©lie i. M. HtU ffitbrarg North (Earaltna *tat? Imnprflitg QH501 R6 NORTH CAROLINA STATE UNIVERSITY LIBRARIES S02 177209 S THIS BOOK IS DUE ON THE DAT INDICATED BELOW AND IS SUl JECT TO AN OVERDUE FINE A POSTED AT THE CIRCULATIO DESK. 2000 I.. THE BRIDGEWATER TREATISES ON THE POWER, WISDOM, AND GOODNESS OF GOD, AS MANIFESTED IN THE CREATION. TREATISE V. ANIMAL AND VEGETABLE PHYSIOLOGY, CONSIDERED WITH REFERENCE TO NATURAL THEOLOGY. BY PETER MARK ROGET, M. D. SEC. R, S. ETC. IN TWO VOLUMES. VOL. II. " And there are diversities of operations, but it is the same God which worketh all in all," 1 Cor. xii. G. ANIMAL AND VEGETABLE PHYSIOLOOY, CONSIDERED WITH REFERENCE TO NATrRAli THEOIiOGY. BY PETER MARK ROGET, M. D. SECRETARY TO THE ROYAL SOCIETY, FULLERIAN PROFESSOR OF PHYSIOLOGY IN THE ROYAL INSTITUTION OF GREAT BRITAIN, VICE PRESIDENT OF THE SOCIETY OF ARTS, FELLOW OF THE ROYAL COLLEGE OF PHYSICIANS, CONS0LTIMQ PHYSICIAN TO THE QOEEN CHARLOTTE'S LYING-IN HOSPITAL, AND TO THE NORTHERN DISPENSARY, ETC, ETC. VOL. II. PHILADELPHIA: CAREY, LEA & BLANCHARD. 1836. GRIGGS & CO., PRINTERS. COIVTEIVTS OF THE SECOND VOLUME. PART II.— THE VITAL FUNCTIONS. Page Chapter I. — Objects of Nutbition ----- 9 Chapter II. — Nutrition ix Vegetables - - . . 19 § 1. Food of Plants - - - . - - 19 2. Absorption of Nutriment by Plants - - - 21 3. Exhalation - - - - - - 27 4. Aeration of the Sap ----- 28 5. Return of the Sap - - - - - 32 6. Secretion in Vegetables - - - - 38 7. Excretion in Vegetables - - - - - 43 Chapter III. — Animal Nutrition in general . . . ^ §1. Food of Animals - - - - - - 47 2. Series of Vital Functions - - - - SS Chapter IV. — ^Nutrition in the lower orders of Animals - 58 Chapter V. — Nutrition in the higher orders of Animals - 80 Chapter VI. — Preparation of Food - - - - 86 § 1. Prehension of Liquid Food - - - - 86 2. Prehension of Solid Food - - - - 89 3. Mastication by means of Teeth - _ - 104 4. Formation and Development of the Teeth - - 114 5. Trituration of Food in Internal Cavities - - 122 6. Deglutition - - - - - - 127 7. Receptacles for retaining Food . - - 130 Chapter VII. — Digestion - * - - - - - 132 Chapter VIIL — Chyhfication - - - - - 148 21991 VI CONTENTS. Chapter IX.— L.UTKAL Absoiiptio.v - ClIAPTEIl X. CincCLATIOX § 1. Diffused Circulation 2. Vascular Circulation - 3. Respiratory Circulation - 4. Distributionof Blood Vessels CHAPTEnXI.-— Rf.si'iiiatiojt . . , _ § 1. Respiration in g-eneral 2. Aquatic Respiration - . . o. Atmospheric Respiration 4. Chemical Chang^cs effected by Respiration Chapter XII.— Seciietiox Chapteii XIII. — AnsoiiPTiojr Chapter XIV.— Nervous Power rage 164 167 167 170 191 201 208 208 210 221 236 243 250 252 PART III.— THE SENSORIAL FUNCTIONS. Chapter I. — Sensatiout - . . , Chapter II. — Touch Chapter III.— Taste Chapter IV.— Sxell Chapter V — Hearing § 1. Acoustic Principles - 2. Physiology of Hearing- in Man 3. Comparative Physiology of Hearing - Chapter Vf.— Vision § 1. Object of the Sense of Vision 2. Modes of accomplishing the objects of Vision 3. Structureof the Eye - 4. Physiology of Perfect Vision 5. Compai-ativc Physiology of Vision . Chapter VII.— Pekceptiox - - 258 268 - 279 281 - 294 294 - 298 308 - 315 315 - 318 325 - 332 ' )7 oot 558 CONTENTS. Vii Page Chapter VIII. — Comparative Phtsiolooy of the Nervous System 378 §1. Nervous System of Invcrteb rated Animals - - 378 2. Nervous System of Vertebrated Animals - - - 388 3. Functions of the Brain .... 395 4. Comparative Physiolog-y of Perception - - - 398 PART IV.— THE REPRODUCTIVE FUNCTIONS. Chapter I. — Reproduction ..... 408 Chapter II. — Organic Development . - • . 420 Chapter HI. — Decline of the System . - . . 433 Chapter IV. — Unity of Design - - . . . 437 Index - - - - - - - * - 449 $ fMfEKTr UBMART SS. C State Colkse ANIMAL AND VEGETABLE PHYSIOLOGY. PART II. THE VITAL FUNCTIONS. CHAPTER I. " OBJECTS OF NUTRITION. The mechanical structure and properties of the organized fabric, which have occupied our attention in the preceding volume, are necessary for the maintenance of life, and tlie exercise of the vital powers. But, however artificially that fabric may have been constructed, and however admirable the skill and the foresight that have been displayed in en- suring the safety of its elaborate mechanism, audi in pre- serving the harmony of its complicated movements, it yet of necessity contains within itself the elements of its own dis- solution. The animal machine, in common with every other mechanical contrivance, is subject to wear and deteriorate by constant use. Not only in the greater movements of the limbs, but also in the more delicate actions of the internal organs, we may trace the operation of miany causes inevita- bly leading to their ultimate destruction. Continued friction must necessarily occasion a loss of substance in the harder Vol. II. 2 10 THE VITAL FUNCTIONS. pnrts of the frame, and evaporation is constantly tending to exhaust the fluids. The repeated actions of the muscles in- duce certain changes in these organs, both in their mecha- nical properties and chemical composition, which impair their powers of contraction, and which, if suffered to con- tinue, would, in no long time, render them incapable of ex- ercising their proper functions: and the same observation applies also to the nerves, and to all the other systems of organs. Provision must accordingly be made for remedy- ing these constant causes of decay by the supply of those peculiar materials, which the organs require for recruiting their declining energies. It is obvious that the development of the organs, and ge- neral growth of the body, must imply the continual addition of new particles from foreign sources. Organic increase consists not in the mere expansion of a texture previously condensed, and the filling up of its interstices by inorganic matter: but the new materials that are added must, for this purpose, be incorporated with those which previously exist- ed, and become identified with the livinjr substance. Thus, we often find structures forming in the bodies of animals of a nature totally different from that of the part from which they arise. In addition to these demands, a store of materials is also wanted for the reparation of occasional injuries, to which, in the course of its long career, the body is unavoidably ex- posed. Like a ship fitted out for a long voyage, and forti- fied against the various dangers of tempests, of icebergs, and of shoals, the animal system, when launched into existence, should be provided with a store of such materials as may be wanted for the repair of accidental losses, and should also contain within itself the latent source of those enerjiies, which may be called into action when demanded by the ex- igencies of the occasion. Any one of the circumstances above enumerated would of itself be sufficient to establish the necessity of supplies of nourishment for the maintenance of life. But there are OBJECTS OF NUTRITION. 11 other considerations, equally important in a physiological point of view, and derived from the essential nature of or- ganization, which also produce a continual demand for these supplies; and these I shall now endeavour briefly to explain. Constant and progressive change appears to be one of the leading characteristics of life; and the materials which are to be endowed with vitality must therefore be selected and arranged with a view to their continual modification, cor- responding to these ever varying changes of condition. The artificer, whose aim is to construct a machine for perma- nent use, and to secure it as much as possible from the de- terioration arising from friction or other causes of injury, would, of course, make choice for that purpose of the most hard and durable materials, such as the metals, or the denser stones. In constructing a watch, for instance, he would form the wheels of brass, the spring and the barrel-chain of steel; and for the pivot, where the motion is to be inces- sant, he would employ the hardest of all materials, — the diamond. Such a machine, once finished, being exempt from almost every natural cause of decay, might remain for an indefinite period in the same state. Far different are the objects which must be had in view in the formation of or- ganized structures. In order that these may be qualified for exercising the functions of life, they must be capable of continual alterations, displacements, and adjustments, vary- ing perpetuall}'^, both in kind and in degree, according to the progressive stages of their internal development, and to the different circumstances which may arise in their exter- nal condition. The materials which nature has employed in their construction, are, therefore, neither the elementary bodies, nor even their simpler and more permanent combi- nations; but such of their compounds as are of a more plastic nature, and which allow of a variable proportion of ingre- dients, and of great diversity in the modes of their combi- nation. So great is the complexity of these arrangements, that although chemistry is fully competent to the analysis of organized substances into their ultimate elements, no hu- 12 THE VITAL FUNCTIONS. man art is adequate to effect their reunion in the same state as that in which they had existed in those substances; for it was by the 'refmcd operations of vitality, the only power that could produce this adjustment, that they have been brought into that condition. We may take as an example one of the simplest of orga- nic products, namely, Sugar; a substance which has been analyzed with the greatest accuracy by modern chemists: yet to reproduce this sugar, by the artificial combination of its simple elements, is a problem that has hitherto baffled all the efforts of philosophy. Chemistry, notwithstanding the proud rank it justly holds among the physical sciences, and the noble discoveries with which it has enriched the arts; notwithstanding it has unveiled to us many of the secret ope- rations of nature, and placed in our hands some of her most powerful instruments for acting upon matter; and notwith- standing it is armed with full powers to destroy, cannot, in any one organic product, rejoin that w^hich has been once dissevered. Through the medium of chemistry we are ena- bled, perhaps, to form some estimate of the value of what we find executed by other agencies; but the imitation of the model, even in the smallest part, is far beyond our power. No means w^hich the laboratory can supply, no process, which the most inventive chemist can devise, have ever yet approached those delicate and refined operations which na- ture silently conducts in the organized texture of living plants and animals. The elements of organic substances are not very nume- rous; the principal of them being oxygen, carbon, hydrogen, nitrogen, sulphur, and phosphbrus, together with a few of the alkaline, earthy, and metallic bases. These substances are variously united, so as to form certain specific com- pounds, wdiich, although they are susceptible, in different instances, of endless modifications, yetf possess such a gene- ral character of uniformity, as to allow of their being ar- ranged in certain classes; the most characteristic substance in each class constituting what is called a proximate orga- ORGANIC CHEMISTRY. ' 13 7iic principle. Thus, in the vegetable kingdom we have Lignin, Tannin, Mucilage, Vil, Sitgar, Fecula, &c. The animal kingdom, in like manner, furnishes Gelatin, Mhu- men, Fihnn, Mucus, Entomoline, Elearin, Stearin, and many others. The chemical constitution of these organic products, formed, as they are, of but few primary elements, is strik- ingly contrasted with that of the bodies belonging to the mi- neral kingdom. The catalogue of elementary, or simple bodies, existing in nature, is, indeed, more extensive than the list of those which enter into the composition of animal or vegetable substances. But in the mineral world they occur in simpler combinations, resolvable, for the most part, into a few definite ingredients, which rarely comprise more than two or three elements. In organized products, on the other hand, although the total number of existing elements may be smaller, yet the mode of combination in each sepa- rate compound is infinitely more complex, and presents in- calculable diversity. Simple binary compounds are rarely ever met with; but, in place of these, w^e find three, four, five, or even a greater number of constituent elements ex- isting in very complicated states of union. This peculiar mode of combination gives rise to a remark- able condition, which attaches to the chemical properties of organic compounds. The attractive forces, by which their several ingredients are held together, being very numerous, require to be much more nicely balanced, in order to retain them in combination. Slight causes are sufficient to disturb, or even overset, this equipoise of affinities, and often pro- duce rapid changes of form, or even complete decomposi- tion. The principles, thus retained in a kind of forced union, have a constant tendency to react upon one another, and to produce, from slight variations of circumstances, a to- tally new order of combinations. Thus, a degree of heat, which would occasion no change in most mineral substances, will at once effect the complete disunion of the elements of an animal or vegetable body. Organic substances are, in 14 ' THE VITAL FUNCTIONS. like manner, unable to resist the slower, but equally destruc- tive agency of water and atmospheric air; and they are also, liable to various spontaneous changes, such as those consti- tuting fermentation and putrefaction, which occur when their vitality is extinct, and when they are consequently aban- doned to the uncontrolled operation of their natural chemi- cal affinities. This tendency to decomposition may, indeed, be regarded as inherent in all organized substances, and as requiring for its counteraction, in the living system, that perpetual renovation of materials which is supplied by the powers of nutrition. It would appear that, during the continuance of life, the progress of decay is arrested at its very commencement; and that the particles, which first undergo changes unfitting them for the exercise of their functions, and which, if suffered to remain, would accelerate the destruction of the adjoining par^are immediately removed, and their place supplied by parWles tliat have been modified for that purpose, and which, when they afterwards lose these salutary properties, are, in their turn, discarded and replaced by others. Hence, the continued interchange and renewal of particles which take place in the more active organs of the system, especial- ly in the higher classes of animals. In the fabric of those animals which possess an extensive system of circulating and absorbing vessels, the changes that are effected are so considerable and so rapid, that even in the densest textures, such as the bones, scarcely any portion of the substance which originally composed them is permanently retained in their structure. To so great an extent is this renovation of materials carried on in the human system, that doubts may very reasonably be entertained as to the identity of any por- tion of the body after the lapse of a certain time. The pe7 riod assigned by the ancients for this entire change of the substance of the body, was seven or eight years: but modern inquiries, which show us the rapid reparation that takes place in injured parts, and the quick renewal of the bones themselves, tend to prove that even a shorter time than this ORGANIC CHEMISTRY. 15 is adequate to the complete renovation of every portion of the living fabric* Imperfect as is our knowledge of organic chemistry, we see enough to convince us that a series of the most refined and artificial operations is required, in order to bring about the complicated and elaborate arrangements of elements which constitute both animal and vegetable products. Thus, in the very outset of this, as of every other inquiry in Phy- siology, we meet with evidences of profound intention and consummate art, infinitely surpassing not only the power and resources, but even the imagination of man. Much as the elaborate and harmonious mechanism of an animal body is fitted to excite our admiration, there can be no doubt that a more extended knowledge of that series of subtle processes, consisting of chemical combinations and de- compositions which are continually going on in the organic laboratory of living beings, would reveal still greater won- ders, and would fill us with a more fervent admiration of the infinite art and prescience which are even now manifest- ed to us in every department both of the vegetable and ani- mal economy. The processes by which all these important purposes are fulfilled, comprise a distinct class of functions, the final ob- ject of which may be termed Nutrition, that is, the repara- tion of the waste of the substance of the organs, their main- tenance in the state fitting them for the exercise of their respective offices, and the application of properly prepared materials to their development and growth. The functions subservient to nutrition may be distinguished according as the processes they comprise relate to seven principal periods in the natural orders of their succession. The first series of processes has for its objects the reception of the materials from without, and their preparation and gradual conversion into proper nutriment, that is, into mat- ter having the same chemical properties with the substance * See the article «' Age " in the Cyclopedia of Practical Medicine, where I have enlarged upon this subject. 16 THE VITAL FUNCTIONS. of the organs with which it is to he incorporated; and their purpose being to assimilate the food as much as possible to the nature of the organic body it is to noiTrish, all these functions have been included under the term Assimila- tion. The second scries of vital functions comprise those which are designed to convey the nutritive fluids thus elaborated, to all the organs that are to be nourished by them. In the more developed systems of organization this purpose is ac- complished by means of canals, called vessels, through which the nutritive fluids move in a kind of circuit: in this case the function is denominated the Circulation. It is not enough that the nutritive juices are assimilated: another chemical process is still required to perfect their ani- malization, and to retain them in their proper chemical con- dition for the purposes of the system. This third object is accomplished by the function oi Respiration. Fourthly, several chemical products, which are wanted in different parts of the economy, are required to be formed by a peculiar set of organs, of which the intimate structure eludes observation; although we may perceive that in many instances among the higher orders of beings, a special appa- ratus of vessels sometimes spread over the surface of a membrane, at other times collected into distinct masses, is provided for that purpose. These specific organs are termed glands, and the office performed by them, as well as by the simpler forms of structure above mentioned, is termed Se- cretion. Fifthly, similar processes of secretion are also employed to carry off from the blood such animal products as may have been formed or introduced into it, and may possess or have acquired noxious properties. The elimination of these materials, which is the office of the excretories, constitutes the function of Excretion. Sixthly, changes may take place in various parts of the body, both solid and fluid, rendering them unfit to remain in their present situation, and measures must be taken for POWERS OF ASSIMILATION. 17 the removal of these useless or noxious materials, by trans- ferring them to the general mass of circulating IdIoocI, so as either to be again usefully employed, or altogether discard- ed by excretion from the system. This object is accom- plished by a peculiar set of vessels; and the function they perform is termed Msorjjtion. Lastly, the conversion of the fluid nutriment into tlic solids of the body, and its immediate application to the purposes of the development of the organs, of their preser- vation in the state of health and activity, and of the repair of such injuries as they may chance to sustain, as far as the powers of the system are adequate to such reparation, are the objects of a seventh set of functions, m.ore especially comprised under the title of Nutrition, which closes this long series of chemical changes, and this intricate but har- monious system of operations. Although the order in which the constituent elements of organized products are arranged, and the mode in which they are combined, are entirely unknown to us, we can ne- vertheless perceive that in following them successively from the simplest vegetables to the higher orders of the animal Idngdpm, they acquire continually increasing degrees of complexity, corresponding, in some measure, to the greater refinement and complication of the structures by which they have been elaborated, and of the bodies to which they are ultimately assimilated. Thus, plants derive their nourish- ment from the crude and simple materials which they ab- sorb from the earth, the waters,, and the air that surround them; materials which consist almost wholly of water, with a small proportion of carbonic acid, and a few saline ingre- dients, of which that water is the vehicle. But these, after having been converted by the powers of vegetable assimila- tion, into the substance of the plant, acquire the character- istic properties of organized products, though they are still the simplest of that class. In this state, and when the fabric they had composed is destroyed, and they are scattered over the soil, they are fitted to become more highly nutn- VoL. 11. 3 18 THE VITAL FUNCTIONS. tive to other plants, which absorb them, and with more fa- cility adapt them to the purposes of their own systems. Here they receive a still higher degree of elaboration; and thus the same materials may pass through several successive series of modifications, till they become the food of animals, and are tlien made to under2;o still farther chansces. New elements, and in j^articulnr nitrogen, is added to the oxy- gen, hydrogen and carbon, which are the chief constituents of vegetable substances:* and new properties are acquired, from the varied combinations into which their elements are made to enter by the more energetic powers of assimilation appertaining to the animal system. The products which result are still more removed from their original state of inorganic matter: and in this condition they serve as the appropriate food of carnivorous animals, which generally hold a higher rank in the scale of organization, than those that subsist only on vegetables. Thus has each created beino; been formed in reference, not merely to its own welfare, but also to that of multitudes of others which arc dependent on it for their support, their pre- servation,— nay, even for their existence. In contemplating this mutual relationship, this successive subordination of the different races to one another, and this continual tendency to increased refinement, we cannot shut our eyes to the mag- nificent unfolding of the great scheme of nature for the pro- gressive attainment of higher objects ; imtil, in the pqrfcct sys- tem, and exalted endowments of man, we behold the last re- sult that has been manifested to us of creative power. • Nitrog-cn, however, frequently enters into the composition of veg-eta- blcs: though, in general, in a much smaller proportion than into the sub- stance of animals, of which lust it always appears to be an essential constitu- ent. ( la ) CHAPTER II. NUTRITION IN VEGETABLES. §1. Food of Plants. The simplest kind of nutrition is that presented to us by the vegetable kingdom, where water may be considered as the general vehicle of the nutriment received. Before the discoveries of modern chemistry, it was very generally be- lieved that plants could subsist on w^ater alone ; and Boyle, and Van Helmont, in particular, endeavoured to establish, by experiment, the truth of this opinion. The latter of these physiologists planted a willow in a certain quantity of earth, the weight of which he had previously ascertained with great care ; and, during five years, he kept it moistened with rain- water alone, which he imagined was perfectly pure. At the end of this period, he found that the earth had scj^cely di- minished in weight, while the w^illow had grown into a tree, and had acquired an additional weight of one hundred and fifty pounds : whence he concluded that the w^ater had been the only source of its nourishment. But it does not seem to have been, at that time, known, that rain-water always con- tains atmospheric air, and frequently, also, other substances, and that it cannot, therefore, be regarded as absolutely pure water: nor does it appear that any precautions were taken to ascertain that the water actually employed was w^holly free from foreign matter, which, it is easy to conceive, it might have held in solution. In an experiment of Duhamel, on the other hand, a horse-chesnut tree and an oak, exposed to the open air, and watered with distilled water alone, the former for three, and the latter for eight years, were kept alive, in- JO THE VITAL FUNCTIONS. «?