel is the most remarkable for the predominance of mineral ingredients ; these amount to no fewer tlian 98 parts in 100, leaving wdien removed only 2 per cent, of organic matter. The softer component of Teeth, known as the Cemcntum, or Crusta jietrosa, possesses the essential characters of true bone ; but when only a thin layer of it is present, we do not find it traversed by medullary canals, its system of lacunse and canaliculi being then in relation to the nearest vascular surface, — as is the case also with very thin laminae of ordinary bone, such as we find in the scapula (blade-bone) of a Mouse. 55. We come, lastly, to the two tissues whicli are of the highest importance in the Animal fabric, and to which all the rest are merely subsidiary ; namely, the Muscular and the Nervous. It is through tlie instrumentality of these, that all ^ the actions are performed which essentially constitute Animal life ; for the nervous apparatus is the medium by which the consciousness of the individual is affected by what takes place around him, or within his own body, and by whicli, in his turn, he originates movements in his body, and through it in things external to it ; whilst the muscles are, so to speak, the servants of the nerves, doing, with a force of their o^\m, the work which the nerves direct. The relation between the two may be likened to that of the rider and his horse, or of the engine-driver and his locomotive ; for the nerves can put forth no motor power by themselves ; whilst, on the other hand, the muscles (with certain excep- tions) remain inert except when stimulated to contract by the agency of the nerves. The muscles use the tendons and the framework of bones, joints, &c., for the mechanical appli- cation of their power, as will be shown hereafter (Chap, xii.); but these parts of the fabric have not the slightest power of originating motion by themselves. Hence, all Animal Force takes its rise in one or other of these two tissues ; and we shall find that the special purpose of the whole apparatus of Organic life, is, by providing materials for their nutrition and renovation, to build them up in the first instance, and then 70 STRUCTURE OF MUSCLE. h-) keep them in working order. For every development of animal force involves a change of state of the Nervo-mus- cular substance : a certain amount of it ceasing to exist as living tissue, and passing into the condition of dead matter ; and its elements resolving themselves, under the influence of the free oxygen brought to them by the blood, into new combi- nations, which are carried forth from the body as quickly as possible. Consequently, if the ISTervo-muscular tissues be not renewed as rapidly as they are used up, their powers must speedily fail from the progressive loss of their substance. In this particular they are on a difi'erent footing from the other elementary parts of the organism ; for although each of these seems to have a certain term of life, the length of which is in some degree related inversely to its functional activity, — those wliicli live the fastest having the. shortest individual duration, and vice versd, — there are none which are called upon to give forth their whole vital energy in one effort, and which may thus have their existence as parts of the living organism terminated at any moment by a demand for their peculiar power. 56. Muscular Fibre presents itself under two forms, which are ordinarily very distinct from each other ; although it is prolmble that they may ultimately prove to be but modifi- cations of one and the same. The first, which is known as the striated fibre, is that of which all those muscles are com- posed, which constitute what is commonly designated as "flesh" or the "lean" of meat. If any "joint" of meat be even cursorily examined, it will be seen that its whole substance is made up of distinct masses, held loosely together by areolar tissue ; and these masses, Avhicli are known as " muscles," are easily isolated from each other by dissection. Every such Muscle is formed by the union of a number of bundles, having a generally parallel arrangement, which are closely bound together by areolar tissue, and are themselves composed of bundles still more minute, united in a similar manner. These, again, may be separated in the same way ; and at last Ave come to the primitive fibres of which this tissue is composed. Each of these primitive fibres termi- nates at either extremity in tendinous fibre, which unites with other fibres to form the tendinous cords or bands, that are attached to the points of the skeleton which the muscle STRIATED MUSCULAR FIBRE. has to bring together. The muscular fibre itself consists of a delicate membranous tube, enclosing a great number of Jlbrillce, or extremely minute fibrils, -which are not capable of further division (lig. 20). The peculiar transverse marking Fi"-. 20.— Striated Musculau Fibri: separ.atii;g into Fibrill;e. or striation by which this form of muscular hbre is characterised, is found, when the fibre is separated into its fibrilla), to be due to the pecuhar markings which every fibril presents. These markings, consisting of alternate light and dark spaces, give to the fibril a beaded appearance ; but this is only an optical deception, since its form is in reality cylindrical, or nearly so. It is easy to see how the correspondence of the hght and dark spaces respectively, throughout the whole bundle of the Jihril, will give rise to the banded appearance which the entire fibre presents. The form and diameter of the fibres vary considerably, both in different tribes, and in different parts of the same animal. In the higher classes, their form usually approaches a cyhnder ; but the parts which press against one another are somewhat flattened, so that it is more or less prismatic. In Insects, on the other hand, the fibrillae are arranged in flat bands, so that the fibre often consists of but a single layer of them. The diameter of the fibres in Man averages about l-400th of an inch, and does not differ very widely in either direction ; in the cold-blooded Yertebrata, however, the average size is greater, and the extremes are also wider ; the diameter of the fibres varying in the Frog from 1-lOOth to 1-lOOOth of an inch, and in the Skate from l-65th to l-300th of an inch. The diameter of tho Jibrils is nearly the same in all classes, seldom departing much from 1-1 0,000th of an inch ; and the average distance of the dark strise from each other is nearly the same. 72 NON-STRIATED MUSCULAR FIBRE. 57. Ilic other form of Muscular Fibre, whicli, from the absence of transverse striation, is distinguished as smooth or non-striated, is found not in large masses, but in thin layers, forming jiart of the wall of various hollow organs, such as the stomach and intestinal canal, the bladder, the principal gland- ducts, and the larger blood-vessels. In all these situations it is so exclusively concerned in the performance of the vege- tative or nutritive functions, and it is so entirely mthdrawn from the influence of the will, that it has been frequently designated as " the muscular fibre of organic life ;" the striated fibre, of which the voluntary muscles are composed, being distinguished as the "muscular fibre of animal life." Eut these designations are not by any means consistent with the facts of the case ; for in a large proportion of the ]\Iolluscous classes, the muscles of animal life are composed of non- striated fibre, whilst the heart of Man and of other Yerte- brata, though a muscle of organic life, is made up of striated fibre. In fact, the employment of the one or of the other kind of fibre would seem to be chiefly determined by the kind of contraction which is required from it (§ 59). The non-striated fibres are arranged, like those of the other muscles, in a parallel manner into bands or bundles; but these bundles, instead of being them- selves grouped into larger ones having a like parallel arrangement, are gene- rally interwoven into a kind of network, having no fixed points of attachment. The form of the indivicbial fibres is much more variable than tliat of the striated kind, being often very much flattened out ; and hence their general dimensions cannot well be estimated. By macerating a portion of this kind of tissue in dilute nitric acid, each fibre may be resolve< ' into bundles of long spindle-shaped bodies, which, contain- ing elongated staff-shaped nuclei (fig. 21), may be regarded as cells, al- though it is diificult to distinguish their walls from their contents, aiiis form of muscular tissue is commonly mingled with a large quantity of the ordinary fibrous Fi-. 21. A, Portion of a band composed of non-striated muscular fibre, showing, a a, the spindle-shaped cells, and, b b, the elonfrated nuclei : B, a single cell isolated, and more highly magnified ; C. a similar cdl treated with acetic acid. MUSCULAR CONTRACTION. 73 structure ; and we find it dispersed in small quantity through the latter in the skin, to which (especially in jDarticular regions) it gives a contractility that is manifested under the influence of cold or of mental emotions, and thus produces that general roughness and rigidity of the surface which is known as cutis anserina, or "goose's skin." 5S. Under the influence of certain exciting causes, or stimuli (Chap, xii.), striated muscular fibres suddenly and for- cibly contract. Their two ends approach one another, and their stria3 become closer ; but they bulge out in the middle to a corresponding degree. This causes a like change in the bundles which are made up of these fibres ; and thus the whole muscle, when shortened by the drawing together of its two ends, is greatly enlarged in diameter, especially towards its middle. Of this any one may convince himself, by bending his fore- arm upon the arm (as when the hand is brought to the mouth), and feeling the fleshy mass upon the front of the latter. The muscle, in fact, does not in the least degree change its ow^n bulk in the act of contraction ; for its enlarge- ment in diameter is exactly equivalent to the shortening of the distance between its extremities. The contraction of a muscular fibre is ordinarily followed, after a short interval, by its relaxation ; of this we have a remarkable illustration in the contractions excited by the electric stimulus. But relax- ation of individual fibres is not incompatible Avitli the con- tinuance of the state of contraction of the muscle as a wdiole. For it appears that wdien an ordinary muscle is thrown into contraction, all its fibres do not usually contract together, but only a small part of them ; and that, as long as its contraction is maintained so as to exert a constant force, a continual in- terchange is taking place in the action of the fibres by which this is kept up — those which have been shortened becoming slack, ancl being replaced (as it were) by others, which pass into the contracted state for a time, and then relax again, being succeeded by another set. I^ow as the ends of those fibres which are actually in a relaxed condition, are brought near together by the contraction of the rest, the fibre is thrown out of the straight line, and assumes a wavy or zigzag form, which was formerly supposed to be the state of con- traction, but is now known to be otherwise. This peculiar arrangement gives place to the straight form, either when the 74 MUSCULAR COXTRACTIOX. NERVOUS TISSUE. fibre passes into the state of contraction, or when, by the relaxation of the whole muscle, its ends are separated again to their full extent. 59. Kow the alternate contraction and relaxation, which is thus made to produce a continued contraction in ordinary- muscles, elsewhere occasions a different effect. Thus in the heart, all the fibres of the ventricles seem to contract to- gether and all to relax together, — those of the auricles contract- ing whilst the others are relaxing, and vice versd; — and in this way the alternate contractions and dilatations of that most important organ are continually kept up. Again, in the muscular coat of the intestinal canal, we observe the contraction of each part to be almost immediately followed by its relaxation ; but the peculiarity of its movement is, that the contraction is jDro- pagated on (as it were) to the succeeding part, which in its turn contracts and then relaxes, producing the same action in the part that follows it, — and so on along the whole canal. This peristaltic motion (§ 215), as it is called, is obviously adapted to propel the contents of the intestinal tube from one ex- tremity of it to the other ; just as the peculiar action of the heart is adapted to receive and propel the blood alternately, or as the mode of contraction of the ordinary muscles enables them to keep up a continued strain for a great length of time. It is much less rapid and energetic than the action of the heart ; for it is the characteristic of the non-striated fibre, that its contraction follows much less closely on the application of the stimulus, and is much less rapidly succeeded by relaxa- tion, than that of the striated fibre. 60. The Nervous tissue consists of two distinct structures, of one of which the trunks of the nerves are entirely made up, whilst the other enters largely into the composition of the ganglia or centres of action (§ 61). The former, termed the ivhiie or fibrous tissue, consists of straight fibres, lying side by side, and bound together by areolar tissue into bundles (fig. 22); these, again, are united with others into a larger group ; and by the union of a considerable number of such groups, the nervous trunks are formed, which are dis- tributed through the body, especially to the skin and muscles. Nervous Fibre, like muscular, presents itself in the higher animals under two forms, of which one may be considered as more completely developed than the other ; these are known STRUCTURE OF TUBULAR NERVE-FIBRES. U) as the tubular and the geJafinous. The " tubular " fibres are so named because each poss.' ^os a distinct tubular sheath of a delicate structureless membrane (fig. 22, a), which encloses the proper nerve-substance, and isolates it completely from the Fig. 22.— Structure of Nerve-Tubes. Tubular Nerve-fibres ; A, from a nerve-trunk; B, from the substance of the brain blood-vessels and other surrounding structures ; this tube does not either l^ranch or unite with others, and there is reason to believe it to be continuous from the origin to the termination of the nerve-trunk. Within the tube is a hollow cylinder of a material known (after its discoverer) as the " white substance of Schwann ;" and tliis encloses a sort of central pith, which is transparent and semi-fluid in the living state, but undergoes a kind of coagulation into a granular sub- stance after death, and under the influence of chemical re-agents. There is reason to believe that this central pith or " axis-cylinder " is the essential component of the nervous fibre, and that the hollow cylinder which surrounds it serves only to isolate it more completely; for we not nnfrequently see the former to be alone contmued, both the tubular sheath and the white substance stopping short ; and this at either extremity of the fibre, where it separates itself from those with which it is bound up in the nerve-trunk. The proper form of the fibre seems always to be truly cylindrical ; though 76 TUBULAR AND GELATINOUS NERVE-FIBRES. it is very liable to be altered by manipulation, a small excess of pressure in one part forcing the contents of the tube towards some other where they are more free to distend it, and thus producing a swelling. The greater delicacy of the tubular sheath in the hbrous substance of the brain and spinal cord, renders its fibres peculiarly susceptible of this kind of alteration, so that they often present under the microscope a somewhat beaded appearance (fig. 22, b) ; when carefully examined, however, without any previous disturb- ance, these fibres are found to be as cylindrical as those of the iicrve-trunks. The diameter of the nerve-tubules is usually between l-2000th and l-4000th of an inch ; but it may be somewhat greater or considerably less than this average. They are larger in the. nerve-trunks than they are near their central termination in the brain ; and it is a remark- able circumstance that the fibres of the nerves of " special sense " are considerably smaller than the average in every part of their course. — The " gelatinous" fibres cannot be shown to consist of the same variety of parts as the preceding ; for neither the tubular sheath nor the white substance of Schwann can be distinguished in them. They are flattened, soft, and apparently homogeneous, sometimes showing a dis- position to split into very delicate fibrillae ; being of a yellowish-grey colour, they are sometimes designated the grey fibres. Their diameter averages between the 1 -4000th and l-6000th of an inch. As these "gelatinous" fibres form a considerable proportion of the trunks of the Sympa- thetic system of nerves (§ 461), they have been supposed to belong properly to it, and to minister exclusively to the organic functions, like the non-striated muscular fibre (§ 57); but-Uhere is no doubt that this is an incorrect notion, and that even the fibres of the ordinary nerve-trunks ma}^ present the " gelatinous " aspect, probably from incompleteness of development. Gl. In the central organs of the Nervous system — namely, the brain and spinal cord of the Vertebrata, the ganglia or knot-like swellings on the nervous cords which take their place in the lower animals, and similar ganglia belonging to the Sympathetic system — we find a form of nervous tissue altogether distinct from the preceding ; which, from its con- sisting ol' lavLC cells or vesicles, is generally known as the VESICULAR OR GANGLIONIC NERVE-SUBSTANCE. 77 vesicular. Tliese nerve-vesicles, sometimes knov/n as gan- giion-giobiiles, may be regarded as originally spherical, or nearly so, in form (fig. 23, a); but they often present one or more prolonged extensions ; and as these when single re- semble tails, and when multiple are like the rays proceeding from a star, the cells are said in the first case to be "caudate," Fig. 23. — Vesicular Nerve-subst.vnce. A, combination of Ganglion-cells (of which one is shown separately at a, more highly magnified), and Nerve-fibres in the grey substance of the brain, which is also traversed by a capillary vessel, b; B B, Ganglionic cells with caudate pro- longations. and in the second to be stellate (b). These prolongations have been traced into continuity, in some instances, with the axis-cylinders of nerve-tubes, whilst in other cases they seem to unite with those proceeding from other vesicles. It is not by any means certain, however, that the nerve-tubes thus connect themselves with the nerve-vesicles in all instances ; since it frequently appears as if the former passed in among the latter, without coming into direct continuity with them. Sometimes a ganglion-cell seems to lie in the course of a tubular fibre, which enlarges to envelope it, and then con- tracts again to its former dimensions. There can be no reasonable doubt, however, that in some way or other the nerve-fibres and the nerve-vesicles come into some kind of communication in the ganglionic centres. The vesicles arc STRUCTURt: OF GANGLIA. — NERVOUS ACTION. filled ^vith a finely-gi^anular substance, wliicli extends into their prolongations ; and in the warm-blooded Yertebrata they contain pigment-granules, which give them a reddish or yellowish-brown colour ; so that the aggregations of vesicular substance which we hnd in the larger nervons centres, are distinguishable by their greyish hue. This "grey matter,"as it is frequently called, is disposed on the surface of the brain; but it occupies the interior of the spinal cord, and holds the same position in the smaller ganglionic centres (fig. 24). It is not only, however, in the central organs that nerve-vesicles are found ; for they present themselves also in certain situa- Fig. 24.— Thin slice of tions at the other extremities of the nerve- o?^THF symi^\tuftic f^^^'^^' Tlius we find a large proportion Systkm, siiowiiig the of the retina (§ 535), which is commonly libSes^'^amonJst^^'Inl described as a mere expansion of the optic giionic cells. ncrvc, to be composed of nerve-vesicles that arc scared}^ distinguishable from those of the brain ; and it is probable that the ultimate branches of other sensory nerves have some such termination. Wherever we meet mth vesicular substance, we find it imbedded in a minute net- work of blood-vessels ; and a copious supply of oxygenated blood is requisite to the due performance of its actions. (j2. There can be no doubt that the special office of the l^ei'VQ-Jlbres is to convey the influence of the changes which are eflectod in one part of the system, to other and remote parts ; just as the Avires of a galvanic battery conduct the electric influence from the instrument which excites it, to some distant point where it is to be applied to some use. The effects of such changes in the state of the iS^ervous System are propagated in two opposite directions ; — the im- pressions made upon the skin and other parts possessed of sensibility, being conveyed towards a portion of the nervous centres called the sensorium, and there giving rise to sensa- tions; — and the influence of the emotions or volitions to which these sensations give rise (§ 7), being propagated /rom the central organs to the muscles, which tliey excite to con- traction. And by the discoveries of Sir C. Bell, hereafter to ACTIONS OF NERVOUS SYSTEM. 79 be described it has been fully proved that these opposite changes are conducted by two different sets of fibres ; — one conveying to the central organs those Avhich originate in the cii'cumference ; — and the other conveying to the circum- ference those which originate in the centre (§ 451), The transmission of these changes is completely interrupted by division of the nervous trunk, or by pressure upon it ; and it sometimes happens that one set of conducting fibres is thus affected, whilst the functions of the other are not impaired; so that a limb may retain its sensibility and yet be totally destitute of the power of motion, or may be completely obedient to the will though totally destitute of sensibility. In Yertebrated animals, w^e find some nerves in which there is only one set of fibres, so that the trunk is only sensory or only motor (§ 459); but in general, the two sets are bound up together in the same sheath. 63. The motor fibres may bo considered as originating in the vesicular substance of the central organs, and as termi- nating in the muscles ; the power which is generated in the former being conveyed by their means to the apparatus through Avhich it operates to produce mechanical motion. When the nerve-trunks reach the muscles, they divide into branches which penetrate their substance, and these again subdivide and ramify minutely, so that at last the fibres may often be observed running singly, passing amongst the muscular fibres, but not appearing to penetrate their tubular sheaths. These terminal fibres seem often to double back upon themselves, so as to form loops, either re-entering the branch from which they issued, or connecting themselves with some neighbour- ing branch ; so that the ultimate distribution of the motor nerves in the muscular substance, is a sort of plexus or net- work. The sensory fibres, on the other hand, may be con- sidered as originating in the sensory surfaces, such as the skin, the interior of the nose, the lining membrane of the cavities of the internal ear, the retina of the eye, &c. ; and as passing towards the central organs, conveying to these the impressions they have received, which impressions may either affect the consciousness, or may excite respondent move- ments, or may act in both modes, through the instrumentality of the vesicular substance to wliich they are transmitted. The immediate dependence of the functional activity of this «0 SIMPLIFICATION OF STRUCTURE IN LOWEST ANIMxVLS. substance upon the supply of blood which it receives, is shown by the fact, that if this supply be temporarily cut off, either by failure of the heart's action (as in fainting), or by pressure on the blood-vessels which convey it, immediate insensibility, with loss of all power of motion, is the result. And the same is the case with regard to the organs of sense ; for if the circulation through them be interrupted, no sensory impression can be made upon the nerve-fibres which originate in them, as we see when the movement of blood in a limb is suspended by pressure upon its artery. G4. The foregoing constitute the principal tissues among the higher animals, in which the principle of division of labour is most fully carried out, every component part having its own peculiar structure and its own special action. As we de- scend in the scale, we find these distinctions less and less obvious, so that when we come down to Zoophytes (§ 121), we meet with but little differentiation either in the textures or in the actions of the several parts of the body ; the whole sub- stance of these animals being composed of a tissue, which very closely resembles that which is first formed in higher animals for the reparation of wounds, having the appearance of a solidified blastema (§ 34), wdtli nuclear particles, in various phases of development into cells and fibres, more or less thickly scattered through it ; and this substance being everyAvhere contractile, and everywhere (at least in many instances) equally capable of participating in the func- tions of nutrition and reproduction. And when we pass still lower, to that simplest type of animal life, which is pre- sented to us in the Rliizopods (§ 129), we do not meet with even this amount of definite structure, but find the entire sub- stance of their bodies composed of an apparently homogeneous jelly, which, like the more organized tissue of the Zoophytes, is everywhere contractile, and which has also the power of performing every operation required for its growth and main- tenance as a living being. In such creatures there is not the sliglitest vestige of a Nervous system ; and it remains a question whether, in consequence of this deficiency, they are altogether destitute of consciousness, or whether this endo\vment is dif- fused, as it were, through the whole substance of their bodies. G-5. Every component part of the fabric must be regarded INDEPENDENT VITALITY OF PARTS OF ORGANISM. 81 as having a life of its own, wliicli it maintains by drawing to itself the nutrient material supplied by the circulating cur- rent ; but as the continuance of its vital activity is dependent upon the continuance of its nutrition, the life of no tissue can be prolonged for any considerable period after the circu- lation has ceased. But after the movement of the blood has come to an end, though the body as a whole is dead, its parU may remain alive for a certain time, and may perform their functions, so long as they are supplied with the necessary materials. Thus, various secretions, the growth of hair, and muscular movements, have been ol3served to take place in dead bodies. But they cannot continue, because the neces- sary conditions are withheld by the stoppage of the circu- lation, — a function which thus binds, as it were, into one whole the scattered elements, and causes the different opera- tions to minister one to another. As every component part has an independent life, so has it a limited duration, quite irrespective of that of the organism as a whole. Thus the cells which float separately in the blood, seem to be con- tinually undergoing change, — dying, and giving place to new ones. We have seen that the cells of the epidermis and of some parts of the epithelium are being constantly thrown off and renewed. The duration of the cells of fat and cartilage appears to be much greater ; in fact, we have no precise knowledge of their term of life. That of the bony tissue is probably greater still ; yet there is adequate evidence that it is by no means indeterminate. But that of the muscular and nervous tissues seems to depend almost entirely on the use that is made of them. Thus we may justly say, — how- ever startling the assertion may seem, — that death and decay are continually going on in every living animal body, and are essential to the activity of its functions. QQ. Many animals are reduced to a state of apparent dec^th by dryness, by cold, or by exclusion of" the air. A curious example of the first kind is furnished by the Tardigrada (Zoology, § 841) ; some species of which may not only be completely dried up, but may even be exposed in that state to a temperature much exceeding that of boiling water, without losing the power of recovery when moistened. A similar power of revival after being dried up is possessed by the common Wheel Animalcule, and probably also by the G 82 SUSPENDED ANIMATION. eggs of many minute Entoiuostracous Crustacea (Zoology, §§ 883, 931). It is unquestionable that many Fishes, especially those of fresh- water lakes, will revive on being thawed after havinut in general the movement of their Fig. 34. — Anatomy of a Coluher bodies is accomplished by their being drawn-up into folds, and then straightened so as to project the head. The pro- longed form of the body in Serpents occasions a curious variation in the arrangement of the principal organs, which is shown in the accompanying figure. The oesophagus or STRUCTURE OF SERPENTS AND BATRACHIA. 97 giiUet, ce, whicli leads from the mouth, to the stomach, is a long and very wide canal, being even larger than the stomach at its commencement ; a portion of it is removed at ce', in order to show the heart, &c., which would otherwise be con- cealed by it. The stomach, i, is long and narrow ; and the intestinal tube, i', after making a few turns or convolutions, passes backwards in a straight line, to terminate in the cloaca, d, which opens externally by the orifice, an. The liver, /, is also much lengthened. From the mouth also proceeds the long ^vindpipo, t t, which conveys air to the lungs, or rather to the single lung ; for the lung on the left side, j)', is scarcely at all developed, whilst that on the right, p, extends along a great part of the body. At o is seen the ovariimi, in which the eggs, o' o', are produced ; and this also is very much lengthened, extending from the cloaca a good way up the body, so as nearly to meet the lung. The other references are to the parts of the heart, and the principal vessels ; the structure and arrangement of which will be explained here- after (§ 284). 86. The Batrachia, or animals of the Frog tribe, are readily distinguished from all the preceding, by their soft naked skins ; even when the form of the body, as in the com- mon Salamander or Water-]N'ewt, resembles that of the lizards. They are also remarkable for the metamorphosis which they undergo in the early part of their lives : for they come forth from the egg in a condition which is, in all essential particu- lars, that of a fish, and undergo a gradual series of changes, by which their form and structure become assimilated to those of the true reptiles. This change is most complete in the !Frogs and Toads ; the early form of which is known as the tadpole. The principal stages of this change are represented in figs. 35 to 39 ; in which, however, the relative sizes are not preserved, the tadpoles being much larger in proportion (for the sake of displaying their form and the gradual development of their legs) than the complete frog. Soon after the young tadpole has come forth from the egg, it pre- sents the form which is shown in fig. 35 ; its head and trunk are large, and the latter is prolonged into a flattened tail, by which the little animal swims freely through the water. There is not the least appearance of limbs or mem- bers. It breathes by gills, which are long fringes, hanging H 98 3IETAM0RPH0SIS OF BATRACHIA. loosely in the water on either side of the head. At a later period, however, these gills, which are merely temporary, disappear ; and the breathing is carried on by another set, which are situated behind the head, and are covered in by a fold of skin; the water gains access to these by passing through the mouth, exactly as in Fishes. The form is then that which is represented in fig. 36. In a short time after- wards, the animal still breathing by its gills, the hind-legs begin to sprout forth, as it were, at the base of the tail ; this Fig. 36. Fig. 37. Fig. 35. Fig. 39. stage is shown in fig. 37. At a still later period, the fore- legs begin to be developed, as seen in fig. 38 ; and from that time they are nourished at the expense of the tail, which gradually disappears, as seen in fig. 39, a, h. During this period, other important changes are taking place in the inte- rior of the body ; the chief of which are the development of the lungs and the gradual disuse of the gills, so that the animal becomes fitted to live on land and breathe air, and is no longer capable of remaining long under water without coming to the surface to respire. 87. The metamorphosis in other members of the group is PERENNIBRANCHIATE BATRACHIA. 99 less complete than in the Frog, being checked at a less advanced stage. Thus in the common Water-Newt, the tail is retained during the whole of life, and the animal continues to be an inhabitant of the water, though breathing air alone. There are some very curious animals, however, in which the change is stopped, as it were, at a much earlier period, so that the gills also are retained ; and in these, the lungs are suffi- ciently developed to enable the animals to breathe air, so that they can live either on land or in water. Such Batrachia are scientifically known as perennihranchiate, this term express- ing the persistency of their gills. In tig. 40 is represented Fig. 40.— AxoLOTL. an animal of this kind, the Axolotl, which inhabits some of the lakes of Mexico. And in fig. 41 is shown the form of a still more remarkable animal, the Lepidosiren, or mud-fish, recently 41. — Lepidosiren. brought from th ; rivers of Africa, the metamorphosis of wliich appears to be checked at a still earlier period, so that it is very difficult to decide whether it should be regarded as H 2 100 STRUCTURE OF FISHES. a Fisli or as a Eeptile, so complete is the mixture of charac- ters which it presents. 