^ pouches, or caica; which, as they are traced along the canal, hecome both wider and longer, so that the tenth pair of caeca (a) extends to the hinder extre- mity of the animal; tlie intestine (i,) which is very short, lying between them.* It has long been known, that if, after the leech has fastened on the skin, a portion of the tail be cut off, the animal will continue to suck blood for an indefinite time: this arises from the circumstance, that the caecal portions of the stomach are laid open, so that the blood received into that cavity flows out as fast as it is swallowed. i^'V'^ A structure very similar to that of the leech is met with in the digestive organs of the Glosso- 264 po7^a tubercxilata, (Ilirudo com- planata, Linn.) of which Fig. 263 represents a magnified view from the upper side. When seen from the under side, as is shown in Fig. 264,*thc cavity of the stomach is distinctly seen, prolonged into several cells, divided by partitions, and directed towards the tail. The two last of these cells (c c) • This figure was engraved from a drawing made, at my request, by Mr. Newport, from a specimen which he dissected, and which he was so oblig-ing as to sliow me. Fig. 261 represents the mouth, within wlilch are seen the three teeth; and Fig. 262, one of the teeth detached. A paper, descriptive of the structure of the stomach of the leech, by Mr. Newport, was lately read NUTRITION IN THE ANNELIDA. 79 are much longer than the rest, and terminate in two blind sacs, between which is situated a tortuous intestinal tube.* at a meeting- of the Royal Society. See the abstracts of the proceedings in the Society, for June, 1833. * In both these figures, t is the tubular tongue, projected from tlie mouth. In Fig, 263, e are the six eyes, situated on tlie extremity whicli coiresponds to the head; and a double longitudinal row of white tubercles is also visible, extending along the back of the animal, e, in Fig. 264, is the entrance into a cavity, or pouch, provided for the reception of the young. See Johnson, Phil. Trans, for 1817, p. 343, ( so ) CHAPTER V. Nutrition in the highe)' oi^dcrs of Animals. In proportion as we rise in the animal scale, we find that the operations of Nutrition become still farther multiplied, and that the organs which perform them are more numerous and more complicated in their structure. The long series of processes requisite for the perfect elaboration of nutri- ment, is divided into different stages; each process is the work of a separate apparatus, and requires the influence of different agents. We no longer find that extreme simplicity which we noticed as so remarkable in the hydra and the medusa, where the same cavity performs, at once, the func- tions of the stomach and of the heart. The manufacture of nutriment, if we may so express it, is, in these lower zoo- phytes, conducted upon a small scale, by less refined me- thods, and with the strictest economy of means; the appara- tus is the simplest, the agents the fewest possible, and many different operations are carried on in one and the same place. As we follow the extension of the plan in more elevated stages of organic development, we find a farther division of labour introduced. Of this we have already seen the com- mencement in the multiplication of the digesting cavities of the Leech and other Annelida: but, in animals which occupy a still higher rank, we observe a more complete separation of offices, and a still greater complication of organs. The principle of the division of labour is carried to a much great- er extent than in the inferior departments of the animal creation. Besides the stomach, or receptacle for the unas- similated food, another organ, the heart, is provided for the COMPLEX APPARATUS FOR NUTRITION. SI uniform distribution of the nutritious fluids elaborated by the organs of digestion. This separation of functions, again, leads to the introduction of another system of canals or ves- sels, for transmitting the fluids from the organs which pre- pare them to the heart, as into a general reservoir. In the higher orders of the animal kingdom, all these processes are again subdivided and varied, according to the species of food, the habits and mode of life, assigned by nature to each in- dividual species. For the purpose of conveying clearer no- tions of the arrangement of this extensive system of vital organs, I have drawn the annexed plan (Fig. 265,) which ^ • exhibits them in their natural order of connexion, and as they might be supposed to appear in a side view of the in- terior of a quadruped. To this diagram I shall make fre- quent reference in the following description of this sys- tem. The food is, in the first place, prepared for digestion by several mechanical operations, which loosen its texture and destroy its cohesion. It is torn asunder and broken down by the action of the jaws and of the teeth; and it is, at the same time, softened by an admixture with the fluid secre- tions of the mouth. It is then collected into a mass, by the action of the muscles of the cheek and tongue, and swal- VOL. II. II S2 THE VITAL rUXCTIOXS. lowed by the regulated contractions of the different parts of the throat. It now passes along a muscular tube, called the Oesophagus, (represented in the diagram by the letter o,) into the stomach, (s,) of which the entrance (c) is called the cardia. In the stomach the food is made to undergo various che- mical changes; after which it is conducted through the aper- ture termed the pi/lninis (r,) into the canal of the intestine (i I,) where it is farther subjected to the action of several fluid secretions derived from large glandular organs situated in the neiglil)ourhood, as the liver (l) and the pancreas; and elaborated into the fluid which is termed Chyle. The Chyle is taken up by a particular set of vessels, called the Lacleals, which transmit it to the heart (ii.) These vessels are exceedingly numerous, and arise by open orifices from the inner surface of the intestines, whence they ab- sorb, or drink up the chyle. They may be compared to internal roots, which unite as they ascend along the mesen- tery (m,) or membrane connecting the intestines with the back; forming larger and larger trunks, till they terminate in an intermediate reservoir (r,) which has been named the Ixeccptacic of the Chyle. From this receptacle there pro- ceeds a tube, which, from its passing through the thorax, is called the Thoracic duct (t;) it ascends along the side of the spine, which protects it from compression, and opens at V, into the large veins which are pouring their contents into tlie auricle, or first cavity of the heart (u.) whence it immediately passes into the ventricle, or second cavity of that organ (ii.) Suc|), in the more perfect animals, is the circuitous and guarded route, which every particle of nou- rishment must take before it can be added to the general mass of circuhiting fluid. By its admixture with the blood already contained in these vessels, and its purification by the action of the air in the respiratory organs (i?,) the chyle becomes assimilated, and is distributed by the heart through appropriate'channels of circulation called arteries (of which the common trunk, COMPLEX APPARATUS FOR NUTRITION. 83 or Aorta^ is seen at A,) to every part of the system; thence returning by the veins (v, v^ v,) to the heart. The various modes in which these functions are conducted in tlie seve- ral tribes of animals will be described hereafter. It will be sufficient for our present purpose to state, by way of com- pleting the outline of this class of functions, that, like the returning sap of plants, the blood is made to undergo far- ther modifications in the minute vessels through which it circulates; new arrangements of its elements take place during its passage through the subtle organization of the glands, which no microscope has yet unravelled: new pro- ducts are here formed, and new properties acquired, adapted to the respec^e purposes which they are to serve in the animal economy. The whole is one vast Laboratory, where mechanism is subservient to Chemistry, where Chemistry is the agent of the higher powers of Vitality, and where these powers themselves minister to the more exalted fiicul- ties of Sensation and of Intellect. The digestive functions of animals, however complex and varied, and however exquisitely contrived to answer their particular objects, yet afford less favourable opportunities of tracing distinctly the adaptation of means to the respective ends, than the mechanical functions. This arises from the circumstance that the processes they efTect imply a refined chemistry, of which we have as yet but a very imperfect knowledge; and that we are also ignorant of the nature of the vital agents concerned in producing each of the chemi- cal changes which the food must necessarily undergo during its assimilation. We only know that all these changes are slowly and gradually effected; the materials having to pass through a great number of intermediate stages before they can attain their final state of elaboration. Hence we are furnished with a kind of scale, whereby, whenever we can ascertain the degrees of difference exist- ing between the chemical condition of the substance taken into the body, and that of the product derived from it, we may estimate the length of the process required, and the S4 THE VITAL FUNCTIONS. amount of power necessary for its conversion into that pro- duct. It is obvious, for example, that the chemical changes which vciictable food must be made to under2;o, in order to assimilate it to blood, must be considerably greater than tiiuse recjuircd to convert animal food into the same fluid, because the latter is itself derived, with only slight modifi- cation, immediately from the blood. We accordingly find it to be an established rule, that the digestive organs of ani- mals which feed on vegetable materials are remarkable for their size, their length, and their complication, when com- pared witli those of carnivorous animals of the same class. This rule applies, indeed, universally to Mammalia, Birds, Reptiles, Fishes, and also to Insects: and below these we can scarcely draw the comparison, because nearly all the inferior tribes subsist wholly upon animal substances. Many of these latter animals have organs capable of extracting nourishment from substances which we should hardly ima- gine contained any sensible portion. Thus, on examining the stomach of the earth-worm, we find it always filled with moist earth, which is devoured in large quantities, for the sake of the minute portion of vegetable and animal materi- als that happen to be intermixed with the soil; and this slen- der nutriment is sufllcient for the subsistence of that ani- mal. Many marine worms, in like manner, feed apparently upon sand alone; but that sand is generally intermixed with fragments of shells, which have been pulverized by the con- tinual rolling of the tide and the surge; and the animal mat- ter contained in these fragments, aflbrds them a supply of nutriment adequate to their wants. It is evident, that when, as in the preceding instances, large quantities of indigestible materials are taken in along with such as are nutritious, the stomach and other digestive cavities must be rendered more than usually capacious. It is obvious also that the structure of the digestive organs must bear a relation to the mechani- cal texture, as well as the chemical qualities of the food; and this we lind to be the case in a variety of instances, which will hereafter be specified. COMPLEX APPARATUS FOR NUTRITION. 85 The activity of the digestive functions and the structure of the organs, will also be regulated by a great variety of other circumstances in the condition of the animal, inde- pendently of the mechanical or chemical nature of the food. The greater the energy with which the more peculiarly animal functions of sensation and muscular action are ex- ercised, the greater must be the demand for nourishment, in order to supply the expenditure of vital force created by these exertions. Compared with the torpid and sluggish reptile, the active and vivacious bird or quadruped requires and consumes a much larger quantity of nutriment. The tortoise, the turtle, the toad, the frog, and the chameleon, will, indeed, live for months without taking any food. Fishes, which, like reptiles, are cold-blooded animals, al- though at all times exceedingly voracious when supplied with food, yet can endure long fasts with impunity. The rapidity of development has also great influence on the quantity of food which an animal requires. Thus, the caterpillar, which grows very quickly, and must re- peatedly throw off its integuments, during its continuance in the larva state, consumes a vast quantity of food com- pared with the size of its body; and hence we find it pro- vided with a digestive apparatus of considerable size. ( 86 ) CHAPTER VI. 1 PREPARATION OF FOOD. § 1. Pi'chcnsion of Liquid Food. In studying the series of processes which constitute assi- mihition, our attention is first to be directed to the mode in which the food is introduced into the body, and to the me- chanical changes it is made to undergo before it is subjected to the chemical action of the digestive organs. The nature of these preliminary processes will, of course, vary accord- ins to the texture and mechanical condition of the food. Where it is already in a fluid state, mastication is unneces- sary, and the receiving organs consist simply of an appara- tus for suction. This is the case very generally with the Entozoa, which subsist upon the juices of other animals, and which arc all provided with one or more sucking ori- fices, often extended in tlic form of a tube or proboscis.* The Hydatid, for instance, has four sucking apertures dis- posed round the head of the animal: the Txnia has orifices of this kind in each of its jointed segments: the Jiscains and the Earih-ivorm have each a simple mouth. The margin of the mouth is often divided, so as to compose lips; of these there are generally two, and in the leech there are three. In some rare cases, as in the Flancuna, there is, besides the • Some species of Fasclolx, or flukes, are furnished with two, three, six, or more sucking- disks, by which they adhere to surfaces: to these animals the names Dlstonia, Trisioma, Hexastoma, and Pohjstnma have been g-iven; but these denominations, implyinjj a plurality of mouths, are evidently in- correct, since tlic suckintj disks are not perforated, and do not perform the office of mouths; and the true mouth for the reception of food is single. Cuvier discovered an animal of this class furnished with above a hundred of these cup-shaped sucking- organs. Sec Edinburgh Philos. Journal, xx. 101. PREHENSION OF LIQUID FOOD. 87 ordinary mouth, a tube also provided for suction, in a dif- ferent part of the body, and leading into the same stomach.* When the instrument for suction extends for some length from the mouth, it is generally termed a proboscis: such is the apparatus of the butterfly, the moth, the gnat, the house fly, and other insects that subsist on fluid aliment. The pro- boscis of the Lejjidoptera, (Fig. 266,) is a double tube, con- structed by the two edges being rolled longitudinally till they meet in the middle of the lower surface, thus form- ing a tube on each side, but leaving also another tube, intermediate to the two lateral ones. This middle tube is formed by the junction of two grooves, which, by the aid of a curious appara- tus of hooks, resembling those of the laminae of a feather already described,! lock into each other, and can be either united into an air tight canal, or be instantly separated at the pleasure of the animal. Reaumur conceives that the lateral tubes are intended for the reception of air, while the central canal is that which conveys the honey, which the insect sucks from flowers, by suddenly unrolling the spiral coil, into which the proboscis is usually folded, and darting it into the nectary.! In the Hemiptera, the proboscis is a tube, either straight or jointed, guarded by a sheath, and acting like a pump. The Diptera have a more complicated instrument for suc- tion, consisting of a tube, of which the sides are Mrong and fleshy, and moveable in every direction, like the trunk of an elephant: it has at its extremity a double fold, resemb- ling lips, which are well adapted for suction. The gnat, and other insects which pierce the skin of animals, have, for this purpose, instruments, termed, from their shape and ofiice, * Phil. Trans, for 1822, 442. •f Volume i. p. 393. \ Kirby and Spcnce's Entomology, vol. ii, p. 390. SS THE VITAL FUNCTIONS. lancets.* In the gnat, they are five or six in number, finer than a hair, exceedingly sharp, and generally barbed on one side. In the Tabamts, or horse-fly, they are flat like the blade of a knife. These instruments are sometimes con- structed so as to form, by their union, a tube adapted for suction. In the flesh-fly, the proboscis is folded like the letter Z, the upper angle pointing to the breast, and the lower one to the mouth. In other flies there is a single fold only. Those insects of the order -^ymc7zo/?/er«, which, like the bee, suck the honey of flowers, have, together with regular jaws, a proboscis formed by the prolongation of the lower lip, which is folded so as to constitute a tube: this tube is protected by the mandibles: and is projected forwards by being carried on a pedicle, which can be folded back when the tube is not in use. The mouths of the Jicephaloits Mol- lusca are merely sucking apertures, with folds like lips, and without either jaws, tongue, or teeth, but having often ten- tacula arising from their margins. Among fishes, we meet with the family of Cydosiomata, so called from tJicir having a circular mouth, formed for suction. The margin of this mouth is supported by a ring of cartilage, and is furnished with appropriate muscles for producing adhesion to the surfaces to which it is applied; the mechanism and mode of its attachment being similar to that of the leech. To this family belong the Myxine and the Lamprey. So great is the force of adhesion exerted by this sucking apparatus, that a lamprey has been raised out of the water with a stone, weighing ten or twelve pounds, adhering to its mouth. Humming birds have a long and slender tongue, which can assume the tubular form, like that of the butterfly or the bee, and for a similar purpose, namely, sucking the juices of flowers. Among the mammalia, the Vampire Bat affords another instance of suction by means of the tongue, which ♦ Kirby and Spcncc's Entomology, vol. ill. p. 467'. PREHENSION OP SOLID FOOD. 89 is said to be folded into a tubular sbapc for that purpose. But suction among the mammalia is almost always performed by the muscles of the lips and cheeks, aided by the move- ments of the tongue, which, when withdrawn to the back of the cavity, acts like the piston of a pump. In the lamprey this hydraulic action of the tongue is particularly remarka- ble. Many quadrupeds, however, drink by repeatedly dip- ping their tongue into the fluid, and quickly drawing it into the mouth. § 2. Prehension of Solid Food. When the food consists of solid substances, organs must be provided; first, for their prehension and introduction into the mouth; secondly, for their detention when so introduced; and thirdly, for their mechanical division into smaller frag- ments. Of those instruments of prehension which are not portions of the mouth itself, and which form a series of variously constructed organs extending from the tentacula of the po- lypus to the proboscis of the elephant, and to the human arm and hand, some account has already been given in the his- tory of the mechanical functions: but, in a great number of instances, prehension is performed by the mouth, or the parts which are extended from it, and maybe considered as its appendices. The prehensile power of the mouth is de- rived principally from the mechanical form and action of the jaws, which open to receive, and close to detain the bodies intended as food; and to this latter purpose, the teeth, when the mouth is furnished with them, likewise materially contribute, although their primary and more usual office is the mechanical division of the food, by means of mastication, an action in which the jaws, in their turn, co-operate. Ano- ther principal purpose effected by the jaws is that of giving mechanical jiower to the muscles, which, by acting upon the sides of the cavity of the mouth, tend to compress and Vol. II. 12 90 THE VITAL FUNCTIONS. propel tlifi contained food. We find, accordingly, that all animals of a highly developed structure arc provided with jaws. Among the animals which are ranked in the class of Zoo- phytes the highest degrees of development arc exhibited by the Kchinodermata, and in them we fmd a remarka- l)le jM'rfection in tlu; organs of mastication. The mouth of the Echinus is surroumled hy a frame-work of shell, con- sisting of five converging pieces, each armed with a long tooth; and for the movement of each part there are provided sei)arate muscles, of whicli the anatomy has been minutely described by Cuvier. In the shells of the echini that arc cast on the shore, this calcareous frame is usually found en- tire in the inside of the outer case; and Aristotle having noticed its resemblance to a lantern, it has often gone by the whimsical name of the lantern of Jiristotlc. In all articulated animals which subsist on solid aliment, the apparatus for the prehension and mastication of the food, situated in the mouth, is exceedingly complicated, and ad- mits of great diversity in the dillcrent tril)es; and, indeed, the number and variety of the parts of which it consists is so great, as hardly to admit of being comprehended in any general description. In most insects, also, their minuteness is an additional obstacle to the accurate observation of their anatomy, and of the mechanism of their action. The re- searches, however, of Savigny,* and other modern entomo- logists, have gone far to prove, that, amidst the infinite va- riations observable in the form and arrangement of the se- veral parts of these organs, there is still ])reserved, in the general plan of their construction, a degree of uniformity quite as great as that which has been remarked in the fabric of the vertebrated classes. Not only may we recognise, in every instance, the same elements of structure, but we may also trace regular chains of gradation, connecting forms ap- • Sec l)is '♦Thcoric cks Organcs de la bouche des Aniniaux invcrtcbrcs et articiiles," which forms the first part of the "Mcnioircs sur Ics Animaux sans vcrtt'brcs." Paris, 181G. JAWS OF THE ECHINUS. 91 parently most remote, and organs destined for widely dif- ferent uses: so that even when there lias l)een a complete change of purpose, we still perceive the same design fol- lowed, the same model copied, and the same uniformity of plan preserved in the construction of the organs of every kind of mastication; and there prevails in tiicm the same unity of system as is displayed in so marked a manner in the conformation of the organs of progressive motion. The jaws, which, in one tribe of insects, are formed for breaking down and grinding the harder kinds of food, are, in another, fitted for tearing asunder the more tough and fd^rous tex- tures; they are fashioned, in a third, into instruments for taking up the semi-fluid honey prepared by flowers; while, again, in a fourth, they are prolonged and folded into a tu- bular proboscis, capable of suction, and adapted to the drink- ing; of fluid aliment. Pursuino; the examination of these or- gans in another series of articulated animals, we And them gradually assuming the characters, as well as the uses, of instruments of prehension, of weapons for warfare, of pillars for support, of levers for motion, or of limbs for quick pro- gression. Some of these remarkable metamorphoses of or- gans have already attracted our attention, in a former part of this treatise.'* Jaws pass into feet, and feet into jaws, through every intermediate form; and the same individual often exhibits several steps of these transitions; and is some- times provided also with supernumerary organs of each de- scription. In the Arachnida, in particular, we frequently meet with supernumerary jaws, together with various ap- pendices, which present remarkable analogies of form with the attennse, and the legs and feet of the Crustacea. The principal elementary parts which enter into the com- position of the mouth of an insect, when in its most perfect state of development, are the seven following: a pair of up- per jaws, a pair of lower jaws, an upper and a lower lip, and a tongue.t These parts in the Locusta vhHdisswia, or com- ♦ Vol. i. p. 206. f All these parts, taken together, were termed by Fabriclus ijistrumcnla 92 THE VITAL FUNCTIONS. mon grasshopper, arc delineated in llieir relative situations^ but detached from one another, in lig. 267. The upper jaws, {m,) which are termed the mandibles, are those prin- cipally employed for the mastication of hard substances; they are, accordingly, of greater strength than the lower jaws, and thcii- edges are generally deeply serrated, so as to act like teeth in dividing and bruising the food. Some of these teeth are pointed, others wedge-shaped, and others broad, like grinders; their form being, in each particular case, adapted to the mechanical texture of the substances to which they arc designed to be applied. Thus, the mandi- bles of some Melolonthx have a projection, rendered rough by numerous deep transverse furrows, converting it into a file for wearing down the dry leaves, w'hich are their natural food."* In most cases, indeed, we are, in like manner, ena- bled, from a simple inspection of the shape of the teeth, to clbarla; and upon their varieties of structure he founded his cclcbnitcd sys- tem of entomological classification. Kirby and Spence have denominated them iropfiL Sec their introduction to Entomology, vol. iii. p. 417. To the seven elements above enumerated, Savigny adds, in the Hcmlptcra, an eighth, whicli he terms the Epv^lotasu. * Knoch, quoted by Kirby. TAWS OF INSECTS. 93 forin tolerably accurate ideas of the kind of food on which the insect naturally subsists.* Above, or rather in front of the mandibles, is situated the labrum, or upper lip (u.) It is usually of a hard or horny texture, and admits of some degree of motion: but its form and direction are exceedingly various in different tribes of insects. The lower pair of jaws (j,) or maxillx, as they have been termed, are behind the mandibles, and between them is situated the labium, or lower lip (l,) which closes the mouth below, as the lahrum does above. In the grass- hopper, each maxilla consists of an outer and an inner plate (o and I.) The jaws of insects are confined, by their arti- culations with the head, to motions in a horizontal plane only, so that they open and close by a lateral movement, and not vertically upwards and downwards, as is the case with the jaws of vertebrated animals. The maxillcC are, in most cases, employed principally for holding the substances on which the dividing or grinding apparatus of the mandibles is exerted. A similar use may be assigned also to the or- gans denominated Palpi, or Aiitennulx (p, q,) which are jointed filaments, or processes, attached to different parts of the mouth, and most usually to the maxillae and the labium; the former (p) being termed the maxillay^y , and the latter (q) the labial palpi. In addition to these parts, another, which, from its supposed use, has been denominated Glossa^ or tongue (g,) is also generally found. For an account of the various modifications which these parts receive in different tribes and species, I must refer to works which treat professedly of this branch of comparative anatomy. I shall content myself with giving a single exam- ple of the conversion of structure here alluded to, in that of the I'ostrum, or proboscis of the Cimex nigricoriiis. This insect belongs l(\ the order Hemiptera, which has been usually characterized as being destitute of both mandibles * See a memoir by Marcel tics Scrrcs, in the Annalcs du MusC'um d'llist. Nat. xiv. 56. 94 THE VITAL FUNCTIONS. and jaws, and as having, instead of these parts, an apparatus of very different construction, designed to pierce the skin of animals, and suck their juices, l^ut Savigny, on applying the j)rinciples of his theory, has recognised, in the proboscis of the Cimex, the existence of all the constituent elements that are found in the mouth of insects formed for the masti- cation of solid food. This proboscis consists of four clon- sated filaments, contained in a kind of sheath: these fila- ments are represented in Fig. 2GS, se- ])arated to a little distance from each other, in order that their respective origins may be distinctly seen; the one set (q) being prolongations of the man- dibles (j,) and the other set (p) being, in like manner, prolongations of the maxillae (m.) Between these filaments, and near their commencement, is seen a pointed cartilaginous body (g,) which is the glossa, or tongue; and the ajier- ture seen at its root is the passage into the oesophagus. The sheath is merely the elongated labium, of which the base is seen at l, in Fig. 2GS; but is represented, in its whole length, in Fis;. 2G9, where the £!;roove for con- P \q W taining the filaments above described, \ is apparent. \ ^^ In the mouths of the Annelida we often meet with hard bodies, which serve the purposes of jaws and of teeth. The"retractile proboscis of the Jlphro- dile, or sea-mouse, is furnished with four teeth of this de- scription. The Leech has, immediately within its lips, three semi-circular teeth, with round and sharp cutting edges: they are delineated in P^ig. 262, in their relative po- sitions; and Fig. 2G3 represents one of the teeth detached from the rest. It is with these teeth that the leech pierces tJie skin of tiie animals whose blood it sucks; and, as soon as JAWS OF INSECTS. 95 the wound is inflicted, tlic teeth, heing moveahle at their base, fall back, leavinp; the openini^ of the mouth free for sucking. The wound thus made is of a peculiar form, being composed of three lines, radiating from a centre, where the three teeth had penetrated. Most of the Mollusca which inhabit univalve shells are provided with a tubular organ, of a cylindric or conical shape, capable of elongation and contraction, by circular and longitudinal muscular fibres, and serving the purpose of a proboscis, or organ of prehension, for seizing and conveying food into the mouth. These tubes are of great size in the Bucciniim, the Miirex, and the Valuta, as also in the Doris, which, though it has no shell, is likewise a gastero- pode. In those mollusca of this order which have not a proboscis, as the Limax, or slug, the Helix, or snail, and the ^plysia, or sea-hare, the mouth is furnished with broad 270 lips, and is supported by an internal cartilage, iSfl^ having several tooth-like projections, which assist in laying hold of the substances taken as food. That of the snail is represented in Fig. 270. All the Sepiss, or cuttle fish tribe, are furnished, at the entrance of the mouth, with two horny jaws, having a re- markable resemblance to the bill of a parrot; excepting that the lower piece is the larger of the two, and covers the up- per one, which is the reverse of what takes place in the parrot. These constitute a powerful instrument for break- ing the shells of the molhisca and Crustacea, which compose the usual prey of these animals. Fishes almost always swallow their food entire, so that their jaws and teeth are employed principally as organs of prehension and detention; and the upper jaw, as well as the lower one, being moveable upon the cranium, they are ca- pable of opening to a great width. The bony pieces which compose the jaws are more numerous than the correspond- ing bones in the higher classes of vertebrata, and tlicy ap- pear, therefore, as if their development had not proceeded 96 THE VITAL FUNCTIONS. t sufTicicntly i-^x to effect tlieir consolidation into more com- pact structures.* Fishes which live upon other animals of the same class having a soft texture, are furnished with teeth constructed merely for seizing their prey, and perhaps also for slightly di- viding it, so as to adapt it to being swallowed. These teeth are of various shapes, though usually sharp at the points, and cither conical or hooked at the extremity, with the points al- wa\'s directed backwards, in order to prevent the escape of the animal which has been seized. Those fishes which subsist on testaceous mollusca have teeth with grinding surfaces, and their jaws arc also adapted for mastication. Every part of the mouth, tongue, and even throat, may afford lodgement for teeth in this class of animals. Almost the whole cavity of the mouth of tlie Anarrhichas lupus, or wolf-fish, may be said to be paved with teeth, a triple row being implanted on each side; so that this fish exerts great power in breaking shells. The Shark has numerous rows of sharp teeth, with serrated margins: these, at first sight, appear to be formidable instru- ments; but as the teeth in the opposite jaws do not meet, it is evident that they are not intended for cutting, like the incisors of mammalia. Among Reptiles we find the Batrachia almost wholly des- titute of teeth. Frogs, indeed, exhibit two rows of very fine points; the one in the upper jaw, and the other passing trans- versely across the palate: they may be considered as teeth existing in a rudimental state; for they are not sufficiently developed to be useful in mastication. There arc about forty of these minute teeth on each side in the frog. In the Salamander, there are sixty above and below; and also thirty on each side of the palate. The tongue of tlic frog is of great length; its root is at- tached close to the fore part of the lower jaw, while its point, • AUcmpts liavc been made to trace analogies between the difTerent seg- ments of the jaws of fishes and covrcsponding parts of the mouths of cms- tacea and of insects: but the justness of these analogies is yet far from being satisfactorily proved. JAWS OP BIRDS. 97 which is cloven, is turned backwards, extending into the throat and acting like a valve in closing the air passage into the lungs. If, when this animal has approached witliin a certain distance of the insect it is about to seize, we watch it with attention, we are surprised to observe the insect suddenly disappear, without our being able to perceive what has become of it. This arises from the froo; bavin"; darted out its tongue upon its victim with such extreme quickness, and withdrawn it with the insect adhering to it, so rapidly, that it is scarcely possible for the eye to follow it in its mo- tion. The Chameleon also has a very long and slender tongue, the extremity of which is dilated into a kind of club or spoon, and covered with a glutinous matter: with this instrument the animal catches insects from a considera- ble distance, by a similar manoeuvre to that practised by the frog.* Serpents and Lizards have generally curved or conical teeth, calculated rather for tearing and holding the food, than for masticating it: like those of fishes, they are affixed partly to the jaws, and partly to the palate. The Chelonian rep- tiles have no teeth, their office being supplied by the sharp cutting edges of the horny portion of the jaws. Birds as well as serpents have a moveable upper jaw; but they are also provided with beaks of various forms, in which we may trace an exact adaptation to the kind of food appro- priated to each tribe; thus, predaceous birds, as the eagle and the hawk tribe, have an exceedingly strong hooked beak, for tearing and dividing the flesh of the animals on which they prey; while those that feed on insects, or on grain, have pointed bills, adapted to picking up minute objects. Aquatic birds have generally fffittened bills, by which they can best select their food among the sand, the mud, or the weeds at the bottom of the water; and their edges are fre- quently serrated, to allow the fluid to filter through, while • Mr. Houston has given a description of the structure of this organ, and of the muscles by which it is moved, in a paper contained in the Transactions of the Royal Irish Academy, vol. xv. p. 177. Vol. II. 13 98 THE VITAL FX^NCTIOXS. the solid portions arc rclalncd in the inoiitli. Tlic duck af- fords an instance of tliis structure; wliich is, however, still more stron«j;ly marked in the genus Mergiis^ or Merganser, where tiie whole Icngtli of tlie inarL!;in of the hill is beset with small sliarp pointed teeth, directed backwards: thc}^ are jiarticularlv conspicuous in the Merf^its sej'ratnr, or red- l)rcastc(l Mcri^nnsor. 'J'he oI)ject of the ])arbs and fringed processes which arc apj^ended to the tongue in many birds, such as that of the Tuvcan and the Pan^akcet^ appears in like manner, to be the detention of substances introduced into the mouth. The beak of the Hxmatopus, or Oyster-catcher, has a wedge shape, and acts like an oyster-knife for opening bi- valve shells. In the Loxia curvirosira, or Cross-])ill, the upper and lower mandibles cross each other when the mouth is closed, a structure which enables this bird to tear open the cones of the pine and fir, and pick out tlie seeds, by insinuating the bill between the scales. It can split cherry stones with the utmost ease, and in a very short lime, by means of this pe- culiarly shajK'd bill.* Birds which dive for the purpose of catching fish have often a bill of considerable length, which enables them to secure their prey, and changes its position till it is adapted for swallowinc;. The li/iijnchnps, or black Skimmer, has a very singular- ly formed beak; it is very slender, but the lower mandible very much exceeds in length the upper one, so that while skimming the waves in its flight, it cuts the water like a plotigh-share, catching the prey which is on the surface of the sea. • The Woodpecker is furnished with a singular apparatus for enabling it to dart out with great velocity its long and jminted tongue, and transfix the insects on which it princi- pally feeds; and these motions arc performed so quickly • Ser .1 papor on the nicclianism of the bill of this bird, by Mr. Yan-cll, in the Zoolojric.il .lonmal, iv. 4.'>9. TONGUE OF THE WOODPECKER. 99 that the eye can scarcely follow them. This remarkable mechanism is delineated in Fig. 271, which represents the head of the woodpecker, with the skin removed, and tlie parts dissected. The tongue itself (t) is a slender sharj)- pointed horny cylinder, having its extremity (b) beset with barbs, of which the points are directed backwards: it is sup- ported on a slender 0.9 Hyoides, or lingual bone, to the ])Osterior end of which the extremities of two very long and narrow cartilaginous processes are articulated.* The one on the right side is shown in the figure, nearly in the whole extent of its course, at c, d, e, f, and a small portion of the left cartilage is seen at l. The two cartilages form, at their junction with the tongue, a very acute angle, slightly di- verging as they proceed backwards; until, bending down- wards (at c,) they pass obliquely round the sides of the neck, connected by a membrane (m;) then, being again in- flected upwards, they converge towards the back of the head, where they meet, and, being enclosed in a common sheath, are conducted together along a groove, whicli ex- tends forwards, along the middle line of the cranium (e,) till it arrives between the eyes. From this point, the groove * Tl\cse cartilages correspond in situation, at the part, at least, where they are jouied to the os hyoklcs, to what arc called the curnuUy or horns of that bone, ui other annuals. 100 THE VITAL FUNCTIONS. and the two cartilages it contains, which arc now more closely conjoined, are deflected towards the right side, and terminate at the edge of the aperture of the right nostril (r,) into which the united cartilages arc finally inserted. In order that their course may be seen more distinctly, these cartilages are represented in the figure (at d,) drawn out of the croove ])rovided to receive and protect them.'* A long and slender muscle is attached to the inner margin of each of these cartilages, and their actions conspire to raise the lower and most bent parts of the cartilages, so that their curvature is diminished, and the tongue protruded to a con- siderable distance, for the purpose of catching insects. As soon as this has been accomplished, these muscles being suddenly relaxed, another set of fibres, passing in front of the anterior portion of the cartilages nearly parallel to them, are thrown into action, and as suddenly retract the tongue into the mouth, with the insect adhering to its barbed ex- tremity. This muscular effort is, however, very materially assisted by the long and tortuous course of these arched cartilages, which are nearly as elastic as steel springs, and effect a considerable saving of m.uscular power.t This was the more necessary, because, while the bird is on the tree, it repeats these motions almost incessantly, boring holes in the bark, and picking up the minutest insects, witli the ut- most celerity and precision. On meeting w^th an ant-hill, the woodpecker easily lays it open by the combined efforts of its feet and bill, and soon makes a plentiful meal of the ants and their eggs. Among the Mamm.alia w-hich have no teeth, the Myrme- cophuf^a, or Ant-eater, practises a remarkable manoeuvre for catching its prey. The tongue of this animal is very long and slender, and has a great resemblance to an earth- worm: tliat of the two-toed ant-eater is very nearly one- third of the length of the whole body; and at its base is • S Is the hrge salivary gland on the right side. f An account of this mechanism is given by Mr. Waller, in the PhiL Trans, for 1716, p. 509. TONGUE OF THE ANT-EATER. 101 scarcely thicker than a crow-quill. It is furnished with a long and powerful muscle, which arises from the sternum, and is continued into its substance, affording the means of a quick retraction, as well as lateral motion; while its elonga- tion and other movements are effected by circular fibres, which are exterior to the former. When laid on the g-round in the usual track of ants, it is soon covered with these in- sects, and being suddenly retracted, transfers them into the mouth; and as, from their minuteness, they require no mas- tication, they are swallowed undivided, and without there being any necessity for teeth. The lips of quadrupeds are often elongated for the more ready prehension of food, as we see exemplified in the Rhi- noceros^ whose upper lip is so extensible as to be capable of performing the office of a small proboscis. The Sorcx mos- chatus, or musk shrew, whose favourite food is leeches, has likewise a very. moveable snout, by which it gropes for, and seizes its prey from the bottom of the nmd. More fre- quently, however, this office of prehension is performed by the tongue, which for that purpose is very flexible and much elongated, as we see in the Camelcojjard, where it acts like a hand in grasping and bringing down the branches of a tree.* In the animals belonging to the genus Fells, each of the papillae of the tongue is armed with a horny sheath termi- nating in a sharp point, which is directed backwards, so as to detain the food and prevent its escape. These prickles are of great size and strength in the larger beasts of prey, as the Lion and the Tiger; they are met with also in the Opossum, and in many species of bats, more especially those belonging to the genus Pteropus: all these horny produc- tions have been regarded as analogous to the lingual teeth of fishes, already noticed. The mouth of the Ornithorhyncus has a form of con- struction intermediate between that of quadrupeds and • Home, Lectures, &c. vi. Plate 33. 102 THE VITAL FUNCTIONS. birds; being furnished, like the former, with grinding teeth at the posterior part of both the ui)per and lower jaws, but they are of a horny substance; and the mouth is terminated in front by a horny bill, greatly resembling that of the duck, or the spoon-bill. The JVfialc is furnished with a singular apparatus de- signed for filtration on a large scale. The palate has the form of a concave dome, and from its sides there descends vertically into the mouth, a multitude of thin plates set pa- rallel to each other, with one of their edges directed towards the circumference, and the other towards the middle of the j)alate. These plates are known by the name of ivhalebone, and their general form and appearance, as they hang from the roof of the palate, are shown in Fig. 272, which represents only six of these plates."" They are connected to the bone by means of a white ligamen- tous substance, to which they arc im- mediately attached, and from which they appear to grow: at their inner margins, the fd^rcs, of which their tex- ture is throughout composed, cease to adhere together; but being loose and detached, form a kind of fringe, calcu- lated to intercept, as in a sieve, all so- lid or even gelatinous substances that may have been admitted into the cavity of the mouth, which is exceedingly ca- pacious; for as the plates of vv'halebonc grow only from the margins of the up- per jaw, they leave a large space with- in, which, though narrow anteriorly, is wider as it extends backwards, and is capable of holding a large quantity of • In ihc Pihed Ulialc the plates of whalebone are placed very near to- gether, not being a (|uartcr of an inch asunder; and there are above three hundred plates in the outer rows on each bide of the mouth. ! h' MOUTH OF THE WHALE. 103 water. Thus, llic whale is cnal)lccl to collect a whole shoal of mollusca, and other small prey, hy taking into its mouth the sea water which contains these animals, and allowing it to drain off through the sides, after passing through the in- terstices of the net work formed hy the fdaments of the whalebone. Some contrivance of this kind was even neces- sary to this animal, because the entrance into its oesophagus is too narrow to admit of the passage of any prey of consi- derable size; and it is not furnished with teeth to reduce the food .into smaller parts. The principal food of the Bahina Mysticetiis, or great whalebone whale of the Arctic Seas, is the small Clio Borealis, which swarms in immense num- bers in those regions of the ocean; and which has been al- ready delineated in Fig. 120.'^ These remarkable organs for filtration entirely supersede the use of ordinary teelh; and, accordingly, no traces of teeth are to be discovered either in the upper or lower jaw. Yet a tendency to conform to the type of the mammalia is manifested in the early conformation of the whale; for rudi- ments of teeth exist in the interior of the lower jaw before birth, lodged in deep sockets, and forming a row on each side. The development of these imperfect teeth proceeds no farther; they even disappear at a very early period, and the groove which contained them closes over, and, after a short time, can no longer be seen. For the discovery of this curious fact we arc indebted to Geoffroy St. Hilaire.t In connexion with this subject, an analogous fact, which has been noticed in the parrot, may here be mentioned. The young of the parrot, while still in the egg, presents a row of tubercles along the edge of the jaw, in external appearance exactly resembling the rudiments of teeth, but without being implanted into regular sockets in the maxillary bones: they are formed, however, by a process precisely similar to that of dentition; that is, by deposition from a vascular pulp, connected with the jaw. These tubercles are afterwards • Vol. i. p. 186. f Cuvicv, Ossemens Fossiles, 3me edition, torn. v. p. 360. 104 THE VITAL FUNCTIONS. consolidated into one piece In each jaw, forming, by their union, the beak of the parrot, in a manner perfectly analo- gous to that which leads to the construction of the compound tooth of the elephant, and which I shall presently describe. The original indentations are obliterated as the beak ad- vances in growth; but they are permanent in the bill of the duck, where the structure is very similar to that above de- scribed in the embryo of the parrot. § 3. Mastication hy means of Teeth. The teeth, being essential instruments for seizing and holding the food, and effecting that degree of mechanical di- vision necessary to prepare it for the chemical action of the stomach, perform, of course, a very important part in the economy of most animals; and in none more so than in the Mammalia, the food of which generally requires considera- ble preparation previously to its digestion. There exist, ac- cordingly, the most intimate relations between the kind of food upon which each animal of this class is intended by na- ture to subsist, and the form, structure, and position of the teeth; and these relations may, indeed, be also traced in the shape of the jaw, in the mode of its articulation with the head, in the proportional size and distribution of the mus- cles which move the jaw, in the form of the head itself, in the length of the neck, and its position on the trunk, and, indeed, in the whole conformation of the skeleton. But since the nature of the appropriate food is at once indicated by the structure and arrangement of the tcetii, it is evident that these latter organs, in particular, will afford to the na- turalist most important characters for establishing a systema- tic classification of animals, and more especially of quadru- peds, where the differences among the teeth are very consi- derable; and these differences have, accordino-lv, been the objects of much careful study. To the physiologist they present views of still higher interest, by exhibiting most OFFICES OF THE TEETH. 