88. The class of Fishes is distinguished from all other Vertebrata, by the adaptation of the animals composing it to breathe by means of water in their adult state, so as to be capable of living in that element only. Like Eeptiles, they are oviparous and cold-blooded ; and in these characters they differ completely from the Whales and other Mammals, which are, like them, inhabitants of the great deep, but which are warm-blooded, viviparous, and air-breathing animals. There is a simple external character, by which the members of the two classes may be at once distinguished. The animals of the Whale tribe are, like fishes, chiefly propelled through the water by means of a flattened tail ; but in the former the tail is flattened horizontally, so that its downward stroke may serve to bring the animal to the surface to breathe; whilst in Fishes it is flattened vertically, that its strokes from side to side may simply propel the fish through the water. A liattenmg or compression of the body is seen more or less in almost all fishes, and is intimately connected with the nature of their motion through the element they inhabit ; as it serves the double purpose of diminishing the resistance which is offered to their progress, and of increasing the extent of the oar-like surface, by the lateral stroke of wliich the body is propelled forwards (Chap. xii.). This stroke is given by a series of muscles of great power, Avhich pass from the prolonged extensions of one vertebra to those of another, and altogether make up the principal part of the bulk of the animal. The fins which represent the limbs are not so much used in propelHng the Fish, as in changing its direction either laterally or vertically. Thus in the lowest group of the Vertebrated series, the act of motion is chiefly performed by the vertebral colimin itself, instead of being committed to the limbs, as in Mammals, Eirds, and most Eeptiles. The larger number of Fishes swim with great activity ; and their lives may be said to be passed in seeking their subsistence and in flying from their enemies. 89. Fishes are for the most part very voracious, and their food consists in great part of the members of their own class. In seeking it, they appear to be chiefly guided by the sight ; fur their eyes are usually large and highly developed, while STRUCTURE OF FISHES. 101 the other organs of sense are formed upon a very inferior t}^e. They swallow it without much division in the mouth ; but it seems to undergo rapid digestion. The blood of some rish, whose muscular activity is pecuHarly great, is rich in red corpuscles, and of a temperature not much lower than that of Mammals ; but, generally speaking, it contains much less solid matter than that of the w^arm-blooded Vertebrata, and its temperature follows that of the surrounding medium. 90. Although Fishes breathe by gills instead of by lungs, these gills are connected with the mouth, so that the water which passes over them is received into it, in the same man- ner as the air is in the higher Vertebrata. This is a character which distinguishes the position of the gills of fishes from that of the corresponding organs of any of the inferior tribes. They are lodged in a cavity on each side of the throat ; and this cavity opens outwardly, either by one large valve-hke aperture on either side, or by several ; through these apertures the streams of water which have been taken in by the mouth, and forced over the gills by the action of its muscles, make their exit. 9 1 . All Fishes are oviparous ; and the number of eggs which they produce is generally prodigious. It is very seldom that after the eggs have been deposited and fertilized, the parents take any further concern in regard to them ; though there are a few instances in which a kind of nest is made, and others in which the egg is retained and hatched within the body, so that the young comes forth alive. This last is the case with the Sharks and Eays, which, notwithstanding that their skeleton is cartilaginous, are higher than Fishes generally in several other parts of their organization. 92. All the animals which are destitute of a vertebral column are called Invertehrata ; and this division into the Vertebrated and Invertebrated groups was formerly regarded as the first step in the classification of the animal kingdom. But it was pointed out by Cuvier, that in the Invertebrated division are comprehended three groups, of which the mem- bers differ as much from one another as they do from Verte- brated animals ; and that each of these ought, therefore, to rank with the first, as a primary division. This is evident to those who are but slightly acquainted with the structure of the animals already named (§ 69) as characteristic speci- 102 GENERAL STRUCTURE OF ARTICULATA. mens of these divisions ; and it will become more apparent as we proceed. 93. In the second division, that of Articulata, or Articu- lated (jointed) animals, we find a conformation very different from that which has been just described. The exterior of the body is still perfectly symme- trical, as in the Yertebrata ; and the interior is even more symmetrical ; for the organs that represent the heart and lungs are equally dis- posed on the two sides of the central line of the body. But the skeleton, instead of being internal, is external; and is composed of a series of pieces jointed together, which form a casing that in- cludes the whole body. In general, these pieces are very similar to each other ; so that the whole body appears like the repetition of a number of similar parts, as we see in the Centipede (fig. 42). The limbs are usually very numerous, where they exist at all ; and they have a jointed cover- ing, like that of the body. But in the lower tribes of this group, such as Leeches and Worms, the limbs or members are but sHghtly developed, or are altogether absent; and in the highest, wliicli approach most nearly to the Yertebrata in their general organization, the number of members is much reduced, — although it is never less than six. The hard matter of which the cen'ti ^^* external skeleton is composed, undergoes little or no change when it is once fuUy formed ; and, in order to accommodate it to the increasing size of the animal, this covering is thrown off and renewed at intervals during the period of growth. 94. The nervous system consists of a series of separate ganglia, which are arranged in a cord or chain along the central line of the body. There is iisuaUy a pair of large gangha in the head, bearing a reseml) lance (in their peculiar connexion with the eyes) to the ganglionic centres of the optic nerves in Yertebrata; and there" is commonly one for each segment or division of the body, from which the nerves pass to supply its muscles, as they do from the spinal cord of Yertebrata. The cord which connects these ganglia is double. GENERAL STRUCTURE OP ARTICULATA. 103 and the ganglia tliemselves are composed of two halves, which have little connexion with each other. The chain thus formed (fig. 43) passes along the nnder-side of the trunk of the animal (as seen at g, fig. 44), not on what seems its back ; and by the presence of tliis double chain of ganglia an Articulated animal may be distin- guished, even when, in its general structure, it should seem to belong to the group of Mollusca (§ 102). 95. The general arrangement of the organs in the Articulata is shown in the accompanying figure of a Cray- fish. The mouth, situated on a pro- jecting head, opens into s, the stomach, from which passes backwards the in- testinal tube, i, i, to terminate at the opposite extremity of the body. Tlie upper part of the tube is surrounded by the liver, /, which is here very large. In the head are seen the ganglia, c; and along the under-side of the body is seen the chain of ganglia, Fig. 43. -Nervous System AN Insect. Fig. 44.— Diagram showing the position of the piuNcirAt, Organs ik THE Articulata. g. The blood is nearly colourless, and is usually impelled through the body not by a single organ or heart, but by a succession of contractile cavities, one for each segment, which open into one other longitudinally, forming what is known as the dorsal vessel; in the Cray-fish and its aUies, however, one part of this, h, is specially enlarged, so as in great degree to serve as a heart for the system generally. The respiratory organs are not connected with the mouth ; and are not usually 104 STRUCTURE OP INSECTS. restricted to one part of the body, but are diffused either on its outside or through its substance. 96. The organs of sense, in this group, are less numerous than in Yertebrata, and are inferior in perfection ; those of sight are the most developed, and are formed upon a very peculiar plan (§ 573); but all organs of special sense appear wanting in the lowest tribes. Yet we find that the muscular power is very great ; for the animals of this group, taken as a whole, can move faster in proportion to their size, and possess greater strength, than those of any other. We observe, too, that -with little or no intelligence, they are prompted to the most remarkable actions by instinct alone. They seem to act like machines, doing as they are prompted, without choice, or knowledge of the end to be gained ; and consequently the dif- ferent individuals of the same species have not that difference of capacity and of disposition, which we see in animals whose endowments are higher. 97. In the highest division of the Articulated series, we easily recognise, as forms quite distinct from each other, the Insects, the Spiders, the Crustaceous animals (crabs, lobsters, &c.), and the Centipedes. The class of Insects is distinguished, for the most part, by the presence of wings ; but to this there are exceptions. It includes those of the higher Articulata, which breathe air by means of air- tubes distributed through the body (§ 320), which have no more than six legs, and whose body, in its perfect form at least, manifests a division into tliree distinct parts — the head, thorax, and abdomen (fig. 45). To the thorax alone are attached the six legs, as well as the wings ; and its cavity is principally occupied by the muscles that move them : the abdomen contains the organs of digestion and reproduction, as in vertebrated animals. In the greater part of this class, the young animal comes forth from the egg in a condition very different from that which it is ultimately to possess ; and it undergoes a complete meta- morphosis, the larva wliich the egg produces bearing a close resemblance in form to the lower Articulata, and only attain- ing the condition of the imago or perfect insect by passing again into a state of inactivity, during which the store of nutriment which it has acquu-ed is appUed to the development of new organs. This p)upa or chrysaUs condition may be considered as a sort of postponed completion of the embryonic STEUCTUKE OF INSECTS AND ARACHNIDxV. 10^ life, which was interrupted at a very early period. In some tribes, however, the genenil form is the same from the first, and the wings are the only parts deficient ; these gradually Antennae Eyes 1st pair of Legs - 1st pair of Wings 2nd pair of Legs 2nd pair of Wings 3rd pair of Legs Tibia Tarsus ^-^^^S^. Head Abdomen Fig. 45.— Skeleton of an Insect. make their ajDpearance, and the insect is then complete. Such is the case with the Grasshopper and Cricket; and a change of this kind is termed an incomjylete metamorphosis. 98. The animals of the class Arachnid a, which includes the spiders, scorpions, and mites, are, hke Insects, articulated, breathing aii', and possessing legs, but the number of these legs is never less than eight; there is an entire absence of wings, and the head is united with the thorax, so that the body seems to bo formed of two principal divisions, — the cephalo-thovax (as it is termed), and the ahdomcn. In fig. 46 we have a representation of the arrangement of the parts con- 106 STRUCTURE OF ARACHNIDA AND CRUSTACEA. tained in these cavities. At c ^ is seen the cephalo-thorax opened from below, and giving attachment to the legs ; at m is shown the place of the mandibles or jaws ; at ^ is seen one ct pa af> pa * Fig. 4G. — Anatomy of of the palpi, which arc appendages to the mouth ; pa is the foremost leg ; t, the large nervous mass, from which the legs arc supplied; a, the collection of ganglia supplying the abdomen ; a b, the abdomen ; p a, the respii-atory chambers ; s s the stigmata or openings into these ; I, the leaf-like folds within them (§ 323); w a, the muscles of the abdomen ; a n, the termination of the intestine ; /, the spinnerets ; o, the ovaries ; and o r, the opening of the oviduct. 99. The class of Crustacea, of which the Crab, Lobster, and Craij-Jish are the best-known forms, differs from both the preceding, in being adapted to breathe by means of gills, and thus to reside in or near water, instead of inhabiting the air. Moreover, the body is inclosed in a hard covering, which generally contains a good deal of carbonate of lime, and which is thrown off at regular intervals. This covering also incloses the members, wliich are never less than ten in number, and are frequently more numerous. There is great variety of form among the animals of this group, which is altogether one of great interest. — In the Crab tribe, the head, thorax, and abdomen are all drawn together, as it were, into one mass ; and the general arrangement of the organs it con- tains is exhibited in the succeeding figure, which shows them nearly as they are found to lie, when the upper part of the shell, or carapace, is removed. At t there is left a portion of STRUCTURE OF CRUSTACEA. 107 the membrane whicli lines the carapace and covers in the ■vascera. On the central line, at c, is seen the heart, which in the Crustacea is large and powerful in its action ; from it there passes forwards the artery a o, which supplies the eyes Fig. 47. — Anatomy of a Cviab. and the front of the body ; whilst the artery a a passes to the lower and hinder parts ; at b are seen the gills of the left side in their natural position ; whilst at b' are seen those of the right side, turned back to show their under-surface, and to disclose the lower portion of the shell, fl. At e is seen the stomach, situated close behind the mouth ; and at ttl are pointed out its powerful muscles, by the action of which the food is ground down. The bulky ovary is seen on either side of the stomach ; and the space between this and the edge of the shell is occupied by the very large liver, / o. 100. In most of the Crustacea, however, the body is more prolonged. In some, as the Lobster, there is an indication of a division of the body into three parts, representing the head, thorax, and abdomen of insects ; whilst in others, as the Sand- 108 STRUCTURE AND METAMORPHOSIS OF CRUSTACEA. Iiopper, the rings or segments are almost as similar to each other as they are in the centipede tribe. There is no class in which we find the same parts exhibiting so great a variety of forms, and rendered subservient to so many uses. _ Thus in the Crab and Lobster the members of the first pair are not used for walking, but form the claws or arms by which the food is seized ; in the Cray-fish, these members may be used either as legs or claws; whilst in the Sand-hopper, they closely resemble the other legs. And the jaws of the higher Crustacea, of which there are several pairs, are really meta- morphosed legs ; as may be seen by comparing them with the corresponding appendages of the Limulus or king-crab, the first joints of which act as jaws, whilst the remaining portions of these members serve either as legs for locomotion, or as claws for prehension. 101. Most of the Crustacea, like insects, come forth fi'om the egg in a state very different from their adult form ; and afterwards undergo a series of changes, which are in some instances so remarkable as to approach the complete metamorphosis of insects, and which end in the production of the complete form. An early form of the common crab, at a time when it is of the minute size indicated on the scroll, is shown in fig. 48. The immature Crustacea of different tribes bear much more resemblance to each other, than do the forms into which they are ulti- mately to be developed ; and the dit- ferences they afterwards present are chiefly due to a variety in the amount of growth which the different parts undergo. 102. It is one of the most remarkable results of modern zoological research, that in immediate connexion with the class of Crustacea, if not as actual members of it, we have to place a group of animals which were for some time asso- ciated with the Mollusca; their bodies being inclosed .in shells, which do not fit closely around them, nor give more than a general protection to theii' members. This group is Fig. 48. — ZOEA, OR LARVAL FORM OP THE Crab. STEUCTUEE OF CIEEHIPEDA. 109 the Barnacle tribe, forming the class Cieehipeda, or tendril- footed animals. They agree with the lower Mollusca, in being fixed to one spot diu-ing all but the earhest period of theii' lives ; the shell being sometimes attached by a long membranous or leathery tube, as that of the Barnacle (fig. 49) j and sometimes being itself fixed on the surface of Fig. 49.— Shell of THE Barnacle. Fig. 50. -Body of THE Barnacle. a rock, or on another shell, as is that of the Balanus or acorn-shell. In both cases, the form and structure of the animal are essentially the same. When taken from the shell (in which it lies doubled up, as it were) and spread out, its articulated nature is evidenced by its division into segments, and by the regularity of the arrangement of their tendril-like appendages. These are not formed like legs, since they could be made no use of, the animal being incapable of moving from place to place ; but they serve to produce currents in the surrounding water, by which food is brought to the mouth, and the blood is submitted to the influence of a fresh supply of air. The nervous system of this group is formed precisely upon the plan of that of the Articulata generally (§ 94) : and if any doubt could have remained as to its true place in the series, it is removed by the knowledge of the fact, that the animals composing it bear a strong resemblance in their early condition to some of the lower Crustacea, possessing eyes and legs, and swimming freely about; and that they attain their adult form by passing 110 STRUCTURE OF MYRIAPODA AND ANNELIDA. through a series of metamorphoses, in which they lose their eyes and legs, and become fixed for the remainder of their lives. 103. 'Wg now pass back to another class of the higher group of Articulata, adapted to breathe air and to inhabit the land, — the Myriapoda or Centipede tribe (fig. 42). Both these names are derived from the great number of legs possessed by these animals, which often amount to 60 pairs or even more. In this class we see a more perfect equahty of the segments or divisions of the body than in any others among the higher Articulata; and the similarity is scarcely less complete in the internal arrangement, than it is in the external form. In its lower tribes (fig. 51), the legs are so Fig. 51. — luLu weak as scarcely to be able to sustain the body, which moves, therefore, partly in the manner of that of a worm. The animals of this class undergo no proper metamorphosis ; but there is a considerable adcUtion to the number of their seg- ments and legs after they have come forth from the egg. 104. AVe now pass to the lower division of Articulata, in which the body possesses no jointed members ; and the animals belonging to this group are for the most part included in the class of Annelida, the Leech and Worm tribe. We here find the body enveloped, — not in a hard casing, formed of distinct pieces united by a flexible membrane, — but in a skin which is altogether flexible, and which gives little indication of a division into segments. This class includes several distinct tribes, which aU agree, however, in the long worm-like form of the body, and in the similarity of the different ganglia oi their nervous system. The Earth-worm and its allies are adapted to live on land and to breathe air ; but the greater number of Annelids arc purely . aquatic ; and these breathe by gills, which form tufts that are disposed on various parts of the body. In the Nereis^ or Sea-centipede (fig. 52), these STRUCTURE OF ANNELIDA AND ENTOZOA. Ill tufts are arranged regularly on tlie several segments, and the animal can swim by the motion that it gives them ; besides these, it has a kind of bristle-shaped aj^pendage, that seems ^^^^Ws^^^ Fig. 52. — Nereis. like a rudimentary leg, which assists it in crawhng. But there are others of these marine-worms, that form a tubular shell, in which they reside during the greatest part of their Lives ; and in these the gills, if disposed along the body, would have been removed from the access of water ; they are therefore arranged round the head, often forming (as in the Serpuloe, fig. 145) tufts of great brilhancy and elegance. lOo. Below the Annelida are other worm-like tribes of yet greater simplicity of conformation, but still presenting the same general plan of structure. Of one of these the common Leech may be taken as an example ; of another, the Tape- Fig. 53.— Tape-worm. worm (fig. 53). This last belongs to a group termed Entozoa, from the circumstance that they inhabit the bodies of other animals. They are remarkable for the very low development of their digestive apparatus, — their nourishment being appa- 112 GENERAL STRUCTURE OF MOLLUSCA. rently imbibed through, the whole surface of their bodies from the juices in the midst of which they live ; whilst, on the other hand, their reproductive apparatus is enormously developed, the multii^Hed segments of the Tape-worm (for example) containing this alone, and the head (as it is com- monly termed, though really the body) being able to repro- duce these to an indefinite extent after they have been throA\Ti off. The group of Eotifera, or Wheel- Animalcules, which is one of great interest to the Microscopist, also belongs to this lower section of the Articulated sub-kingdom. lOG. The general character of the animals composing the group or division Mollusc a, is, in many respects, the very opposite of that which prevails in the Articulated animals. The body is soft (whence the name of the group is derived), neither possessing an internal skeleton, nor any proper ex- ternal skeleton. In some of the most characteristic specimens of the group, such as the Slug, there is no hard frame- work or skeleton whatever, the body being alike destitute of support and protection. In most MoUusks, however, the body has the power of forming a shelly covering, which serves for its protection ; but this does not give any assistance in its movements by affording fixed points for the attachment of the muscles ; in fact, when the animal puts itself in motion, it is obhged to make its locomotive organs project beyond the shell. AVe must not regard the shell as an essential part of the Molluscous animal ; because there are many tribes entirely destitute of it ; and also because some of the Articulata have the power of forming a shell (§ 102), which bears a close resem- blance to that produced by the animals of this group. N"ot un- Fig. 54.— Testacei.la. frequently we see that, of two animals whose general structure ifl almost exactly the same,— as that of the Simil and Slug,— STRUCTURE OF MOLLUSCA. 113 one possesses a shell into whicli it can withdraw its whole body for the sake of protection, whilst the other has none; and several intermediate forms exist, in which the shell bears a larger or smaller proportion to the body, sometimes being able to contain nearly the whole of it, and sometimes being a mere rudiment, as in the Testacella (hg. 54). 107. The external form of the body of Mollusks is subject to great variation ; and generally has a good deal to do with the degree in wliich the organs of sense and the instruments of motion are developed in the particular animal. For these are almost always symmetrical, being arranged ^vith equality on the two sides of a middle line ; whilst the rest of the body, containing the organs of nutrition, is often unequal on the two sides. But in the lower Mollusca, which have little or no power of moving from place to place, even this degree of symmetry is altogether lost. Few of the Mollusca have any powers of active movement ; in fact, the term sluggish- ness, derived from a characteristic member of the group, very well expresses their general habit. The Gasteropods, which may be regarded as the types of the whole series, crawl upon a flesh}^ disk, by the successive contractions and relaxations of which they advance slowly along the surface over which they move ; this kind of action is easily studied, by causing a Snail or Slug to crawl- upon a piece of glass, and by looking through this at the under side of its foot. Hence, there is a great contrast between the inertness of the Mollusca, and the liigh activity of the Articulata, This contrast shows itself in the structure of their bodies ; for whilst the chief part of the interior of an Insect is made up of the muscles which move its legs and wings, — the apparatus of nutrition being small, — the chief part of the hulk of a Slug or Snail is given by its very complex apparatus for nutrition — there being no other muscles (except some small ones connected with the mouth and head) than the fleshy disk already mentioned. The blood of the Mollusca is nearly colourless, as it is in the Articulata ; but the organ by which it is circulated through the body is much more powerful and complete, bearing more resemblance to the heart of Vertebrated animals. The skin is usually thick and spongy in its texture ; having muscular fibres inter- woven in its substance, so that it can contract or extend itself in any part ; and having the power of exuding shelly matter I lU STRUCTURE OF MOLLUSCA. from its surface, in those si3ecies wliicli form such, a pro- tection. This envelope, which is called the mantle^ is very loosely applied round the parts which it contains; and it frequently extends itself into folds or duplicatures, which wrap round the gills, and sometimes meet and adhere so as to inclose them within a cavity of their own. In the Cuttle- fish, the water within this cavity is renewed from time to time by the muscular movements of its walls ; but usually a current of fluid is kept up over the surface of the gills, by the action of the c'dla (§ 45) with which they are covered. m vb ah b ov Fig. 55.— Anatomy of Turbo Pica. 108. Tlie accompanying figure of the interior of a Turho \s\S\ show the very large size of the digestive apparatus, and of the other organs of nutrition. The muscular disk or foot is seen at/); and this carries the operculum o, which serves to close the mouth of the shell when the body of the animal is dra^vn within it. At t is sho^vn the proboscis, on either side of which are the tentacula or feelers, ta, bearing the eyes at y. Just behind the tentacula is seen the large cephalic ganglion, sending nerves to the eyes ; and behind this again are the saUvary glands. The mantle, m, is opened and folded hack to show the respiratory cavity, in which lie the gills STRUCTURE OF MOLLUSCA. 115 hh; to this cavity, water has access by means of a wide slit, of which the edge, t\ of the mantle forms one part of the border, whilst at d is seen a fringed membrane that forms another part. At c is seen the heart, which receives the l)lood from the gills by v b, the branchial vein, and then transmits it to the body generally ; at e, far np in the spire, are the stomach and liver ; at a, the anal orifice of the intes- tine within the branchial cavity, and at ov the oviduct, which opens in the same situation. 109. Thus it is seen that, — whilst the body of an Articu- lated animal may be compared to that of a man in whom the apparatus of nutrition (contained in the chest and abdomen) is of the smallest possible size, but whose limbs are strong, and his movements agile, — the body of a Mollusk resembles that of a man " whose god is his belly," his digestive appa- ratus becoming enormously developed, whilst his limbs ar© feeble, and his movements heavy. Such varieties, in a greater or less degree, are continually presenting themselves to our notice. 110. The nervous system of the MoUusca generally consists of a single ganglion or pair of ganglia, which are placed in the head, or (when that is deficient) in the neighbourhood of the mouth ; and of two or more separate ganglia, which are found in different parts of the body, and are connected with the preceding by nervous cords. The former correspond to those con- tained in the head of Insects ; but of the latter, one only is connected with the foot or organ of motion, the remainder having for their function to regulate the action of the gills, and to perform other movements connected with the operations of nutrition. In fig. 56 is represented one of the simpler forms of this nervous system, — that of the Pecten or Scallop-shell ; A A are the ganglia near the mouth, Fig- sc- from which the organs of sense are supplied ; b is the ganglion connected with the gills ; and c is that from wliich power is given to the foot. The " I 2 -Nervous System of Pecten. 116 • GENERAL STRUCTURE OF MOLLUSCA. — CEPHALOPODS. two first lie wide apart, but are connected by an arched band that passes over the gullet, e. The organs of sense in the higher forms of Mollusca are more developed than those of motion. They serve to direct the animal to its food, and to warn it of danger ; but there seems an absence, in all save the highest species, of that ready and acute sensibility which is so remarkable in the preceding groups ; and the variety of impressions which they can receive appears to be but small. In no instance has a special organ of smell been certainly dis- covered ; the organ of hearing is always imperfect, and fre- quently absent altogether ; and the eyes are very often wanting. In the lower Mollusca there are no certain indications of the existence of any organs of special sense ; and there is probably but a limited amount of general sensibility. 111. As the Articulata are divided into two subordinate groups, according to the presence or absence of articulated limbs or meml^ers, so may we arrange the Mollusca in two subdivisions, according to the presence or absence of a dis- tinct head^ that is, a projecting part of the body, containing the mouth or entrance to the digestive cavity, and also bearing the organs of sense which guide the animal in the discovery and selection of its food. In the higher Mollusca, there is a distinct head, furnished with eyes, and sometimes with im- perfect ears ; but in the lower, the entrance to the digestive cavity or stomach is buried deep among other parts, and is guarded ])y no other organs of sense than the tentacula or sensitive lips. These are termed acephalous, or headless Mollusca: and among the lowest of them (§ 114), we meet with composite fabrics, formed by the process of multiplica- tion by budding, which was formerly regarded as peculiar to Zoophytes. — The highest group of Mollusca, in regard to the approach of several parts of its structure to that of Yerte- brated animals, is the class of Cephalopoda, or Cuttle-fish Ifibe : which receives its name from the peculiar arrangement of the arms or feet around the mouth, which is the cha- racteristic of its members (fig. 57). The common Cuttle-fish and its allies are destitute of any external protection ; but they usually have a flat sheh, commonly known as the cuttle- fish bono, inclosed in a fold of the mantle, and lying along the back. In the Calamary, this is horny in its texture, and is sufficiently flexible to ofler no resistance to the action of STRUCTURE OF CEPHALOrODS A^'D I'TEROPODS. 117 the fin-like tail, By which the animal is propelled through the water very much in the manner of a fish. The Pearly Nautilus is the only type now existing of an inferior order of Fig. 58. — Ammonite. Fig. 57. — Calamary. Cephalopods, which approaches the Gasteropoda in many parts of its organization. The body is inclosed in the last chamber of a shell (usually spiral in form), the cavity of which is divided by numerous transverse partitions ; and such shells, the fossihzed remains of very numerous forms of this group that existed in the ancient seas, con- stitute the nautilites, ammonites, helemnites, Szo,., wliicli abound in many rocks (fig. 58). Tlie Cuttle- fish are animals of considerable activity; their mouth is furnished with a horny beak, strongly reseni- bling that of the parrot ; and their arms are provided with a series of very curiously constructed suckers, by the action of which they can take a very firm hold of anything which they desire to grasp. 112. The class of Pteropoda, or wing-footed MoUusks, consists of but few species, and the animals which it contains arc all of them of small size ; but the individuals are often very numerous, whole fleets of them being sometimes seen covering the ocean, especially in the Arctic and Antarctic regions, where they constitute one of the ^'^- 59--Htal;ea. principal articles of food to tlie Whale. The general form of the body usually difi"ers but little from that represented in fig. 59. 118 STRUCTURE OF GASTEROPODS AND BIVALVES. On cither side, a little behind the head, the mantle is extended into a fin-like expansion, by the aid of which the animal can swim through the water. The hinder part of the body is usually inclosed, more or less completely, in a shell, which is conmionly of extreme thinness and delicacy. The head is not furnished with long arms, to grasp the food ; but it has a number of minute sucking disks, by which it can lay firm hold of whatever it attacks : whilst its powerful rasp-like tongue is set to work upon it. — The class Gasteropoda con- tains those animals which, hke the Snail and Slug, crawl upon a fleshy disk on the under side of their bodies ; and the number of distmct forms which it includes is very large. The greater part of them are inhabitants of the sea-shore^ rivers, lakes, &c. ; some have the power of swimming freely through the open sea; and the proportion of those that breathe air and live on land, is comparatively small. The general structure of the animals of this group has been already described (§ 108). Some of them form shells, wMst others are destitute of them. The shells are composed of a smglo piece, or are univalve, except in one tribe ; and they have usually more or less of a spiral formation (fig. GO). The animals of this class all ])ossess a distinct head ; and this is generally I'urnished with eyes, as well as with tentacula. They have often a powerful masticating ap- paratus, and are voracious in their habits ; Fir. oo.-siiell some of them feed upon vegetable matter, others OF PALumsA. ^^^^^^^ animals. 1 1 .'5. The Acephalous MoUusca are divided into two groups, — those which form shells, and those which do not. The former are termed Conchifera, or shell-hearing animals ; and tliis class includes all the MoUusca that form a shell composed of two parts or valves fitted together (which shell is termed hivalve), as well as some others whose general structure is the same, but whose shell is formed in several pieces, or multivalve. The two valves of a bivalve shell (fig. 61) are connected by a hinge, where they arc united by a "ligament, which, by its elasticity, keeps them apart wliile it holds them together. Tliis is their usual condition when the annual is alive ; and in this manner the water which is required for their respiration, and also to convey their supply of food, has free access to the internal STRUCTURE OF CONCHIFERA, OR BIVALVES. 119 parts. But when any alarm or Kritation causes tlie animal to so by means of a muscle (sometimes close its sliell, it does Fig. 61.