105 striking evidences of the provident care with which every part of the organization of animals has been constructed, in exact reference to their respective wants and destinations. The purposes answered by the teeth are principally those of seizing and detaining whatever is introduced into the mouth, of cutting it asunder, and dividing it into smaller pieces, of loosening its fibrous structure, and of breaking down and grinding its harder portions. Occasionally, some particular teeth are much enlarged, in order to serve as weapons of attack or defence; for which purpose, they ex- tend beyond the mouth, and are then generally denominated tiisks; this we see exemplified in the Elephant, {\\e Narwhal^ the Walrus^ the Hlppopotainus, the Boar, and the Babi- roussa. Four principal forms have been given to teeth, which ac- cordingly may be distinguished into the conical, the sharp- edged, the flat and the tuberculated teeth; though we occa- sionally find a few intermediate modifications of these forms. It is easy to infer the particular functions of each class of teeth, from the obvious mechanical actions to which, by their form, they are especially adapted. The conical teeth, which are generally also sharp-pointed, are principally em- ployed in seizing, piercing, and holding objects: such arc the offices which they perform in the Crocodile, and other Saurian reptiles, where all the teeth are of this structure; and such are also their uses in most of the Cetacea, where similar forms and arrangements of teeth prevail. All the Dolphin tribe, such as the Porpiis, the Grampus, and the Dolphin, are furnished with a uniform row of conical teeth, set round both jaws, in number amounting frequently to two hundred. Fig. 273, which represents the jaws of the Por- pus, shows the form of these simply prehensile teeth. The Cachalot has a similar row of teeth, which are, how- ever, confined to the lower jaw. All these animals subsist upon fish, and their teeth are therefore constructed very much on the model of those offish; while those Cetacea, on the other hand, which are herbivorous, as the Manatus and Vol. II. • 14 106 THE VITAL FUNCTIONS. the Diigong, or Indian Walrus, liave teeth very differently formed. Tiic tusks of animals must necessarily, as respects their shape, be classed among the conical teeth. 273 The sharp-edged teeth perform the office of cutting and dividing the yielding textures presented to them; they act inilividually as wedges or chisels, but when co-operating with similar teeth in the opposite jaw, they have the power of cutting like shears or scissors. The flat teeth, of which the surfaces are generally rough, are used in conjunction with those meeting them in the opposite jaw for grinding down the food by a lateral motion, in a manner analogous to the operation of mill-stones in a mill. The tuberculated teeth, of which the surfaces present a number of rounded eminences, corresponding to depressions in the teeth op- posed to them in the other jaw, act more by their direct pressure in breaking down hard substances, and pounding them, as they would be in a mortar. The position of the teeth in the jaws has been another ground of distinction. In those INIammalia which exhibit the most complete set of teeth, the foremost in the row have the sharp-edged or chisel shape, constituting the blades of a cutting instrument; and they are accordingly denominated iiicisors. The incisors of the upper jaw are always im- planted in a bone, intermediate between the two upper jaw bones, and called the intermaxillary bones.* The conical • Tliose teeth of the lower juw which coiTespond with the incisors of the upper j:iw, are also considered as incisors. In Man, and in the species of qiiadrumana that most nearly resemble him, the sutures which divide the in- tennaxillary from the maxillary bones are obliterated before birth, and leave in the adult no trace of their former existence. TEETH OP CETACEA. 107 teeth, immediately following the incisors, are called cuspi- date, or canine teeth, from their being particularly conspi- cuous in dogs; as they arc, indeed, in all the purely carnivo- rous tribes. In the larger beasts of prey, as the lion and the tiger, they become most powerful weapons of destruction: in the boar they are likewise of great size, and constitute the tusks of the animal. All the teeth that are placed farther back in the jaw are designated by the general name of inoUn^ teeth, or grinders, but it is a class which includes several different forms of teeth. Those teeth which are situated next to the canine teeth, partake of the conical form, having pointed eminences; these are called the false molar teeth, and, also, from their having generally two points, or cusps, the bicuspidate teeth. The posterior molar teeth are diffe- rently shaped in carnivorous animals, for they are raised into sharp and often serrated ridges, having many of the proper- ties of cutting teeth. In insectivorous and frugivorous ani- mals their surface presents prominent tubercles, either point- ed or rounded, for pounding the food; while in quadrupeds that feed on grass or grain they are flat and rough, for the purpose simply of grinding. The apparatus for giving motion to the jaws is likewise varied according to the particular movements required to act upon the food in the diflferent tribes. The articulation of the lower jaw with the temporal bone of the skull, approaciics to a hinge joint; but considerable latitude is allowed to its motions by the interposition of a moveable cartilage between the two surfaces of articulation, a contrivance admirably an- swering the intended purpose. Hence, in addition to the principal movements of opening and shutting, which are made in a vertical direction, the lower jaw has also some de- gree of mobility in a horizontal or lateral direction, and is likewise capable of being moved backwards or forwards, to a certain extent. The muscles which effect the closing of the jaw are principally the temporal and the masseter mus- cles; the former occupying the hollow of the temples, the latter connecting the lower angle of the jaw with the zygo- 108 THE VITAL FUNCTIONS. matic arch. The lateral motions of the jaw are effected by muscles placed internally between the sides of the jaw and the basis of the skull. In the conformation of the teeth and jaws, a remarkable contrast is presented between carnivorous and herbivorous animals. In the former, of which the Tiger, Fig. 274, may be taken as an example, the whole apparatus for mastication is calculated for the destruction of life, and for tearing and dividing the fleshy fibres. The molar teeth aj-e armed with pointed eminences, which correspond in the opposite jaws so as exactly to lock into one another, like wheel-work, when the mouth is closed. All the muscles which close the jaw are of enormous size and strength, and they imprint the bones of the skull with deep hollows, in which we trace marks of the most powerful action. The temporal muscles occupy the whole of the sides of the skull (t, t;) and by the continuance of their vigorous exertions, during the growth of the animal, alter so considerably the form of the bones, that the skulls of the young and the old animals arc often with difficulty recognised as belonging to the same species.* The process of the lower jaw (seen between t and t,) to which this temporal muscle is attached, is large and promi- nent; and the arch bone (z,) from which the masseter arises, takes a wide span outwards, so as to give great strength to • This is remarkably the case with the Bear, the skull of which exhibits, in old aiiiinuls, a large vertical crest, not met with at an early period of life. JAWS AND TEETH OF HERBIVORA. 109. the muscle. The condyle, or articulating surface of the jaw (c) is received into a deep cavity, constituting a strictly hinge joint, and admitting simply the motions of opening and shutting. In herbivorous animals, on the contrary, as may be seen in the skull of the Jintelope^ Fig. 275, the greatest force is bestowed, not so much on the motions of opening and shut- ting, as on those which are necessary for grinding, and which act in a lateral direction. The temporal muscles, oc- cupying the space t, are comparatively sm.all and feeble; the condyles of the jaw are broad and rounded, and more loosely connected with the skull by ligafnents; the muscles in the interior of the jaw, which move it from side to side, are very strong and thick; and the bone itself is extended downwards, so as to afford them a broad basis of attachment. The surfaces of the molar teeth are flattened and of great extent, and they are at the same time kept rough, like those of mill-stones, their office being in fact very similar to that performed by these implements for grinding. AH these circumstances of difference are exemplified in the most marked manner, in comparing together the skulls of the larger beasts of prey, as the tiger, the wolf, or the bear, with those of the antelope, the horse, or the ox. The Rodentia, or gnawing quadrupeds, which I have al- ready had occasion to notice, compose a well-marked fiimily of Mammalia. These animals are formed for subsisting on 110 THE VITAL FUNCTIONS. dry and tough materials, from which but little nutriment can be extracted; such as the bark, and roots, and even the woody fibres of trees, and the harder animal textures, which would appear to be most diflicult of digestion. They are all animals of diminutive stature, whose teeth are expressly formed for gnawing, nibbling, and wearing away by conti- nued attrition, the harder tex- tures of organized bodies. The 7i«/, whose skull is de- lineated in Fig. 216, belongs to this tribe. They are all furnished with two incisor teeth in each jaw, generally very long, and having the exact shape of a chisel; and the molar teeth have surfaces irregularly marked with raised zig zag lines, rendering them very perfect instruments of tritura- tion. The zygomatic arch is exceedingly slender and fee- ble; and the condyle is lengthened longitudinally to allow of the jaw being freely moved forwards and backwards, which is the motion for which the muscles are adapted, and by which the grinding operation is performed. The Beaver, the Baty the Marmot, and the Porcup'uie, present exam- ples of this structure, among the omnivorous rodentia: and the Hare, the Babbit, the Shrew, among those that are principally herbivorous. The Quadrumajia, or Monkey tribes, approach nearest to the human structure in the conformation of their teeth, which appear formed for a mixed kind of food, but are especially adapted to the consumption of the more esculent fruits. The other orders of Mammalia exhibit intermediate gradations in the structure of their teeth to those above de- scribed, corresponding to greater varieties in the nature of their food. Thus, the teeih and jaws of the Hyena are formed more csj)ccially for breaking down bones, and in so doing exert prodigious force; and those of the Sea Otter have rounded eminences, which peculiarly fit them for breaking shells. %' STRUCTURE OF TEETH. Ill The teeth, though composed of the same chemical ingre- dients as the ordinary hones, differ from them by having a greater density and compactness of texture, whence they derive that extraordinary degree of hardness which they re- quire for the performance of their peculiar office. The sub- stances of which they are composed are of three different kinds: the first, which is the basis of the rest, constituting the solid nucleus of the tooth, has been considered as the part most analogous in its nature to bone, but from its much greater density, and from its differing from bone in the mode of its formation, the name of ivory has been ge- nerally given to it. Its earthy ingredient oonsists almost entirely of phosphate of lime, the proportion of the carbo- nate of that earth entering into its composition being very small; and the animal portion is albumen, with a small quan- tity of gelatin. A layer of a still harder substance, termed the enamel, usually covers the ivory, and, in teeth of the simplest struc- ture, forms the whole of their outer surface: this is the case with the teeth of man and of carnivorous quadrupeds. These two substances, and the direction of their layers, are seen in Fig. 277, which is the section of a simple tooth, e is the outer case of enamel, o the osseous portion, and p the cavity where the vascular pulp which formed it was lodged. The enamel is composed almost wholly of phosphate of lime, containing no albumen, and scarcely atrace of gelatin; 112 THE VITAL FUNCTIONS. it is the hardest of all animal substances, and is capable of striking fire with steel. It exhibits a fibrous structure, ap- proaching to a crystalline arrangement, and the direction of its fibres, as shown by the form of its fragments when bro- ken, is every where ])crpendicular to the surface of the ivory to whicli it is applied. Tlie ends of the fibres arc thus alone exposed to the friction of the substances on which the teeth are made to act; and the effect of that friction in wearing the enamel is thus rendered the least possible. In the teeth of some quadrupeds, as of the Rhinoceros , the Hippopotamus, and most of the Rodentia, the enamel is intermixed with the ivory, and the two so disposed as to form jointly the surface for mastication. In the progress of life, the layers of enamel, being the hardest, are less worn down by friction than those of the ivory, and therefore form prominent ridges on the grinding surface, preserving it always in that rough condition, which best adapts it for the bruising and comminuting of hard substances. The incisors of the rodentia are guarded by a plate of en- amel only on their anterior convex surfaces, so that by the wearing down of the ivory behind this plate, a wedge-like form, of which the enamel constitutes the fine cutting edge, is soon given to the tooth, and is constantly retained as long as the tooth lasts (Fig. 2S0.) This mode of growth is admi- rably calculated to presefve these chisel teeth fit for use during the whole lifetime of the animal, an object of greater consequence in this description of teeth than in others, which continue to grow only during a limited period. The same arrangement, attended with similar advantages, is adopted in the structure of the tusks of the Hippopotamus. In teeth of a more complex structure, a third substance is found, uniting the vertical plates of ivory and enamel, and performing the office of an external cement. This substance has received various names, but it is most commonly known by that of the Criistd pctrosa: it resembles ivory both in its composition and its extreme hardness; but is generally more opaque and yellow than that substance. STRUCTURE OF TEETH. 113 Other herbivorous quadrupeds, as the horse, and animals belongnig to the ruminant tribe, have also complex teeth com- posed of these three substances; and their grinding surfaces present ridges of enamel intermixed in a more irregular manner with the ivory and crusta pctrosa; but still giving the advantasfe of a very rougrh surface to the trituration. Fig. 278 represents the grinding surface of the tooth of a horse, worn down by long mastication, e is the enamel, marked by transverse lines, showing the direction of its fibres, and enclosing the osseous portion (o,) which is shaded by interrupted lines. An outer coating of enamel {e) is also visible, and between that and the inner coat, the substance called crusta petrosa (c,) marked by waving lines, is seen. On the outside of all there is a plate of bone, which has been left w^hite. In ruminants, the plates of enamel form cres- cents, which are convex outwardly in the lower, and in- wardly in the upper jaw; thus providing for the crossing of the ridgeS'Of the two surfaces, an arrangement similar to that which is practised in constructing those of mill-stones. The teeth of the lower javv fall within those of the upper jaw, so that a lateral motion is required in order to bring their surfaces opposite to each other alternately on both sides. Fig. 279 shows the grinding surface of the tooth oi d. Sheep, where the layers of bone are not apparent, there being only two layers of enamel (e,) and one of crusta petrosa (c.) These three component parts are seen to most advantage in a vertical and longitudinal section of the grinding tooth of the elephant, in which they are more completely and equally intermixed than in that of any other animal. Fig. 281 presents a vertical section of the grinding tooth of the Asiatic elepiiant, in the early stage of its growth, and highly polished, so as to exhibit more perfectly its three component structures. The enamel, marked e, is formed of transverse fibres; the osseous, or innermost structure is composed of longitudinal plates. The general covering of crusta pe- trosa, c, is less regularly deposited, p is the cavity which had been occupied by the pulp. In this tooth, which is still Vol. II. 15 114 THE VITAL FUNCTIONS. rn a growing state, the fangs are not )-et added, but they are, at one part, beginning to be formed. The same tooth, in its '^<'^^ usual state, as worn by mastication, gives us a natural and horizontal section of its interior structure, in which the plates of white enamel are seen forming waved ridges. These con- stitute, in the Asiatic Elephant, a series of narrow transverse bands, (Fig. 2S3,) and in tlie African Elephant, a series of loy.enge-shapcd lines, (Fig. 2S2,) having the ivory on their interior, and the yellow crusta petrosa on their outer sides; which latter substance also composes the whole circumfe- rence of the section. § 4. Formation and Development of the Teeth. Few processes in animal development are more remarka- ble than those which arc employed to form the teeth; for they are, by no means, the same as those by which ordina- ry bone is constructed; and being commenced at a very ear- ly period, they aftbrd a signal instance of Nature's provident anticipation of the futiu'e necessities of the animal. Tiic teeth, being the hardest parts of the body, require a peculiar DENTITION. 115 system of operations for giving them this extraordinary- density, which no gradual consolidation could have impart- ed. The formation of the teeth is, in some respects, analo- gous to that of shell; inasmuch as all their parts, when once deposited, remain as permanent structures, hardly ever ad- mitting of removal or of renewal by the vital powers. Un- like the bones, which contain within their solid substance vessels of different kinds, by which they are nourished, mo- dified, and occasionally removed, the closeness of the texture of the teeth is such as to exclude all vessels whatsoever. This circumstance renders it necessary that they should ori- ginally be formed of the exact size and shape which they are ever after to possess: accordingly, the foundation of the teeth, in the young animal, are laid at a very early period of its evolution, and considerable progress has been made in their growth even prior to birth, and long before they can come into use. A tooth of the simplest construction is formed from blood- vessels, which ramify through small masses of a gelatinous appearance; and each of these pulpy masses is itself enclosed in a delicate transparent vesicle, within which it grows till it has acquired the exact size and shape of the future tooth. Each vascular pulp is farther protected by an investing membrane of greater strength, termed its capsule^ which is lodged in a small cavity between the two bony plates of the jaw. The vessels of the pulp begin at an early period to deposite the calcareous substance, which is to compose the ivory, at the most prominent points of that part of the vesi- cle, which corresponds in situation to the outer la3'er of the crown of the tooth. The thin scales of ivory thus formed increase by farther depositions made on their surfaces next to the pulp, till the whole has formed the first, or outer layer of ivory: in the mean time, the inner surface of the capsule, which is in immediate contact with this layer, se- cretes the substance that is to compose the enamel, and tie- posites it in layers on the surface of the ivory. This double operation proceeds step by step, fresh layers of ivory being 116 THE VITAL FUNCTIONS. deposited, and building up the body of the tooth, and in the same projiortion cncroachinii; upon tlie cavity occupied by the pulp, which retires before it, until it is shrunk into a small compass, and fdls only the small cavity which remains in the centre of the tooth. The ivory has by this time re- ceived from the capsule a complete coating of enamel, which constitutes the whole outer surface of the crown; after which no more is deposited, and the function of the capsule having ceased, it shrivels and disaj^pears. But the formation of ivory still continuing at the part most remote from the crown, the fangs are gi'adually formed by a similar process from the pulp; and a pressure being thereby directed against the bono of the socket at the part where it is the thinnest, that portion of the jaw is absorbed, and the ])rogress of the tooth is onl}^ resisted by the gum; and the gum, in its turn, soon yielding to the increasing pressure, the tooth cuts its way to the surface. This process of successive deposition is beautifully illustrated by feeding a young animal at dif- ferent times with madder; the teeth which are formed at that period exhibiting, in consequence, alternate layers of red and of white ivory.* / The formation of the teeth of herbivorous quadruj:)eds, which have three kinds of substance, is conducted in a still more artificial and complicated manner. Thus, in the ele- phant, the pulp which deposites the ivory is extended in the form of a number of parallel plates; while the capsule which invests it, accompanies it in all its parts, sending down du- plicatures of membrane in the intervals between the plates. Hence the ivory constructed by the pulp, and the enamel deposited over it, are variously intermixed; but besides this, the crusta petrosa is deposited on the outside of the enamel. Cuvier asserts that this deposition is made by the same cap- sule which has formed the enamel, and which, previously to this change of function, has become more spongy and vascular than before. Hut his brother, M. Frederic Cuvier, • Cuvier. Dictionnnirc dcs Sciences Medicalcs, t. viii. p. 320. DENTITION. 117 represents the depositc of crusta pctrosa, as pcrforined by a third membrane, wholly distinct from the two others, and exterior to them all, although it follows them in all their folds. In the horse and the ox, the projecting processes of the pulp, have more of a conical form, with undulating sides; and hence the waved appearance presented by the enamel, on making sections of the teeth of these animals. The tusks of the elephant are composed of ivory, and are formed precisely in the same manner as the simple conical teeth already described, excepting that there is no outer cap- sule, and therefore no outer crust of enamel. The whole of the substance of the tusk is constructed by successive depo- sites of layers, having a conical shape, from the pulp which occupies the axis of the growing tusk; just as happens in the formation of a univalve shell which is not turbinated, as, for instance, the patella. Hence, any foreign substance, a bul- let, for example, which may happen to get within the cavity occupied by the pulp, becomes, in process of time, encrusted with ivory, and remains embedded in the solid substance of the tusk. The pulp, as the growth of the tusk advances, re- tires in proportion as its place is occupied by the fresh de- posites of ivory. The young animal requires teeth long before it has attained its full stature; and these teeth must be formed of dimen- sions adapted to that of the jaw, wdiile it is yet of small size. But, as the jaw enlarges, and the teeth it contains admit not of any corresponding increase, it becomes necessary that they should be shed to make room for others of larger di- mensions, formed in a more capacious mould. Provision is made for this necessary change at a very early period of the growth of the embryo. The rudiments of the human teeth begin to form four or five months before birth: they are contained in the same sockets with the temporary teeth, the capsules of both being connected together. As the jaw en- larges, the second set of teeth gradually acquire their full dimensions, and then, by their outward pressure, occasion 118 THE VITAL FUNCTIONS. the absorption of the fangs of the temporary teeth, and, push- ing them out, occupy tlieir places.