— Shell of Tridacne. single, sometimes double), wliicli stretches across from one valve to the other, and which, by contracting, draws them together. Each valve is hned by an extended fold or lobe of the mantle. In the higher tribes of the class, these lobes are united along their edges, leaving apertures for the ingress and egress of water (which are sometimes prolonged into tubes, fig. 150), and another for the foot. But in the Oy&ter and its allies, which have no foot, or a very small one, the mantle-lobes are quite disunited. The accompanying diagram (fig. 62) gives a general idea of the arrangement of the organs in one of the higher acephalous Mollusca, the Mactra, which is among those having two muscles for the drawing together of the valves. The upper end, as represented in this figure, is that wliich is considered as the anterior end or front of the animal, being that nearest which the mouth lies ; and the posterior extremity (the loAvest in the figure) is that at which the intestinal canal terminates, and at which the respiratory tubes are formed. Kear the anterior muscle, we find the mouth, or entrance to the stomach ; it is furnished with four riband-shaped tentacula, of which one is seen in the figure ; and these seem to possess peculiar sensitiveness. Near the mouth lie the anterior ganglia of the nervous system, which represent the brain of higher animals ; and these are connected by long cords with the posterior ganglion, which Hes near the posterior muscle. The stomach, intes- tines, and liver occupy the central portion of the cavity of the shell ; and the intestinal tube is seen to pass backwards, 120 STRUCTURE OF COXCHIFERA OR BIVALVEri. terminating near one of the canals or siphons, wliich also carries out the water that has been taken in through the other for the purposes of respiration. The figure also shows the large fleshy foot, by which this animal can move itself along Intestine Mantle (Anterior \Ganglia Respiratory Tubes, Fig. 62.— Anatomy of Mactra. the ground, or bore into sand or mud. The heart and circu- iutrng system are less complete than in the Gasteropoda ; but arc far higher in character (as arc most of the other parts STRUCTURE OP TUNICATA. 121 of tlie nutritive apparatus) than the corresponding parts in Articulated animals, in which the apparatus for locomotion so much predominates. 114. The group"'of Acephalous Mollusks wliicli are desti- tute of the powerJof forming a shell, includes two classes, of Avhich one does not depart widely from the general Molluscan iype, whilst the other presents 'so strong a general resem- blance to Zoophytes, that until recently it has been universally ranked with it. The first of these classes receives its name TuNiCATA from the circumstance that the mantle, instead of secreting a shell, is very commonly condensed into a tough leathery or cartilaginous tunic. Many of these animals live separately, and have the power of freely moving through the water. Others are associated in comj^ound masses, of which, however, the individuals are not connected by any internal union. But others form really composite structures, like those of Zoophytes (§ 124); each individual being able to live by itself alone, but being connected by a stem and vessels with the rest. The general structure of the individuals is the same, however, in the single and in the composite animals of this class, and may be understood from the accom- b a Fig. 63. — Social Ascidiax panying lig-ure (fig. 63). The cavity of the mantle possesses, as in the former instance, two orifices ; by one of which, 6, a current of water is contmually entering, whilst by the other, a, it is as continually flowing out. These orifices lead into a large chamber, the lining of which, folded in various ways, constitutes the gills ; and at the bottom of this chamber lie the stomach, e, and the intestinal canal, i, which terminates near the aperture for the exit of the Avater. All these parts 122 STRUCTURE OF TUNICATA AND POLYZOA. are covered with cilia, by the action of wliicli a continual stream is made to flow over the gills and to enter the stomach ; and the minute particles which the water brings with it, and which are adapted to serve as food, are retained and digested in the stomach. Even these animals, fixed to one sjDot during all but the early part of their hves, and presenting but very slight indications of sensibility, possess a regular heart and system of vessels ; and these vessels form part of the stem, t, by which the compound species are connected. A single nervous ganglion is found between the two orifices ; this seems to receive sensory fibres from tentacula situated around the oral orifice, and to transmit motor filaments to the mus- cular coat wliicli underhes the outer tunic, so that any irrita- tion applied to the former occasions a contraction of the latter, which tends to expel the ofiending particle. — This class is one of particular interest to the naturahst, since we see in it the tendency to the formation of compound struc- tures, by a process resembling that of the hudding of plants, which is essentially characteristic of Zoophytes; this ten- dency, however, is more fully manifested in the succeeding class. 115. The animals forming the class Polyzoa (more com- monly known as Bryozoa) are seldom or never found solitary ; since, in consequence of their universal tendency to multiply by gemmation, they form clusters or colonies of various kinds. The body of each individual is inclosed in a sheath or " cell," which is somcthncs horny, sometimes calcareous ; and the composite skeleton formed by the aggregation of these, which has sometimes a branching or leaf-like form, but sometimes possess the compactness of a stony coral, is known as the " polyzoary." In their general structure the animals of this class possess considerable analogy to the Tunicata ; but the Molluscan type presents itself under a more degraded aspect, no vestige of a heart or of blood-vessels being here dis- cernible, and the general structure being so simplified as to manifest no great degree of elevation above that of Polypes. The typical structure of these animals may be understood from that of the Bowerhankia (fig. 64), which is one of those whose cells are not in contact with each other, but grow forth at intervals from a creeping stem. The mouth, a, is situated in tlio midst of a circle of arms fringed with cilia : these STRUCTURE OF POLYZOA AND RADIATA. 123 arms do the food not serve, however, like those of polypes, to grasp but the vibration of their cilia produces a powerful current which brings both food and oxygen. The mouth leads by a large funnel-shaped oesophagus or gullet, to a gizzard, h ; in which the particles of food that enter it are ground down, by the action of its muscular walls and of the tooth-like processes that line it. Below this gizzard is the true digestive stomach, c, around which the rudiment of a liver may be traced ; and from tliis stomach there passes upwards an intestinal tube, which terminates by a distinct orifice at d, on the outside of the circle of arms. The digestive apparatus is evi- dently formed, therefore, upon a much higher plan in these animals than it is in the true polypes, which have no true anal orifice. The Molluscan character of these animals is further shown by the presence of a single nervous ganglion, situated between the two orifices, as in the Tunicata ; this acts uj^on a complex apparatus of muscles, by which the animal ?'^orifice^of"Sti- can be either drawn into its cell or projected tine. forth from it, with great rapidity. 116. The fourth subdivision, that of Eadiata, includes those animals which have the parts of the body arranged in a circular manner around a common centre, so as to present a radiated or rayed asj)ect. This arrangement is well seen in the common Star-fisic (fig. Q5), which has five such rays, all having a precisely similar structure, and thus repeating each other in every respect. The mouth of this animal is in the centre ; and it opens into a stomach, which occupies the cen- tral disk, and sends prolongations into the rays. The nervous system is, in like manner, composed of a repetition of similar parts. A plan of it is seen in fig. 6Q ; where a shows the position of the mouth, which is surrounded by a ring or nervous cord, having five ganglia, corresponding to the five arms. From each of these ganglia proceeds a branch along its arm, terminating in a little organ at its extremity, which is believed to be an imperfectly-developed eye. 'No other organs of special sense can be cletected in any of these ani- Fig. 64. BOWERBANKIA a, cesop]iag;us ; b, s 12-i STRUCTURE OP RADIATA. Dials ; and it is only in a few that even these imperfect can ho discovered. ~ ^ ^^^^ -~^ ^ In the inferior Eadiata, not the sUghtest Fig. 65.— SuKLL OF Star Fish. traces of a nervous system liave yet been discovered; and it is very doubtful whether any such structure exists in them. It is only among the higher liadiata that any locomotive power exists ; and this is usually so feeble that the animals remain in the same locality during the greater portion of their lives. Generally speaking, there is a period in the history of each species, in which there is a more active movement, that serves to prevent the accumulation of indi- viduals in one spot ; but this move- ment is of a purely automatic ••haracter, rather resembling that of the " zoospores " of plants, tlian the intentional change of place of the higher animals. Fig. 6G. — Nervoxts System OF Star Fish. STRUCTURE OF RADIATA. ECHINODERMATA. 125 117. The circular arrangement of the organs of Eadiatcd animals is a striking point of resemblance to the Vegetable kingdom ; and it has frequently caused mistakes to be made in regard to the Sea- Anemones and other large polypes, which, when their mouths are open and their arms spread out, look so much hkc the blossoms of some of the Com- posite tribe of plants, as to have received the name of animal fiowers. But there is yet a stronger analogy between the lower members of the Eadiated group and the Vegetable kingdom ; for among the former, as in the latter, we find a union of many individuals, which are capable of existing separately, into one compound structure, having a plant-like form. This is the nature of the stem of Coral (fig. 76); which is, in fact, the skeleton of one of these compound systems, consisting of a number of polypes united by a jelly- like flesh ; just as the woody stem of a tree is the skeleton that supports a vast number of buds, each of which is capable of living by itself. This aggi-egation results from the in- definite multiplication of parts by the process of gemmation or budding, and from the persistence of the connexion between these parts, notwithstanding that, if separated, they can maintain an independent existence. To the tree- like fabrics thus produced, the name Zoophytes (animal plants) is commonly given; and ordinary observers often find it difficult to get rid of the idea of their vegetable origin. The animals that formed them are, of course, fixed to one spot during all but the earliest periods of life ; and the amount of movement which they perform, for the pur- pose of obtaining and secui'ing their food, is very little greater than that which is witnessed in the Sensitive plant and Venus's fly-trap. 118. The class of Echinodermata receives its name from the prickly character of its covering, which is evident enough in the Echinus or Sea- Urchin, and in the Star-fish; but there are other animals, sufficiently resembling these in general structure to be united in the same class, which have a body entirely soft, — namely, the Holothurioe (fig. 67), commonly termed Sea-Cucumbers. This class ranlis as the highest among the Eadiata, in regard to general complexity of struc- ture. The skeleton of the Sea-lJrchin, Star-fish, and other animals resembling them, is a box-Uke shell or " test," formed 126 STEUCTURE OF ECHINODERMATA. of a great nimiber of pieces, very regularly arranged and united together (fig. 69, e); but these pieces are for the most Fig. 67. — HoLOTiiURiA. part only repetitions of one another ; and the different portions have not that variety of uses which we see in higher animals. With the exception of the tribe of Encrinites or lily-like animals (fig. 68), of which there are very few now existing, but which were very abundant in former ages, all the animals belonging to this class are unattached, and are capa- ble of moving freely from place to place. Their motions are very sluggish, however, and are princi- pally effected by means of an im- mense number of minute tubular feet (fig. 6S, c), furnished with suckers at their extremities, which can be projected from ahnost any part of the body. These are seen in rows on the under side of each arm of the Star-fish ; they are put forth through rows of very minute aper- tures in the shell of the Sea- TJrohin (commonly termed the Sea- STRUCTURE OF ECHINODERMATA. 127 Egg) ; and they are also arranged in rows on the surface of the body of the Holothuria, as seen in fig. 67. 119. The radiated arrangement is very evident in the whole bodies of the Star-Fish (fig. 65), and Echinus or Sea- Urchin (fig. 69); but in the Holothuria (fig. 67) it is nearly confined to the parts about the mouth; which, however, exhibit it so completely, that such an animal cannot be mis- taken for one of the Articulated series, even though, as some- times happens, the body is prolonged into a worm-like form. The digestive apparatus in this class has usually a high degree of complexity, as will be seen by the accompany- Fig. 69.— Interior op Echinus ing figure (fig. 69), which shows the interior of an Echinus^ whose globular shell has been sawn across its equator, so as 128 STRUCTURE OF ECHIXODERMATA AND ACALEPH^. to alloAV of the separation of its two halves. The mouth, A-, situated at one of the poles of the shell, is surrounded by a very curious apparatus of jaivs and teeth (fig. 69), which forms what is termed the "lantern;" from the mouth com- mences the long narrow oesophagus, m, that leads to the stomach, n, which is merely a dilated portion of the alimen- tary tube ; the continuation of this, o, q, r, forms the intestinal canal, which winds once round the shell, and then doubles back and winds in the opposite direction, terminating at the anal orifice, s, which is situated at the opposite pole. The intestine is held in its place by a double fold of" the mem- brane lining the shell, resembling the mesentery of higher animals ; the blood is distributed over this membrane, to be exposed to the aerating influence of the water admitted into the canity of the shell ; and the water is kept in movement by the cilia with which the membrane is clothed. Eound the anus, s, are seen the five branching ovaries, each of wliich discharges its contents by a distinct orifice. The circulating apparatus is imperfect, the blood not being impelled by a distinct heart ; still, however, it moves in great part of its course through proper vessels, and not through mere chan- nels in the tissue. — In the Star-fish, however, the body is very much flattened ; and the stomach, instead of having a separate intestinal tube with a distinct orifice, is a mere bag with a single aperture, which serves both to take in food and to cast forth the indigestible remains. This character will be Ibund to prevail among all the inferior Eadiata. 120. The radiated structure is also w^ell seen in the greater number of animals forming the group of Acaleph^, or Sea^ Nettles. Their bodies are entirely soft and jelly-hke ; contain- ing so small a quantity of solid matter, that, when upon being taken out of the water their fluid drains away, there is scarcely anything left; hence they are commonly termed Jelhj-Fish. They derive their other name of Sea-Nettles from the stinging power wliich most of them possess. They are formed to float freely in the water; but they do not in general possess any means of actively propelling themselves through it. The radiated arrangement is very regularly pre- served in some of this group, whilst it is less evident in others. The accompanying figure (fig. 70) represents one of the Medusa tribe, as seen floating in water. The umbrella- STRUCTURE OF ACALEPHiE. 129 shaped disc above contains the stomacli, which is placed in the centre, and which opens by a single oritice or mouth, directed downwards. Around the stomach are four chambers, in wliich the eggs are prepared. The mouth is surrounded by four large tentacula, which bring to it the necessary supply of 70. — Pelagia. food; and other tentacula are seen, in this species, to be hanging from the edge of the disc. In the edge of this disc, the nutritious fluid, which flows in channels prolonged from the stomach and excavated out of the soft tissues, seems to be exposed to the influence of the surrounding water ; but nothing like a heart or a regular circulation exists. — Eecent discoveries in regard to the developmental history of the Medusce and their allies, have rendered it very doubtful whether the Acalephce should continue to take rank as a dis- tinct class ; since many of them constitute only a jiarticular phase in the life of the Hydroid Zoophytes (§ 124). 121. The class of Polypifera, or coral-forming animals, commonly known as Zoophytes, includes two principal tribes, which differ from one another in structure to such a degree as to 130 STRUCTUKE OF HYDRA. Fig. 71. Hydra, or Fresh-water Polype. require separate notice. The group of Hydrozoa, or Hydroid Zoopliytes, so named from the little Hydra, or fresh- water polype, which may be regarded as its type, will be first described on account of its near con- nexion with the preceding. The Hydra (fig. 71) is a solitary polype, not at all uncommon in ponds or other collections of fresh water, where it is found attached to aquatic plants, or to floating sticks, straws, &c., by means of a kind of sucker at its lower extremity, stretching out its tentacles in search of its food, which consists of minute aquatic worms and insects. These are securely laid hold of by one or more of the tentacles, and are drawn into the mouth, a, which leads to the stomach or general cavity of the body, in which they are digested, and from the walls of which the nutritious portions are absorbed, the portions of the food which are not capable of being digested being cast out through the mouth. 122. The Hydra multiplies in two ways ; namely, by gem- mation or budding, and by a proper generative process. Little bud -like processes are developed from various parts of the walls of the stomach, which gradually assume the form of the parent, possessing a mouth surrounded by tentacles, and a digestive cavity which is at first in connexion with that of tlie parent ; the communication is gradually cut off, how- ever, by the closure of the canal of the footstalk of the young polype ; and ere long the footstalk itself separates, and the young polype henceforth leads an entirely independent Hfe. Not uniVequently, however, the young polype itself puts forth buds before its separation ; and as many as nineteen young IIydra>, in difrercnt stages of development, have been seen to be thus connected with one and the same stock. Another very curious endowment of the Hydra depends upon the same facihty of developing the whole structure from any part of it ; HYDRA, AND HYDROID ZOOPHYTES. 131 for into whatever number of parts its body may be cut up, each, under favourable circumstances, can give origin to a new and entire polype, so that tliirty or forty individuals may thus be produced by the division of oue. 123. The proper generative process, here reduced to its utmost simplicity, consists in the development of a germ-cell and of sperm-cells in the substance of the w^all of the stomach, the former being produced near the footstalk, the latter just beneath the arms. The egg which is evolved from the former, being fertilized by the products set free from the latter, gives origin to a young Hydra, which resembles its parent. The two reproductive processes, however, are perlbrmed under very ditferent conditions ; for whilst multiplication Ijy gemmation is favoured by warmth and a copious sup- ply of food, the true gene- rative process seems to be brought about by a lower- ing of the temperature, and to have for its object the perpetuation of the race through the winter, the egg being capable of enduring a degree of cold which would be fatal to the polype itself. 124. The group of Hy- drozoa is for the most part made up of composite fabrics more or less resembling the Gampanularia (fig. 72), which may be likened to a Hydra whose buds do not detach themselves, but re- main in connexion with the stock that produced them ; the whole plant-like struc- Fig. 72.— Camj ture, moreover, being strengthened by the consolidation of its external layer into a horny sheath, which retains its form after the destruction of the soft parts. Thus each comes to consist of a stem and branches, on the sides or ends of which k2 132 REPRODUCTION OF HYDROID ZOOPHYTES. are a number of little cells or bell-shaped chambers, with their mouths upwards, every one of them containing a polype that bears a strong resemblance to the Hydra. Each of these polypes is capable' of living independently of the rest, obtains its nourishment by means of its own arms, and digests it in its own stomach ; but all are connected by a canal that passes along the stem and branches, in which a kind of cii'culation takes place, that strongly reminds us of that of the compound Tunicata (§ 114). This plant-like structure extends itself by budding; new branches are formed from those previously existing ; and these are enlarged at a certain point into cells, in which after a time polypes make their appearance. 125. Besides the cells containing the polypes, however, we find capsules in which are evolved buds of a different nature, that form within themselves the generative products. These buds in some instances assume the form of Medusae, and, becoming detached from the stalk that put them forth, swim about freely, living upon food obtained by themselves, and setting free either sperm-cells or germ-cells, by the concur- rence of whose contents eggs are formed, from which new polype-growths arise. In otlier instances the Medusoid bodies give forth their generative products, without ever leaving the capsules in which they were themselves developed. And in other cases, again, it does not seem that any Medusoid form intervenes at all, the germ-cells and sperm-cells being evolved from the Zoophytic structure itself But since it is also known that even the most characteristic Medusan forms are evolved as buds from a Zoophytic stock (Chap, xv.), and since those comj)osite forms of Acalephse whose structure has until lately been most obscure, turn out to be, as regards their essential characters, Ilydrozoa organized for floating, there seems to be no longer any sufiicient ground for ranking the Acalephse as a separate class. 12G. It is not, however, by animals of this very simple structure, that the massive stony fabrics are built up, which constitute the coral islands of the Pacific Ocean, and of which a large portion of our limestone rocks seems to be composed. These are constructed by animals belonging to the group of Anthozoa, and formed upon the same general plan with the Sea- Anemone,— di plan which is higlier than that of the Hydra, inasmuch as we find the interior uf the body containing other STRUCTUBE OP ANTHOZOA : SEA- ANEMONE. 133 cavities around the stomach, which are destined to pre- pare the generative products. In lig. 73, we have a repre- sentation of the Sea- Anemone, as seen from above ; showing its mouth in the centre, surrounded by its numerous radi- ating tentacula ; these are often brightly coloured, and give to the animal the appearance of a beautiful flower. In fig. 74, a siniihir animal is represented, cut open to sho\v its interior. Fig. 73.— Sea-Anemone, seen from above. Fig. 74. — Section of Sea-Anemone. a, cavity of stomach; b, surrouuding chambers. The mouth is seen to open into a rounded stomach, a, which has no other orifice outwards ; and round this stomach there is a series of radiating membranous partitions, which divide the space intervening between it and the outer covering of the body into numerous chambers, b. Witliin tliese chambers, and attached to their partition- walls, are found the bodies which are commonly designated ovaries, but which contain sperm- ceUs or germ-cells according to the sex. It is doubtful whether these two products are ever formed by the same individual, as they are in the Hydra. The Sea- Anemone does not usually multiply itself by budding, though some species do so ; but large numbers of young are produced from the eggs, which are fertilized and partly developed whilst still within the ovarian chambers, and these make their Avay into the stomach through an aperture at its deepest point, and finally escape by the mouth. 127. The Sea- Anemone itself, like the Hydra, is a solitary animal, capable of shifting its place at will ; and it forms no stony skeleton or support. But there are other animals of the same general structure, which have the power of depositing stony matter in the membrane of their base or foot, and in the membranous partitions between the chambers ; and this stony deposit forms a Coral or Madrepore, such as is shown 134 ANTHOZOA : — STONY CORALS. in the accompanying figure (lig. 75). The particular arrange- ment of the radiating plates of the Madrepore (shown at the top of each stem) is the result of the form of the soft structures by which it was deposited; aiid wherever we see a structure of this kind in coral, whether upon a large or a small scale, we may infer that it was formed by an animal nearly allied in structure to the Sea- Anemone. Of the stone depositing coral- animals, a large number are often associated in a com- pound structure, as in fig. 76 ; this consists of a stony tree-like stem and branches ; but instead of the soft ani- Fig. 75.-CAKYOPHYLLIA. ^^^^ mnUev bciug contained in its interior, as in tlie Hydrozoa, it usually forms a kind of flesh Fipf. 76.— Stem of Cural. tliat clothes the surface, and connects together the diilereut STRUCTURE OF PROTOZOA. 135 polypes ; and new branches, are formed either by the sub- division of the polypes, or by gemmation from the connecting substance. 128. When we pass from Zoophytes to animals of still simpler organization, we lose all trace of definite symmetry, and find ourselves amid forms which cannot be referred to any particular plan of growth. These, moreover, are for the most part distinguished by an extreme simplicity of structure ; no such differentiation of parts exhibiting itself among them, as is shown in the " organs " of even the simplest Zooj^hyte or Worm. Hence they are appropriately designated Protozoa. They may, in fact, be considered as essentially consisting of homogeneous particles of a jelly-like substance, to which the name of Sarcode has been given ; and the chief modification this undergoes, consists in the consolidation of certain parts of it by the deposit of horny, calcareous, or siliceous matter, so as to form a skeleton. This may take place on the outer surface only, so as to form shells very like those of MoUusks in miniature, as we see amoiig For am in if era (fig. 78); or it may occur in the midst of the fleshy substance, so as to form an internal network, such as presents itself in the Sponge. The endowments of the " sarcode " are very extra- ordinary ; and will be best understood by observation of the life-history of one of those simplest Protozoa, in which the whole body consists of but a minute particle of it. Fig. 77.— RiiizopoDA :— A, Amceba ; B, Actiiiopliiys. 129. Such an example is afforded by the Amoiha (fig. 77 a), —a creature frequently to be met with in great abundance in fresh and stagnant waters, vegetable infusions, &c. Its 136 RHizoroDA : — amoeba; actinophrts. organization is so low, that there is not even that distinct differentiation into containing and contained parts which is necessary to constitute a cell (§ 32) ; for although the super- ficial layer of the sarcode possesses more consistence than the interior, it is nevertheless obvious that it has not the tenacity of a membrane, since (as will be presently seen) it does not oppose the passage of solid particles into the interior. How- ever inert this creature may seem when first glanced at, its possession of vital activity is soon made apparent by the movements which it executes and the changes of form it imdergoes ; these being, in fact, parts of one and the same set of actions. For the shapeless mass puts forth one or more finger-like prolongations, which are simply extensions of its gelatinous substance in those particular directions ; and a continuation of the same action, first distending the prolongation, and then, as it were, carrying the whole body into it, causes the entire mass to change its place. After a short time another prolongation is put forth, either in the same or in some different direction ; and the body is again absorbed into it, so as to shift its place still more. It is by means of this movement that the creature obtains its supplies of food ; for when, in the course of its progress, it meets with a particle appropriate for its nutriment, its gelatinous body spreads itself over this, so as to envelope it completely ; and the substance (sometimes animal, sometimes vegetable), thus taken into this extemporized stomach, undergoes a sort oi digestion therein, the nutrient material passing into the sub- stance of the sarcode, and any indigestible portion making its way to the surface, from some part of which it is (as it were) finally squeezed out. 130. Many other forms of this group, which has received the designation of Bhizopoda, ha^-e less power of movmg from place to place, but obtain their food by a modification of the same arrangement : of this we have an example in Actinophrys (lig. 77 b). The body being stationary, i! ; gelatinous substance extends itself into long filaments, ten;.-, I pseudopodia : these ollcn divide themselves again like the roots of a tree (whence the designation of the group), so as to form threads of ex- treme tenuity; and sometimes these threads meet again and coalesce, so as to form a sort of irregular network. When any niinutc animal or vegetable organism happens to come in contact RHIZOrODA : FORAMINIFERA. 137 with one of these thrccads, it is usually held by adhesion to it, and the filament forthwith begins to retract itself; as it shortens, the surrounding filaments also apply themselves to the captive particle, bending their points together, so as gra- dually to inclose it, and themselves retracting until the prey is brought to the surftice of the body ; and the substance of the threads being itself drawn into that of the body, the entrapped particle is embedded along with this, and under- goes digestion in the surrounding sarcode, any indigestible particle being subsequently extruded from the surface of the body, just as in the Amoeba. The reproduction of these creatures, so far as is yet known, is effected by self-division, like that of the Infusoria (§ 135); but there is reason to believe that a "conjugation," or reunion of two individuals, sometimes occurs, and that this is to be looked on as repre- senting the sexual propagation of higher animals. Fig. 78. — FORAMINIFERA. A, Oolina; B, C, Nodosaria; D, rristellaria ; E, Polystomella ; F, Dendritina, G, Globigerina; l\, Textularia; I, Quinqueloculina. 131. This Ehizopod type of animal hfe is manifested in two ga^oups of great interest, which are characterised by the possession of hard shells, formed by the consolidation of the external layer of sarcode. The Foraminifera have calcareous shells, which often bear a strong resemblance to those of Nautili, &c. in miniature (fig. 78), but which really have an entirely different relation to the animals that form them. For whilst the Nautilus occupies only the last or outer chamber of its shell, the chambers previously formed 138 FORAMINIFERA AND TOLYCYSTINA. beiii'-' empty and deserted, each cliainber of tlie B,otalia, or any other i^oraminiferous sliell, is occupied by a segment of sarcode, which is to a great degree independent of the rest, and is only connected with those on either side of it by dehcate threads of the same substance ; and the extension of the shell is due to the formation of an additional segment of sarcode on the outside of the last-formed chamber. Each segment has usually the power of gutting forth its own "pseudopodia" through minute apertures in the shell, and thus of drawing in its own nourishment through these ; but even when (as sometimes happens) these food-collecting threads are put forth from the last chamber alone, the nutri- ment there obtained is transmitted to the segments within by percolation through the substance of the sarcode, and not through any tubular canal. — The accumulation of the shells of Foraminifcra in some parts of the existing sea-bottom is very remarkable ; and similar accumulations in past ages have formed no unimportant part of the crust of the earth — a large part of the Chalk-formation having had its origin in theni, as well as nearly the whole of the Nummulitic limestone by which it was succeeded. 132. But animals whose essen- a b tial structure seems to be nearly the same, may form siliceous in- stead of calcareous shells ; and thus are produced those beautiful organisms, known under the name of Polycystina (fig. 79), which arc occasionally found in the existing seas, but whose re- mains are met with under a far greater variety of forms in certain of the newer marine deposits. Tliere is not in these the same tendency to form composite sti-uctures by the multiplication of segments, as in the Foraminifera ; but the complication of the individual form is often much greater. Yet, however complex the form, the essential composition of these crea- tures seems to retain the same attrilmte of simplicity, which cannot bu conceived capable of further reduction. Fig. Id. — Polycystina. A, Podocyrtis ; B, Rhopalocanium. INFUSORY ANIMALCULES. 139 133. The Animalcules to which the name of Infusoria may be properly restrictcil (the Eotifera, or Wlieel- Animal- cules, § 105, whose organization is much higher, together with many organisms whose true nature is vegetable, being ex- cluded), present an advance upon the simplicity of the Rliizo- poda in this, — that whilst their bodies consist for the most part of sarcode, and present scarcely anything that can be termed a distinction of organs, their external surface is con- densed into a membrane too firm to admit either of indehnite extension into pseudopodia, or of the passage of alunentary particles through it ; and consequently the form of the body, although not insusceptible of being temporarily changed by pressure, possesses a considerable degree of constancy for each species (fig. 80). A mouth, or definite aperture for the in- gestion of food, is provided; with an additional orifice in some instances, through which indigestible or effete matters may be discharged from the interior. Into this moutli, ali- Infusory Animalcules. I. Monads ; ii. Trachelis anas; iii. Enchelis, discharging fecal matter, iv. Para- ma'cium ; v. Kolpoda ; vi, Trachelis fasciolarls. mentary particles are drawn by the agency of the cilia with which some part of the surface of the body is provided ; these cilia being always so disposed as to serve at the same time for the general locomotion of the animalcule, and for the production of currents that shall bring food to its interior. 