^ As the jaw bone, during its growth, extends principally backwards, the posterior portion, being later in forming, is comparatively of a larger size than either the fore or the la- teral parts; and it admits, therefore, of teeth of the full size, which, consequently, are permanent. The molar teeth, which are last formed, are, for want of space, ratlier smaller than the others, and are called the wisdom-teeth, because they do not usually make their appearance above the gum till the person has attained the age of twenty. In the negro, however, where the jaw is of greater length, these teeth have sufficient room to come into their places, and are, in gene- ral, fully as large as the other molars. The teeth of carnivorous animals are, from the nature of their food, less liable to be worn, than those of animals living on grain, or on the harder kinds of vegetable sub- stances; so that the simple plating of enamel is sufficient to preserve them, even during a long life. But in many herbi- vorous quadrupeds we find that, in proportion as the front teeth are worn away in mastication, other teeth are formed, and advance from the back of the jaw to replace them. This happens, in a most remarkable manner, in the elephant, and is the cause of the curved form which the roots assume; for, in proportion as the front teeth are worn away, those imme- diately behind them are pushed forwards by the growth Of a new tooth at the back of the jaw; and this process goes on continually, giving rise to a succession of teeth, each of which is larger than that which has preceded it, during the whole period that the animal lives. A similar succession of teeth takes place in the wild boar, and, also, though to a less extent, in the Sus Q^thiopicusA This mode of dentition • It Is stated by Rousseau that the shedding- of the first molar tooth both of the Guinea-pig, and the Capihara, and its replacement by the permanent tooth, take place a few days before birth. Anatomic Compai-ce du systeme dentaire, p. 164-. t Home, Phil. Ti-ans. for 1799, p. 237; and 1801, p. 319. DENTITION. 119 appears to be peculiar to animals of great longevity, and which subsist on vegetable substances containing a large pro- portion of tough fibres, or other materials of great hardness; and requiring for their mastication teeth so large as not to admit of both the old and new tooth being contained, at the same time, in the alveolar portion of the jaw. An expedient of a different kind has been resorted to in the Rodentictf for the purpose of preserving the long chisel- shaped incisors in a state fit for use. By the constant and severe attrition to which they are exposed, they wear away very rapidly, and would soon be entirely lost, and the ani- mal would perish in consequence, were it not that nature has provided for their continued growth, by elongation from their roots, during the whole of life. This growth proceeds in the same manner, and is conducted on the same princi- ples, as the original formation of the simple teeth already described: but, in order to effect this object, the roots of these teeth are of great size and length, and are deeply em- bedded in the jaw, in a large bony canal provided for that purpose; and their cavity is always filled with the vascular pulp, from which the continued secretion and deposition of fresh layers, both of ivory and enamel, take place. The tusks of the Elephant and of the Hippopotamus exhibit the same phenomenon of constant and uninterrupted growth. In the Shark, and some other fishes, the same object is attained in a different manner. Several rows of teeth arc lodged in each jaw, but one only of these rows projects and is in use at the same time; the rest lying flat, but ready to rise in order to replace those that have been broken or worn down. In some fishes, the teeth advance in proportion as the jaw lengthens,* and as the fore teeth are worn away: in other cases, they rise from the substance of the jaw, which presents on its surface an assemblage of teeth in different stages of growth: so that, in this class of animals, the great- est variety occurs in the mode of the succession of the teeth,. The teeth of the Crocodile, which arc sharp-pointed hoi- 120 THE VITAL FUNCTIONS. low cones, composed of ivory and enamel, are renewed by the new tooth (as is shown at a, in Fig. 2S4,) being formed 234 in the cavity of the one (b) wliich it is _____ to replace, and not being enclosed in any C ,/ sejiarate cavity of the jaw bone (c.) As this new tooth increases in size, it press- es against the base of the old one, and entering its cavity, acquires the same conical form; so that when the latter is shed, it is already in its place, and fit for immediate use. This succession of teetli takes place several times during the life of the animal, so that they are sharp and perfect at all ages. The fangs of serpents are furnished, like the stings of nettles, with a receptacle at their base for a poisonous li- quor, which is squeezed out by the pressure of the tooth, at the moment it inflicts the wound, and conducted along a canal, opening near the extremity of the tooth. Each fang is lodged in a strong bony socket, and is, by the interven- tion of a connecting bone, pressed forwards whenever the jaw is opened sufficiently wide; and the fang is thus made to assume an erect position. As these sharp teeth are very liable to accidents, others arc ready to supply their places when wanted: for which purpose there are commonly pro- vided two or three half-grown fangs, which arc connected only by soft parts with the jaw, and are successively moved forwards into the socket to replace those that were lost.* The tube through which the poison flows is formed by the folding in of the edges of a deep longitudinal groove, extending along the greater part of the tooth; an interval being left between these edges, both at the l^ase and extre- mity of the fang, by which means there remain apertures at both ends for the passage of the fluid poison. This struc- ture was discovered by jNlr. T. Smith in the Coluber naia, * Home, Lectures, &c. I. 333. FANGS OF SERPENTS. 121 or, Cobra de Capcllo;'^- and is shown in Fig. 285, which represents the full grown tooth, where the slight furrow, in- dicating the junction of the two sides of the original groove, maybe plainly seen; as also the two apertures (a and b) above mentioned. This mode of formation of the tube is farther illustrated by Fig. 286, which shows a transverse 291 section of the same tooth, exhibiting the cavity (p) which contains the pulp of the tooth, and which surrounds that of the central tube in the form of a crescent. Figures 287 and 288 are delineations of the same tooth in different stages of growth, the bases of which, respectively, are shown in Figures 289 and 290. Figures 291 and 292 are magnified representations of sections of the fangs of another species of serpent, resembling the rattle-snake. Fig. 291 is a section of the young fang taken about the middle: in this stage of growth, the cavity which contains the pulp, almost entirely surrounds the poison tube, and the edges of the depression, which form the suture, are seen to be angular, and present so large a surface to each other, that the suture is complete- ly filled up, even in this early stage of growth. Fig. 292 is a section of a full-growni fang of the same species of ser- pent, at the same part as the preceding; and here the cavity * Philosophical Transactions, 1818, p. 471. Vol. II. 16 122 THE VITAL FUNCTIONS. of the pulp is seen much contracted from tlie more advanced staf];e of growth. It is a remarka])le circumstance, noticed by Mr. Smith, that a similar lonii;itudinal furrow is perceptible on every- one of the teeth of the same serpent; and that this appear- ance is most marked on those which are nearest to the poi- sonous fangs: these furrows, however, in the teeth that are not venomous, are confmed entirely to the surface, and do not influence the form of the internal cavity. No trace of these furrows is discernible in the teeth of those serpents which are not armed with venomous fangs. Among the many instances in which teeth arc converted to uses widely different from mastication, may be noticed that of the Squalus pristis, or Saw-fish, where the teeth are set horizontally on the two lateral edges of the upper jaw, which is prolonged in the form of a snout (seen in a, Fig. 293,) constituting a most formidable weapon of offence, b is a more enlarged view of a portion of this instrument, seen from the under side. §. 5. Tr 'duration of Food in Internal Cavities. The mechanical apparatus, provided for triturating the harder kinds of food, does not belong exclusively to the GASTRIC TEETH. 123 mouth, or entrance into the alimentary canal, for in many- animals we find this office performed by interior organs. 294 Among the inferior classes, we find examples of this conforma- tion in the Crustacea, the Mol- lusca, and above all in Insects. Thus, there is found in the sto- mach of the Lobster, a cartilagi- nous frame-work, in which are implanted hard calcareous bodies, having the form, and performing the functions of teeth. They are delineated in Fig. 294, which presents a view of the interior of the stomach of that animal. The tooth a is situated in the middle of this frame, has a rounded conical shape, and is smaller than the others (b, c,) which are placed one on each side, and which resemble in their form broad molar teeth. When these three teeth are brought together by the action of the surrounding muscles, they fit exactly into each other, and are capable of grinding and completely pulverizing the shells of the mollusca introduced into the stomach. These teeth are the result of a secretion of calca- reous matter from the inner coat of that organ, just as the outer shell of the animal is a production of the integu- ment: and at each casting of the shell, these teeth, together wnth the whole cuticular lining of the stomach to which they adhere, are thrown off, and afterwards renewed by a fresh growth of the same material. In the Craw-fish, the gastric teeth are of a different shape, and are more adapted to divide than to grind the food. Among the gasteropodous Mollusca, se- veral species of Bullx have stomachs armed with calcareous plates, which act as cutting or grinding teeth. The Bulla aperta has three instruments of this description, as may be seen in Fig. 295, which shows the interior of the stomach of that species. Similar organs are found in the Bulla lig- 124 THE VITAL PUNCTIOXS. nciria. The ^^p^ysia lias a considerable number of these gas- tric teeth. An apparatus of a still more complicated kind is provided in most of the insects belonging to the order of Orthoptera; but I shall not enter at present in their de- scription, as it will be more convenient to include them in the sreneral account of the alimentary canal of insects, which will be the subject of future consideration. The internal machinery for grinding is exemplified on the largest scale in granivorous birds; where it forms part of the stomach itself, and is termed a Gizzard. It is shown in Fig. 298, representing the interior of the stomach of a Swan. Both the structure and the mode of operation of this organ bear a striking analo- gy to a mill for grinding corn, for it consists of two powerful mus- \ cles (g,) of a hemispherical shape, '] with their flat sides applied to each other, and their edges united by a strong tendon, which leaves a va- cant space of an oval or quadrangular form between their two surfaces. These surfaces are covered by a thick and dense horny substance, which, when the gizzard is in ac- tion, performs an office similar to that of mill-stones. In most birds, there is likewise a sac, or receptacle, termed the Craw, (represented laid open at c) in which the food is collected for the purpose of its being dropped, in small quantities at a time, into the gizzard, in proportion as the latter gradually becomes emptied.* Thus, the analogy be- tween this natural process and the artificial operation of a corn-mill is preserved even in the minuter details; for while the two flat surfaces of the gizzard act as mill-stones, the craw supplies the place of the hopper, the office of which is • The gastric glands, wliich are spread over the greater part of the inter- nal surface of the craw, and which prcjjare a secretion for macerating the grain, are also seen in this part of the fig^ire. ACTION OF THE GIZZARD. 125 to allow the grain to pass out in small quantities into the aperture of the upper mill-stone, which brings it within the sphere of their action. Innumerable are the experiments wliich have been made, particularly by Reaumur and Spallanzani, with a view to ascertain the force of compression exerted by the gizzard on its contents. Balls of glass, which the bird was made to swallow with its food, were soon ground to powder: tin tubes, introduced into the stomach, were flattened, and then bent into a variety of shapes; and it was even found that the points of needles and of lancets fixed in a ball of lead, were blunted and broken off by the power of the gizzard, while its internal coat did not appear to be in the slightest degree injured. These results were long the subject of ad- miration to physiologists; and being echoed from mouth to mouth, were received with a sort of passive astonishment, till Hunter directed the powers of his mind to the inquiry, and gave the first rational explanation of the mechanism by which they are produced. He found that the motion of the, sides of the gizzard, w^hen actuated by its muscles, is lateral, and at the same time circular; so that the pressure it exerts, though extremely great, is directed nearly in the plane of the grinding surfaces, and never perpendicularly to them; and thus the edges and points of sharp instruments are either bent or broken off by the lateral pressure, without their having an opportunity of acting directly upon tlfose sur- faces. Still, however, it is evident that the effects we ob- serve produced upon sharp metallic points and edges, could not be accom.plished by the gizzard without some assistance from other sources; and this assistance is procured in a very singular, and, at the same time, very effectual manner. On opening the gizzard of a'bird, it is constantly found to contain a certain quantity of small pebbles, which must have been swallowed by the animal. The most natural rea- son that can be assigned for the presence of these stones, is, that they aid the gizzard in triturating the contained food, and that they, in fact, supply the office of teeth in that ope- 126 THE VITAL FUNCTIONS. ration. Spallanzani, however, has called in question the souiuhicss of this explanation, and has contended that the pebbles found in the gizzard are swallowed merely by acci- dent, or in consequence of the stupidity of the bird, which mistakes them for grain. But this opinion has been fully and satisfactorily refuted both by Fordyce and by Hunter, whose observations concur in establishing the truth of the common opinion, that in all birds possessing gizzards, the presence of these stones is essential to perfect digestion. A greater or less number of them is contained in every giz- zard, when the bird has been able to meet with the requi- site supply, and they are never swallowed but along with the food. Several hundred w^ere found in the gizzard of a turkey; and two thousand in that of a goose: so great an accumulation could never have been the result of mere ac- cident. If the alleged mistake could ever occur, w^e should expect it to take place to the greatest extent in those birds which are starving for want of food; but this is far from be- • ino- the case. It is found that even chickens, which have been hatched by artificial heat, and which could never have been instructed by the parent, are yet guided by a natural instinct in the choice of the proper materials for food, and for assistinsi its digestion: and if a mixture of a large quan- tity of stones with a small proportion of grain be set before them, they will at once pick out the grain, and swallow alono- with it only the proper proportion of stones. The best proof of the utility of these substances may be derived from the experiments of Spallanzani himself, who ascer- tained that grain is not digested in the stomachs of birds, when it is protected from the effects of trituration. Thus, the gizzard may, as Hunter remarks, be regarded as a pair of jaws, whose teeth are taken in occasionally to assist in tliis internal mastication. The lower part of the gizzard consists of a thin muscular bag, of which the office is to digest the food which has been thus triturated. Considerable differences are met with in the structure of the gizzards of various kinds of birds, corresponding to dif- SALIVARY APPARATUS. 127 ferences In the texture of their natural food. In the Tin^keyy the two muscles which compose the gizzard are of unequal strength, that on the left side being considerably larger than that on the right; so that while the principal cifort is made by the former, a smaller force is used by the latter to restore the parts to their situation. These muscles produce, by their alternate action, two effects; the one a constant tritura- tion, by a rotatory motion; the other a continued, but oblique, pressure of the contents of the cavity. As this cavity is of an oval form, and the muscle swells inwards, the opposite sides never come into contact, and the interposed materials are triturated by their being intermixed with hard bodies. In the Goose and Swan, on the contrary, the cavity is flat- tened, and its lateral edges are very thin. The surfaces ap- plied to each other are mutually adapted in their curvatures, a concave surface being every where applied to one which is convex: on the left side, the concavity is above: but on the right side, it is below. The horny covering is much stronger, and more rough, than in the turkey, so that the food is ground by a sliding, instead of a rotatory motion, of the parts opposed, and they do not require the aid of any intervening hard substances of a large size. This motion bears a great resemblance to that of the grinding teeth of ruminating animals, in which the teeth of the under jaw slide upwards, within those of the upper, pressing the food between them, and fitting it, by this peculiar kind of tritu- ration, for being digested.'^ ^ 6. Beglulition, The great object of the apparatus which is to prepare the food for digestion, is to reduce it into a soft pulpy state, so as to facilitate the chemical action of the stomach upon it: for this purpose, solid food must not only be subjected to mechanical trituration, but it must also be mixed with a cer- * Home, Phil. Trans, for 1810, p. 188. 128 THE VITAL FUNCTIONS. tain proportion of fluid. Hence, all animals that masticate their food are provided with organs wliich secrete a fluid, called the Saliva, and which pour this fluid into the mouth as near as possible to the <;;rinding surfaces of the teeth. These organs are glands, ])laced in such a situation as to he compressed hy the action of the muscles which move the jaw, and to pour out the fluid tliey secrete in greatest quan- tity, just at the time when the food is undergoing masti- cation. Saliva contains a large quantity of water, together with some salts and a little animal matter. Its use is not only to soften the food, but also to luhricate the passage through which it is to be conveyed into the stomach; and the quantity secreted has always a relation to the nature of the food, the degree of mastication it requires, and the mode in which it is swallowed. In animals which subsist on ve- getable materials, requiring more complete maceration than those which feed on flesh, the salivary glands are of large size: they are particularly large in the Rodent ia, which feed on the hardest materials, requiring the most complete tritu- ration: and in these animals we find that the largest quantity of saliva is poured out opposite to the incisor teeth, which are those principally employed in this kind of mastication. In Birds and Reptiles, which can hardly be said to masti- cate their food, the salivary glands are comparatively of small size: the exceptions to this rule occurring chiefly in those trlhes which feed on vegetables, for in these the glands are more considerable.* In Fishes there is no structure of this kind provided, there being no mastication performed: and the same observation applies to the Cetacea. In the cephalopodous and gasteropodous Mollusca, we find a sali- vary apparatus of considerable size: Insects, and the t^nne- lida,\ also, generally present us with organs which appear to perform a similar oflice. • The l.'irge salivary gland in the woodpecker, is seen at s, Fig-. 271, page 99. t The bunch of filaments, seen at s, Fig. 260 (p. 78) are the salivary or- gans of tliclccch. DEGLUTITION. 