134. Although most Infusoria move freely through the water in which they live, yet certain kinds of them attach themselves by footstalks to marine plants or other floating bodies, during at least a part of their lives ; and in this con- dition bear no slight resemblance to Zoophytes, tho^igh of far simpler organization. It is in these sessile forms that the agency of the cilia in creating currents which l)ring food to 140 INFUSORIA. — PORIFERA OR SPONGES. the mouth, becomes most conspicuous. The alimentary par- ticles introduced into the mouth commonly have to pass down a short canal before they enter the general cavity of the body ; and within this cavity a number of minute par- ticles are commonly aggregated into a sort of little pellet, as may be Avell seen when Infusoria are fed with carmine or indigo. One after another of these pellets being thus intro- duced into the interior, which is occupied by a soft sarcode, each seems to push onwards its predecessors ; and a kind of circulation is thus occasioned in the contents of the cavity. The pellets that first entered make their way out after a time (their nutritive materials having been yielded up), generally by a distinct anal orifice, sometimes, however, by any jiart of the surface indifferently, and sometimes by the mouth. 1 35. The multiplication of Infusoria ordinarily takes place by spontaneous fission, precisely after the manner of the multiplication of ordinary cells (§ 33). This process, under favourable circumstances, may be performed with such rapidity, that, according to the computation of Ehrenberg, no fewer than 268 millions might be produced in a month by the repeated subdivision of a single Paramecium. Sometimes, instead of undergoing subdivision into two equal parts, the Animalcule puts forth a bud, which gradually increases, and then detaches itself from the parent stock. Whether any- thing equivalent to the sexual generation of higher animals occurs among Infusoria, is yet uncertain ; but recent re- searches aftbrd a probability in the affirmative. 13G. In the tribe of Porifera, or SjmiigeSj we seem to have the connecting link between Protozoa and Zoophytes. For their animality does not lie so much in the individual particles, as in those aggregations which begin to shadow forth that distinction into organs which is carried out more completely among Zoophytes : and there is a large section of the last-named group, in which the polypes are connected together, not by a regular stony or horny stem, but by a sponge-like mass ; while the extension of the flibric is provided for -by the budding out of this spongy portion of it, the orifices of whose canals after a time become fu;'nished mth polype-mouths. The true Sponge (fig. 81) consists of a fleshy sub.stiince, composed of an aggregation of particles of sarcode, supjM.rtcd upon a skeleton which usually consists of a net- PORIFERA OR SPONGES. 141 work of horny fibres, strengthened by spicules of mineral matter, sometimes calcareous, but more commonly siliceous. The entire mass is traversed by a great number of canals, which may be said to commence in the small pores upon its surface, and which discharge themselves into the wide canals that terminate in the large orifices, or vents, that usually pro- ject more or less from the surface of the Sponge. Through this sys- tem of canals, there is continually taking place, during the living state of the animal, a circulation of water, which is drawn in from without through the minute pores, then passes into the large canals, and is ejected in a constant stream from the vents. The immediate cause of this movement seems to lie in the vibration of cilia so extremely minute that their existence can only be detected by the most careful micro- scopic examination. Its purpose is evidently to convey to the animal the nutriment which it requires, and to carry off the matter which it has to reject. No distinct indications of sensation, or of power of locomotion, have been seen in the Sponge : but changes in the form of its projecting vents may be seen to take place from time to time, if it be watched sufficiently long. 137. The reproduction of the Sponge is commonly effected by the budding forth of little particles of sarcode, from the layer which lines the larger canals ; these become furnished with cilia, and, when detached and carried out by the current that issues from the vents, swim freely about for some time ; so as, before fixing themselves and beginning to develope into Sponges, to spread the race through distant localities. But it appears that Sponges are also reproduced by a true sexual process ; " sperm-cells " and " germ-cells " being pro- duced (as in the Hydra, § 123) in different parts of the organism, and a true embryo taking its origin in the action of the contents of the former upon those of the latter. 142 NATURE AND SOURCES OF ANIMAL FOOD. 138. We thus conclude our general survey of the Animal Kingdom ; which, it is hoped, will be found to answer the puj'pose for which it was designed, — that of giving such an amount of preparatory knowledge respecting the principal types of animal structure, as may enable even the beginner to comprehend what will hereafter be stated of their physiological actions. It has not been attempted to observe any proportion in the notice of these several types ; the higher forms having been slightly passed over, because the details of their vital phenomena wdll constitute the principal subject of the follow- ing pages ; whilst some among the lower have been more fully treated, because the ordinary reader cannot be expected to liave even that outline-acquaintance with their nature and actions, which he can scarcely help possessing in the case of animals that are familiar to him. CHAPTEE III. NATURE AND SOURCES OF ANIMAL TODD. 139. Before we examine the nature of the process by which the food of animals is prepared for absorption into their bodies, it will be desirable to consider the characters of the aliment itself, and the purposes to which it is to be appro- priated. The term food or aliment may be applied to all those substances which, when introduced into the living body, serve as materials for its growth, or for the repair of the losses which it is continually sustaining (§ 55). When animals are deprived of these materials, we see their bodies ]irogressively diminishing in bulk, their strength decreases, and death at last takes place, after sufferings more or less prolonged. In warm-blooded animals, however, a yet more urgent demand for food is created by the requirements of the licat-producing process ; and many substances are fitted to supply this, which cannot serve for the nourishment of the tissues. UO. The demand of the body for food is made known by a peculiar sensation, which has its seat in the stomach, namely, hunger. It is increased by mental and bodily exercise, and NATURE AND SOURCES OF ANIMAL FOOD, 143 by everything which augments the general energy of the system ; whilst, on the contrary, everytliing which tends to retard the operations of life, such as bodily and mental inac- tivity, sleep, or depression of spirits, tends also to render the demand for food less imperious. Thus, cold-blooded animals, particularly Eeptiles, can sustain a very prolonged abstinence, when the general activity of their functions is kept down by a low temperature; and hybernating Mammals, which pass the winter in a state of torpidity, require no food during the continuance of their lethargy. But with this exception, warm-blooded animals require a constant supply of nutriment, not merely for the maintenance of their proper heat, but also for the repair of the waste resulting from that continuous activity which the uniform temperature of their own bodies enables them to keep up. This is the case with Man and the MammaUa generally, and still more with Birds, whose temperature is higher, and whose movements are more active and energetic. It is also more the case with young animals than with adults ; since in the former the changes in the tissues, in consequence of the increase they are undergoing, take place with much more rapidity than in the latter, the bulk of whose bodies remains stationary. Hence, if children, young persons, and adults be shut up together, and deprived of food, the younger will usually perish first, and the adults will survive the longest. The Itahan poet Dante has given a terrible picture of such an occurrence, in his history of the imprisonment of Count Ugolino and his children. 141. The difference in the demand for food between the young growing animal and that which has arrived at maturity, is very remarkable in the case of Insects. There are no animals more voracious than the larva or caterpillar; and there are none that can sustain abstinence, with little dimi- nution of their activity, better than the imago or perfect insect. The larva3 of the Flesh-fly, produced from the eggs laid in carrion, are said to increase in weight 200 times in the course of 24 hours ; and theu^ voracity is so great as to have caused Linnceus to assert, that three individuals and their immediate progeny (each female giving birth to at least 20,000 young, and a few days sufficing for the production of a third generation) would devour the carcase of a horse with greater celerity than a lion. The larva of the Silk-worm 144 NATURE AND SOURCES OF ANIMAL FOOD. weighs, ^vllCIl hatched, about 1-lOOth of a grain ; previously to ils first metamorphosis it increases to 95 grains, or 9,500 times its original weight. The comparative weight of the full-grown caterpillar of the Goat-moth to that of the young one just crept out of the egg, is said to be as 72,000 to 1. For this enormous increase a very constant supply of material is necessary, and many larva3 perish if left unsupplied with food for a single day. On the other hand, a black beetle (Melasoma) has been known to live seven montlis, pinned down to a board ; and another beetle (Scarabseus) has been kept three years without food,— and this ^vithout manifesting any inconvenience or loss of activit}^ There are many perfect insects which never eat after their last change, but die as soon as they have performed their part in the propagation of the race. 142. The nature of the food of animals is as various as the conformation of their different tribes. It always consists, however, of substances that have previously undergone organ- ization. There are some apparent exceptions to this, in the case of animals which seem to derive their support, in part at least, from mineral matter. Thus, the Spatangus (an animal allied to the Echinus, § 119) hlls its stomach with sand ; but it really derives its nourishment from the minute animals which this contains. The Earthworm and some kinds of Beetles are known to swallow earth ; but only to obtain from it the remains of vegetable matter that are mixed with it. By some races of Man, too, what seems to be mineral matter is mixed with other articles of food, and is said to be nutri- tious ; this may be beneficial, in part, by giving bulk to the aliment, and thus exciting the action of the stomach (§ 205); but it has been found, in one case at least, that the supposed earth consists of the remains of animalcules, and contains no inconsiderable portion of organic matter. 143. There arc many instances in which, no obvious sup- plies of food bemg afforded, the mode of sustenance is obscure ; and it has been frequently supposed that, in such cases, the animals are sustained by air and water alone. But it will always be foTind that, wliere food is taken in no other way, a supply of tlic microscopic forms of animal or vegetable life is introduced by ciliary action (§ 45); and it is on these, indeed, that a large proportion of the lower forms of aquatic annuals depend entirely for their sui)port. NATURE OF THE FOOD OF ANIMALS. 145 144. The first division of aliments is naturally into those which are derived from the Animal and Vegetable kingdoms respectively. Wherever plants exist, we find animals adapted to make use of the nutritious products they furnish, and to restrain their luxuriance within due limits. Thus among Mammals, the Dugong (an animal having the general form and structure of the whale, but adapted to a vegetable diet) browses upon the sea-weeds that grow beneath the surface of the tropical ocean ; the Hippopotamus roots up with his tusks the plants growing in the beds of the African rivers, and fills his huge paunch, not only with these, but with the decaying vegetable matter which he finds in the same situa- tion ; the Antelopes, Deer, Oxen, and other Euminants, crop the herbage of the plains and meadows ; the Giraffe is enabled by his enormous height to feed upon the tender shoots which are above the reach of ordinary quadrupeds ; the Sloths, living entirely in trees, and hanging from their branches, strip them completely of their leaves ; the Squirrels extract the kernels of the hard nuts and seeds ; the IMonkeys devour the soft pulpy fruits ; the Boar grubs up the roots and seeds buried under the soil ; the Eeindeer subsists during a large part of the year upon a lichen that grows beneath the snow ; and the Chamois finds a sufficient supply in the scanty vegetation of Alpine heights, Not less is this the case among Birds ; but in the classes of Reptiles and Fishes, the number of vegetable-feeders, and consequently the variety of their food, is much less. 145. Among Insects, a very large proportion derive their food entirely from Plants, and many from particular tribes of plants only; so that, if from any cause these should fail, the race may for a time disappear. There is probably not a species of plant which does not furnish nutriment for one or more tribes of insects, either in their larva state or their per- fect condition ; and in this manner it is prevented from mul- tiplying to the exclusion of others. Thus, on the Oak no less than two hundred kinds of caterpillars have been estimated to feed ; and the Nettle, which scarcely any beast will touch, supports fifty different species of insects, — but for which check it would speedily annihilate all the plants in its neigh- bourhood. The habits and economy of the different races existing on the same plant, are as various as their structure. L 146 VORACITY OF INSECTS. Some fcGcl only upon the outside of the leaves ; some upon the internal tissue ; others upon the flowers or on the fruit ; a few wiil cat nothing but the bark ; while many derive their nourishment only from the woody substance of the trunk. 146. The excessive multiplication of certain tribes of Insects has sometimes had the effect of devastating an entire coiuitry. Tims the " plague of locusts " is not unfrequently repeated in tropical countries, and is dreaded by the inhabi- tants even more than an earthquake. These insects are of such extreme voracity that no gi-een thing escapes them; and when their nimibers are so increased that they fly in masses which look like dark clouds, and cover the ground where they aUght for miles together, it may be easily con- ceived that the devastation they create must produce incal- culable injury. The north of Africa and the west of Asia are the countries most infested by these pests. It is related by Augiistin, that a plague, induced partly by the famine they had created, and partly by the stench occasioned by theii' dead bodies, carried off 800,000 inhabitants from the kingdom of i!^umidia and the adjacent parts. They occasionally attack the south of Europe. It is recorded that Italy was devastated by them in the year 591 ; and that a prodigious number both of men and beasts perished from similar causes, — no less than 30,000 persons in the kingdom of Venice alone. These tremendous swarms usually advance towards the sea ; and being there checked, and having completely exhausted the country behind them, they themselves die of famine, or are blo^vn into the sea by a gale. In 1784 and 1797, they de- vastated Southern Africa ; and it is stated by Mr. Barrow (in his Travels in that country) that they covered a surface of 2,000 square miles ; that, when cast into the sea by a strong wind from the north-east, and washed upon the beach, they formed a line fifty miles long, and produced a barrier along the coast three or four feet high ; and that, when the wmd again changed, the stench created by the putrefaction of their bodies was perceived at a distance of 150 miles inland. A similar event occurred in the Earbary States in 1790, and was followed, as in the other cases, by a plague. 147. We have occasionally an example of similar devasta- tion in our own country, thougli on a smaller scale. Thus, a few years ago, the turnip-crops of some parts of England VORACITY OF INSECTS. 147 were almost entirely destroyed by the larva' of an insect called the " turnip-fly." The parent insects were seen buzzing over the fields, and depositing their eggs in the plants, which they do not themselves employ as food ; and in a few days all the soft portions of the leaves were destroyed, and nothing but the skeletons and stalks were left. — Some kinds of timber occasionally suffer to no less an extent from the devastations of insects, whose operations are confined to the wood, and do not manifest themselves externally, until the tree is seen to languish and at last to die. The pine-forests of the Hartz mountains in Germany have been several times almost de- stroyed by the ravages of a single species of beetle, less than a quarter of an inch in length. The eggs are deposited beneath the bark ; and the larvae, when hatched, devour the sap- wood and inner bark (the parts most concerned in the func- tions of vegetation) in their neighbourhood. It was estimated that, in the year 1783, a million and a half of pines were destroyed by tliis insect in the Hartz alone ; and other forests in Germany were suffering at the same time. The wonder is increased, when it is stated that as many as 80,000 larvse are sometimes found on a single tree. 148. But every class in the Animal Kingdom has its car- nivorous tribes, which are adapted to restrain the too rapid increase of the vegetable-feeders (by which a scarcity of their food would soon be created), or to remove from the earth the decomposing bodies that might otherwise be a source of dis- ease or annoyance. The herbivorous races, being for the most part very prolific, would very rapidly increase to such an extent as to produce an absolute famine, if not kept in check by the races appointed to limit their multiphcation. Thus, the myriads of Insects which find their subsistence in our forest-trees, if allowed to increase without restraint, w^ould soon destroy the hfe that supports them, and must then all perish together ; but another tribe (that of the insectivorous Birds, as the woodpecker) is adapted to derive its .subsistence from them, and thus to keep their numbers wdtliin salutary bounds. Their occasional multiplication to the enoimous extent mentioned in the preceding paragraphs, is probably due in general to the absence of the races that should keep them in check. This may occur from accidental causes, or may be produced by the interference of Man. Thus, a set of l2 148 BALANCE AMONG DIFFERENT RACES. ignorant farmers have imagined that a neighbouring rookery was injurious to them, because they saw the rooks hovering over the newly-sown corn-fields, and seeming to pick the grains out of the ground ; and ha^dng extirpated the rookery, they have found in the course of a year or two that they have done themselves an immense injury, — the roots of their corn and grasses being devoured by the grubs of cockchafers and other insects, the multiplication of which was before prevented by the rooks, whose natural food they are. 149. On the other hand, by an intelligent application of this principle, the excessive multiplication of insects has been prevented where it had already commenced. Thus, no means of extirpating the larvse of the turnip-fiy was found so suc- cessful, as turning into the fields a number of ducks, which quickly removed them from the plants. And in the island of Mauritius, the increase of locusts, which had been accidentally introduced there, and which were becoming quite a pest, was checked by the introduction from India of a species of bird, the grakle, which feeds upon them. 150. Of the carnivorous tribes themselves, however, the increase might be so great as to destroy all the sources of their food, were it not that they are kept in check by others, larger and more powerful than themselves, which, not being prolific, are not likely ever to gain too great a power. Thus, among birds, the eagles, falcons, and hawks rear only two or three young every year, whilst many of the smaller birds produce and bring up four or five times that number. — The following is a curious instance of the system of checks and counter- checks, by which the "balance of power" is maintained amongst the different races. A particular species of moth having the fir-cone assigned to it for the deposition of its eggs, the young caterpillars, coming out of the shell, consume the cone and superfluous seed ; but, lest the destruction should be too great, another insect of the ichneumon kind lays its eggs in the caterpillar, inserting its long tail in the openings of the cone until it touches the included insect, its own body being too large to enter. Thus it fixes upon the caterpillar its minute egg, which, when hatched, destroys it. 151. The peculiarity of the agency of Insects, in the economy of nature, has been justly remarked to consist in their power of very rapid multiplication, in order to accomplish a VARIATIONS IN POWER OP ABSTINENCE. 149 certain object, and then in their as rapidly dying off. In tliis re- spect they resemble the Fungi among plants. ( Botany, § 789.) 152. There are great variations in the degree of power possessed by animals of different species to sustain abstinence from food, which appear to be related to their respective habits of life ; such as most easily obtain a constant supply of food being immediately dependent upon it, and vice versd. Thus, among the larv?e of Insects, those that feed upon vege- tables or dead animal matter (in the neighbourhood of which theii' eggs are usually deposited by the parent) speedily die if placed out of reach of their aliment ; whilst those that lie in wait for living prey, the supply of which is uncertain, are able to endure a protracted abstinence, even to the extent of ten weeks, without injury. Again, carnivorous Birds and Mam- mals are generally able to exist for some time without food ; their natural habits leading them to glut themselves upon the carcase of the animal they have destroyed, in such a manner as to prevent them from requiring any new supply for some time : thus the wild cat has been kept twenty days without food, the dog has lived for thirty-six days in the same circum- stances, and the eagle for a similar period. But some herbi- vorous animals, such as the camel and the antelope, whose habits are such as to keep them out of the reach of food for several days together, are able to endure a similar abstinence ; whilst among the insectivorous Mammals, which naturally take food often, and but little at a time, the powxr of absti- nence is much less, — the mole, for instance, perishing in confinement, if not fed once a day, or even more frequently. 153. We have next to consider the different substances used as food, in regard to their chemical composition ; and to inquire for what purposes in the nutrition of the body they are respectively destinecl. The Vegetable tissues are chiefly made up of the three components, oxygen, hydrogen, and carbon ; the oxygen and hydrogen having the same proportions as in water. Their composition being thus nearly the same as that of starch, gum, and sugar (into which, indeed, they may for the most part be converted by a simple chemical process), alimentary substances of this kind form a natural group to which we may give the name of Saccharine (sugary). — But in many vegetable substances used as food, there is a considerable quantity of oily matter, stored up in cells ; and the same kind loO ORGANIC CONSTITUENTS OP ANIMAL FOOD. of matter constitutes the principal part of the fat of animals. Of these oily and fatty matters, also, the chemical elements, oxygen, hydrogen, and carbon, are the only ingredients ; but they are combined in i)roportions different from the last, the two latter predominating considerably. Hence they consti- tute another group of alimentary materials, to which the term Oleaginous maybe given. — Lastly, most Vegetables con- tain, in greater or less amount, certain compounds M-hich consist of the four elements, oxygen, hydrogen, carbon, and nitrogen, of which the animal tissues are composed. These compounds exist most largely in the corn-grains, and also in the seeds of the pea and bean tribe ; but there are few vege- table substances used as food by animals, that do not contain them in some small amount. The gluten of wheat, the legu- min of peas, and other vegetable substances of this kind, together v/ith the flesh of animals, the composition of which (§ 13) is identical with theirs, are united into a third group, 10 which the name Albuminous is given. — "We cannot pro- perly include in this group, however, the gelatinous portions of the animal tissues, which exist largely in gristle, bone, the skin, and other ])arts ; because gelatin (the substance that forms glue), though it agrees with albumen in being made up of WiQj'our ingredients just named, differs from it extremely in the proportions of tliose elements (§ 19) ; so that, although there is good reason to believe that gelatin may be formed out of albumen, it does not seem that any albuminous compound can be formed out of gelatin. Hence we must consider the gelatinous compounds separately. 154. Of these four groups, the last two are distinguished as azotized compounds, or substances that contain azote or nitro- gen ; whilst the first two are spoken of as non-asotizecl, being destitute of this element. The distinction is a very important one ; and must be kept steadily in view in considering the ulti- mate destination of each kind of food. It is obvious from what has'been already stated as to the composition of the animal tis- sues (§§13—21), that azotized compounds must supply the chief materials for their nutrition and re-formation. The non-azotized substances must be for the most part destined, unless converted mto azotized compounds within the living body, either to be sniiply deposited in its interstices, or to be thrown ofl' from it again witliout ever actually forming part of its organised structure. DESTINATION OF NON-AZOTIZED ALIMENTS. 151 155. Now, in regard to the non-azotized, or the saccharhie and oleaginous groups of aUmentary substances, it aj^pears to be an established fact, that none of the higher animals can be permanently supported upon them alone. Thus, dogs that have been fed on sugar and starch only, do not survive long ; and it is evident, before their death, that their tissues are gradually undergoing decay. It has been thought that such results might be partly explained upon the fact, that animals fed upon one simple substance soon become disgusted with it, and will even refuse it altogether ; but the experiments have been repeated with a combination of various non-azotized sub- stances, and the same result has occurred. Still it is too much to afiirm, as some have done, that these substances do not con- tribute in any degree to the nutrition of the animal tissues; since there is ample evidence that the presence of fatty matter in the blood is a condition essential to the production of newly forming tissue ; and we find that either oleaginous substances, or substances belonging to the saccharine group which can be readily converted into fat within the bod}^, constitute an im- portant part of the food of Man, and of animals generally.^ 156. That such a conversion can take place, has been de- monstrated by experiments carefully conducted upon bees, which have been found to generate wax when fed upon sugar only ; and also upon cows, which give off in their milk so much larger a quantity of butter than can be produced at the expense of the fat contained in their food, that there is no other mode of accounting for its 2)resence, than by regarding it as generated from the starchy portion of their diet. And the fattening power of starchy and saccharine articles of diet is well known to breeders of cattle ; though the articles which contain oily matter in addition seem to possess a higher value in this respect. 157. But if these non-azotized compounds, which exist so largely in the food of herbivorous animals, are not destined to form any other permanent part of the anunal organism than the oleaginous contents of the fat-cells (§ 4G), the ques- tion again arises, — what becomes of them 1 It is not enough ^ The value of cod-liver oil, which is now so extensively used in the treatment of diseases of imperfect nutrition, seems to depend upon the readiness with which it can be digested and assimilated, so as to furnish the supply of fat required by the formative processes. 152 DESTINATION OF NON-AZOTIZED ALIMENTS. to say that tlicy are deposited as fat ; since it is only when a largo quantity of them is taken in, that there is any in- crease in the ijuantity of fat already in the body. We shall hereafter see that they are used up in the process of respira- tion, one great object of which is, to produce a certain amount of heat, sufficient to keep up the temperature of the body, in ^^•arm-blooded animals, to a high standard. We might almost say mth truth, that a great part of the oleaginous and sac- charine principles is burned within the body, for this pur- |)Ose. The process will be hereafter considered more in detail (§§ 412, 413) ; and at j)resent we need only stop to remark upon the adaptation between the food provided for animals in different climates, and the amount of heat which it is necessary for them to produce. Thus the bears, and seals, and whales, from which the Esquimaux and the Green- lander derive their support, liaA^e an enormous quantity of fat in their massive bodies : this fat is as much esteemed as an article of food amongst these people, as it would be thought repulsive by the inhabitants of southern climates ; and by the large quantity of it they consume, they are able to support the bitterness of an Arctic winter, without appearing to suffer more from the extreme cold than do the residents in more temperate climes during their winter. On the other hand, the antelopes, deer, and wild cattle, which form a large pro- l)ortion of the animal food of savage or half-cultivated nations inhabiting tropical regions, possess very little fat; and the comparatively small supply of carbon and hydrogen, of which the combustion is required to keep up the bodily temperature of the inhabitants of those regions, is derived from i\ie flesh of these animals, in the manner that will be presently explained. 158. The application of the substances forming the albu- minous group, to the support of the animal body, by affording the materials for the nutrition and re-formation of its tissues, needs little explanation. The proportions of the four ingre- dients of which they are all composed, are so nearly the same, that no essential difference appears to exist among them ; and it is a matter of little consequence, except as far as the gra- tification of the palate is concerned, whether we feed upon the llcsh of animals (syntonin, § IG), upon the white of agg (albumen, § 1 ;^), the curd of milk (casein, § 15), the grain of wheat (gluten), or the seed of the pea (legumin). All these DESTINATION OF NON-AZOTIZED ALIMENTS. 153 substances are reduced in the stomach to the form of albumen ; which is the raw material out of which the various fabrics of the body are constructed. But the rule holds good with re- gard to these also, that by being made to feed constantly on the same substance, — boiled white of q,^^^ for instance, or meat deprived of the principle that gives it flavour, — an animal may be effectually starved ; its disgust at the food being such, that even if it be swallowed it is not digested. It is very interest- ing to remark that, in the only instance in which Nature has provided a single article of food for the support of the animal body, she has mingled articles from all the three preceding groups. This is the case in Milk ; which contains a consider- able quantity of the albuminous substance, casein., that forms its curd ; a good deal of oily matter, the butter ; and no in- considerable amount of sugar, which is dissolved in the whey. The proportions of these vary in different Mammalia, being- related as it would seem to the habits of the young animal thus sustained, while they depend in part upon the nature of the food supplied to the animal that forms the milk ; but the three substances are thus combined in every instance. 