129 • Tho passage of, the food along the lliroat is facilitated by the mucous secretions, which are poured out from a multi- tude of glands interspersed over the whole surface of the membrane lining that passage. The Cmiid, which is formed for traversing dry and sandy deserts, where the atmosphere as well as the soil is parched, is specially provided with a glandular cavity placed behind the palate, and which fur- nishes a fluid for the express purpose of moistening and lu- bricating the throat. In the structure of the (Esopha^iis^y^X^xoXx is the name of the tube along which the food passes from the mouth to the stomach, we may trace a similar adaptation to the particular kind of food taken in by the animal. When it is swallowed entire, or but little changed, the cosophagus is a very wide canal, admitting of great dilatation. This is the case with many carnivorous birds, especially those that feed on fishes, where its great capacity enables it to hold, for a considera- ble time, the large fish which arc swallowed entire, and which could not conveniently be admitted into the stomach. Blumenbach relates that a sea-gull, which he kept alive for many years, could swallow bones of three or four inches in length, so that only their lower ends reached the stomach, and were digested, while their upper ends projected into the CESophagus, and descended gradually, in proportion as the former were dissolved. Serpents, which swallow animals larger than themselves, have, of course, the oesophagus, as well as the throat, capable of great dilatation, and the food occupies a long time in passing through it, before it reaches the digesting cavity. The turtle has also a capacious oeso- phagus, the inner coat of which is beset with numerous firm and sharp processes, having their points directed towards the stomach; these are evidently intended to prevent the re- turn of the food into the mouth. Grazing quadrupeds, who, while they eat, carry their heads close to the ground, have a long oesophagus, with thick muscular coats, capable of ex- erting considerable power in ])ropelling the food in the di- rection of the stomach, which is contrary to that of gravity. Vol. II. 17 130 THE VITAL FUNCTIONS. § 7. Bccepfaclcsfor retaining Food, Provision is often made for the retention of the undigest- ed food in reservoirs, situated in difTercnt parts of the mouth, or the ccsopliagus, instead of its being immediately intro- duced into the stomach. These reservoirs are generally em- ployed for laying in stores of provisions for future consump- tion, ^lany quadrupeds have cheek pouches for this purpose: this is the case with several species of Monkeys and Ba- boons; and, also, willi the Miis cricclus, or Hamster. The Mus buscu'ius, or Canada rat, has enormous check pouches, ■which, when distended with food, even exceed the bulk of the head. Small cheek pouches exist in that singular ani- mal, the Ornil/ior/iT/ncns. The Scitirus palmariim, or palm squirrel, is also provided with a pouch for laying in a store of provisions. A remarkable dilatation, in the lower part of the mouth and throat, answering a similar purpose, takes place in the Pelican; a bird which displays great dex- terity in tossing about the fish with which it has loaded this bag, till it is brought into the proper position for being swal- lowed. The JV/iale has also a receptacle of enormous size, extending from the mouth to a considerable distance under the trunk of the body. Analogous in design to these pouches are the dilatations of the oesophagus of birds, denominated c?'ops. In most birds which feed on grain, the crop is a capacious globular sac, placed in front of the throat, and resting on the furcular bone. The crop of the Pam^ot is represented at c. Fig. 299; where also, s indicates the cardiac portion of the stomach, and g the gizzard, of that l)ird. The inner coat of the crop is furnished with numerous glands, which pour out considerable quantities of fluid for macerating and softening the dry and hard texture of the grain, which, for that purpose, remains there for a considerable time. Many birds feed their young from the contents of the crop; and, at those seasons its glands arc RECEPTACLES FOR RETAINING FOOD. 131 much enlarged, and very active in preparing tlieir peculiar secretions: this is remarkably the case in the Pigeon (Fig. 300,) which, instead of a single sac, is provided with two (seen at c, c, Fig. 300,) one on each side of the ccsophagus (o.) The pouting pigeon has the facul- ty of filling these cavities with air, which produces that distend- ed appearance of the throat from which it derives its name. Birds of prey have, in general, very small crops, their food not re- quiring any previous softening; but the Vulture, which gorges large quantities of flesh at a single meal, has a crop of considerable size, form- ing, when filled, a visible projection in front of the chest. Birds which feed on fish have no separate dilatation for this purpose, probably because the great width of the oesophagus, and its having the power of retaining a large mass of food, render the farther dilatation of any particular part of the tube unnecessary. The lower portion of the oesophagus ap- pears often, indeed, in this class of Animals, to answer the purpose of a crop, and to effect changes in the food which may properly be considered as a preliminary stage of the digestive process. ( 132 ) CHAPTER VII. Digestion. All the substances received as food into the stomach, whatever be their nature, must necessarily undergo many changes of chemical composition before they can gain ad- mission into the general mass of circulating fluids; but the extent of tiie change required for that purpose will, of course, be in proportion to the difference between the (qualities of the nutritive materials in their original, and in their assimi- lated state. The conversion of vesretable into animal mat- ter necessarily implies a considerable modification of proper- ties; but even animal substances, however similar may be their composition to the body which they are to nourish, must still pass through certain j)rocesses of decomposition, and subsequent recombination, before they can be brought into the exact chemical state in which they are adapted to the pur])oses of the living system. The preparatory changes we have lately been occupied in considering, consist chiefly in the reduction of the food to a soft consistence, which is accomplished by destroying the co- hesion of its parts, and mixing them uniformly with the fluid secretions of the mouth; eficcts wdiich may be considered as wholly of a mechanical nature. The first real changes in its chemical state are produced in the stomach, where it is con- verted into a sul)stance termed Chyme; and the process by which this first step in the assimilation of the food is pro- duced, constitutes what is proj^erly termed Digestion. Nothing has been discovered in the anatomical structure of the stomach, tending to throw any light on the means by which this remarkable chemical change is induced on the DIGESTION. 133 materials it contains. The stomacli is, in most animals, a simple sac, composed of several membranes, enclosing thin layers of muscular fibres, abundantly supplied with blood- vessels and with nerves, and occasionally containing struc- tures which appear to be glandular. The human stomach, which is delineated in Fig. 301, exhibits one of the simplest form^of this organ; c being the cardiac portion, or part where the oesophagus opens into it; and p the pyloric por- tion, or that which is near its termination in the intestine. At the pylorus itself, the diameter of the passage is much constricted, by a fold of the inner membrane, which is sur- rounded by a circular band of muscular fibres, performing the office of a sphincter, and completely closing the lower orifice of the stomach, during the digestion of its contents. The principal agent in digestion, as far as the ordinary chemical means arc concerned in that operation, is a fluid secreted by the coats of the stomach, termed the Gastric juice. This fiuid has, in each animal, the remarkable pro- perty of dissolving, or, at least, reducing. to a pulp, all the substances which constitute the natural food of that particu- lar species of animal; w'hile it has comparatively but little solvent power over other kinds of food. Such is the con- clusion which has been deduced from the extensive re- searches on this subject, made by that indefatigable experi- mentalist, Spallanzani, who found, in numberless trials, that the gastric juice taken from the stomach, and put into glass 134 THE VITAL FUNCTIONS. vessels, produced, if kept at the usual temperature of the animal, changes, to all appearance, exactly similar to those which take place in natural digestion.* In animals which feed on flesh, the gastric juice was found to dissolve only animal substances, and to exert no action on vegetable mat- ter; while, on the contrary, that taken from herbivorous ani- mals, acted on grass and other vegetable substances, without producing any eflect on flesh; but in those animals, which, like man, are omnivorous, that is, partake indiscriminately of both species of aliment, it appeared to be fitted equally for the solution of both. So accurate an adaptation of the chemical powers of a solvent to the variety of substances employed as food by diflcrent animals, displays, in the most striking manner, the vast resources of nature, and the re- fined chemistry she has put in action for the accomplishment of her difierent purposes. In the stomachs of many animals, as also in the human, it is impossible to distinguish with any accuracy the organi- zation by which the secretion of the gastric juice is effected: but where the structure is more complex, there may be ob- served a number of glandular bodies interspersed in various parts of tlic internal coats of the stomach. These, which are termed the Gastric glands, are distributed in various ways in different instances: they are generally found in greatest number, and often in clusters, about the cardiac ori- fice of the stomach; and they are frequently intermixed with glands of another kind, which prepare a mucilaginous fluid, serving to protect the highly sensible coats of the stomach from injurious impressions. These latter are termed the * The accuracy of this conclusion has been lately contested by M. Ue Montcgrc, whose report of the effects of the gastric juice of animals out of the body, docs not accord with that of Spallanzani; but the difference of cir- cumstances in which liis experiments were made, is quite sufficient to ac- count for the discrepancy in the results; and those of M. De Montcgre, there- fore, by no means, invalidate the general facts stated in the text, which have been estabhslicd by the experiments, not only of Spallanzani, but also of Reaumur, Stevens, Leuret, and Lassaigne. See Alison's Outlines of Phy- siology and Pathology, p. 170. DIGESTION. 135 mucous glands, and they are often constructed so as to pour their contents into intermediate cavities, or small sacs, which are denominated follicles, where the fluid is collected before it is discharged into the cavit}^ of the stomach. The o-astric dands of birds arc larger and more conspicuous than those of quadrupeds: but, independently of those which are situated in the stomach, there is likewise found, in almost all birds, at the lower termination of the ocsophngus, a large glandular organ, which has been termed the bulbulus glan- dulosus. In the Oslrich, this organ is of so great a size as to give it the appearance of a separate stomach. A view of the internal surface of the stomach of the African ostrich is given in Fig. 302; where c is the cardiac cavity, the coats 305 303 304 of which are studded with numerous glands; G, G, are the two sides of the gizzard. Fig, 303 shows one of the gas- tric glands of the African ostrich; Fig. 304, a gland from the stomach of the American ostrich, and Fig. 305, a sec- tion of a gastric gland in the beaver, showing the branching of the ducts, which form three internal openings. In birds that live on vegetable food, the structure of the gastric glands is evidently different from that of the corresponding glands in prcdaceous birds; but as these anatomical details have not as yet tended to elucidate in any degree the pur- . 136 THE VITAL FUNCTIOB^S. poses to which they are subservient in the process of diges- tion, I pass them over as being foreign to the object of our present inquiry.* It is essential to the perfect performance of digestion, that every part of tlie food received into the stomach sliould be acted upon by tlic gastric juice; for which purpose provi- sion is made that each portion shall, in its turn, be placed in contact with tlic inner surface of that organ. This is the more necessar}', as many facts render it ])robable, as will be noticed more particularly hereafter, that, besides the chemi- cal action of the gastric juice, an influence, derived from the nerves, essentially contributes to the accomplishment of the chemical changes which the food undergoes in the stomach. For this reason it is that the coats of the stomach are pro- vided with muscular fibres, passing, some in a longitudinal, others in a transverse, or circular direction; while a third set have an oblique, or even spiral course.t When the greater number of these muscles act together, they exert a considerable pressure upon the contents of the stomach; a pressure which, no doubt, tends to assist the solvent action of the gastric juice. When different portions act in succes- sion, they propel the food from one part to another, and thus promote the mixture of every portion with the gastric juice. We often fnid that the middle transverse bands con- tract more strongly tlian the rest, and continue contracted for a considerable time. The object of this contraction, which divides the stomach into two cavities, appears to be to separate its contents into two portions, so that each may be subjected to different processes; and, indeed, the differ- ences in structure, which are often observable between these two portions of the stomach, would lead to the belief that their functions are in some respects different. • These structures have been examined M'ith great care and minuteness by Sir Everard Home, who lias given the results of his inquiries in a scries of papers, read from time to time to the lioyal Society, and published in their Tmnsactions. t Sec rig. 51, vol. i. p. lOG, and its description, p. 107. DIGESTION. 137 During digestion the exit of the food from the stomach into the intestine is prevented by the pylorus being closed by the action of its sphincter muscle. It is clear that the food is required to remain for some time in the stomach in order to be perfectly digested, and this closing of the j)ylo- rus appears to be one means employed for attaining this end; and another is derived from the property which the gastric juice possesses of coagulating, or rendering solid, every animal or vegetable fluid susceptible of undergoing that change. This is the case with fluid albumen; the white of an egg, for instance, which is nearly pure albumen, is very speedily coagulated when taken into the stomach; the same change occurs in milk, which is immediately curdled by the juices that are there secreted, and these efi^ects take place quite independently of any acid that may be present. The object of this change from fluid to solid appears to be to detain the food for some time in the stomach, and thus to allow of its being thoroughly acted upon by the digestive powers of that organ. Those fluids which pass quickly through the stomach, and thereby escape its chemical ac- tion, however much they may be in themselves nutritious, are very imperfectly digested, and consequently afibrd very little nourishment. This is the case with oils, with jelly, and with all food that is much diluted.* Hunter ascertained * A diet consisting" of too large a proportion of liquids, although it may contain much nutritive matter, yet if it be incapable of being coagulated by the stomach, will not be sufficiently acted upon by that organ to be proper- ly digested, and will not only afford comparatively little nourishment, but be very liable to produce disorder of the alimentary canal. Thus, soups will not prove so nutritive when taken alone, as when they are united with a certain proportion of solid food, capable of being detained in the stomach, during a time sufficiently long to allow of the whole undergoing the pro- cess of digestion. I was led to this conclusion, not only from theory, but from actual observation of what took place among the prisoners in the Mil- bank Penitentiary, in 1823, when on the occasion of the extensive preva- lence of scorbutic dysentery in that prison, Dr. P. M. Latham and myself were appointed to attend the sick, and inquire into the origin of the disease. Among the causes which concurred to produce this formidable malady, one of the most prominent appeared to be an impoverished diet, consisting of a Vol. II. 18 13S THE VITAL FUNCTIONS. that this coagulating power belongs to the stomach of every animal which he examined for that purpose, from the most perfect down to reptiles;* and Sir E. Home has prosecuted the inquiry with the same result, and ascertained that this property is possessed by tlie secretion from the gastric glands, which communicates it to the adjacent membranes.t The gastric juice has also the remarkable property of cor- recting putrefaction. This is particularly exemplified in animals that feed on carrion, to whom this property is of great importance, as it enables them to derive wholesome nourishment from materials which would otherwise taint the whole system with their poison, and soon prove de- structive to life. It would appear that the first changes wdiich constitute digestion take place principally at the cardiac end of the stomach, and that the mass of food is gradually transferred towards the pylorus, the process of digestion still continuing as it advances. In the Babbit it has been ascertained that food newly taken into the stomach is always kept distinct from that which was before contained in it, and which has bco-un to undergo a change: for this purpose the new food is introduced into the centre of the mass already in the sto- mach; so that it may come in due time to be applied to the coats of that organ, and be in its turn digested, after the same change has been completed in the latter. :j: As the flesh of animals has to undergo a less considera- ble change than vegetable materials, so we find the stomachs of all the purely carnivorous tribes consisting only of a mem- branous bag, which is the simplest form assumed by this or- largc proportion of soups, on which the prisoners had subsisted for the pre- ceding; cii^ht months. A very full and perspicuous account of that disease has been drawn up, with great abihty, by my friend Ur. P. M. Latliam, and published under the title of "An account of the disease lately prevalent in the General Penitentiary." London, 1825. • Observations on the Animal Economy, p. 172. I Phil. Trans, for 1813, p. 9G. \ Sec Dr. Philip's ExperimcnUd Inquiry into the Laws of the Vital Func- tions, 3d edition, p. 122. STOMACHS OF MAMMALIA. 13a gan. But in other cases, as we have already seen, the sto- mach exhibits a division into two compartments by means of a slight contraction; a condition which, as Sir E. liome has remarked, is sometimes found as a temporary slate of the human stomach;* while, in other animals, it is the na- tural and permanent conformation. The Rodcntia furnish many examples of this division of the cavity into two dis- tinct portions, which exhibit even differences in their struc- •ture: this is seen in the Dormouse, (Fig. 306) the Beaver, the Hare, the Rabbit, and the cape Hyrax, (Fig. 307.) The first or cardiac portion is often lined with cuticle, while the lower portion is not so lined; as is seen very conspicuously in the stomachs of the Solipeda. The stomach of the Horse, in particular, is furnished at the cardia, with a spiral fold of the inner, or cuticular membrane, which forms a complete valve, offering no impediment to the entrance of food from the oesophagus, but obstructing the return of any part of the contents of the stomach into that passage.! This • The figure given of the human stomach, p. 182, shows it in the state of partial contraction here described. f The total inabihty of a horse to vomit is probably a consequence of tlie impediment presented by this valve. Sec Mem. du Museum d'llist. Nat. viii. 111. 140 THE VITAL FUNCTIONS. valve is shown in Fig. 311, wliich represents an inner view of the cardiac portion of the stomach of the horse; o being the termination of the oesophagus. The stomach of the Water Rat is composed of two dis- tinct cavities, having a narrow passage of communication: the first cavity is lined with cuticle, and is evidently in-* tended for the maceration of the food before it is submit- ted to the agents which are to efl'ect its digestion; a process which is completed in the se- cond cavity, provided, for that purpose, with a glandular sur- face. In proportion as nature allows of greater latitude In diet, we find her providing great complication in the digestive appara- tus, and subdividing the stomach into a greater numijer of ca- vities, each having probably a separate office assigned to it, thougli concurring in one general effect. A gradation in this respect mf»y be traced through a long line of quadrupeds, such as the Hog, the Peccari, the Porcupine, (Fig. 308,) and the Hippopotamus, where we find the number of separate pouches for digestion amounting to four or five. Next to these we may rank tlie very irregular stomach of the Kan- gnroo, (Fig. 309) composed of a multitude of cells, in which the food probably goes through several preparatory processes: and still greater complication is exhibited by the stomachs of the Cctacea, as, for example, in that of the Porpus (Fig. 310.) As the fishes upon which this animal feeds are swal- lowed wliole, and have large sharp bones, which would injure any surface not defended by cuticle, receptacles are provided, in which they may be softened and dissolved, and even con- verted into nourishment, by themselves, and without inter- fering with the di;rrstion of the soft parts. The narrow com- STOMACHS OF MAMMALIA. 141 munications between these several ^stomachs of the cetacea are probably intended to ensure the thorough solution of their contents, by preventing the exit of all such portions as have not perfectly undergone that process. Supernumerary cavities of this kind, belonging to the stomach, are more especially provided in those animals which swallow food either in larger quantity than is imme- diately wanted, or of a nature which requires much prepa- ration previous to digestion. The latter is more particularly the case with the horned ruminant tribes that feed on the leaves or stalks of vegetables, a kind of food, which, in pro- portion to its bulk, affords but little nutriment, and requires, therefore, a long chemical process, and a complicated diges- tive apparatus, in order to extract from it the scanty nutri- tious matter it contains, and prepare it for being applied t^ the uses pf the system. This apparatus is usually considered as consisting of four stomachs; and, in order to convey a distinct idea of this kind of structure, I have selected for re- presentation, in Fig. 312, that of the Sheep, of which the four stomachs are marked by the numbers 1, 2, 3, 4, re- spectively, in the order in which they occur, when traced from the oesophagus (c) to the intestine (p.) 142 THE VITAL FUNCTIONS. The orass, which is devoured in large quantities by these animals, and which undergoes but little mastication in the mouth, is hastily swallowed, and is received into a capacious reservoir, marked 1 in tlu; figure, called ihc paunch. This cavity is lined internally with a thick membrane, beset with numerous flattened ])a])illa^, and is often divided into pouches by transverse contractions. While the food remains in this bail, it continues in rather a diy state; but the moisture with which it is surrounded contributes to soften it, and to pre- pare it for a second mastication; which is effected in the fol- lowing manner. Connected with the paunch is another, but much smaller sac (2,) which is considered as the second sto- mach; and, from its internal membrane being thrown into numerous irregular folds, forming the sides of polygonal tells, it has been called the honcy-comh stomach, or reticule. Fig. 313 exhibits this reticulated appearance of the inner surface of this cavity. A singular connexion exists between this stomach and the preceding; for, while the asophagus ap- pears to open naturally into the paunch, there is, on each side of its termination, a muscular ridge, wdiich projects from the orifice of the latter, so that the two together form a chan- nel leading into the second stomach; and thus the food can readily pass from the oesophagus into either of these cavi- ties, according as the orifice of the one or the other is open to receive it. It would appear, from the observations of Sir E. Home, that liquids drank by the animal pass at once into the second stomach, the entrance into the first being closed. The food contained in the paunch is transferred, by small portions at a time, into this second, or honey-comb stomach, iil wiiich there is always a supply of water for moistening the portion of food introduced into it. It is in this latter stomach, then, that the food is rolled into a ball, and thrown up, through the oesophagus, into the mouth, where it is again masticated at leisure, and wliile the animal is reposing; a process which is well known by the name of chewing the cud, or rumi- 7iation. STOMACHS OF RUMINANTS. 143. When the mass, after being thoroughly ground down by the teeth, is again swallowed, it passes along the esophagus into the third stomach (3,) the orifice of which is brought forward by the muscular bands, forming the two ridges al- ready noticed, which are continued from the second sto- mach, and which, wheti they contract, effectually prevent any portion of the food from dropping into either of the pre- ceding cavities. In the ox, this third stomach is described by Sir E. Home, as having the form of a crescent, and as containing twenty-four septa, or broad folds of its inner membrane. These folds are placed parallel to one another, like the leaves of a book, excepting that they are of unequal breadths, and that a narrower fold is placed between each of the broader ones. Fig. 314 represents this plicated struc- ture in the interior of the third stomach of a bullock. What- ever food is introduced into this cavity, w^hich is named, from its foliated structure, the maiiy-plies stomach, must pass between these folds, and describe three-fourths of a cir- cle, before it can arrive at the orifice leading to the fourth stomach, which is so near that of the third, that the distance between them does not exceed three inches. There is, how- ever, a more direct channel of communication between the oesophagus and the fourth stomach (4,) along which milk taken by the calf, and which does not require to be either macerated or ruminated, is conveyed directly from the oeso- phagus to this fourth stomach; for, at that period, the folds of the many-plies stomach are not yet separated, and adhere closely together; and, in these animals, rumination does not take place, till they begin to eat solid food. It is in tliis fourth stomach, which is called the reed, that the proper di- gestion of the food is performed, and it is here that the coa- gulation of the milk takes place; on which account the coats of this stomach are employed in dairies, under the name of rennet, to obtain curd from milk. A rciiular jiradation in the structure of ruminatino; sto- machs may be traced in the different genera of this family of quadrupeds. In ruminants with horns, as the bullock 144 THE VITAL FUNCTIONS. and the sheep, tlicre are two preparatory stomachs for re- taining the food previous to rumination, a third for receiving it after it lias undergone this process, and a fourth for effect- ing its digestion. Ruminants without horns, as tiic Camel, Dromedar}", and Lama, have only one preparatory stomach before rumination, answering the purpose of the two first sto- machs of the bullock; a second, which I shall presently no- tice, and which takes no share in digestion, being employed merely as a reservoir of water; a third, exceedingly small, and of which the office has not been ascertained; and a fourth, which both receives and dij2;csts the food after rumination. Those herbivorous animals which do not ruminate, as the horse and ass, have only one stomach; but the upper portion of it is lined with cuticle, and appears to perform some pre- paratory office, which renders the food more easily digesti- ble by the lower portion of the same cavity.* The remarkable provision above alluded to in the Camel, an animal which nature has evidently intended as the inha- bitant of the steril and arid regions of the East, is that of reservoirs of water, which, when once filled, retain their contents for a very long time, and may minister not only to the wants of the animal that possesses it, but, also, to those of man. The second stomach of the Camel has a separate compartment, to which is attached a series of cellular ap- pendages; (exhibited, on a small scale, in Fig. 315:) in these the water is retained by strong muscular bands, which close the orifices of the cells, while the other portions of the sto- mach are performing their usual functions. By the relaxa- tion of these muscles, the water is gradually allowed to mix with the contents of the stomach, and thus the Camel is ena- bled to sujDport long marches across the desert, without re- ceiving any fresh supply. The Arabs, who traverse those extensive plains, accompanied by these useful animals, are, it is said, sometimes obliged, when faint, and in danger of perishing from thirst, to kill one of their camels, for the sake * Home, rhil. Trans. 8vo. 1806, p. 370. DIGESTION. 145 of the water contained in these reservoirs, which they al- ways find to be pure and wholesome. It is stated by those who have travelled in Egypt, that camels, when accustomed to go journeys, during which they are for a long time de- prived of water, acquire the power of dilating tlie cells, so as to make them contain a more than ordinary quantity, as a supply for their journey.* ^ When the Elephant, while travelling in very hot weather, is tormented by insects, it has been observed to throw out from its proboscis, directly upon the part on which the flies fix themselves, a quantity of water, with such force as to dislodge them. The quantity of water thrown out is in pro- portion to the distance of the part attacked, and is common- ly half a pint at a time: and this, Mr. Pierard, who resided many years in India, has known the elephant to repeat eight or ten times within the hour. The quantity of water at the animaPs command for this purpose, observes Sir E. Home, cannot, therefore, be less than six quarts. This water is not only ejected immediately after drinking, but six or eight hours afterwards. Upon receiving this information, Sir E. Home examined the structure of the stomach of that animal, and found in it a cavity, like that of the camel, perfectly well adapted to afford this occasional supply of water, which may, at other times, be employed in moistening dry food for the purposes of digestion.* In every series of animals belonging to other classes, a correspondence may be traced, as has been done in the jNIam- malia, between the nature of their food and the conformation of their digestive organs. The stomachs of birds, reptiles, and fishes, are, with certain modifications, formed very much upon the models of those already described, according as the food consists of animal or of vegetable materials, or presents more or less resistance from the cohesion of its texture. As it would be impossible, in this place, to enter into all the de- * Home, Lectures on Comparative Anatomy, vol. i. p. 171. t Supplement to Sir E. Home's Lectures on Comparative Anatomy, vol. vi. p. 9. Vol. II. 19 146 THE VITAL FUNCTIONS. tails necessary for fully illustrating this proposition, I must content myself with indicatino; a few of the most general re- suits of the inquiry.* As tiie food of birds varies, in difierent species, from the softest animal matter to the hardest grain, so we observe every gradation in their stomachs, from the membranous sac of the carnivorous tribes, which is one extreme, to the true gizzard of granivorous birds, which occupies the other extremity of the series. This gradation is established by the muscular fibres, which surround the former, acquiring, in difTcrent tribes, greater extent, and forming stronger mus- cles, adapted to the corresponding variations in the food, more especially as it partakes of the animal or vegetable character. In all the cold-blooded vcrtebrata, where digestion is not assisted by any internal heat, that operation proceeds more slowly, though in the end not less effectually, than in ani- mals where the contents of the stomach are constantly main- tained at a high temperature. They almost all rank as car- nivorous animals, and have accordingly stomachs, which, however they may vary in their form, are alike simply membranous in their structure, and act by means of the sol- vent power of their secretions. Among reptiles, only a few exceptions occur to this rule. The common sea-turtle that is brought to our tables, is one of these; for it is found to feed exclusively on vegetable diet, and chiefly on the sea- weed called zostira maritima, and the structure of its sto- mach corresponds exactly to the gizzard of birds. Some tortoises, also, which eat grass, make an approach to the same structure. In fishes, indeed, although the membranous structure of * The comparative anatomy of the stomach has been investig-atcd with great diligence by the late Sir E. Home, and the results recorded in the pa- pers he communicated, from time to time, to the I{oyul Society, and which have been republished in his splendid work, entitled *' Lectures on Compara- tive Anatomy," to which it will be seen that I have been largely indebted for the ficts and observations relating to this subject, detailed in the text. DIGESTION IN PISHES. 147 the stomach invariably accompanies the habit of preying upon other fish, yet there is one species of animal food, namely, shell-fish, which requires to be broken down by powerful means before it can be digested. In many fish, which consume food of this kind, its trituration is effected by the mouth, which is, for this purpose, as I have already noticed in the wolf-fish, armed with strong grinding teeth. But in others, an apparatus similar to that of birds is em- ployed; the office of mastication being transferred to the stomach. Thus, the Mullet has a stomach endowed witli a degree of muscular power, adapting it, like the gizzard of birds, to the double office of mastication and digestion; and the stomach of the Gillaroo trout, a fish peculiar to Ireland, exhibits, though in a less degree, the same structure. The common trout, also, occasionally lives upon shell-fish, and swallows stones to assist in breaking the shells. | Among the invertcbrated classes we occasionally meet with instances of structures exceedingly analogous to a giz- zard, and probably performing the same functions. Such is the organ found in the Sepia; the earth-worm has both a crop and a gizzard; and insects offer numerous instances, presently to be noticed, of great complexity in the structure of the stomach, wdiich is often provided, not only with a mechanism analagous to a gizzard, but also with rows of gastric teeth. ( 14S ) CIIx\PTEll VIII. Chrjl'ificalion. The formation of Chyle, or the Huid which is the imme- diate and cxchisivc source of nutriment to the system, takes ])lace in the intestinal tube, into which the chyme ])repared by the stomach is receiv'ed, and where farther chemical changes are effected in its composition. The mode in which the conversion of chyme into chyle is accomplished, and indeed the exact nature of the changes themselves, being, as yet, very imperfectly known, it is consequently impos- sible to trace distinctly the correspondence which, in all cases, undoubtedly exists between the objects, to be answered and the means employed for their attainment. No doubt can be entertained of the importance of the functions that are performed by structures so large and so complicated as are those composing the alimentary canal, and its various appendages. We plainly perceive that provision is made in the interior of that canal, for subjecting its contents to the action, first, of an extensive vascular and nervous sur- face; and secondly, of various fluid secretions, derived from different sources, and exercising powerful chemical agencies on the digested aliment; that a muscular power is su])plied, by means of the layers of circular and longitudinal fibres, contained between the outer and inner coats of the intes- tine,* for exerting a certain pressure on their contents, and for propelling them forwards by a succession of contractions, which constitutes what is termed their pcrisiallic motion; and lastly, that contrivances arc at the same time resorted to for retarding the progress of the aliment in its passage • See vol. i. p. 106. CHYLIFICATION. 149 i along the canal, so that it may receive tlie full action of these several agents, and yield the utmost (quantity of nutri- ment it is capable of affording. The total length of the intestinal tube differs much in dif- ferent animals, being in general, as already stated, smaller in the carnivorous tribes, than those which iced on substances of difficult dige stion, or affording but little nourishment. In these latter animals, the intestine is always of great length, exceeding that of the body many times; hence it is obliged to be folded into a spiral oi* serpentine course, forming many convolutions in the abdominal cavity. Sometimes, probably for greater convenience of package, instead of these nume- rous convolutions, a similai' effect of increasing the suri\ice of the inner membrane is obtained by raising it into a great number of folds, which project into tl>c cavity. These folds are often of consiflerable breadth, contributing not only to the extension of the surface foi' secretion and absorption, but also to the detention of the materials, with a view to their more complete elaboration. Remarkable examples of this kind of structure occur in most of the cartilaginous fishes, when the inner coat of the large in- testine is expanded into a broad fold, which, as is seen in fig. 316, representing this struc- ture in the interior of the intestine of the sliark, takes a spiral course; and this is con- tinued nearly the whole length of tlie canal, so that tlie internal surface is much augment- ed without any increase in the length of the intestine.* When the nature of the assimilatory pro- cess is such as to require the complete detention of the food, for a certain time, in particular situations, we find this ob- ject provided for by means of casca, or separate pouches • Structures of ttiis description have a particular claim to attention, from the light they tlirow on the nature of several fossil remains, lately investi- gated with singular success by Dr. Duckland. 150 THE VITAL FUNCTIONS. opening laterally from the cavity of the intestine, and having no otlicr outlet. Structures of this description have already hccn noticed in the infusoria,! and they are met with, indeed, in animals of every class, occurring in various parts of the alimentary tube, sometimes even as high as the pyloric por- tion of the stomach, and frequently at the commencement of the small intestine. Their most usual situation, however, is lower down, and especially at the part where the tube, af- ter having remained narrow in the first half of its course, is dilated into a wider cavity, which is distinguished from the former by the appellation of the great intestine, and which is frequently more capacious than the stomach itself. It is exceedingly probable that these two portions of the canal perform different functions in reference to the assimilation of the food: but hitherto no clew has been discovered to guide us through the intricacies of this difficult part of physiology; and we can discern little more than the existence already mentioned, of a constant relation between the nature of the aliment and the structure of the intestines, which are longer, more tortuous, and more complicated, and are furnished with more extensive folds of the inner membrane, and with larger and more numerous ca:ca, in animals that feed on ve- getable substances, than in carnivorous animals of the same class. The class of insects supplies numberless exemplifications of tlie accurate adaptation of the structure of the organs of as- similation to the nature of the food which is to be convert- ed into nutriment, and of the general principle that vegeta- ble aliment requires longer processes and a more compli- cated apparatus for this purpose, than that which has been al- ready animalized. In the herbivorous tribes, we find the oeso- phagus cither extremely dilatable, so as to serve as a crop, or receptacle for containing the food previous to its digestion, or having a distinct pouch appended to it for the same object: to this there generally succeeds a gizzard, or apparatus for • Page 73, of this volume. CHYLIFICATION. 151 trituration, furnished, not merely with a hard cuticle, as in birds, but also with numerous rows of teeth, of various forms, answering most effectually the purjDOse of dividing, or grind- ing into the minutest fragments, all the harder parts of the food, and thus supplying any deficiency of power in the jaws for accomplishing the same object. Thence the aliment, properly prepared, passes into the cavity appropriated for its dio-estion, which constitutes the true stomach.* In the lower part of this organ a peculiar fluid secretion is often in- termixed with it, which has been supposed to be analogous to the bile of the higher animals. It is prepared by the coats of slender tubes, termed hepatic vessels, which are often of great length, and sometimes branched or tufted, or beset, like the fibres of a feather, with lateral rows of fila- ments, and which float loosely in the general cavity of the body, attached only at their termination, where tliey open into the alimentary canal. t In some insects, these tubes are of larger diameter than in others: and in many of the or- thoptera, as we shall presently see, they open into large re- ceptacles, sometimes more capacious than the stomach itself, which have been supposed to serve the purpose of reservoirs of the biliary secretion, pouring it into the stomach on those occasions only when it is particularly wanted for the com- pletion of the digestive process. J * It is often diflficult to distinguish the portions of the canal, which cor- respond in their functions to the stomach, and to the first division of the in- testines, or duodenum; so that different naturalists, according to the views they take of the peculiar office of tliese parts, have applied to the same ca- vity the term of chjllferous stomach, or of du'odmum. See the memoir of Leon IJufour, in the Annales des Sciences Naturelles, ii. 473. f The first trace of a secreting structure, corresponding to hepatic vessels, is met with in the Asierlas, where the double row of minute lobes attached to the cxcal stomachs of those animals, and discharging their fluid into these cavities, are considered by Cams, as performing a similar office. The floc- culent tissue which surrounds the intestine of the Ilolothuria, is probably, also, an hepatic apparatus. \ A doubt is suggested, by Lcon Dufour, whether the liquid found in these pouches is real bile, or merely aUmcnt in the progress of assimilation, Ann. Sc. Nat. ii. 478. 152 THE VITAL FUNCTIONS. Tlic distinction into small and great intestine is more or less marked, in different insects, in proportion to the quan- tities of food consumed, and to its vegetable nature; and in herbivorous tribes, more especially, the dilatations in the lower part of the canal are most conspicuous, as well as the duplicatures of llie inner membrane, which constitute im- perfect valves for rctardinij; the progress of the alim.ent. It is generally at the j)oint where this dilatation of the canal commences, that a second set of hepatic vessels is inserted, having a structure essentially the same as those of the first set, iKit generally more slender, and uniting into a small number of ducts before they terminate. The number and complication of both these sets of hepatic vessels, appear to liave some relation to the existence and development of the gizzard, and consequently, also, to the nature and bulk of the food. Vessels of this description are, indeed, constantly found in insects; but it is only where a gizzard exists, that two sets of these secreting organs are provided; and in some larvae, remarkable for their excessive voracity, even three orders of hepatic vessels are met with.* A muscular power has also been provided, not only for the strong actions exerted by the gizzard, but, also, for the necessaiy propulsion, in different directions, of the contents both of the stomach and intestinal tubes. The muscular fibres of the latter are distinctly seen to consist of two sets, the one passing in a transverse or circular, and the other in a longitudinal direction. Glandular structures, analogous to the mucous follicles of the higlicr animals, are also plainly distinguishable in tlie internal coat of the canal, more espe- cially of herl)ivorous insects.! The whole tract of the ali- mentary canal is attached to the sides of the containing ca- vity by a fine membrane, ov ])eritoncuin, containing nume- rous air-vessels, or truchece.X • Sec the IMemoirs of Marcel dcs Serres, in the Annalcs du Museum, XX. 48. ■j- Lyonet. 4^ It has been stated by Malpighi and by Swammcrdam, and the statement DIGESTIVE ORGANS OF INSECTS. 153 To engage in a minute description of the endless varia- tions In the structure of the digestive organs, presented in the innumerable tribes which compose this class of animals, would be incompatible with the limits of this treatise. T shall content myself, therefore, with giving a few illustrations of their prin- cipal varieties, selected from those in which the leading characters of struc- ture are most strongly marked. I shall, with this view, exhibit first one of the simplest forms of the alimentary or- gans, as they occur in the Mantis reli- giosa, (Linn.) which is a purely carni- vorous insect, belonging to the order of Orthoptera. Fig 317 represents those of this insect, freed from their attach- ments, and separated from the body. The whole canal, as is seen, is perfect- ly straight: it commences by an oeso- phagus (o,) of great length, which is succeeded by a gizzard (g;) at the low- er extremity of this organ the upper hepatic vessels (b, b,) eight in number, and of considerable diameter, are in- serted: then follows a portion of the canal (d,) which may be regarded either as a digesting stomach, or a chyliferous duodenum: farther downwards, the second set of hepatic vessels (h h,) which are very numerous, but as slender as hairs, are received: and after a small contraction (n) there is again a slight dilatation of the tube (c) before it termi- nates. has been repeated by every succeeding- anatomist; that almost all tlie insects belong-ing" to the tribe of Grylll, possessed the faculty of ruminating- tiieir food; but this error has been refuted by Marcel des Serres, wlio luis offered satisfactory evidence that in no insect is the food subjected to a true rumina- tion, or second mastication, by the organs of the mouth. See Annales du Mu- seum, XX. 51 and 364. Vol. II. 20 154 THE VITAL FUNCTIONS. The alimentary canal of the Cicinchla campcstris, (Lin.) which preys on other insects, is represented in Fi^^. 31S; where we see that the lower part of the oesophagus (o,) is dilated into a crop (p,) succeeded by a small gizzard (g,) which is provided for the purpose of bruising the elytra, and other hard parts of their victims: but, their mechanical division being once efiected, we again find the true digesting stomach (s) simply membranous, and the intestine (i) very short, but dilated, before its termination, into a large colon (c.) The hepatic vessels (ii,) of which, in this insect, there is only one set, terminate in the cavity of the intestine by four ducts, at the point where that canal commences. A more complicated structure is exhibited in the alimen- tary tube of the Meloloniha vidf^aris^ or common cock- chafier, which is a vegetable feeder, devouring great quanti- /lies of leaves of jjlants, and consequently requiring a long DIGESTIVE ORGANS OF INSECTS. 1^5 and capacious canal for their assimilation; as is shown in Fig. 319, which represents them prepared in a similar man- ner to the former. In this herbivorous insect, the oesopha- gus (o) is, as might be expected, very short, and is soon di- 320 lated into a crop (*p;) this is followed by a very long, wide, and muscular stomach (s,) ringed like an earth-worm, and continued into a long and tortuous intestine (i, i,) which presents in its course several dilatations (c, c,) and re- ceives very elongated, convoluted, and rami- fied hepatic vessels (h,h.) Fig. 320 is a highly magnified view of a small portion of one of these vessels, showing its branched, form. In the alimentary canal (Fig. 321"^) o^ihe ^crida aptera (Stephens,) which is a species of grass- hopper, feeding chiefly on the dewberry, we observe a long oesophagus (o,) which is very dilatable, enlarging occasionally into a crop (i,) and succeeded by a round- ed or heart;shaped gizzard (g,) of very o 325 complicated structure, and connected with two remarkably large biliary pouches (u and B,) which receive, at their anterior 324 extremity, the upper set of hepatic ves- sels (v V.) A deep furrow in the pouch (b,) which, in the horizontal position of the body, lies underneath the gizzard, divides it apparently into two sacs. The intestinal canal is pretty uniform in its diameter, receives in its course a great K number of hepatic vessels (h ir,) by se- parate openings, and after making one convolution, is slightly constricted at n, and is dilated into a colon (c,) on the * The figures relating- to this insect were engraved from the drawings of Mr. Newport, who was also kind enough to supply me with the description of the parts they represent. Fig. 321 is twice the natural size. 156 THE VITAL FUNCTIONS. coats of which the longitudinal muscular bands are very dis- tinctly seen. Fig. 322 is a magnified view of the gizzard laid open, to show its internal structure. It is furnished with six longitudinal rows of large teeth, and six intermedi- ate double rows of smaller teeth; the total number of teeth being 270. One of the rows of large teeth is seen, detached, and still more magnified, in Fig. 323; it contains at the up- per part, five small hooked teeth (f,) succeeded below by four broad teeth (d,) consisting of quadrangular plates, and twelve tricuspid teeth (t;) that is, teeth having three cusps, or points at their edges. Fig. 324 shows the profile of one of these teeth; a, being the sharp point by which the ante- rior acute angle of the base terminates. Fig. 325 exhibits the base of the same tooth seen from below, e, e, e, being the three cusps, and m, the triangular hollow space for the insertion of the muscles which move them, and which com- pose part of the muscular apparatus of the gizzard. The smaller teeth, which are set in double lines between each of the larger rows, consist of twelve small triangular teeth in each row. All the teeth contained in this organ are of a brown colour and horny texture, resembling tortoise shell. The same insect, as we have seen, often exhibits, at dif- ferent periods of its existence, the greatest contrast, not only in external form, but also in its habits, instincts, and modes of subsistence. The larva is generally remarkable for its voracity, requiring large supplies of food to furnish the m,a- terials for its rapid growth, and frequently consuming enor- mous quantities of fibrous vegetable aliment: the perfect in- sect, on the other hand, having attained its full dimensions, is sufliciently supported by small quantities of a more nu- tritious food, consisting cither of animal juices, ^r of the fluids prepared by flowers, which are generally of a saccha- rme quality, and contain nourishment in a concentrated form. It is evident that the same apparatus, which is ne- cessary for the digestion of the bulky food taken in during the former period, would not be suited to the assimilation of that which is received during the latter; and that in order DIGESTIVE ORGANS OP INSECTS. 