159. But although the greater part of the organised tis- sues of animals have a composition nearly allied to that of albumen, many of them also contain a large quantity of gelatin (§ 19). It seems certain that this gelatin may be pro- duced out of albuminous substances ; since in animals that are supported on these alone, the nutrition of the gelatinous tissues does net seem to be impaired. But it appears equally certain, that gelatin cannot be applied to the nutrition of the albuminous tissues. Many series of experunents have been made on this subject, with a view of determining how far gelatin-soup made from crushed bones (such as that which long constituted a principal article of diet in the hospitals of Paris) is adequate for the support of the body in health. The result of these has been uniformly the same, — namely, that although gelatin may be advantageously mixed with albumen, fibrin, gluten, &c., and those other ingredients which exist in meat-soup and bread, yet that, when taken alone, it has little (if any) more power of sustaining life, than sugar or starch possesses. Although it might have been thought hkely that gelatin employed as food might be applied witliin the body to the nutrition of its gelatinous tissues, yet there 1,')4 SOURCES OF DEMAND FOR ALIMENT. is strong reason to believe that these, like the albuminous, are formed at the expense of the albuminous matter of the blood, and that gelatin thus introduced undergoes a rapid decomposition, yielding up a considerable part of its carbon and hydrogen to the combustive process, which is the only function to wliich it affords any substantial imhidum. Con- sequently the current idea regarding the nutritive value of jdliea of various kinds, has little or no real foundation. IGO. It has been already stated (§ ^^) that all the living tissues of the body are continually undergoing a sort of death and decay ; and that they do this the more rapidly, in pro- ])ortion as they are called upon for the discharge of their functions. The need of material capable of rejDlacing that Avhich has been lost, is consequently the chief source of the constant demand for aliment. Even in young, actively growing animals, the quantity required for the increase of their bodies constitutes but a very small proportion of that which is taken in ; of the remainder, a part is at once re- jected as indigestible ; and the rest is appro j)riated to the ix'pair of the loade which is continually going on. This wast^e is much greater in young animals than in adults ; for all their vital i)rocesses are more actively and energetically performed : their movements are quicker in proportion to their size ; and injuries are more speedily repaired. To remove the products of this decomposition is the special object of the various pro- cesses of excretion ; and among these, the resjnration, by which a large quantity of carbon and hydi-ogen is carried off in the form of carbonic acid and water, is of the most constant importance, on account of the heat which it thus enables the animal body to maintain. This temperature, in Carnivorous animals, appears to be sufficiently kept up by the combustion of the carbon and hydrogen set free by the decay (or metamorphosis, as it may be termed) of their tis- sues ; but this combustion goes on with much more rapidity, in consequence of their almost unceasing acti^dty, than it does in the Herbivorous animals, which lead comparatively inac- tive lives. Every one who has visited a menagerie must have noticed the continual restlessness of the Tigers, Leopards, Hyenas, &c., which keep pacing from one end of their narrow cages to the other ; and it would seem as if this restlessness were a natural instinct, impelling them to use muscular exer- NUTRITION OF CARNIVOROUS ANIMALS. 155 tion sufficient for the metamorphosis of an adequate amount of tissue, that enough carbon and hydrogen may be set free for the support of the respiratory process. And we see a cor- responding activity in the Human hunters of the swift-footed antelope and agile deer, which answers a similar purpose ; and which is remarkably contrasted with the stupid inertness of the inhabitants of the frigid zone, that is only occasionally interrupted by the necessity of securing the supplies of food afforded by the massive tenants of their seas. 161. The nutrition of the Carnivorous races may, then, be thus described. The bodies of the animals upon which they feed, contain flesh, fat, &c., in nearly the same proportion as their own ; and all, or nearly all, the aliment they consume, goes to supply the waste in the fabric of their own bodies, being converted into its various forms of tissue. After having remained in this condition for a certain tune, varying ac- cording to the use that is made of them, these tissues un- dergo another metamorjDhosis, which ends in restoring them to the condition of inorganic matter ; and thus give back to the mineral world the materials wliich were drawn from it by plants. Of these materials, part are burned off, as it were, within the body, by union with the oxygen of the air taken in through the lungs, from which organs they are discharged in the form of carbonic acid and water : the remainder are carried off in the liquid form by other channels. Hence we may briefly express the destination of their food in the following manner : — l^ Carbonic acid and ^ , . ^. . ,. \ water, tlirown ofT ^0"^,^""^}^*"^^ ) Livin- ) and this (by respiration, of albuminous I converted \ ^r^anised metamorphosed / Uren and biliary ami other com- r into ( -^^^^^^ | i„^|, matter, &c., P°""^s J ' ^1 thrown off by ' other excretions. 162. But ill regard to the Herbivorous animals, the case is different. They perspire much more abundantly, and their temperature is thus continually kept down (§ 372). They consequently require a more active combustion, to de- velop sufficient bodily heat ; and the materials for this are supplied, as we have seen, by the non-azotized constituents of their food, rather than by the metamorphosis of their own tissues, which takes place with much less rapidity than in the carnivorous tribes. Hence we may thus express the 156 NUTRITION OF IIERBIVORA AND OF MAN. destination of this part of tlieir food ; that of the albuminous matters, here mueh smaller in amount, being the same as in the preceding case : — Starch, oil, and 'i partly ( Fatty and -j but chiefly j' Carbonic acid and water, other lion -azo- > converted J other animal [ thrown otf < disengaged by the respi- tized compounds j into ( tissues, ; directly as Uatory process. The proportion of the food deposited as fat, will depend in part upon the surplus which remains, after the necessary sup- ply of materials has been afforded to the respiratory process. Hence, the same quantity of food being taken, the quantity of fat will be increased by causes that check the perspiration, and otherwise prevent the temperature of the body from being lowered, so that there is need of less combustion within the body to keep up its heat. This is consistent with the teach- ings of experience respecting the fattening of cattle ; for it is well known that this may be accomplished much sooner, if the animals are shut up in a warm dwelling and are covered with cloths, than if they are freely exposed in the open air. 163. Now the condition of Man may be regarded as inter- mediate between these two extremes. The construction of his digestive apparatus, as well as his own instinctive pro- pensities, point to a mixed diet as that which is best suited to his wants. It does not appear that a diet composed of ordinari/ vegetables only, is favourable to the full develop- ment of either his bodily or his mental powers ; but this cannot be said in regard to a diet of which the corn-grains furnish the chief ingi-edient, since the gluten they contain appears to be as well- adapted for the nutrition of the animal tissues, as is the flesh of animals. On the other hand, a diet composed of animal flesh alone is the least economical that can be conceived ; for, since the greatest demand for food is created in him (taking a man of average habits in regard to activity and to the climate under which he lives) by the ne- cessity for a supply of carbon and hydrogeii to support his respiration, this want may bo most advantageously fulfilled by the employment of a certain quantity of non-azotized food, in which these ingredients i)redominate. Thus it has been calculated that, since lifteen pounds of flesh contain no more cai-]jon than four pounds of starch, a savage with one animal and an equal weight of starch, could support life for the same length of time during which another restricted to annual COMPOSITION OF ARTICLES OF HUMAN FOOD. 157 food would require five such animals, in order to procure the carbon necessary for respiration. Hence we see the immense advantage as to economy of food, which a fixed agricultural population possess over the w\andering tribes of hunters which still people a large part both of the Old and Xew Continents. 164. The following Table exhibits the proportions of albu- minous, starchy or saccharine, fatty, and saline substances, contained in various articles ordinarily used as food by Man ; together with the proportion which ivater bears in each case to the solid constituents of the food, which becomes a most important element of consideration when the nutritive value of diiferent kinds of food is compared : — Substances, 100 parts. Human Milk Cow's Milk Skimmed Milk ... Butter Milk Beef and Mutton Veal Poultry Bacon Cheese (Cheddar) ,, (Skimmed) Butter Eggs White of Egg Yolk of Egg White Fish Salmon Eel Wheat Flour Barley-meal Oat-meal Rye-meal Indian-meal Rice Haricots Peas Beans Lentils , Wheat-bread Rye-bread , Potatoes Green Vegetables . Arrow-root 1 ^ III 73 X ^ i 89 3.5 4.2 3.0 8(5 4.5 5.0 4.1 87 4.5 5.0 2.7 87 4.5 5.0 0.5 73 19.0 5.0 77 19 1.0 74 21.0 3.0 20 0.8 70.0 .•!6 29.0 30.0 44 45.0 6.0 15 830 74 14.0 10.5 78 20.0 52 Ib.O 30.0 79 19.0 ... 1.0 78 17.0 4.0 80 10.0 .., 8.0 15 11.0 70.0 2.0 15 10.0 70.0 2.4 15 12.0 620 6.0 15 9.0 66.0 2.0 14 9.0 65.0 8.0 14 7.0 76.0 0.3 19 23.0 45.0 3.0 13 22.0 58.0 2.0 14 24.0 44.0 1.4 14 29.0 44.0 1.5 44 9.0 49.0 1.0 48 5.3 4'i.O 1.0 74 2 23.0 0.2 8G 2.0 4.0 0.5 18 ... 82.0 ... 0.2 0.7 0.7 0.7 2.0 0.6 1.2 1.3 4.5 5.0 2.0 1.5 1.6 1.3 1.2 1.4 1.3 1.7 2.0 3.0 1.8 1.7 0.3 3.G 3.0 3.6 23 2.3 1.4 0.7 0.7 i3 3 11.4 14.8 11.5 6.0 12.0 2.4 7.2 168.0 72.0 14.4 199.0 25.0 72.0 2.4 9.6 19.2 74.8 75.8 76.4 70.8 84.2 76.7 52.2 C2.8 47.4 47.6 51.4 48.4 23.5 5.0 82.0 3.5 4.5 4.5 4.5 19.0 19.0 21.0 0.8 29.0 45.0 14.0 20.0 16.0 19.0 17.0 10.0 11.0 10.0 12.0 9.0 9.0 7.0 23.0 22.0 24.0 29.0 9.0 5.3 2.0 2.0 14.9 19.3 16.0 10.5 31.0 21.4 28 2 168.8 101.0 59.4 199.0 39.0 20.0 88.0 21.4 26.6 29.2 85 8 85.8 88.4 79.8 93.2 83.7 75.2 84.8 71.4 76.6 60.4 53.7 25.5 7.0 82.0 * The value of the Fat is stated in this cohimn according to its heating equivalent of starch, which is larger in the ratio of 2,4 to 1. Hence, in the last column, the proportion of nutriment in aliments containing fat, comes to be greater than the weight of their solids would indicate. 158 ECONOMY OF HUMAN DIET. Those articles of food in which tlie nitrogenous compounds predominate, are especially fitted for the maintenance of the soHd fabric of the body ; whilst those in which the carbon- aceous compounds are in largest excess, are those which are most efi'ective as supplying materials for the combustive pro- cess. Conspicuous among the former are the various kinds of animal flesh, as also the white of eggs ; whilst among the latter the most noticeable are bacon and butter, rice and potatoes, the former consisting almost wholly of fat, the latter being chiefly composed of starch. Of all single articles of food, good wheaten bread, in which the proportion of nitro- genous to carbonaceous components is about as 5.7 to 1, seems to be the one best suited to the ordinary wants of Man ; but this acquires much additional value from the con- current use of a moderate amount of fatty matter in the form of butter, 165. If the more highly azotized forms of food be em- ployed exclusively, a great excess of them must be consumed to supply the carbon needed for respiration ; whilst if the more carbonaceous kinds of food be used as the sole susten- ance, unless the quantity ingested be large enough to afford the requisite supply of azotized material for the maintenance of the tissues, their nutrition must be imperfectly effected, and the strength must fail. I^ot only in the instance just cited, but in a variety of others, the instincts of mankind have led to such a combination of different articles of diet, as includes in their appropriate proportions the albuminous, the saccharine, and the oleaginous principles. Thus with meat we eat potatoes ; and with the white meats which are deficient in fat, we eat bacon. We use melted butter with most kinds of fish, or fry them in oil ; whilst the herring, the salmon, and the eel, are usually fat enough in themselves, and are dressed and eaten alone. A similar adjustment is made when we mix eggs and butter with sago, tapioca, and rice ; when we add oil and the yolk of an egg to salad ; when we boil rice with milk, and combine cheese with maccaroni. Bacon and greens, and pork and pease-pudding, again, are combinations founded in taste, which approve themselves to the judgment; as is also the Irish dish termed kolcannon, con- sisting of potatoes and cabbage, with a little bacon or fat pork. So arc the mixture so common in Ireland and Alsace, of butter- ECONOMY OF HUMAN DIET. 159 milk or curdled-milk with potatoes ; and the combination of rice and fat, which is the staple of the diet of many Eastern nations. Even the morsel of butter or the bit of cheese which the English labourer eats with his hard-earned bread, are not matters of luxury, but have a positive importance ; and the existence of these tastes and habits shows how by long experience Man has at last learned to adjust the com- position of his food, so as best to maintain the health and vigour of his body. With a difference of requirement comes a difference of tastes. Thus men who are going through a very laborious course of exertion, prefer meat to bread or vegetables, feeling it to be more sustaining to their strength. On the other hand, those who are continuously exposed to the severity of an Arctic winter, eat with relish large masses of fat, on which they would look with disgaist under other circumstances. The quantity of work which a man can do, and his power of sustaining extreme cold, both depend in gi-eat part, as has now been abundantly proved, upon the adequacy of the sustenance he takes : the demand, in the first case, being for albuminous material to supply the waste of his tissues ; whilst in the second it is for combustive material suitable to generate heat iii large measure, — a purpose which is far more efficiently answered by oleaginous substances, than by those of a starchy or saccharine nature. Experience fur- ther shows that the healthy condition of the blood of Man can only be maintained by the use of fresh vegetables as part of his ordinary diet. When these are withdrawn for any length of time, the disease known as Scurvy is certain to appear, unless lemon -juice or some other efficacious anti- scorbutic be employed as a substitute. This is a fact of the utmost importance in provisioning ships for long voyages ; the tendency to scurvy being increased by confinement and insufficient ventilation, and by the exclusive use of salt provisions. 166. Besides these organic substances, there are certain Mineral ingredients, which may be said to constitute a part of the food of Animals ; being necessary to their support, in the same manner as other mineral substances are necessary to the support of Plants. Of this kind are common salt, and also phosphorus, sulphur, lime, and iron, either in combina- tion or separate. — The uses of Salt are very numerous and I GO MINERAL INGREDIENTS OF FOOD. iiiiportaiit. It exists largely in the blood, and in the various aninial fluids which are secreted from it ; and it is also an essential ingi-edient of most of the solid tissues. Its presence obviously tends to prevent that spontaneous decomposition to whicli organic substances are liable. Fhosphoi^us is chiefly required to be united with fatty matter, to serve as the material of the nervous tissue ; and to be combined with oxygen and lime, to form the bone-earth by which the bones are consolidated. Sulphur exists in small quantities in several animal tissues ; but its part seems by no means so important as that performed by phosphorus. Lime is required for the consolidation of the bones, and for the production of the shells and other hard parts that form the skeletons of the Invertebrata. Of the limestone rocks of which a great part of the crust of our globe is composed, a very large proportion is made up of the remains of animals that formerly existed in the ocean. Thus some almost entirely consist of masses of Coral, others of beds of Shells, and others of the coverings of minute Foraminifera (§131). To these mineral ingredients we may also add Iron, which is a very important elem.ent in the red blood of Vertebrated animals. 167. These substances are contained, more or less abun- dantly, in most articles generally used as food ; and where they are deficient, the animal suffers in consequence, if they be not supplied in any other way. Common Salt exists, in no inconsiderable quantity, in the flesh and fluids of animals, in milk, and in the egg : it is not so abundant, however, in plants ; and the deficiency is usually supplied to herbi- vorous animals by some other means. Thus salt is purposely mingled with the food of domesticated animals ; and in most parts of the world inhabited by wild cattle, there are spots Avherc it exists in the soil, and to which they resort to obtain it ; such are the " buffalo-licks" of North America. Fhos- phorus exists also in the yolk and white of the egg, and in jnilk, — the substances on which the young animal subsists < hiring the period of its most rapid gi-owth ; it abounds not only in many animal substances used as food, but also (in the state of phosphate of lime or bone-earth) in the seeds of many plants, especially the grasses ; and in smaller quantities it is found in the ashes of almost every plant. When flesh, bread, fruit, and husks of grain, arc used as the chief articles MINERAL INGREDIENTS OF ANIMAL FOOD. 161 of food, more phosphorus is taken into the body than it requires ; and the excess has to be carried out in the excre- tions. 6'ulphur is derived aUke from, vegetable and animal substances. It exists in flesh, eggs, and milk ; also in the azotized compounds of plants ; and (in the form of sulphate of lime) in most of the river and spring water that we drink. Iron is found in the yollc of egg, and in milk, as well as in animal flesh ; it also exists, in small quantities, in most vegetable substances used as food by man, — such as potatoes, cabbage, peas, cucumbers, mustard, &c. ; and probably in most articles from which other animals derive their support. 168. Lirne is one of the most universally diffused of all mineral bodies ; there being very few animal or vegetable substances in which it does not exist. It is most commonly taken in, among the higher animals, combined with phos- phoric acid, so as to form bone-earth, in which state it exists largely in the seeds of most grasses. A considerable quantity of lime exists, moreover, in the state of carbonate and sul- phate, in all hard water. 169. When an unusual demand exists for lime, however, for a particular purpose, an increased supply must be aflbrdecl. Thus a hen preparing to lay, is impelled by her instinct to eat chalk, mortar, or some other substance containing the car- bonate of lune which is required for the consolidation of the shell ; and if this be withheld, the egg is soft, its covering being composed of animal matter alone, not consolidated by the deposit of earthy i)articles. The thickness of the shells of aquatic Mollusks depends greatly upon the quantity of hme in the surroundmg water. Those which inhabit the sea, find in its waters as much as they require ; but those that dwell in fresh-water lakes, which contain but a small quan- tity of lime, form very thin shells ; whilst, on the other hand, those that inhabit lakes in which, from peculiar local causes, the water is loaded with calcareous matter, form shells of remarkable thickness. 170. The mode in which the Crustacea, whose calcareous shell is periodically thrown ofi' (§ 99), are able to renew it with rapidity, is very curious. There is laid up in the walls of their stomachs a considerable supply of calcareous matter, in little concretions, which are commonly known as " crabs' eyes." When the shell is cast, this matter is taken up by M 1C2 DIGESTION AND ABSORPTION. the Llood, and is thrown out from the surface, mingled with animal matter. This hardens in a day or two, and the new covering is complete. The concretions in the stomach are then found to have disappeared ; but they are gradually replaced, before the supply of lime they contain is again required. CHAPTER IV LIGESTION AND ABSORPTION. 171. Having now considered the nature of the food of Animals, and the sources from Avliich it is obtained, we have next to consider the process by which the aliment is received into their bodies, and prepared to form a part of their own fabric. This process, termed Digestion, is naturally divided, among the higher animals at least, into various stages. In the first place, there is the prehension or laying hold of the aliment, and its introduction into the mouth or entrance to the digestive cavity. In the mouth it usually undergoes a preparation ; which consists partly in its being cut, ground, or crushed, by mechanical action, into minute pieces ; and partly in the working-up of these pieces with a fluid that is poured into the mouth, — the saliva. These two processes are termed mastication and insaUvation ; similar processes are performed, in some animals, in a part of the digestive tube intermediate between the mouth and the stomach, and even in the latter itself. The stomach is usually situated at some distance from the mouth, and is connected with a tube called the oesophagus or gullet ; and the passage of the food into this, constituting the act of swallowing, is termed deglutition. T]ie food, having arrived in the stomach, is acted-upon by a peculiar fluid which it contains, and much of its alimentary portion is dissolved, so that a pulpy mass is formed which is termed chyme ; hence this process, which is the first stage of digestion properly so called, is termed clujmification or the manufacture of chyme. The chyme, which passes into the intestines, is further acted-on by secretions that are poured into thorn ; and a certain nutritive combination of albuminous PREHENSION OF FOOD. 163 and fatty matters, termed chyle, is sejoarated from the matters that are to be thrown off : this process, which is the second stage of true digestion, is termed chylification. The rejected portions of the food, with secretions poured into the alimen- tary canal, find their way out through the intestinal tube ; and are voided at its terminal orifice by the act of defecation. And lastly, the nutritive materials are taken up by absorption into vessels that are distributed upon the walls of the diges- tive cavity, and undergo a gradual change, by which they are converted into blood. These two processes are called absorp- tion and sanguification (or manufacture of blood). Each of the foregoing stages will now be separately considered. Prehension of Food. 172. The introduction of aliment within the entrance to the digestive cavity is accomphshed in various methods in dif- ferent animals. In the Mammalia in general, the aperture of whose mouth is guarded by fleshy lips, these, with the jaws and teeth, are the chief instruments of this operation. But ill Man and the Monkey tribe the division of labour is carried further ; the food being laid hold of by the anterior members, or hands, and by them carried to the mouth. Where the hand has the power of grasping, and especially where the thumb can be opposed to the fingers, the action of a single member is sufficient ; but there are several animals which, lilvC the Squirrel, use both limbs conjointly to hold their food, the extremity not having itself the power of grasp- ing. The Ant-eaters, Woodpeckers, Chameleons, and other insect-eating annuals, obtain their food by means of a long extensible tongaie ; this either serving to transfix the insect, or being covered with a viscid saliva which glues it to the surface. The Giraffe uses its long tongue to lay hold of the young shoots on which it browses ; and the Elephant employs its trunk, which is nothing else than a prolonged nose, for every kind of prehension (fig. 82). Many of the Invertebrata are furnished with ^ ''^'^^ Tiji 1 ^ A^ • ill Head of Elephant. little appendages round their mouths by which the food is conveyed into th(^m ; such are the palpi m2 164 BECEPTION OF SOLID AND LIQUID ALIMENT. of Insects, of which a pair is attached to each jaw (fig. 84); the tentacula of Mollusks, which are sornetiiues extremely prolonged, as in the Cuttle-fish tribe (fig. Sb) ; and the similar organs of the polypes (fig. 71). Carabos. Fig. 84.— Jaws of the SAME Insect. 173. The reception of liquids is accomplished in two ways. Sometimes the liquid is made to fall into the mouth, simply by its own weight (fig. 8G) ; in other instances it is drawn or pumped up into this cavity, — cither by the expansion of the chest, which causes a rush of air towards the lungs, — or by the movement of the tongue, which, being drawn back like a piston, produces the action of sucking. Some of the lower animals are destined to be entirely supported by liquids which they find in plants, or which they draw from the bodies of other animals whereon they live as parasites. This is the case with many Insects ; and their mouth, instead of present- ing the ordinary structure, is formed into a sort of tube or trunk, very much extended, through which the juices arc drawn up according to the wants of the animal. Such a conformation exists in the butterfly and moth tribe, whose SUCTION OF LIQUIDS. 165 trunk, wlien not in use, is coiled up in a spiral boneath the head ; as is shown in fig. 87, representing the head of a Fig. 87.— Trunk of a Butterfly. Fig. 86. — Chimpanzee dkinking. Butterfly, or, of which the eye is seen at c, the base of the antennae at 6, the palpi at e, and the trunk at d. In some of the Fly tribe, the trunk attains a length several times greater than that of the body, as shown in fig. 88, representing a Fig. 88. — Nemestrina longirostris. dipterous (two-winged) insect from the Cape of Good Hope, which sucks the juices of a single kind of flower, the length of whose tube just equals that of its long proboscis. 16G DEVELOPMENT OF TEETH. Mastication. 174. The act oi Mastication, or the mechanical division of the ahmcntary matter, is effected in most of the higher animals, by the Teeth ; which are implanted in the jaws, and are so fixed as to act ao-ainst one another, with a cntting, crushing, _ or power, according Grinding Fig. 89. — Devklopment the gum ; h, the lower jaw ; jaw ; d, dental capsules 01' Teeti to the nature of the food on which they have to oj^erate. The manner in which they are form.ed is worthy of note. In Man, who may be taken as a fair example, each tooth is developed in the interior of a little mem- angieofthe brauous sac, wliich is lodged in the thickness of the jaw- bone ; as seen in the accompanying figure, wliich represents half the lower jaw of a very young infant, from which the outside has been removed. This sac, which is named the dental capsule («, tig. 90), is composed of two membranes, abundantly furnished with blood-vessels ; and it encloses in its interior a little bud-like protuberance, h, in which ramify a great number of nervous filaments and minute vessels, c. The matter composing this little body, which is termed the j)idp, is gradually converted into the dentine (§ 54) of the tooth, d d which in Man constitutes nearly its whole structure ; this conversion takes place first at its highest points, d, d. The crown or ui)pcr portion of the tooth receives a (•overing of enamel (§ 54). Gradually the process of conversion extends more and more to the interior of the pulp ; and at last the whole is changed into dentine, with the exception of a small portion occupying what is termed the cavity of is frequently laid open by decay of its Fig. 90.— L)KK'r.vr, Capsi LE. that still the tooth reuuiin> , which external wall. The fang of the tooth, which is the part last formed, receives an envelope of cementum (§ 54), wliich invests it up to the part at which the enamel begins. As the DEVELOPMENT OF TEETH. 167 root of the tooth is developed, the crown is gradually pushed upwards, so as to 23ress against the upj)er portion of the capsule and the gum by which this is covered. These parts yield slowly to the pressure ; and the tooth makes its way to the surface ; or, in common language, is cut. 175. The process of "cutting teeth" is usually not a severe one in the healthy and well-managed infant ; but it occasions the death of vast numbers of children who are injudiciously treated ; and it is especially fatal to those who have a ten- dency to disease of the nervous system. The irritation caused by the pressure of the tooth against the gum, is liable to excite, in such cases, convulsive actions of various kinds, on the principles hereafter to be explamed (§ 473) ; and, as the removal of the source of irritation is of the most urgent importance, the lancing of the gums, — doing that in an instant which the pressure of the tooth might not accomplish for days, — is a measure of most obvious utility ; however unnecessary it may seem, in ordinary cases, to in- terfere with the course of nature. But it is of the utmost importance at the same time to bring the nervous system into a less excitable condition ; and no measure is commonly more efficacious in this respect, than removal into a fresh and pure atmosphere. 176. At the same time that the development of the tooth is thus taking place, the bone of the jaw is becoming liardened, and closes round its root, formuig a complete socket. This partly interrupts the passage of vessels and nerves to the tooth, which, when once fully formed, seems to acquire no further growth, and to possess but little power of repairing injuries occasioned by disease or accident. Hence a tooth which is broken or decayed, is not restored as a bone would be. Still, however, its root or fang is penetrated by a small nerve and artery, which are distributed to the membrane that lines the cavity ; and it is to the action of air upon the former, when the cavity is laid open by decay, that the pain of tooth-ache is chiefly due. The remedies which are most effectual in removing this pain, such as kreosote, nitric acid, or a heated wire, are those which destroy the vital jwwer of the nerve. 177. But there are teeth, in many animals, which never cease to grow, and in which the central cavity is always filled 168 TEETH OF RODENTS. MOTION OF JAWS. with pulp. Such have no proper root ; for additional matter is being continually formed at their base, and thus the whole tooth i^ pushed upwards. This is the case with the Elephant's tusks ; and also with the large teeth that occupy the front of the iaw in Eabbits, 8(iuirrels, Rats, and other gnawing ani- mals (fig. 91). The upper edges of these teeth are being con- stantly worn away by use : and they are kept up to their proper level by the growth of the tooth Fig. 91. -Jans AND Teeth OF RAniur. from beloW. But it sometimes happens that one of these teeth is broken off ; and the one opposite to it in the other jaw is then thrown into dis- use. It continues, however, to grow up from below ; but, not being worn down at the top, its length increases greatly, so that it may l)ccome a source of great inconvenience to the animal. 178. The teeth are but passive instruments in the act of mastication. They are put in movement by the jaws in which they are fixed ; and these are made to act against each other by various muscles. The upper jaw is usually fixed to the head; and has not, therefore, any power of moving inde- pendently of it. But the lower jaw is connected with the skull by a regular joint on either side ; and is so moved by the muscles attached to it, as to cut, crush, or grind the food, according to the nature of the teeth. 179. There is considerable variety, in different animals, as to the extent of motion which the lower jaw possesses. In the purely Carnivorous quadrupeds, it has merely a hinge-like action, that of opening and shutting ; and by the sharpness of the edges of the molar teeth, it is thus rendered a powerful cutting instrument. But in the Herb-rorous animals, which have to grind or triturate their food 1 . Iween the roughened surfaces of their molars, such a limited motion would be of no avail ; and we accordingly notice, if we watch an ox or a horse whilst masticating its food, that the lower jaw has con- siderable power of motion from side to side. On the other hand, in the Ilodents, or gnawing animals furnished with MOVEMENTS OF LOWER JAW. 169 two large front teeth, tlie lower jaw has no power of moving from side to side, but is rapidly drawn backwards and for- wards ; and, as the ridges of the molar teeth are arranged in the opposite direction, they become very powerful filing in- struments, by which the toughest vegetable sulxstances are quickly reduced. 180. In the Human jaw, there is a moderate power of motion in all these different directions ; and it is furnished with all the muscles by which they are effected in the different animals that perform them ; but these are not so large or strong. The most powerful of the muscles of the lower jaw, in all animals, is that by wdiich it is drawn up against the upper, so as to close the mouth. This arises from the side of the skull in the region of the temple, and is hence called the temporal muscle. It covers at its origin a large surface of bone ; but its fibres approach one another as they descend, and pass under a bony arch (which may be felt between the cheek and the ear), to attach themselves to a process or projection of the lower jaw^ (cr, fig. 92), about an inch in front of the joint. As the distance from the ful- crum of the point a, at which the powder is applied, is thus much less than that of the front of the jaw h, where chiefly the resistance is encountered, the powder of the muscle is applied at a mechanical disadvantage ; and, to overcome a given resist- ance, the muscle must itself be several times more powerful. Thus the Tiger and Lion, which can lift and carry away the bodies of animals weighing several hundred pounds, must possess temporal muscles that shall contract wdth a force of two thousand, or even more. 181. In Man, as in most of the other Mammalia, there are three kinds of teeth, adapted for different purposes. The first terminate in a thin cutting edge, and are intended simply to divide the food introduced into the mouth ; these are termed inciso?' teeth (fig. 93). Others have more of a conical form. Fig. 92.— Human Skull. 170 DIFFERENT KINDS OP TEETH. and ill many animals (especially those of carnivoi'ous liabits) project far beyond the former ; they are adapted not to cut the food, but, by being deeply fixed in it, to enable the animal to tear it asunder : these are termed ca^iine teeth. The teeth of the third kind have large irregular flattened surfaces, and are adapted to bruise and grind the food ; these are called mola?- (or mill-like) teeth. The manner in which these different teeth are implanted in the jaw, varies with the ibrm of their crowns, and is in accordance with their several uses. The incisors, whose action tends as much to bury them in their sockets as to cb^aw them forth, have but a single root or fang of no great length. The canine teeth, on which there is often considerable strain, penetrate the jaw more deeply than the incisors ; esj)ecially when they are large and Molars. Bicuspid. Canine. Fig. 93.— Human Teeth. long, as in the Cat tribe (fig. 94). And the molars, whose action requires great firmness, have two, three, or even four roots or fangs, which spread out from each other ; and these at the same time increase the solidity of their attachment to the jaw, and prevent the teeth from being forced into their sockets by any amount of pressure. 182. The arrangement of the dental apparatus varies, in different Mammalia, accordmg to the nature of the aliment on which they are destined to feed ; and this correspondence is so exact, that the anatomist can generally determine by the simple inspection of the teeth of an animal, not only the nature of its food, but the general structure of the body, and even its ordinary habits. Thus, in those that feed exclusively on annual lU'sh, the molar teeth are so compressed as to form DIFFERENT KINDS OF TEETH. 171 cutting edges, which work against each other Hke the blades of a pair of scissors (fig. 94) ; whilst in animals that live on insects, these teeth are raised into conical points, which lock Fig. 94. — Teeth of Carnivorous Animai Fig. 05. Teeth of Insectivorous Animal. into corresponding depressions in the teeth of the opposite jaw (fig. 95), When the nourishment of the animal con- sists principally of soft fruits, these teeth are simply raised into rounded elevations (figs. 97, 98) ; and when they are Fig. 06. Teeth of Herbivorous Animal. Fig. 97.— Teeth of Frugivorous Animal. destined to grind harder vegetable substances, they are termi- nated by a large flat and roughened surface (figs. 96, 99). The roughness of this surface is maintained by the peculiar arrangement of the three substances of which the tooth is composed. The enamel, instead of covering its crown, is arranged in upright plates, which are dispersed through the tooth ; and the space between them is filled up by plates of ivory and of cementum (§ 54). These last, being softer than the enamel, are worn down the soonest ; and thus the plates of enamel are left constantly projecting, so as to form a rough surface admirably adapted to the giinding action which the tooth is destined to perform. The mode in which these plates are disposed, afibrds a most characteristic distinction between the two species of Ele2:)hant at present existing, DIFFERENT KINDS OF TEETH. namely, the African and tlie Indian ; as also between each of these and the great extinct species known as the Mam- moth (tig, 99). In the great gnawing teeth of the Eabbit, Fig. 98. — Molar Tooth of Mastodon. Fig. 99.— Molar Teeth of Elephants. 1 , African Elephant ; 2, 1 ndian Elephant ; 3, Mammoth. &c., the front surfoce only is covered with enamel ; and as this is worn away more slowly than the ivory, it stands uj) as a sharp edge (fig. 91), which is always retained, however much the tooth may be worn away. 183. Of all the teeth, the molars may be regarded as the most useful. They are seldom absent in the Mammalia ; and their office is usually essential to the proper digestion of the food. Animal flesh (the most easily digested of all substances) needs but to be cut in small pieces ; but the hard envelopes of beetles and other insects must be broken up ; and the tough woody structure of the grasses, and the dense coverings ^ /Ji^^^^^-xjiXMSb^r^^" J *^'^ ^^^^ seeds and fruits on which V^V^iJ^-^^-^^'-^^^^^^'/^ ^^^^ herbivorous animals are supported, must be ground down. The incisors and canines are chiefly employed among purpose of seizing their living prey, and are never delicient in them ; but they are less re- quired in Herbivorous animals ; and either or both kinds are not uufro(piently dcificient. Sometimes, however, they are not Fig. 100. —Skull of Boar. Carnivorous animals for the SUCCESSION OF TEETH. WHALEBONE. 173 only present in the latter, but are largely developed, serving as weapons of attack and defence ; as in the Boar (fig. 100). 184. In the Mammalia in general, as in Man, the teeth are not much developed at the time of birth, that they may not interfere with the act of sucking; and they do not make theii- appearance above the gum, until the time approaches when the young animal has to prepare its own food, instead of smiply receiving that wdiich has been prepared by its parent. The teeth which are first formed are destined to be shed after a certain period, and to be replaced by others. They are called milk-teeth ; and in Man they are twenty in number, — namely, four incisors in the front of each jaw, and two canines and four molars on each side. These begin to fall out at about the age of seven years ; previously to which, however, the first of the permanent molars appears above the gum, behind those of the first set. The incisors and canines of the first set are replaced by incisors and canines respec- tively ; but the molars of the first set are replaced by teeth like small molars, having only two fangs ; these are called false molars, or, more properly, hicuspid teeth (fig. 93). The second of the true molars does not make its appearance until all the milk-teeth have been shed ; since it is only then that the jaw becomes long enough to hold any additional teeth. The third does not usually come up until the growth of the jaw is completed ; and as this time corresponds w-ith that at wdiich the mind as well as the body is matured, they are commonly known as wise or wisdom teeth. There are then thirty- two teeth in all, or sixteen in each jaw ; — namely, four incisors, two canines, four bicuspid, and six true molars. — In extreme old age, these teeth fall out like those of the first set ; but they are not replaced by others, and their sockets are gradually obliterated. 185. There are a few Mammalia which do not possess teeth. This is the case with the common A¥hale, in which they are replaced by an entirely different structure. From the upper jaw (fig. 102) there hang down into the mouth a number of plates of a fibrous substance (fig. 101), to which we give the name of whaleho7ie, though it is really analogous to the gum of other animals. The fibres of these plates are separate at their free extremities, and are matted (as it were) together, so as to form a kind of sieve. Through this sieve the "Whale 174 ABSENCE OF TEETH IN WHALE, ANT-EATER, ETC. draws water in enormous quantities, whenever it is in want of food ; and in this manner it strains out, as it w^ere, the minute gelatinous animals upon wdiich it lives, from the w^ater of the seas it inhabits. The water thus taken in is expelled from the nostrils or blow-holes, which are situated at the top Fig. 102.— Skull of Whale, Fig. 101. — WnALKlioNE. of the head. ^Most of the Whale tribe have short fringes of this kind in the roof of the mouth ; but in none, except the Balosna, or Greenland Whale, is it long enough to make it w^orth separating ; all the other sjDecies havmg teeth, either in one or both jaws. — It is a curious fact, that the rudiments of teeth may be discovered in both jaws of the young Greenland whale, although they are never to be developed. And the rudiments of incisor teeth in the upper jaw, and of canine teeth in both jaws, may also be discovered in the young of the Euminant quadrupeds (oxen, sheep, &c.), though they never show themselves above the gum. 18G. The Ant-eaters, also, are destitute of teeth, and usually '»btain their food by means of their long extensible tongues, which are covered with a viscid saliva ; this being pushed into the midst of an ant-hill, and then drawn into the mouth, brings into it a large number of Fig. 103.— Skill oi- the Ant-eater, these insects, which are sufficiently bruised between the toothless jaws (fig, 103). Lastly, may be mentioned as a curious exception to the general rules respecting the teeth of Mammalia, the remarkable Orni- thorhyncus of New Holland (Zoology, § 317), which feeds, TEETH OF RErTILES AND FISHES. 17^ like the duck, upon the water-insects, shell-fish, and aquatic plants, that it obtains from the mud, into which it is continu- ally plunging its singular bill ; and its jaws, entirely destitute of teeth, are furnished with horny ridges, by which it can in some degree masticate its food. 187. Among Birds, there is an entire absence of teeth; and the mechanical division and the reduction of food is per- formed in the stomach, in the manner hereafter to be men- tioned (§ 200). The mouths of almost all Reptiles, excepting the Turtle tribe, are furnished with numerous teeth (fig. 104) ; but these are not adapted for much variety of purposes, being prmcipally destined to prevent the escape of the prey which the animals have secured ; and their shape is conse- quently nearly uniform, being for the most part simply conical. There are some Lizards, however, which are herbivo- rous ; and these have large rough teeth, somewhat resembling the molars of Mammalia. The Igiianodon, an animal of this tribe, attained a gagantic size in past ages of the world. 188. In Fishes, the teeth are commonly very numerous (fig. 105), but they have for their object only to separate and retain ■Head of Gavi of the Ganges.) (Crocodile Fig. 105.— Head of Shark. their food ; and there is little variety in their form. Fre- quently they have no bony attachment, being only held by the gum, as in the Shark ; and they are consequently often torn away, but they are as readily replaced. Sometimes, how- 17G MASTICATING INSTRUMENTS OF INVERTEBRATA. ever, the tooth seems like a coutiiiiiation of the ])one of the jaw, not being in any way separated from it, and the tubular structure of the latter being continued into it without any interruption. The teeth of fishes are often set, not only upon the proper jaw-bones, but upon the surface of the palate, and even in the 'pharynx or swallow. 189. In the Invertebrata there are generally no proper teeth ; in the Articulated and sometimes in the Molluscous series, however, we meet with firm horny jaws, which are often furnished with projections that answer the same pur- pose ; and in most Gasteropods we find a very curious organ, commonly designated as the tongue^ more correctly the imlate, the surface of which is beset with innumerable tootli- like points (fig. 106), by whose rasping action the food is reduced. These teeth present great varieties of form and arrangement in the different genera andsjDecies of this group ; and these varieties appear to bear some relation to the nature of the food on which the animals respectively live. It is remark- able that in an animal so low in the scale as the Echinus or Sea- Urchin (§ 1 19), a very com- plex dental apparatus should exist. This consists of five long hard teeth, which surround the mouth ; and these are fixed in p. ,„, ,. ^ ,, a framework which is worked Fig.ioe.—DKNTAL Organ OF Nerixa. ■, ^ ^ , i^ ^ by a power! ul set oi muscles, and thus serve effectually to grind down the food. Insalivation. 190. The act of mastication is connected with another; which is also of great importance in preparing for the sub- sequent process of digestion. This is the blending of the sahva with the food, during its reduction between the teeth, —an act which is termed inmUmtion. The saliva is separated Irom the blood, by glands which arc situated in the neigh- SECRETION OP SALIVA, AND ITS USES. 177 l30iirliood of the mouth ; of these there are three pair in Man, two beneath the tongue (fig. 107), and one in the cheek, each pouring-in its secretion by a separate canal. The salivary fluid is prmcipally composed of water, in which a small quantity of animal matter and some saline substances (chiefly common salt) are dissolved ; the whole amount of these, however, is not more than 1 part in 100. The secretion of saliva is not constantly going on ; but the fluid is formed as it is wanted. The stimvlus by which the gland is set in action may be simply the motion of the jaws ; thus, on first waking in the morning, the mouth is usually dry, but it is soon rendered moist by the movements which take place in speaking. The contact of solid substances with the membrane lining the mouth appears also to excite the How ; hence dryness of the mouth may often be remedied for a time, when no water is at hand, by taking a pebble into its interior, and moving this from side to side. There are certain substances, however, whose presence in the mouth has a special influence in provoking an increased secretion of saliva ; and every one knows, too, that the simple idea of savoury food will excite an increased flow, making the "mouth water" as it is popularly termed. These are instances of the power of the nervous system, through which such impressions are conveyed, over the act of secretion. 191. In the case of farinaceous or starchy food, the admix- ture of saliva occasions the commencement of that chemical change in which its digestion consists, namely, its conversion into sugar ; but in general, the benefit derived from this pro- cess of insalivation is just that which is obtained by the chemist, when he bruises in a mortar, with a small quantity of fluid, the substances he is about to dissolve in a larger amount of the same. If the preliminary operations of masti- cation and insalivation be neglected, the stomach has to do the whole of the work of preparation, as well as to accomplish the digestion ; thus more is thrown upon it than it is adapted to bear ; it becomes over- worked, and manifests its fatigue by not being able to discharge even its own proper duty. Thus the digestive function is seriously impaired, and the general health becomes deranged in consequence. A malady of this kind is very prevalent in the United States ; and is almost universally attributed by medical B'^en, in part at least, to the N DEGLUTITION OR SWALLOWING. general habit of very rapidly eating or rather " bolting " the meals. There is another evil attendant on this practice, — that much more food is swallowed than is necessary to supply the wants of the system ; for the sense of hunger is not so readily abated by food which has not been prepared for digestion ; and thus the feehng of satiety is not produced, until the stomach has already received a larger supply than it is well able to dispose of. Imperfect mastication of the food is very apt to occur, in persons who are losing their teeth by old age or decay; and where these are not replaced by artificial means, the next best remedy is to cut the food into very small por- tions, before it is taken into the mouth, and to masticate it there as thoroughly as possible. Deglutition. 192. In the Mammalia, the cavity of the mouth is guarded behind by a sort of moveable curtain, which is known as the veil of the palate (fig. 107) ; and this hangs down during Veil of the palate Nose Pharynx (Esophagus Tongue Salivary glands Os hyoides Larynx Thyroid gland Trachea Fig. 107.-PKnPKNDICULAR SeCTIOK of TlIK MoUTH AND THROAT. mastication, in such a manner as to prevent any of the food trom pas.-iiig backwards. Tliis partition, which does not exist DEGLUTITION OR SWALLOWING. 179 in Birds and other animals that do not masticate their food, hangs from the arch and sides of the paUite, so as to touch the tongue by its lower border ; but it can be lifted in such a manner as to give the food free passage beneath it, into the top of the gullet. When mastication is completed, the food is collected on the back of the tongue into a kind of ball ; and this, bemg carried backwards by the action of its muscles, presses against the partition just mentioned, and causes it to o^eii. The food thus passes into a sort of funnel, formed by the expansion of the top of the oesophagus or gullet ; this cavity, termed the phari/nx, communicates above with the nostrils, and in front with the larynx, which is at the top of the trachea or windpipe. The oesophagus is a long and narrow tube, which descends from the pharynx to the stomach, lying just in front of the vertebral column, and behind the heart and lungs. It is surrounded by muscular fibres, disposed in various ways ; by the action of which the food that has once passed into the pharynx is propelled downwards to the stomach. 193. But in order to reach this tube, the alimentary ball must pass over the glottis or aperture of the larynx. With a view to prevent its falling-in, the larynx is drawn, in the very act of swallowing, beneath the base of the tongue ; and this action presses down a little valve-like flap, the epiglottis, upon the aperture, so as in general effectually to prevent any solid or fluid particles from entermg it. But it sometimes happens that, if the breath be drawn-in at the moment ot swallomng, a small particle of the food, or a drojD of fluid, is drawn into the glottis ; and this action (commonly termed " passing the wrong way,") excites a violent coughing, the object of which is to drive up the particle, and to prevent it from finding its way into the lower part of the windpipe. It may also happen that a larger substance may slip backwards, by its own weight, into the glottis, when there was no intention of swallowing, and when the larynx was conse- quently not drawn forwards beneath the tongue. The presence of such a substance in the windpipe excites a violent and fre- quently ahnost suffocating cough (§ 342) ; the effect of which is sometimes to drive it up through the glottis, and thus to get rid of the source of irritation. 194. The act of swallowing is itself involuntary, and may n2 180 MOVEMENTS OF DEGLUTITION. be even made to take place against the will. ^ This may seem contrary to every one's daily experience ; but it is nevertheless true. The movement by which the food is carried back, beneath the arch of the palate, into the pharynx, is effected by the will ; but when the food has arrived there, it is laid hold of, as it were, by the muscles of the pharynx, and is then carried down involuntarily. It has several times happened, that a feather, with which the back of the mouth was being tickled to excite vomiting, having been introduced rather too far, has been thus grasped, by the pharynx, and has been swallowed. Moreover, we cannot perform the act of SAvallowing, without carrying something backwards upon the tongue ; and it is the contact of this something, even if it be only a little saliva, A\'ith the membrane lining the pharynx, that produces the muscular movement in question. 195. This action is one of the kind now denominated rejlex (§ 430). It is produced through the nervous system ; for if tlie nerves supplying the part be divided, it will not take place. But it does not depend upon the Brain ; for it may be performed after the brain has been removed, or when its power has been destroyed by a blow. It is caused by the conveyance to the top of the Spinal Cord, of the impression made on the lining of the pharynx ; this impression, brought thither through one set of nerves, excites in the spinal cord a motor impulse ; which, being transmitted thence through another set of nerves, calls the muscles into action. 196. This action is, therefore, necessarily connected with tlie impression, so long as this portion of the spinal cord, and the nerves proceeding from it, are capable of performing their functions : and it is one of those to which we may give the name of instinctive, to distinguish it from those which are eifected by an effort of the Will, intentionally directed to accomplish a certain purpose. It may even take place without the animal being aware of the contact of any substance to be swallowed with the lining of the pharynx ; for there is good reason to believe that when the brain has been destroyed, or ])aralyzed by a blow, all sensibility is destroyed ; and we have also sufficient reason to consider it as suspended in profound sleep or apoplexy, in which states swallowing is still per- formed. In the severest cases of apoplexy, however, the power of swallowing is lost ; and this is a symptom of gi-eat DEGLUTITION DIGESTIVE APPAllATUS. 181 danger, since it shows that not the brain alone, but the ujiper part of the spinal cord, is suffering from the pressure ; and that the movements of respiration, which depend upon a sunilar action of the nervous system (§ 340), will probably- soon cease, so that death must ensue. Digestive AppcnxUiis. 197. Tlie food, thus propelled downwards by the action of the muscles of the pharynx and of the esophagus (gullet), Gall-Bladder ■ Large Intestine Spleen Colon - Small Intestine • Colon Small Intestine Rectum Fig. 108.— Digestive AppAnAxus of Man. arrives, in Man and the Mammalia, at the stomach ; which is a large membranous bag, placed across the upper part of the 232 FORM OF THE STOMACH. abdomen (fig. 108). The form of this stomach varies much, according to the nature of the ahmcnt to be digested. Wliere tlie food is animal flesh, which is easily dissolved, the stomach is small, and appears like a mere enlargement of the alimentary tube ; this is the case in the Cat tribe, for example. In Her- bivorous animals, on the contrary, the stomach is very large, the food being delayed there a long time on account of the difficulty Avith which it is digested ; and the principal part of its cavity is not a simple enlargement of the alimentary tube, but a bag or sac that bulges out, as it were, on the left side of that canal. By the degree of this bulging, we can judge of the nature of the food on which the animal is destined to live. Thus in Man (fig. 108), the large end of the stomach, situated on tlie left side (the y^iglit side of the figure as w^e look at it), is moderately developed ; showing, as we might expect from the form of his teeth, as w^ell as from his natural tastes, that he is adapted for a diet in which animal and vegetable food are mixed. In the purely carnivorous tribes, this large end of the stomach is almost deficient ; whilst in the herbivorous races, it is enormously developed, and some- times forms a distinct pouch. (Esophagus Intestine Pylorus 4tliStom. 2d Stoni. 1st Stem. Fig. 109.— Stomachs of the Siif.ep. 108. The most eom[)lcx form of the stomach among Mam- mals, is that which we find in the animals that ruminate or chew the cud. It possesses, in fact, no less tlian four distinct cavities, through all of Avhich the food has to pass during the STOMACH OF RUMINANTS. 183 process of digestion. The external appearance of the stomach of the Sheep is seen in fig. 109 ; and its interior is displayed in fig. 110. The food of the Euminant animals is not chewed by them before it is first swallowed. In their wild state, they are pecuKarly exposed to the attacks of their car- nivorous enemies, w^hen they come down from their rocky heights to browse upon the rich pastures of the valleys. If they were then obliged to masticate every mouthful, they would be subjected to long-continued danger at every meal ; but, by the curious construction of the digestive apparatus, this is spared to them ; for they are enabled to swallow their food as fast as they can crop it, and afterwards to return it to their mouths, so as to masticate it at their leisure, when they have retreated to a place of safety. The crude unmasticated food, wliich is brought-down by the oesophagus, first enters the large cavity on the left side, which is commonly termed the paunch. It is there soaked, as it were, in the fluid secreted Intestine Honeycomb Paunch Fig. 110. — Section of the Stomachs of the Shf.ef. by its walls ; and is then transmitted to the second cavity, which, from the sort of network produced by the irregular folding of its lining membrane, is called the reticulum or honey-comb stomach. This stomach also has a direct commu- nication with the a^sophagus, and appears destined especially to receive the fluid that is swallowed ; for this passes im- mediately into it, without going into the first stomach at all. The folds of its lining membrane present a large surface, through which fluid may be absorbed into the system. It is 134 ACT OF RUMINATION. here that we find the curious arrangement of water-cells in the stomach of the Camel, by which that animal is enabled to retain a supi^ly of water for several days. These cells corre- spond with the little pits which are seen in the honey-comb stomach of the Sheep, but are much deeper, and their orifices may be closed by the action of a set of muscular fibres wdiicli pass in every direction round each, so as to form a net- work including these orifices in its meshes. 199. After the food has been macerated in the fluids of the lirst and second stomachs, it is returned to the mouth by a reversed peristaltic action of the oesophagus, which brings it up as a succession of globular pellets, that are formed by compression in a sort of mould at the lower end of the oeso- l)hagus. These pellets are subjected within the mouth to mastication and insalivation ; and the food is then ready for the real process of digestion. It is this mastication which is commonly known as the " chewing of the cud ; " and the animal, whilst performing it, seems the very picture of placid enjoyment. When again swallowed, the food is directed, by a pecuhar valvular groove at the bottom of the oesophagus, into the third stomach, commonly termed the manyplies, from the pecuhar manner in which its lining membrane is arranged. This presents a number of folds, lying nearly close to one another, like the leaves of a book, but all directed, by their free edges, towards the centre of the tube, — a narrow fold intervening between each pair of broad ones. The food has, therefore, to pass over a large surface, before it can reach the outlet of the cavity ; and this leads to the fourth stomach, commonly termed the reed. This is the seat of the true digestive process,^ the gastric juice (§ 204) being formed here only ; and it is from this that the rennet is taken, which is used in making clieese to cause the milk to coagulate or curdle. In tlie sucking animal, the milk passes directly into this fourth stomach, without entering either the first or second stomachs, and without being delayed in the third, the folds of which adhere together so as to form a narrow undi^•ided tube. The paunch is at that time comparatively small, being of less size than the reed ; and its dimensions increase, as soon as the young animal begins to distend it by swallowing solid vegetable matter. 200 lu the digestive apparatus of P.irds, we find a con- DIGESTIVE APPARATUS OF BIRDS. 185 siderable modification of form, resulting from the fact that, as these animals do not masticate their food, they require somQ (Esophagus Ventriculus\ Succenturiatusi Gizzard Pancreas Duodenum Liver Gall-bladder 13ile ducts Cceca Large Intestine Ureter Oviduct Cloaca Fig. 111. — Digestive Apparatus of Fowl. other means of reducing it. lliis means is provided for them in their stomach. In the tribes whose food is of such a nature as to require being moistened before it is rubbed down, and especially in those which feed upon grains, the oesopha- gus has a pouch-like dilatation, termed the crop or craw (iig. HI) ; in iliis it is retained, and exposed to the action 286 TRITURATING ACTION OF GIZZARD. of fluid secreted by its walls, just as it is in the pauncli of ruminant quadrupeds. This crop is of enormous size in some of the granivorous (grain-eating) birds, such^ as the Turkey. The second stomach (or ventriculus succenturiatus) is the one in which the gastric juice is secreted ; but this is seldom large enough to retain the food, which passes-on through it to the gizzard, a hollow muscle, furnished with a hard tendi- nous lining. In the granivorous birds this is extremely strong and thick ; and pieces of gravel are swallowed by them, which, being worked-up with the food by the action of the gizzard, assist in its reduction. In the rapacious flesh- or flsh-eating birds, however, no such assistance is required, the food being easy of solution ; the walls of their gizzard are thin, possessing but few tendinous fibres ; and the three cavities of the stomach are almost united into one. 201. Various experiments have been made to test the mechanical powers of the gizzard of Birds. Balls of glass which they were made to swallow with their food, were soon ground to powder ; and the points of needles and of lancets, fixed in a ball of lead, were blunted and broken-off by the power of the gizzard, whilst its own internal coat did not appear to be in the least injured. On the other hand it has been ascertained, that grain enclosed in metal balls which protected it from the mechanical action of the gizzard, but which were perforated so as to afford the gastric fluid free access to their contents, was not in the least digested ; so that the utility, and even the necessity of this operation, become evident. 202. As there are few animals, save the Mammalia, that perform any proper masticaton in their mouths, the grinding down of their food (where it is of such a nature as to require it) must bo performed in the stomach ; and accordingly we find many tribes, belonging to different divisions of the animal kingdom, in which a gizzard, or something analogous to it, exists. It is possessed by almost all Cephalopods, and by many of the Gasteropoda. In the walls of the stomach of some of these last, there is a considerable amount of mineral matter deposited, intermixed with the hard tendinous fibres of which they chiefly consist. A powerful gizzard is also found in many Insects, but here it is placed above the diges- tive stomach (fig. 112, c). The accompanying figure exhibits 187 the alimentary canal of a Beetle, from its commencement to its termination. At a is seen the head, bearing the jaws, &c. ; from this the gullet passes straight backwards, and is dilated into a crop at 6, below which is the gizzard, c. This opens at its lower end into the true digestive stomach, d; which is surrounded by an immense number of little follicles or bags, by which the secretion of the gasUic juice is effected (§ 204). Into the lower end of this, the long vessels, e, open, which constitute in Insects ths only rudiment of a liver (§ 358). In many of the Crustacea, the walls of the stomach are beset with re- gular rows of teeth, which are moved by the action of powerful muscles. These teeth are cast or shed at the same time with the shell. In the Wheel-Aniinalcules, the place of the gizzard is occupied by a curious pair of jaws, armed with teeth ; by the working of which, the food is effectually crushed. In the Bryozoa, a gizzard exists between the oesopha- gus and the true digestive stomach ; and the stomach itself is surrounded by the little follicles which secrete the bile, and pour it into that cavity (§ 115). 203. In animals which subsist exclusively on flesh, how- ever, no such complicated apparatus exists. Thus in Serpents (fig. 34), the stomach is but a slight dilatation of the alimen- 112.— Digestive Apparatus of Beetle. 188 DIGESTIVE APPAEATUS — GASTRIC DIGESTION. tary tube ; and it is not easy to say where it commences and terminates. In Spiders and Scorpions, too, which live upon the juices they suck from other animals, the alimentary tube is very simple ; and it is scarcely dilated into a proper sto- mach. And in most of the Eadiated classes, we find the stomach to possess only one orifice, through which the undi- gested residue of the food is cast out, as well as fresh sup- plies taken in. But this stomach is not always a simple bag ; thus in the Star-fish it sends prolongations into the rays, the use of which is at present un- determined. There are certain animals in which no digestive cavity exists : their sustenance being derived either from the juices jDrepared by other animals, in whose tissues or cavities they are im- bedded, and being introduced by absorp- tion through the whole surface, as is the case in the lower Entozoa (fig. 53) ; or from particles which are drawn into the midst of the soft gelatinous substance of their bodies, and undergo a sort of diges tion there, as is the case with the lihko- poda (§ 129). Gastric Blgestion : — Chymification. 204. The food which has been re- chtred. in the mouth by the action of the teeth, or in the stomach itself by the movement of its own tendinous walls, is prepared for the real process of digestion; by which it is converted into a fluid, and thus made fit to be truly received into the system, by being absorbed into its vessels. The chief agent in the digestive process is a fluid termed the gastric juice, which is secreted or sepa- 38 seen in a vL-rticai section rated from the blood bv a vast number of thcStoniacli, maRnified , !• ^•ll^ \ o ^^• i /n t i .->\ • three diameters at a, and ^^ ^itflc bagS Or lolllcleS (fig. 113), IIU- tventy diameters at 1). bcddcd in thc walls of the stomacli. When the cavity is empty, this fluid is secreted in very small quantities ; but, like the sahvary secretion, it is poured out Fig. 113. Gastkic Follicles SENSE OF HUNGER — SECRETION OF GASTRIC JUICE. 189 in abundance when the lining membrane is stimulated by the contact of food, especially solid food. Only a limited (quantity is secreted at any one time ; and this quantity is just that which is sufficient to dissolve food enough for the supply of the natural wants of the system. The contact of any solid substances with the interior of the stomach, is suffi- cient to produce a flow of this fluid into its cavity ; but the secretion soon ceases if the substance be not of an alimentary nature. 205. The sense of hunger appears due to the distension of the blood-vessels of the stomach, which takes place in pre- paration for the secretion of the gastric fluid. This deter- mination of blood towards the stomach seems to occur when- ever the body needs a fresh supply of nourishment ; and it ceases as soon as a sufficient amount of gastric fluid has been drawn off. Hence it is, that hunger is relieved by eating ; and hence it is, also, that hunger is for a time relieved by taking solid substances into the stomach, even though they contain no nourishing matter. It is from having experienced this, that savage nations are in the habit of mixing indiges- tible solid matter with the fluids that sometimes constitute their principal articles of food. Thus the Kamschatdales mix earth or saw-dust with the train-oil on which alone they are frequently reduced to live ; and the Yeddahs, or wild hunters of Ceylon, mix the pounded fibres of soft or decayed wood with the honey on which they feed when meat is not to be had. One of them being asked the reason of the practice, replied, " I cannot tell you, but I know that the belly must be filled." It has been found by experunent, that soups and other forms of liquid aliment are not alone fit for the support of the system, even though they may contain a large amount of nutritious matter ; and the medical man well knows, that many persons have stomachs too weak and irritable to retain "slops" (as they are commonly termed), who can yet digest solid food of a simple kind. All these instances show, that the contact of a solid substance with the walls of the stomach, is the proper stimulus or excitement to the secretion of the gastric fluid. 206. This fluid, when poured upon the food, is thoroughly mixed-up with it by a peculiar movement of the walls of the stomach, which is continually bringing fresh portions of the 190 TROPERTIES OF GASTRIC JUICE. alimentary mass into contact with its sides, so that the whole is after a time equally exi^osed to the influence of the gastric secretion. If this movement Avere not to take ^^lace, only the outside of the mass would be digested, and the central portion would remain but httle affected. 207. The nature of the gastric fluid, and the mode of its operation upon the food, have been studied by withdrawing a portion of it from the stomach, and by observing its pro- perties and actions out of the body. A sufficient quantity for this purpose cannot be easily procured. Spallanzani, an Italian physiologist of the last century, contrived to obtain it, by causing birds and other animals to swallow sponges to which pieces of thread were attached j these, when they had remained long enough in the stomach to cause a secretion of the gastric juice, were drawn up again ; and the fluid they had absorbed was pressed out into vessels, in which its pro- perties could be examined. More recently, however, an advantageous opportunity has presented itself for obtaining supplies of gastric fluid in a less objectionable manner. A young man, named Alexis St. Martin, received a very severe wound in his left side, by the bursting of a gun ; and al- though this wound laid open the cavity of his stomach, he recovered his health completely, and subsequently married and had a family. There remained, however, an aperture in his stomach, which would not close up ; and through this orifice, which was usually covered by a bandage, the contents of the stomach could be drawn out. The gastric juice was obtained by introducing an India-rubber tube into the sto- mach when it was empty, and by moving it about within the (cavity ; the contact of the tube then excited the folHcles to .secretion (on the principle already mentioned, § 204) ; and the fluid thus poured into the stomach was drawn off through the tube. 208. The Gastric Juice is very like saliva in its appearance, Imt it is distinctly acid to the taste ; and it is found, by chemical examination, to contain a considerable quantity of muriatic acid '^ in an uncombined state. Besides this, it con- tains a considerable quantity of a peculiar animal substance which seems like altered albimien, and which has been desig- nated pepsin; as well as other ingredients of less importance. * Muriatic acid is commonly known as spirit of salt. ACTION OF GASTRIC JUICE. 191 This fluid possesses the power of dissolving albuminous sub- stances of various kinds, when these are submitted to its action at the constant temperature of 100° (which is about that of the stomach), and are frequently shaken-up with it. The solution appears to be in all resjoects as jDcrfect as that which naturally takes place in the stomach, but requires a longer time. It does not seem, however, that the gastric juice has a special solvent power for any other than albummous substances. Gelatinous and saccharine matters are taken-up by it, as by other watery fluids ; but neither starchy nor oleaginous substances undergo any other change by its action, than consists in the separation of their particles by the solu- tion of the membranes and fibres which held them together. There is every reason to believe that what is true of artificial is true of natural digestion ; and that so far from the whole operation being performed in the stomach, as was formerly supposed, gastric digestion is limited to the solution of the albuminous, gelatinous, and saccharme constituents of the food. 209. With regard to the precise mode in which the gastric fluid acts in dissolving albuminous substances, there is yet some uncertainty ; although there can be no longer any rea- sonable doubt, that the operation is of a purely chemical nature. An artificial gastric fluid, capable of eflecting all that can be done by that which is secreted in the liviaig stomach, may be made, by macerating (or soaking) a portion of the membrane lining the stomach of a pig, or of the fourth stomach of a calf (even after it has been washed and dried) in water, which dissolves a portion of the pepsin ; and by then acidulating this solution with muriatic or acetic acid. It has been proved that both the acid and the pepsin are essential to the process of solution ; for the acidulated fluid without the animal matter acts extremely slowly upon pieces of meat, hard-boiled qq^, &c., submitted to it ; and water in which the stomach has been macerated, but which contains no acid, will not act at all. But the acidulated water alone will readily dissolve the substances just mentioned, at a higher temperature ; and thus it appears that the acid is the real sol- vent ; and that the pepsin has for its oflice to produce some change in the albuminous substances, by whicli they are more readily dissolved, The recent inquiries of Liebig and other 192 GASTRIC DIGESTION : CHTMIFICATION. Chemists, render it probable that this change is of the nature of fermentation. 210. It is a fact of gi-eat practical importance, that a cer- tain quantity of the gastric fluid can act only upon a limited amount of alimentary matter ; so that, if more food be taken into the stomach than the gastric fluid can dissolve, it remains there undigested. Isow it has been already mentioned, that the quantity of the gastric fluid secreted at any one time, is proportional, not to the amount of food in the stomach, but to the wants of the system ; so that, if more food be swal- lowed than is required to repair the waste of the body, it Ues for some time unchanged in the stomach, and becomes a source of irritation which prevents the due discharge of its functions ; and the evil goes on increasing with every addi- tion to the contents of the cavity. This may not be felt by the individual at the time ; but it leaves permanent effects, which manifest themselves sooner or later in derangement of the general health. The habit of taking more food than is really necessary, and of irritating the stomach by stimulating substances or fluids (such as pepper, mustard, spirits, &c.), is a fertile source of disease. The injurious effects of these are manifested by the thirst which is the consequence of their use, and which is a call (as it were) on the part of the stomach, to prevent their irritating action by diluting them with water. 211. By the solution of its albuminous portion, and the separation of its other component particles, the food is re- duced in the stomach to a kind of pulp, which is termed chyme. The consistence of this will of course vary accord- ing to the nature of the food, and the quantity of fluid in the stomach ; but in general it is grayish, semi-fluid, and uniform tliroughout. When the food has been of a rich character, the aspect of the chyme resembles that of cream ; but when the food has consisted of farinaceous substances (rice, potatoes, &c.), the chyme is more like gruel. At the point where the stomach opens into the intestinal canal, which is called the 'pylorm, there is a kind of valve, which permits the chyme to pass as fost as it is formed, but closes against the portions of the food which are yet solid and undigested ; and thus the chyme escapes from the stomach in successive waves, slowly at iirst, but afterwards more rapidly, as the digestive process approaches its completion. INTESTINAL DIGESTION. 193 Intestinal Digestion; Chylification. 212. The process of digestion is by no means completed in the stomach ; for much of the matter which escapes from it in the chyme, is destined to undergo a further cliange whilst "passing through the intestinal canal ; especially in the her- bivorous tribes, whose food, being less digestible than that of the carnivorous races, requires to be longer delayed in the intestinal canal, in order that it may yield up its nutritious portion. Hence we find this canal of enormous extent in most animals whose food is vegetable, being in the Sheep about twenty-eight times the length of the body ; in the purely carnivorous animals, on the other hand, it is compara- tively short, being in the Lion only about three tunes the length of the body, while in the Serpent it runs almost straight from one extremity to the other ; and in animals which hve on a mixed diet, it is of medium length, being in Man about six times as long as his body. The intes- tinal tube is usually distinguished into the small and the large intestine ; of which the small is the first portion, and the large the second. The former, as shown in fig. 108, is disposed in a convoluted or twisted manner, so that a great extent of it may be packed within a small compass ; it usually forms about three-fourths of the whole length of the canal. It is held in its place by a serous membrane termed the peritoneum, which forms an immense number of folds that suspend it (as it were) from the vertebral column ; but these stiU allow it a considerable power of movement. 213. Soon after passing from the stomach into .the intes- tinal canal, the food is mingled with three secretions, which have an important influence on the changes it is further to undergo ; these are the Bile, the Pancreatic fluid, and the In- testinal juice. The two former are prepared by two large glan- dular masses, the Liver and the Pancreas (or sweetbread), which, in all the higher animals, are completely detached from the alimentary canal, and send their secretions into it through special ducts ; the latter, like the gastric juice, is formed in little follicles lodged in the wall of the canal itself. The peculiar matter which forms the chief solid constituent of hile, is essentially a soap formed by the union of two resinoid acids, with soda as a base (§ 364). The composition of the o 194 BILIARY, PANCREATIC, AND INTESTINAL SECRETIONS. pancreatic fluid closely corresponds with that of saliva, which it much resembles in appearance. The intestinal juice, like the gastric, is a nearly colourless, somewhat viscid fluid, con- tainilig an organic compound not far removed from albumen ; but it°differs from the gastric juice in being alkaline instead of acid. The relative offices of these three fluids have not yet been determined with certainty ; but there appears good reason to believe : (1) that the bile, by its alkalinity, neutralizes the acidity which the chyme derives from the gastric juice, and that this neutralization favours the metamorphosis of starch into sugar, which has been almost suspended in the stomach ; (2) that the bile aids the pancreatic fluid in re- ducing the oleaginous particles to the condition of an emul- sion, that is, in bringing them into a state of very minute division, in wliich tliey remain suspended in the albuminous solution ; (3) that the pancreatic fluid aids the salivary mat- ter which was swallowed with the food, in the transforma- tion of starch into sugar ; (4) that the intestinal juice has a solvent power for albuminous substances which is scarcely inferior to that of the gastric juice, with a power of converting starch into sugar which is scarcely inferior to that of saliva or pancreatic fluid. The fluid of the Small Intestine, com- pounded of the salivary, gastric, intestinal, biliary, and pan- creatic secretions, appears to possess a far greater digestive power than that of the stomach, being capable of dissolving, or at any rate of reducing to an absorbable condition, nutri- tious substances of every class. This process goes on during the passage of the alimentary mass along the small intes- tine ; and the nutritious materials are progressively with- drawn by absorption, partly into the blood-vessels, which appear to receive whatever are in a state of perfect solution (§ 218), and i)artly into the lacteal absorbents, which take up nothing but that peculiar emulsion of albumen and fatty matter which is termed chyle (§ 222). 214. At the extremity of the Small Intestine, there is a kind of pouch, called the coecum ; which in some animals seems almost like a second stomach, and which is furnished with one or more little appendages, termed coeca."^ This is very small in Man, and does not seem to perform any important * The word coecum is used in Anatomy to denote a tube closed at one extremity. PERISTALTIC MOVEMENT DEFECATION. 195 fimction ; but in most herbivorous animals it is larger (as in the Monkey, fig. 30) ; and it is found to secrete an acid fluid, which resembles the gastric juice, and which may have for its office to perform a second digestion upon the sub- stances which have escaped the first. These ca^ca are some- times very large in the intestinal canal of Birds (fig. 111). — From the coecum, the Large Intestine ascends as high as the liver, crosses the upper part of the abdomen, and then descends again, as shown in fig. 108 ; this portion is termed the colon ; and it terminates in the rectum^ which forms the extremity of the intestinal tube. 215. The alimentary mass is propelled along the first part of the intestinal canal, — and the residue left after the absorp- tion of the nutritive materials is carried along the continua- tion of it, — by the contraction of its muscular coat, producing what is termed the perutaltic motion of the bowels. The fibres of this muscular coat are chiefly arranged in a ring-like manner around the tube ; so that, when they contract, they narrow the diameter of the tube. They are stimulated to contract by the contact of the solid or liquid matter passing through it (Chap, xii.) ; and their contraction forces this matter onwards, into the succeeding portion of the tube. This con- tracts in its turn, so as to propel its contents further ; and thus the mass is gradually diiven from one extremity of the canal to the other. The peristaltic movement does not seem to depend (as do the contractions of the muscles concerned in swallowing, § 195) upon the nervous system ; for it will take place after the intestinal tube has been completely separated from the principal nervous centres ; and also after the death of the animal, if this have been produced by a sudden cause. Thus, if a Kabbit be killed by a smart blow at the top of the neck, and the abdomen be immediately opened, the peristaltic movement will be seen in vigorous action, especially if the animal have eaten a full meal an hour or two previously. Defecation. 216. In passing through the large intestine, the undigested residue is still more completely deprived of the nutritive matter it may contain ; and its fluid portion is absorbed, so that it becomes more solid. It is allowed to accumulate in the rectum, until its bulk occasions inconvenient pressure upon o2 196 DEFECATION LACTEAL ABSORPTION. the surrounding parts ; and it is kept-in by a circular muscle or sphincter, which surrounds the outlet of the alimentary canal. Eut when the accumulation has taken place beyond this amount, it excites a reflex action (§ 195) in the muscles that surround the abdomen ; and these make pressure suf- ficient to overcome the resistance of the sphincter, and to force out the contents of the rectum. Absorption of Nutritive Material. 217. We have only now to inquire into the mode, by which the nutritive matter extracted from the food is taken-up from the alimentary canal and applied to the nutrition of the body. In all Vertebrated animals, there exists a special set of vessels termed Ahsorhents: of which those formino; one division. Thoracic Mesenteric Aorta Duct Glands Origins of --'' Lacteal Vessels • — Intestine Lymphatic Mesentery Vessels Fig. 114,— Chyle-vessels, known as Lacteals, from the milk-like character of their con- tents, originate in tlie numberless villi or minute projections Avith which tlie mucous membrane that lines the small intes- tmo IS covered (§ 41). During the act of digestion, the ABSORPTION BY LACTEALS AND BLOOD-VESSELS. 197 epithelmm-cells, which clothe the extremity of each vilhis (iig. 9), become distended with an opalescent fluid, the chyle (§ 222), which they select from the contents of the small intestine ; and this is subsequently given up by them to a lacteal tube, which, without any open mouth, commences in the midst of each villus. The vessels which thus originate, unite into minute trunks, and these again into larger ones ; and these pass between the two layers of the mesentery (or fold of peritoneum by which the intestines are suspended, § 212) towards the lower part of the spinal column : where they deliver their contents into a sort of reservoir, which thus becomes the receptacle for all the chyle that has been collected from the alimentary canal (fig. 114). In traversing the me- sentery, the lacteals of the higher animals pass through little knot-like bodies of a peculiar nature, which are called mesen- teric glands. These appear to afford the means for the per- formance, within a more concentrated space, of the assimi- lating action which is carried on during the passage of the chyle through the lacteal system ; for in Reptiles, in which these glands do not exist, the absorbent vessels are much more extended and spread out than they are in Birds and Mammals. 218. ]!^ear the surface of each of these villi, moreover, lies a minute network of Blood-vessels ; and there is now no longer any doubt that these receive, by simple imbibition,* any substances, whether alimentary or otherwise, which exist in a state of perfect solution in the contents of the intestinal canal. For a great variety of such substances have been detected, by chemical analysis, in the blood which is returned from the walls of the intestines by the mesenteric veins ; whilst it is seldom that anything is found in the lacteals, save the proper constituents of chyle. It is through this channel that poisonous substances are taken into the circula- tion ; and these may be absorbed from the walls of the stomach (on which there are no villi or lacteals), without ever passing from it into the intestinal tube. Hence it is a great * That tendency — called Endosmose—v^hich. thinner liquids have to pass-towards and mix-with such as are more viscid, even through an intervening membrane, seems to be the physical cause (as experi- ment indicates) of this imbibition ; which is greatly promoted by the movement of blood in the vessels. 198 ABSORPTION BY LYMPHATICS. mistake to characterise the lacteals (with the lymphatics) as Absorbents in any exclusive sense ; the fact being that their function is limited to a special selective absorption, whilst the more general action is performed by the blood-vessels. 219. But the reservoir above-mentioned receives, not only the lacteal vessels that bring nutritious matter from the intes- tinal tube, but also lymphatics, which are absorbent vessels of similar character, that originate in every part of the body. These, also, pass through a set of (so-called) glands, in their way towards this receptacle ; and the structure of these glands, of wliicli many are seated in the neck, some in the arm-pit, others in the groin, &c., is exactly the same as that of the mesenteric glands. The fluid they convey, which resembles very dilute liquor sanguinis (§ 229), seems evidently destined to be again a2)plied to the purposes of nutrition. There is some obscurity as to its source ; but it seems probable that it may partly consist of the residual fluid, which, having escaped from the blood-vessels into the tissues, and having furnished the latter with the materials of their nutrition, is now to be returned to the former ; and partly of those par- ticles of the body, which, though they have lost their vitality in the course of the change it is continually undergoing, have not undergone a degree of decay that unfits them for serving, like the dead bodies of other animals, as a material for reconstruction by the organizing process. The lymphatics, being copiously distributed in the true Skin, absorb substances which are introduced into its tissue ; and if these substances be of an irritating nature, they may occasion an inflammatory action in the absorbents and their glands. Thus when poisoned wounds in the hand have been received, as in opening the bodies of men or animals that have died of particular diseases, the cfl'ect is usually manifested at first by heat and pain in the arm, along which the inflamed absorbents can be traced as hard cords ; and the glands in the arm-pit swell and become tender. 220. The lymphatics do not appear destined, however, to absorb from the surface of the skin ; this function being per- formed by the blood-vessels which are distributed abundantly in its substance. It is a fact now well established, that when the quantity of fluid in the body has been greatly reduced, absorption of water through the skin may take place to a ABSORPTION THROUGH SKIN THORACIC DUCT. 199 considerable amount. Thus there is a case recorded by Dr. Currie, of a patient who suffered under obstruction of the gullet, of such a kind that no nutriment, either solid or fluid, could be received into the stomach ; and who was supported for some weeks by immersion of his body in milk and water, and by the introduction of nutritive liquids into the lower end of the intestine. During this time, his weight did not diminish ; and it was calculated by Dr. Currie, that from one to two pints of fluid must have been daily absorbed through, the skin. The patient's thirst, which had been very trouble- some previously to the adoption of this plan, was removed by the bath, in which he experienced the most refreshing sensa- tions. — It is well known that shipwrecked sailors and others, who are suffering from thirst owing to the want of fresh water, find it greatly alleviated, or altogether relieved, by dipping their clothes into the sea, and putting them on whilst still wet. 221. From the receptacle into which the chyle, and a con- siderable proportion of the contents of the Ijonphatics, are delivered, a tube passes upwards in front of the spine (fig. 114) ; and this tube, called the Thoracic Duc% conveys these nutritious fluids to the point where they are to be delivered into the current of blood. This dehvery takes place at *he angle where two great veins unite, — a point at which there is less resistance than in any other part of their walls. These veins are the Jugular, which brings the blood from the neck, and the Subclavian, which conveys it from the arm, of the right side (fig. 122) ; on the left side there is a smaller duct, which receives some of the lymphatics of the left side, and opens into the blood-vessels at a corresponding point between the left jugular and subclavian veins. Sanguification. 222. The Chyle of Vertebrated animals, as taken-up by the lacteals, may be regarded as blood in an early stage of its formation, with a large excess of fatty matter. It contains about 90 parts of water in 100 ; about 3| j)arts of albumen, and the same of fat ; and about 3 parts of other animal and saline matter. Its appearance and characters differ, according to the part of the lacteal system from which it is drawn. If obtained near the surface of the intestines, before it has passed 200 PROPERTIES OF CHYLE SANGUIFICATION. through the glands, it is entirely destitute of that power of spontaneously coagulating, or clotting, which is so remarkable in blood : and when examined with a microscope, it is seen to present a number of oily globules of various sizes ; together with an immense number of very minute particles or mole- cules, which also seem of a fatty nature ; and to these last, whose diameter is between l-24,000th and l-36,000th of an inch, the milky whiteness which characterises chyle appears principally due. But the chyle drawn from the lacteals, after they have passed through the mesenteric glands, possesses the power of coagulating slightly ; hence it would seem that some of its albumen has undergone a transformation into fibrin (§ 17). At the same time, a great increase is observed in the number of certain floating corpuscles, which are occa- sionally to be noticed in the first chyle, but which are very abundant in the fluid drawn from the glands and from the lacteals that have passed through them ; of these, which bear a strong resemblance to the colourless corpuscles of the blood (§ 234), the average diameter is about 1-4, 600th of an inch. — By the time that the chyle reaches the central receptacle, its power of coagulating has still further increased ; so that its resemblance to blood, except in regard to colour, is much stronger. The proportion of hbrin and albumen which it contains, is much greater than that which existed in the first chyle, whilst the amount of oily matter is less. 223. There can be little doubt that the change which the chyle undergoes in its passage through the lacteals, is partly due to the influence of the living walls of these vessels upon the fluid in contact with them, and partly to that of the colourless corpuscles which float in the fluid, and which form the principal constituents of the absorbent glands. The whole apparatus, indeed, may be looked upon as one great Assimi- lating Gland, having for its function to make blood out of crude nutriment ; provided-for in the higher Vertebrata by the convolution of the lacteals in the mesenteric glands, and in the lower, by the simple extension of the vessels themselves. It is probable that, by being brought into very close neighbourhood with the blood in these glands, the chyle may be made to undergo some further change ; although, as each fluid is con- tained in its own tubes, which do not communicate, there can be no proper intermixture. ASSIMILATING GLANDS ABSORPTION IN INVERTEBRATA. 201 224. There are certain glandular bodies, disposed in various parts of the system, which seem to discharge a similar office ; withdrawing the raw material (so to speak) from the general current of the circulation, and returning it again in a state of higher elaboration. Such are the Spleen, the Thyroid and Thymus glands, and the Supra-Eenal capsules. Besides these, the Liver probably exerts an assimilating action upon the crude materials which are made to pass through its substance, almost immediately after having been received into the blood-current, and before they are allowed to pass into the general circula- tion ; the whole of the blood returned by the gastric and mesenteric veins from the walls of the alimentary canal, being conveyed through the liver by the portal system, in its way to the heart (§ 267). 2'25. In the Invertebrated animals, neither lacteals nor lymphatics exist; and the blood-vessels, whose absorbent powers are to a certain extent restricted in the higher animals, have to perform the functions of these. There are animals, however, which are destitute not only of lacteal and lymphatic vessels, but even of blood-vessels ; and in these, as in the CeUular Plants, there is but little transmission of fluid from one part of the body to the other ; for every portion, both of the internal surface (or lining of the stomach), and of the external surface which is bathed in the surrounding fluid (for most of these animals are aquatic), seems equally to possess the power of absorption ; and the parts to w^hose nourishment the fluid thus received into the body is to be appropriated, are in the immediate neighbourhood of those which have absorbed it. This is the case, for example, in the Hydra and Sea-Anemone, and, more or less, in all the Polypes ; as well as in the lower Worms. Between these, therefore, and the Cellular Plants, a remarkable analogy exists in regard to the mode in which the nutriment is absorbed and applied ; the diff"erence being, that the Animal possesses a digestive cavity, lined by an inward extension of the external surface, which does not exist in Plants (§ 8). And it is upon the walls of this cavity, that the absorbent vessels of the higher Animals (whether lacteals or blood-vessels) are distributed, collecting the nourishment in contact with them ; just as the roots of a Plant, spread through the soil, draw up that which it contains. But among those lowest animals in which the digestive cavity altogether 202 OF THE BLOOD, AND ITS CIRCULATION. disappears (§ 203), the function of absorption is not in any wa}' limited ; since every part seems to have the power of re- » ceiving from without, and of assimilating to its own substance, I the nutrient materials which it needs. CHAPTER V. OF THE BLOOD, AND ITS CIRCULATION. 226. The processes that have been already explained, have for their object to prepare the nutritious fluid, which suppUes the materials for the growth of the several parts of the body, and which is conveyed through them by the apparatus to be presently described. In Man and the higher animals, this fluid, which is known as tlie Blood, has a red colour, and con- tains a large quantity of solid matter. The redness of the blood has been mentioned as a distinctive character of the Verte- brated classes (§ 75) ; it exists in Mammalia, Birds, Eeptiles, and Fishes, and in these alone. In the Molluscous classes, as also in most of the Articulated, the nutritious fluid is nearly colourless ; and it will hereafter appear that this fluid bears, in some respects, a stronger resemblance to the chyle and lymph of the Vertebrata, than to their blood (§ 234). There is an apparent exception in the case of certain marine Worms, the fluid circulating in whose vessels has a reddish hue ; this does not depend, however, upon the presence of any red par- ticles, such as are characteristic of the blood of Vertebrata (§ 229), but upon a reddish tinge in the fluid itself, which does not seem altogether to answer to the character of blood (§ 294). 227. The blood of all the higher animals exists in two difl'crent states. Wlien it is drawn from a slight scratch or other wound of the skin, it is of a bright red hue ; whilst that which is drawn in bleeding from the arm, is of a dark purple. The former is termed arterial blood, because it is contained, for the most part, in the tubes which are called Arteries, and which are conveying it from the heart to the tissues it has to nourish. The latter is called venous blood, because it is drawn from the Veins, by which it is returned from the tissues to the heart, after having performed its part in them. Hence it VENOUS AND ARTERIAL BLOOD. 203 is evident that this change of character has been produced during the passage of the blood through the tissues ; and so important is the alteration, that the blood which has been subjected to it is not fit to pass again into the arteries of the body, until it has been renewed by exposiu-e to air in the Lungs, In their vessels, the contrary change — of which the nature will be presently explained (§ 253) — is effected, the dark hue of venous blood giving place to the bright red of the arterial fluid ; this is again changed during the passage of the blood through the body, to be again restored in the lungs. The same is the case in regard to Fishes, whose gills perform the same function as the lungs of air-breathing Vertebrata. And among the Invertebrated classes, although the deteriora- tion of the blood in its passage through the body is not made manifest by any change of colour, yet its renewal by exposure to air in the respiratory organs is not less requisite. 228. Hence the continual movement of the blood is neces- sary for two purposes in particular ; — first, to convey the nutritive materials from the place where they are received and prepared, to that in which they are appropriated, and thus to afford to every organ a constant supply of the materials which it requires ; — and, second, to carry this fluid, at regular intervals, to certain organs by whose instrumentality it may be exposed to the influence of the air, so as to regain the qualities it has lost, and part with what it has taken-up to its prejudice. But there are many other objects fulfilled by it, which will unfold themselves as we proceed. Properties of the Blood. 229. Wlien the circulating blood of a red-blooded animal is examined with a microscope, it is seen to consist of two distinct parts ; — a clear and nearly colourless fluid, to whicli the name of liquor sanguinis (or liquor of the blood) is given ; and of an immense number of rounded particles floating in this fluid, which are often termed the glohides of the blood. The shape and size of these particles are, for the most part, very uniform in animals of the same species ; but in no instance are they globular ; and it is better, therefore, to term them corpuscles. In Man and most other Mammals, they are nearly flat discs, resembling pieces of money, but usually exliibiting a slight depression towards the centre (fig. 115). 204 BLOOD-DISCS OF MAN AND MAMMALS. No nucleus can be distinguislied in tliem, but they present a dark central spot, which is an optical effect of their bi-concave form ; and this spot may be made to disappear by the addition Fig. 115. — Red Cokpuscles of Human Blood. Seen separately at a, a a showing the front view, b the profile or edge view, and * a three-qi:arter view; at b united with each other so as to form columns likepiljs of money; at c in a state of alteration such as exposure to air will produce; D shows a colourless corpuscle, or lymph-globule. of water to the liquid in which they are suspended, the discs first becoming flat, then bulging- out on either side, and at last swelling so as to burst. The reason of this will be pre- sently explained (§ 231). In Man and Mammals generally, the diameter of these blood-discs varies from about l-2800th to 1 -4000th of an inch ; but in the small Musk-deer, it is less than 1-1 2,000th. In the Camel tribe, the discs are oval, as in the lower Vertebrata. 230. In Birds, Eeptiles, and Fishes, the blood-particles present some curious differences from those of Mammalia. In the first place, they are much larger ; their form, also, is oval instead of being round ; and instead of being depressed in the centre, they bulge-out on each side. Tliis bulging is ^ Fig. 116. — Blood Corpuscles of Pigeok. At A are seen the red corpuscles a, 6, and the colourless, or lymph globules c, c; at D, a red corpuscle treated with acetic acid ; and at c, the same treated with water, 80 as to render the nucleus more distinct. ovidently occasioned by the presence of a nucleus which is more sohd than the rest ; the nucleus, however, is not so well BLOOD-DISCS OF BIRDS AND REPTILES. 205 seen in the corpuscles of circulating or of freshly-drawn blood, as it is in that of blood which has been drawn for some little time ; and it is best brought into view by treating the blood either with water or with acetic acid. The long diameter of the oval discs of Birds (fig. 116) varies from about l-1700th to 1 -2400th of an inch ; and the short diameter from about Fig. 117.— Blood Corpuscles of Frog. At A are seen the red corpuscles a, b, and the colourless corpuscle c ; at b, a red corpuscle treated with acetic acid. 1 -300th to l-4800th. Thus the discs, though much longer than those of Man, are not in general much broader. In Eeptiles, Fig. 118. — Blood Corpuscles of Proteus. a, b, red corpuscles ; a*, corpuscle showing the nucleus ; c, colourless corpuscle ■ d, red corpuscle treated with water. 206 BLOOD-DISCS OF REPTILES AND FISHES. there is considerable diversity as to the size of the discs ; but the largest particles are found in the group of Amphibia, and especially in those species which retain their gills through life. The oval discs of Frogs (fig. 117) have a long diameter of about 1-1 000th of an inch, and a transverse diameter of about 1-1 800th. Those of the perennibranchiate Amphibia (§ 87) may even be distinguished by the naked eye ; those of the Siren having a long diameter of about 1 -435th of an inch, whilst in the Proteus (fig. 118) the long diameter is stated occasionally to reach 1 -337th of an inch. In Fishes, also, the size of the blood-discs is variable ; they are sometimes smaller (fig. 119), though generally larger, than those of the Frog ; but they never approach those of the last-named remarkable ani- mals. Hence the great size of the Fig. 119.-BL00D Corpuscles OP blood-discs of the CUrioUS Lcpido- RoACH. si^^n (fig. 41) is strongly indicative a, a, b, red corpuscles ; c, colour- £ ^j Ecptilian affinities of that less corpuscle; d, red corpuscle '^^ , ^ J^ treated with water. SpecicS. 231. It is by observing the large blood-discs of the Frog, and still better those of the Proteus and Siren, that we can obtain the best information as to their structure. They are exidenilj fattened cells, having an envelope or cell-wall, which consists of an extremely delicate membrane, and which con- tains a fluid. The nucleus consists of an assemblage of minute granules, which seem adherent to each other and to the wall of the cell ; and it corresponds, in all essential particulars, to the nuclei of the cells of other Animal tissues (§ 32). The fluid contained in the cells has a red colour.; and it is to this that the peculiar hue of the blood of Yertebrata is owing. When we are looking at a single layer of blood-discs, how- ever, their red colour is not apparent, but they have rather a yellowish tint; and it is only when we look through a number at once, that the characteristic hue is seen. The fluid is of about the same density as that in which the par- ticles float ; and thus neither will have a tendency to pass towards the other. T3ut, if Ave dilute the liquor sanguinis with water, the fluid outside the cells will have a tendency to pass towards their interior, according to the law of Endos- mose. The cells Avill in consequence be first distended, and STRUCTURE AND COMPOSITION OF RED CORPUSCLES. 207 will then burst ; and their contents will be diffused through the surrounding fluid, whilst their membranous walls will subside to the bottom. On the other hand, if the liquor sanguinis be rendered denser than the fluid in the blood- discs, as by the admixture of gum or syrup, the latter will pass towards it, and the cells will become still more flattened, and more or less completely emptied. The flexibility and elasticity of the blood-discs are well seen, in watching (with a microscope) its flow tlirough the minute vessels ; for if one of them meets with an accidental obstruction to its progress, its form becomes accommodated to that of the space left for it to pass, and it makes its way tlirough a very small ajDcrture, recovering its usual form immediately afterwards. 232. The Red Corpuscles differ considerably in chemical composition from the liquid in wliich they float. Of the solid residue obtained by drying, about one-eighth is formed by their cell-walls, the remainder being yielded by the cell- contents. The latter portion seems to consist chiefly of a mixture of two components, which have been named glohulin and h(Ematin. The former is a colourless substance, nearly allied to albimien in composition, but differing from it in some of its reactions ; its most characteristic peculiarity, how- ever, being its power of crystallizing. Its crystals, the form of which varies in different animals, are usually tinged deeply with haematin, from which they cannot easily be freed. The composition of hsematin, to which alone the colour of the red corpuscles (and consequently of the whole mass of the blood) is due, is notably diflerent from that of the albuminoid compounds ; the proportion of carbon to the other components being much greater, and a definite quantity of iron being an essential part of it. This iron, in a certain state of oxidation, has been supposed to be the source of the red colour; but such is certainly not the case ; and this hue must be, like the colours of Plants, a peculiar attribute of the organic compound which presents it. — Besides their globulin and haematin, the red corpuscles contain a certain proportion of fatty and mineral matters. The former, which are united with phos- phorus, are of a kind which are scarcely traceable in the liquor sanguinis ; and the latter are remarkable as having potass for their principal base, whilst the base of the salts of the liquor sanguinis is chiefly soda. Hence it appears that 208 PROPORTION OF RED CORPUSCLES. the Eed Corpuscles draw into themselves nearly the whole of the iron, phosphorus, and potass, which the chyle pours into the circulating current; and that they modify a large pro- portion of the solid matter of the blood, that which they con- tain being notably different in composition from that of the liquor sanguinis, which does not differ, save in the proportion of its components, from the liquid portion of Chyle or Lymph. 233. The proportion of Eed Corpuscles to the whole mass of the blood varies greatly in different animals, and even in different states of the same animal. It is greatest in those which have the highest muscular vigour and activity, and which consume the largest quantity of oxygen by respiration ; hence these particles are rather more numerous in the blood of Birds than in that oi Mammals, and far more abundant in these last than in Eeptiles or Fishes. Again, they are more numerous in Men of ruddy complexion, strong pulse, and active habits, than in those of pale skins, languid circu- lation, and comparatively feeble powers. In a healthy Man they seem to constitute about half the mass of the circulating blood ; but they contain as much as three-fourths of its solid matter, the proportion of dri/ corpuscles being about 150 in 1000 parts of blood, whilst that of the other solid matters is about 50. A very marked decrease occasionally presents itself in disease ; the proportion of dry corpuscles being some- times reduced as low as 27. When too abundant, they pro- duce what is known as the plethoric condition of the body, in which haemorrhage from the bursting of a blood-vessel is liable to occur. Their number is effectually reduced by bleed- ing ; and the aspect of those who have suffered from extreme loss of blood, gives sufficient evidence that the deficiency is not made-up for a long period. The most effectual means of restoration, in cases where the proportion of blood-corpuscles is too low, is a highly nutritious diet, with the administration of iron as a medicine ; for this substance seems to have the power of hastening the reproduction of the corpuscles, being itself an essential ingredient in their contents ; and there arc facts which show its remarkable power of increasing their amount in proportion to the mass of the blood. 234. It appears that the red corpuscles, like other cells, have a certain allotted term of life ; and as they are con- tii.ually dying, they must be as continually reproduced. The COLOURLESS CORPUSCLES — USES OF RED CORPUSCLES. 209 mode in which this reproduction is effected has not yet been clearly made out ; hut there is strong reason to believe that the red corpuscles are developed from the corpuscles of the chyle and lymph (§ 222) which are continually being poured into the circulating current, and of which isolated examples, known as the white or coloitrless corpuscles, are met with in every drop of blood that is examined under the microscope. The size of these is pretty much the same in all Vertebrata, their diameter being usually about l-3000tli of an inch. In the blood of Maji and the Mammalia in general (fig. 115, d) they are not easily distinguished from the red particles ; their diameter being nearly the same, while the colour of single discs of the two kinds is not very dissimilar. But in the lower Vertebrata, whose blood has large oval red particles, the differ- ence between the two kinds is very obvious ; and the resem- blance which the colourless globule!^ (c, figs. 116-119) bear to those of the chyle and lymph, is very striking. Similar colour- less particles exist, to a variaoie amount, in the nutritive fluid of Invertebrated animais ; so that in this, as in some other respects, that fiaid bears a stronger resemblance to the chyle and lymph of the Vertebrata, than it does to their blood, which is characterised by the presence of the red particles. 235. Physiologists are now generally agreed, that one of the functions of the Eed Corpuscles is to convey oxygen from the lungs to the tissues and organs through which the blood circulates, and to bring back the carbonic acid which is set free in these, so as to deliver it at the lungs. For although it is certain that the liquor sanguinis can also convey these gases, yet experiment shows that the red corpuscles can take up, bulk for bulk, a much larger proportion of them ; and that the blood w^hich is richest in these particles is, therefore, most fit to serve as the medium for the transmission between the respiratory organs and the body at large. I^^ow it is in the nervo-muscular apparatus that there is the greatest demand for oxygen ; for this apparatus is not capable of vigorous action, unless oxygen be freely supplied to it. The quantity of this it requires, however, depends upon the exercise of its powers ; for when at rest, it needs little or no more than is made use of by the other tissues ; but whilst in activity, it needs a greatly-increased supply. The quantity of oxygen which the animal takes-in by its lungs, and the amount of p 210 USES OF RED CORPUSCLES LIQUOR SANGUINIS. carbonic acid whicli it gives-off by the same channel, vary, therefore, with the muscular exertion it makes. This variation is most easily observed and measured in Insects ; and it is found in them to be enormous (§ 308). As, however, the blood of the Invertebrata does not contain these red particles, to which so important a function has been assigned, it may be asked, how the conveyance of oxygen to their tissues is provided for. The reply is very simple. In Insects, and other Arti- CULATA which have active powers of motion, the air is con- veyed to the tissues, not through the medium of the blood, but directly through air-tubes which convey it to every part of the body (§ 321). And in the Molluscous classes, as among the Crustacea also, the nervo-muscular system forms so subordinate a part of the general mass of the body, and its movements are so sluggish, that the quantity of oxygen which the fluid part of the blood conveys to them, is sufficient for their need. 236. Of the properties of the Liquor Sanguinis, whilst it is circulating in the vessels, the microscope tells us nothing ; since it constantly remains in the state of a transparent fluid. Eut if the blood be withdrawn from the living body, it soon imdergoes a very curious and important change. A large portion of it passes into the solid state, forming the crassa- mentum or clot ; whilst there remains a transparent liquid of a yellowish hue, which is termed the serum. It is evident that the clot contains all the red particles ; but it is easily proved that its coagulation is not due to them. For the blood of a Frog, or of any other animal having blood-discs suffi- ciently large, may be caused to pass through filtering-paper, which will retain and collect its blood-discs, allowing the liquor sanguinis to flow through it ; and this fluid will coagu- late just as completely as if these particles were retained in it. Again, in certain conditions of the blood (generally result- ing from disease), even when the coagulation is allowed to take place in the ordinary manner, the fibrin and the red ])articles separate from one another, — the latter gradually su})siding, whilst the former are left at the surface ; and the upper part of the clot is then nearly colourless, exhibiting what is commonly known as the huffy coat or crust ; whilst the lower part of it includes the red particles, and has a very deep colour. The bufly coat, being composed almost exclu- LIQUOR SANGUINIS COAGULATION. 211 sively of the fibrous network, is very firm in its texture, being sometinjes almost leathery in its character ; whilst the lower part of the clot, which is chiefly composed of the red particles, loosely bound together by scattered fibres, is very soft, and easily broken asunder. This effect may be also produced, by acting on healthy blood with certain substances which retard its coagulation, such as a strong solution of Glauber's salt ; for if sufficient time is allowed, the red par- ticles will subside in consequence of their greater specific gravity, leaving a colourless layer of fibrin above them. — It is of the liquor sanguinis, in a concentrated form, that those exudations consist, which are poured out from the blood for the repair of injuries, and which pass spontaneously into the condition of a simple form of tissue (§ 393). 237. When a very thin slice of the clot is examined with a microscope, it is found to be made up of a net-work of an imperfectly fibrous character, interlacing in every direction, and including the blood-discs in its meshes. These fibres are produced by the spontaneous change in the fibrin of the blood, from the fluid to the solid form. So long as the blood is circulating in the vessels of the living body, so long does its fibrin remain dissolved in the watery part of it ; but so soon as it is withdrawn from these, and is allowed to remain at rest, it undergoes this remarkable change. If fresh-drawn blood be continually stirred with a stick or beaten w4th twigs, the fibrin coagulates in irregular strings, which adhere to the stick or twigs ; and it does not then include the red particles, which are left behind in the fluid. In this maimer it may be completely separated from the other elements of the blood, which have not in themselves the least tendency to coagulate spontaneously. Although forming a large proportion of the substance of the clot, the fibrin, when dried, does not consti tute more than from 2 to 3 parts by weight in 1000 of blood. This proportion is augmented to 6, 8, or even 10 parts, in severe inflammatory diseases. 238. When the fibrin and the red particles have both been separated from the blood, there remains a fluid, the serum, in which a good deal of albumen is dissolved, together wlvii fatty matter, and other organic substances ; with the addition of saline matter, of which a considerable pr( portion is chloride of sodium, or common salt. The proportion which the solid p2 212 SERUM USES OF BLOOD. matter of the serum bears to the whole mass of blood, in health, is about 53 parts in 1000 ; and of these about 40 parts are albumen, 8 parts saline matter, and 5 parts fat, with certain ill-defined substances, of which some appear to be organic compounds that are undergoing metamorphosis into sohd tissues, whilst others are the products of the decay of the tissues, which are being progressively withdrawn and ehminated by the excretory organs. 239. The influence of the Blood as a whole upon the anunal as well as on the nutritive functions, is easily proved. When an animal is bled largely, it is gradually weakened as the flow proceeds, and at last it seems to lose all consciousness and power of movement. If allowed to remain in this con- dition, it seldom or never recovers of itself. But if we inject into its veins, by small quantities at a time, blood similar to that which it has lost, the apparent corpse becomes as it were reanimated, and all its functions are completely re-established. The importance of the red particles is manifestly seen in the (tfifect of this remarkable operation, which is called the tra7is- /usion of blood ; for if, instead of blood freshly obtained from another living animal, we inject serum without these particles, the effect is but httle greater than if so much water were introduced, and the animal dies of the hasmorrhage. By this operation, practised on the Human subject, many valuable hves have been saved, that would otherwise have been de- stroyed by loss of blood. Again, if, by mechanical means, as by tying the principal blood-vessel going to any organ, we cause a permanent diminution to any considerable extent, in the quantity of blood with which it is supplied, a decrease in its size is soon apparent, and it may even shrink almost to nothing. On the other hand, we observe that, the more active the function of a part, the larger is the quantity of blood with which it is supplied. Thus, when the antlers of the Stag, which fall off every year, are being renewed, the arteries that supply the parts of the skull from which they spring, are greatly increased in size ; but they shrink again, as soon as the growth of the horns is completed for that year. A similar increase takes place among animals tliat suckle their young, in the size of the arteries that supply the mammary glands, by which the milk is formed ; and these also shrink, when this liquid is no longer required. USES OP SEPARATE CONSTITUENTS OF BLOOD. 5l3 240. The following appear to be tlie chief uses of the principal constituents of the Blood, considered separately, in the general economy : — The fibrin is the material which is most assimilated to the condition of the solid tissues, having the power of passing from the Hquid state into a low and simple form of organization. It was formerly supposed to be the nutritive material at the expense of which the solid tissues generally are immediately produced ; the muscular substance, in particular, being regarded as chemically identical with it. But there is now good reason to think that the greater part of the tissues form themselves at the expense of the albumen of the serum and perhaps of the globulin of the red corpuscles ; and that the purpose of the fibrin is chiefly to give origin to those simple forms of fibrous or connective substance, the production of which is the first step in the reparation of injuries. Were it not for its power of coagula- tion, the slightest cut or scratch might become fatal, from the gradual draining-away of the blood ; and such, in fact, has actually happened, in cases of disease in which the fibrin is deficient. The presence of fibrin also gives a degree of vis- cidity to the blood, which, as experiment proves, favours (instead of resisting, as might have been expected) its passage through capillary tubes ; and thus, when there is a deficiency in this ingredient, local stagnations and obstructions in the circulation of the blood are very liable to occur. The albumen of the blood may be considered, like that of the egg, as the raw material, at the expense of which (in combination with fat) every other organic compound in the body is generated. It is, as we have seen, the substance to which all the tissue- forming elements of the food are reduced in the process of digestion ; and in this condition it seems to be continually appropriated by the acts of self-formation that are taking place, with varying rapidity, throughout the body, just as the albumen of the egg is appropriated by the self-formative operations of the embryo. There is strong reason to believe that a large proportion of the solid tissues regenerate themselves by the direct appropriation of this material ; and if (as has been already stated to be probable) the simple fibrous tissues find their material in the fibrin, and the muscular substance in the globulin of the red corpuscles, it is from the albumen that these substances are themselves elaborated, both of thein 214 USES OF SEPARATE CONSTITUENTS OF BLOOD. being, as it were, in process of organization. The albumen of the blood further serves to supply the albuminoid matters which are required as constituents of various secretions, espe- cially those which are concerned in the digestive process, as the saliva, the gastric juice, and the pancreatic fluid. A large amoimt is daily drawn-ofF for the production of the peculiar ferments contained in these secretions, whose action upon the food is necessary for its reduction to the form in which alone it can be received into the circulating current. Hence the making of new blood involves a considerable ex- penditure of the old. 241. The liquid in which the fibrin and albumen are dis- solved, has a considerable power of absorbing gases ; and this is greatly increased by the presence of the saline matters which it holds in solution. Hence the liquor sanguinis not only sustains the nutrition of the body, but can also serve, to a considerable extent, as a medium of communication between the lungs and the tissues. In this kind of activity, however, it is completely surpassed by the red corpuscles (§ 235). Independently of their use in ministering to the function of Respiration, there seems reason to believe that the red cor- puscles are also subservient to that of Nutrition ; for a certain conformity which exists between the organic and mineral sub- stances they contain (§ 232), and the composition of Muscle and Nerve, taken in connexion with the manifest relation between their number and the activity of the Nervo-muscular apparatus, makes it probable that they have it for their especial office to prepare the materials which are to be used in its pro- duction and renewal of those tissues. The saline matter of the blood has many important offices : thus it furnishes the mineral ingredients which are requisite for the production of the tissues and secretions ; it helps to preserve the organic substances from decomposition ; and, in conjunction with the albumen, it keeps up the density of the scrum to the point at which it is equi- valent to that of the contents of the red corpuscles, without which balance the condition of the latter would be seriously impaired (§ 231). Finally, the fattij matters of the blood are subservient to two very important functions — the maintenance of heat, and the formation of tissue. They maintain the combustive process, whenever there is a deficiency of more readily combustible material 3 and they also take part with ASSIMILATING AND SELF-PURIFYING POWER OF BLOOD. 215 albumen in the formation of all new tissue, its nuclear par- ticles being always found to include fat-granules. 242. The presence of a due proportion of the foregoing substances in the blood is an essential condition of health ; and we find it provided-for in the marvellous power w^hich the blood, like any solid tissue, seems to have of making itself from the materials supplied to it, and of getting rid of what is superfluous or unsuitable. Thus an excess of albuminous matter in the food does not seem to produce more than a very limited increase in the quantity of albumen in the blood, the surplus being made to undergo changes within the body, which issue in its being removed by the excretory organs. An excess of any of the saline compounds is very speedily strained off (as it were) into the urine. And an excess of fatty matters is draT\Ti oif either by the formation of fat as a tissue, or by the augmented activity of the liver in producing bile. This conservative power is still more remarkably shown in the completeness with which the poisons that are generated in the body by the decay of its tissues, and which are received into the current of the circulation for the purpose of being conveyed to the several excreting organs, are drawn off from it, so as to leave the blood pure. Thus, carbonic acid is being continually produced in such large quantities, that its accu- mulation in the blood, even for five minutes, would be fatal ; yet by the aerating process to which the limgs are subservient, it is got rid of as fast as formed, so that the blood is restored to its previous purity. In like manner, the urea, which is one of the products of the wear and tear of the muscles consequent upon their use, is so perfectly and constantly eliminated by the kidneys, that its detection in the circulating current is a matter of difficulty, although we know that it must always be passing through this. 243. Thus the circulating current may be likened to a tidal river running through the midst of a large town, and supplying it with the water needed for the drink of its human and other inhabitants, as well as with that w^hich is required for the various manufacturing and cleansing opera- tions carried on within its precincts ; the same stream also receives the drainage of the town, and consequently becomes charged with the products of animal and vegetable decompo- sition, and the foul refuse of manufactories j and as the flow 216 CIRCULATION OF THE BLOOD. of the tide brings back a large proportion of what is carried down at ebb, the waters speedily become so contaminated with hurtful and offensive matters as to be unfit for use, unless means be provided for getting rid of these as fast as they are poured in. The perfection with which tliis requirement is fullilled in the Animal body, while it excites our admiration, should also incite us to imitation, so far as the art of Man can hope to imitate the works of the Divine Artificer. Circulation of the Blood. 244. In some of the lower tribes of Animals, the blood appears to circulate in channels which are merely excavated in the substance of their tissues and organs. But among all the Vertebrata, and even in most of the Invertebrated classes, the movement of the blood takes place in a very complicated apparatus, which is composed, 1st, of a system of tubes or canals wliich serve to convey it through every part of the structure, and 2d, of a special organ for the purpose of giving motion to that liquid. These canals are known as the blood-vessels ; and this special organ is the heart. 245. The Zleart is the centre of the circulating apparatus. It is a kind of fleshy bag, communicating with the blood- vessels : and it alternately dilates to receive the blood, which is conveyed to it by one set of these ; and then contracts so as to force it out into another set of tubes. In this manner a continual current is kept up. All but tlie lowest animals have a heart, or something which represents it. Such an organ exists, not merely among all the Vertebrated classes, but in all the Mollusca, and in the higher Articulata. But, as will presently appear, there is a great diversity in its form, and in the complexity of its construction ; for whilst, in its simplest condition, it possesses but one cavity, communicating with both sets of vessels, — it contains, in its highest forms, four different chambers, each of which has its own peculiar function. 246. The two sets of blood-vessels just adverted-to are, 1st, the Arteries, which convey the blood from the heart into the several parts and organs of the body ; and 2d, the Veins, which collect the blood that has been distributed through these, and return it to the heart. The Ai'terial system, as it issues from i\w heart, consists of one or more large trunks, which divide into branches, very much in the manner of the. DISTRIBUTION OF ARTERIES AND VEINS. 217 stem of a tree ; these branches again subdivide into othei-s more numerous but smaller, and these again into twigs still more nmnerous and more minute ; until almost every portion of the body is so penetrated with them, that not even a trifling scratch, cut, or prick, can be made, without wounding some one of these small divisions (fig. 120). — The Venous system presents a corresponding distribution, but it is destined for an opposite purpose ; and we must regard it as commencing in the tissues by the minuter canals, which run together like the Fig. 120.— Distribution of thk smaller Blood-Vessels in the Membrane BETWEEN two OF THE TOES OF THE HINI) FOOT OF THE CoMMON FrOG ; « U, veins; b b, arteries. little rivulets that form the origin of a mighty river, or like the smallest fibres of which the roots of a tree are made up. The larger canals thus formed gradually unite with each other as they approach the heart, towards which they all tend, just as the various tributary streams pour their contents into one prin- cipal channel : and at last all the veins empty into the heart, by one or two large trunlvs, the blood which they have conveyed from the several parts of the body ; just as all the tributaries which have arisen over a wide extent of country, pour into the ocean the water they have collected, by one mouth which is thus common to all of them. 247. Although the number of the Arterial branches increases 218 AREAS OF ARTERIAL TRUNKS AND BRANCHES. SO vastly, as we proceed from their origin towards their termi- nation, yet their capacity does not, at least in any considerable (jegree ; — that is, the first or main trunk will allow as much fluid to pass through it in a certain time, as will the whole of the first set of branches into which it divides, or the still more numerous subordinate branches into which these diverge. Or, to put this fact in another form, if we cut across the main trunk, and compare the area, or space included within its circular walls, with the sum of the areas of all the branches it supplies at a certain distance — say a foot — from the heart, we shall find them precisely equal ; and the same will hold good, if the comparison be made with the sum of the areas of the more nimierous but smaller branches at a greater distance from the main trunk. It is quite true that, when an artery divides into branches, the combined size of these seems to be greater than that of the trunk ; but this is only because the compa- rison is made, not between the areas of their circles, but their diameters. Thus, an artery of 10*1 lines in diameter, may divide into three branches, two of them having a diameter of 7 lines, and the third a diameter of 2 lines ; — and yet these will convey no more blood than the single trunk. For, according to a simple rule in geometry, the areas of circles are to each other as the squares of their diameters. The area of the trunk is expressed, therefore, by the square of lO'l, which is almost exactly 102. The area of each of the two large branches, in like manner, is expressed by the number 49, which is the square of 7 ; and that of the smaller one by 4, the square of 2 ; and the sum of these (49+49+4) is ex- actly 102, making the combined areas of the branches the same as that of the trunk. In like manner, one of the branches of 7 lines diameter might subdivide into two branches of a little less than 5 lines each ; for, as the square of 5 is 25, and twice that number is equal to 50, the combined areas of the two branches of 5 lines each, exceed by very little the area of the trunk of 7 lines. — Hence it results, that the pressure of the blood upon the walls of the arteries will be everywhere almost exactly the same ; — a conclusion which is confirmed by experiment. 248. Tliere are certain differences in the structure and dis- tribution of the Arteries and Veins, which it is desirable to mention. The Arteries receive the blood pressed out from STRUCTURE AND DISTRIBUTION OP ARTERIES AND VEINS. 219 the heart, and must be strong enougli to resist the force of its contraction ; otherwise, as there is a considerable impediment to its onward flow, produced by the minuteness of the tubes through which it has to pass, and the friction to which it is subjected against their sides, their walls would give way, and they would burst. They have, accordingly, a tough elastic fibrous coat, which contains also more or less of non-striated muscular fibre. On the other hand, the Veins receive the blood after the heart's power over it has been almost ex- pended in forcing it through the capillary system, and when it is consequently moving much more slowly. They are very large in proportion to the arteries ; so that, if we were to cut across a limb at any place, and to estimate the respective areas of all the veins and arteries, we should find that of the veins two or three times as great as that of the arteries. Hence the pressure on their walls is much less ; and their strength does not require to be so great. Accordingly we find their walls much thinner, and the tough elastic fibrous coat almost entirely wanting. 249. The difference in the force with which the blood presses on the walls of the arteries and veins, is seen when these vessels are wounded. If a small incision be made into an artery, the blood spouts from it to a great distance ; but if a similar incision be made in a vein, the blood merely flows out, unless we stop its passage to the heart, by making pres- sure on the vein above the orifice, as in ordinary blood-letting (§ 277). Hence much greater pressure is requisite to check bleeding from an artery, than to stop bleeding from a vein ; and it frequently happens that no amount of pressure can prevent the continued drain of blood from the former, so that it becomes necessary to stop the flow of blood through the artery altogether, by tying a ligature tightly round it. 250. The Arteries are for the most part so distributed, that their trimks lie at a considerable distance from the surface of the body, so as to be secluded from injury ; and they are often specially protected by particular arrangements of the bony parts. Of the Veins, on the other hand, a large proportion he near the surface, and they are consequently more liable to be injured ; but, for the reason just stated, wounds in them are of comparatively little consequence. 251. The ultimate ramifications of the Arteries are conti- 220 CAPILLARY BLOOD-VESSELS. nuous with the commencing twigs of the Venous system. The communication is established by means of a set of extremely minute vessels, which are termed Capillaries.'^ These capil- laries form a network, Avhich is to be found in almost every part of the body (fig. 121). It is in them alone that the blood ministers to the opera- tions of nutrition and secre- tion. Even the walls of the larger blood-vessels are inca- pable of directly imbibing nourishment from the blood which passes through them ; but are supplied with minute branches, which proceed from neighbouring trunks, and form a capillary network in their substance. The diameter of the capillaries must of course bear a certain proportion to that of the blood-discs which have to pass through them : in Man they are commonly from about 1 - 2500th to 1-1 600th of an inch in dia- meter. In the true capilla- ries, it would seem that only one row or file of these particles can pass at a time ; but we frequently see vessels passing across from the arteries to the veins, which will admit several rows. There seems, however, to be a considerable difference in the diameter of the same capillary at different tim.es ; a change sometimes taking place from causes which are not yet understood. f The rate at which the blood moves * From the Latin capilla, hair ; so named on account of their being, like hairs, of very minute size. Their diameter is really, however, far less than that of ordinary hairs. t The circulation of the blood in the Frog's foot, the tail of the Tadpole, the gilis of the larva of the Water-Newt, the yolk-bag of embryo Fish, and other appropriate subjects for the observation, is one of the most beautiful and interesting spectacles that the Microscope can open to us. Details of the various modes of exhibiting it will be found in the Author's treatise on " The Microscope and its Eevelations," Chap, xviii. Fig. 121.— Portion of thic Membrane BETWEEN THE TUES OF THli HIND FOOT