157 to accommodate it to this altered condition of its function, considerable changes must be made in its structure. Hence, it will be interesting to trace the gradual transitions in the conformation of the alimentary canal, during the progressive development of the insect, and more especially while it is undergoing its different metamorphoses. These changes are most conspicuous in the Lepidoptera, where we may observe the successive contractions which take place in the immensely voluminous stomach of the ca- terpillar, while passing into the state of chrysalis, and thence into that of the perfect insect, in which its form is so changed that it can hardly be recognised as the same organ. I have 328 327 326 I given representations of these three different states of the en- tire alimentary canal of the Sjjhinx ligiislri, or Privet Ilawk- moth, in Figures 326, 327, and 32S;* the first of which * Tliesc figures also have been engraved from the drawings of Mr. New- port, which he was so obliging as to make for mc, from preparations of his own, the result of very careful dissections. 158 THE VITAL FUNCTIONS. is that of tlic caterpillar; the second, that of the chrysalis; and the third, that of the moth. The whole canal and its appendages, have been separated from their attachments, and spread out so as to display all their parts; and they are de- lineated of the natural size, and in each case, so as to show their comparative dimensions in these three states. In all the figures, A is the oesophagus; b, the stomach; c, the small intestine; d, the ccecal portion of the canal; and e, the colon, or large intestine. The hepatic vessels are shown at f; and the gizzard, which is developed only in the moth, at g, Fig. 328. It will be seen that in the caterpillar, (Fig. 326,) the sto- mach forms by far the most considerable portion of the ali- mentary tube, and that it bears some resemblance in its struc- ture and capacity to the stomachs of the Annelida, already described.* This is followed by a large, but short, and per- fectly straight intestine. These organs in the pupa (Fig. 327) have undcgone considerable modifications, the whole canal, but more especially the stomach, being contracted both in length and width if the shortening of the intestine not being in proportion to that of the whole body, obliges it to be folded upon itself for a certain extent. In the moth, Fig. 328,) the contraction of the stomach has proceeded much farther; and an additional cavity, which may be consi- dered as a species of crop or gizzard (g,) is developed: the small intestine takes a great many turns during its course, and a large pouch, or cxcum, has been formed at the part where it joins the large intestine. The hepatic vessels are exceedingly numerous in the Crus- tacea, occupying a very large space in the general cavity; and they compose by their union an organ of considerable size, which may be regarded as analogous in its functions to • Sec the figures and description of those of the Nais and the Leech, p. 102 and 103. f Cams states tliat he found tlie stomach of a pupa, twelve days after it had assumed that state, scarcely half as long, and only one-sixth as wide as it had been in the caterpillar. DIGESTIVE ORGANS OF MOLLUSCA. 159 the Liver of the higher classes of animals. This organ ac- quires still greater size and imporUnce in the Mollusca, where it frequently envelops the stomach, pouring the bile into its cavity by numerous ducts.* As the structure and course of the intestinal canal varies greatly in diflcrcnt tribes of Mollusca, they do not admit of being comprised in any general description. The only examples I think it necessa- ry to give, in this class, are those of the Patella, or Limpet, and of the Pleiiro- branchiis. The intestinal tube of the Patella is delineated in Fig. 329; where M is the mouth; t, the tongue folded back; o, the oesophagus; and s, the sto- mach, from which the tortuous intestinal tube is seen to be continued. All the convolutions of this tube, as well as the stomach itself, are enclosed, or rather imbedded, in the substance of the liver, which is the largest organ of the body. The Pleurobranchiis Peronii (Cuv.) is remarkable for the number and complication of its organs of digestion. They are seen laid open in Fig. 330; where c is the crop; g, the gizzard; p, a plicated stomach,resemblingthc third stomach of ruminant quadrupeds; and d, a fourth ca- vity, being that in which digestion is com- pleted. A canal of communication is seen at T, leading from the crop to this last cavity: b is the point where the biliary duct enters. In the Cephalopoda, the structure of these organs is very complicated; for they are pro- vided with a crop, a muscular gizzard, and a caecum, which has a spiral form. In these animals we also discover the rudiment of another auxiliary * Transparent crystalline needles, the nature ami uses of which are quite unknown, are frequently found in the biliary ducts of this class of animals. 160 THE VITAL FUNCTIONS. / organ, namclvj the Panaceas, which secretes a fluid contri- buting to tlie assimihitfon of the food. This organ becomes more and more developed as we ascend in the scale of ani- mals, assuming a glandular character, and secreting a watery fluid, which resembles the saliva, both in its sensible and chemical properties. It has been conjectured that many of the vessels, which are attached to the upper portion of the alimentary canal of insects, and have been termed hepatic, may, in fact, prepare a fluid having more of the qualities of the pancreatic than of the biliary secretion. The alimentary canal of fishes is in general characterized by being short; and the continuity of the stomach with the intestines is often such as to. oflcr no well marked line of distinction between them. The ca3ca are generally large and numerous; and a number of tubular organs, connected more especially with the pyloric appendices, arc frequently met with, resembling a cluster of worms, and having some analogy, in situation at least, to the hepatic or pancreatic vessels of insects. Their appearance in the Sabyion is re- presented at p, in Fig. 331. The pancreas itself is only met with, in this class of ani- mals, in the order of cartilaginous fishes, and ire especially in the Ray and the Shark tribes. A distinct gall-bladder, or reservoir, is also met with in some kinds of fish, but is by no means general in that class. In the class both of Fishes and of Reptiles, which are cold-blooded animals, the processes of digestion are conducted more slowly than in the more energetic sys- tems of Birds and of Mammalia; and the comparative length of the canal is, on the whole, greater in the former than in the latter: but the chief diflcrenccs in this respect depend on the kind of food which is consumed, the canal being always shortest in those tribes that are most carnivo- rous.^ As the Frog, in the different stages of its growth. • See Home, Lectuies, &c. I. 401. DIGESTIVE ORGANS OF MAMMALIA. 161 lives upon totally different kinds of food, so we find that the structure of its alimentary canal, like that of the moth, undergoes a material change during these metamorphoses. The intestinal canal of the tad-pole is of great length, and is collected into a large rounded mass, composed of a great number of coils, which may easily be distinguished, by the aid of a magnifying glass, through the transparent skin. During its gradual transformation into a frog, tliis canal be- comes much reduced in its length; so that when the animal has attained its perfect form, it makes but a single convolu- tion in the abdominal cavity. A similar correspondence exists between the length of the canal, and the nature of the food in the class of Birds. At the termination of the small intestine there are usually found two caeca, which, in the gallinaceous and the aquatic fowls, are of great length: those of the ostrich contain in their interior a spiral valve. Sir E. Home is of opinion that in these animals the functions of the pyloric portion of the stomach are performed by the upper part of the intes- tine. In the intestines of the Mammalia contrivances are em- ployed with the apparent intention of preventing their con- tents from passing along too hastily: tliese contrivances are most effectual in animals whose food is vegetable, and con- tains little nourishment, so that the whole of what the food is capable of yielding is extracted from them. Sir E. Home observes that the colon, or large intestine of animals which live upon the same species of food, is of greater length, in proportion to the scantiness of the supply. Tluis, the length of the colon of the Elephant, which inhabits the fertile woods of Asia, is only 26^ feet; while, in the Dromedary, which dwells in the arid deserts of Arabia, it is 42. This con- trast is still more strongly rwarked in birds. The Cassowa- ry of Java, which lives amidst a most luxuriant suppl}' of food, has a colon of one foot in length, and two caica, each of which is six inches long, and one quarter of an inch in Vol. II. 21 162 THE VITAL FUNCTIONS. diameter. The African ostrich, on the other hand, which inhabits a country where the supply of food is very scanty, has the colon forty-five feet long; each of the cseca is two feet nine inches in length, and, at the widest part, three inches in diameter; in addition to which, there arc broad valves in the interior of both these cavities.* On comparing the structure of the digestive organs of Man with those of other animals belonging to the class Mam- malia, we find them holding a place in the series intermedi- ate between those of tlie purely carnivorous, and exclusively herbivorous tribes, and, in some measure, uniting the cha- racters of both. The powers of the human stomach do not, indeed, extend to the digestion either of the tough woody fibres of vegetables, on the one hand, or the compact texture of bones on the other; but, still, they are competent to ex- tract nourishment from a wider range of alimentary sub- stances, than the digestive organs of almost any other animal. This adaptation to a greater variety of food may also be in- ferred from the form and disposition of the teeth, which coml)inc those of different kinds more completely than in most mammalia, excepting, perhaps, the Quadrumana, in whicii, however, the teeth do not form, as in man, an unin- terrupted series in both jaws. In addition to these pecu- liarities, we may also here observe, that the sense of taste, in the human species, appears to be affected by a greater variety of objects than in the other races of animals. All these arc concurring indications that nature, in thus render- ing man omnivorous, intended to qualify him for maintain- ing life wherever he could procure the materials of subsist- ence, whatever might be their nature, whether animal or vegetable, or a mixture of both, and in whatever soil or * Lectures, &c. I. 470. In the account above given of the digestive or- gans I have pui-poscly omitted all mention of the spleen; because, although it is probably in some way related to digestion, the exact nature of its func- tions has not yet been determined with any certainty. DIGESTIVE ORGANS OP MAN. 1G3 climate they may be produced; and for endowing liini with the power of spreading his race, and extending liis dominion over every accessible region of the globe. Thus, then, from the consideration of the peculiar structure of the vitiil, as well as the mechanical organs of his frame, may be de- rived additional proofs of their being constructed with re- ference to faculties of a higher and more extensive range than those of any, even the most favoured species of the brute creation. ( 164 ) CHAPTER IX. LACTEAL ABSORPTION. The Chyle, of which wc have nowtraccd the formation, is a fluid of uniform consistence, perfectly hland and unirri- tating in its properties, the elements of which have been brought into that precise state of chemical composition which renders them fit to be distributed to every part of the sys- tem for the purposes of nourishment. In all the lower or- ders of animals it is transparent; but the chyle of mammalia often contains a multitude of globules, which give it a white colour, like milk. Its chemical composition appears to be very analogous to that of the blood into which it is afterwards converted. From some experiments made by my late much valued friend Dr. INIarcet, it appears that the chyle of dogs, fed on animal food alone, is always milky, whereas, in the same animals, when they are limited to a vegetable diet, it is nearly transparent and colourless.* The chyle is absorbed from the inner surface of the intes- tines by the Lacteals, which commence by very minute ori- fices, in incalculable numbers, and unite successively into larger and larger vessels, till they form trunks of considera- ble size. They pass between the folds of a very fine and delicate membrane, called the Tuesentery^ which connects the intestines to the spine, and which appears to be inter- posed in order to allow them that degree of freedom of mo- tion, which is so necessary to the proper performance of their functions. In the mesentery, the lacteals pass through several glandular bodies, germed the vicsenleric glands, where it is probable that the chyle undergoes some modifi- cation, preparatory to its conversion into blood. • Medico-Chii'urgical Transactions, vi. 630. LACTEAL ABSORPTION. 1G5 The mesenteric glands of the Whale contain large spheri- cal cavities, into which the trunks of the lacteals open, and where the chyle is probahly blended with secretions proper to those cavities; but no similar structure can be detected in terrestrial mammalia. It is only among the Vertebrata that lacteal vessels are met with. Those of Fishes are simple tul)es, either wholly without valves, or if there be any, they are in a rudimcntal state, and not sufficiently extended to prevent the free pas- sage of their fluid contents in a retrograde direction. The lacteals of the Turtle are larger and more distinct than those of fishes, but their valves are still imperfect, though they present some obstruction to descending fluids. In Birds and in Mammalia these valves are perfectly eflcctual, and arc exceedingly numerous, giving to the lacteals, when distend- ed with fluid, the appearance of strings of beads. The ef- fect of these flood-gates, placed at such short intervals, is that every external pressure made upon the tube, assists in the propulsion of the fluid in the direction in which it is intend- ed to move. Hence it is easy to understand how exer- cise must tend to promote the transmission of the chyle. The gflands are more numerous and concentrated in the Mammalia, than in any other class. From the mesenteric glands the chyle is conducted, by the continuation of the lacteals, into a reservoir, which is termed the receptacle of the chyle; whence it ascends through the thoracic diict^^ which passes along the side of the spine, in a situation afibrding the best possible protection from in- jury or compression, and opens into the great veins leading directly into the heart. In invertebrated animals having a circulatory system of vessels, the absorption of the chyle is performed by veins instead of lacteal Vessels. The sanguification of the chyle, or its conversion into blood, takes place, during the course of the circulation, and • This duct Is occasionaUy double. IGG THE VITAL FUNCTIONS. is principally cfTectcd by the action of atmospheric air in certain organs, hereafter to be described, where that ac- tion, or aeration as it may be termed, in common with an analogous process in vegetables, takes place. In all verte- brated animals the blood has a red colour, and it is also red in most of the Annelida; but in all other invertebratcd ani- mals, it is cither white or colourless.* We shall, for the present, then, consider it as having undergone this change, and j)roceed to notice the means employed for its distribu- tion and circulation throughout the system. • Vauquclin has observed that the chyle has often a red tinge in animals. ( 1G7 ) CHAPTER X. Circulation. § 1. Diffused Circulation. Animal life, implying mutual actions and reactions be- tween the solids and fluids of the body, requires for its maintenance the perpetual transfer of nutritive juices from one part to another, corresponding in its activity to the ex- tent of the changes which are continually taking place in the organized system. For this purpose we almost con- stantly find that a circulatory motion of the nutrient fluids is established; and the function which conducts and regu- lates their movements is emphatically denominated the Cir- culation. Several objects of great importance are answered by this function; for in the first place, it is through the cir- culation that every organ is supplied wiih the nutritive particles necessary for its development, its growth and the maintenance of its healthy condition; and that the glands, in particular, as well as the other secreting organs, arc fur- nished with the materials they require for the elaboration of the products, which it is their peculiar ofiice to prepare. A second essential object of the circulation, is to transmit the nutritive juices to certain organs, where they are to be sub- jected to the salutary influence of the oxygen of the atmo- sphere; a process which in all warm-blooded animals, com- bined with the rapid and extensive distribution of the blood, difl'uses and maintains throughout the system the high tem- perature required by the greater energy of their functions. Hence it necessarily follows that the particular mode in which the circulation is conducted in each respective tribe, 168 THE VITAL FUNCTIONS. must influence every other function of the economy, and must, therefore, constitute an essential element in deter- mining the physiological condition of the animal. We fmd, accordingly, that among the characters on which systematic zoologists have founded their great divisions of the animal kingdom, the utmost importance is attached to those de- rived from differences of structure in the organs of circula- tion. A comprehensive survey of the different classes of ani- mals with reference to this function, enables us to discern the existence of a regular gradation of organs, increasing in complexity as we ascend from the lower to the higher or- ders; and showing that here, as in other departments of the economy of nature, no change is made abruptly, but always by slow and successive steps. In the very lowest tribes of Zoophytes, the modes by which nutrition is accomplished can scarcely be perceived to differ from those adopted in the vegetable kingdom, where, as we have already seen, the nu- tritive fluids, instead of being confined in vessels, appear to permeate the cellular tissue, and thus immediately supply the solids with the materials they require; for, in the sim- pler kinds of Polypi, of Infusoria, of JVIedusee, and of Ento- zoa, tlie nourishment which has been prepared by the di- gestive cavities is apparently imbibed by the solids, after having transuded through the sides of these organs, and without its being previously collected into other, and more general cavities. This mode of nutrition, suited only to the torpid and half vegetative nature of zoophytes, has been de- nominated nourishment by imbibition, in contradistinction to that by circulation^ a term, which, as we have seen, im- plies, not merely a system of canals, such as those existing in Medusae, where there is no evidence of the fluids really circulating, but an arrangement of ramified vessels, composed of membranous coats, through which the nutrient fluid moves in a continued circuit. The distinction whicli has thus been drawn, however, is one on which wc should be careful not to place undue reli- DIFFUSED CIRCULATION. 1G9 nnce, for It is founded, perhaps, more on our Imperfect means of investigation, tlian on any real diflercnces in the ])ro- cedurcs of nature relative to this function. When the juices, either of plants, or of animals, are transparent, their motions are imperccptihlc to the eye, and can be judged of only by other kinds of evidence; but when they contain globules, differing in their density from that of the fluid, and there- fore capable of reflecting light, as is the case with tlie sap of the Chara and Caulinia, we have ocular proof of the ex- istence of currents, which, as long as the plant is Jiving and in health, pursue a constant course, revolving in a regular and deflned circuit; and all ])lants which have milky juices exhibit this phenomenon. Although the extent of each of these vegetable currents is very limited, compared with the. entire plant, it still presents an examj)lc of the tendency which the nutrient fluids of organized structures have to move in a circuit, even when not confined within vessels or narrow channels; for this movement of rolalion, or cf/clo.siSj as it has been termed,* whatever may be its cause, appears always to have a definite direction. The current returns into itself, and continues without intermission, in a manner much resembling the rotatory movements occasionally pro- duced in fluids by electro-magnetism. t Movements, very similar in their appearance and cha- racter to those of vegetable cyclosis, have been recently dis- covered in a great number of polyferous Zoophytes, by Mr. Lister, who has communicated his observations in a paper which was lately read to the Royal Society, and of which the following are the principal results. In a specimen of the Tubulai'ia indivisa, when magnified one hundred times, a current of particles was seen within the tubular stem of the polype, strikingly resembling, in the steadiness and continuity of its stream, the vegetable circulation in the * See pages 41 and 42 of this volume. f So great is this resemblance, that it has led several physiologists to as- cribe these movements to the agency of electricity; but there docs not, as yet, appear to be any substantial foundation for this hypothesis. Vol. II. 22 170 THE VITAL FUNCTIONS. Chara. Its general course was parallel to the slightly spiral lines of irregular spots on the surface of the tube, ascending on the one side, and descending on the other; each of the opposite currents occu])ying one-half of the circumference of the cylindric cavity. At the knots, or contracted parts of the tube, sliglit eihlies were noticed in the currents; and at each end of the tube the particles were seen to turn round, and pass over to the otiier side. In various species of *S'er- tiilariic the stream does not flow in the same constant di- rection; but, after a time, its velocity is retarded, and it then either stops, or exhibits irregular eddies, previous to its return in an op])osite course; and so on alternately, like the ebb and flow of the tide. If the currents be designedly obstructed in any part of the stem, those in the branches go on without inteiTuptlon, and independent!}^ of the rest. The most remarkable circumstance attend In 2; these streams of fluid is that they appear to traverse the cavity of the sto- mach itself, flowing from the axis of the stem into that or- gan, and returning into the stem without any visible cause determining these movements. Similar phenomena were observed by Mr. Lister in Campanxdarhe and Flumula- rix. In some of the minuter species of Crustacea the fluids have been seen, by the aid of the microscope, moving with- in the cavities of the body, as if by a spontaneous impulse, without the aid of a propelling organ, and apparently with- ' out being confined in membranous channels, or tubes of any sort. This kind of diff'used circulation is also seen in the embryos of various animals, at the earliest periods of their development, and before any vessels are formed. § 2. Vascular Circulation. The next step in the gradation of structures consists in the presence of vessels, within which the fluids arc confined, and by which their course and their velocity are regulated j VASCULAR CIRCULATION. 171 and, in general, these vessels form a complete circuit. The first rudiments of a vascular organization arc those observed and described by I'iedemann, in the Jl^tcrivc, which are si- tuated higher in the animal scale than McduscT; but whether any actual circulation takes place in the channels constituted by these vessels, which communicate both with the cavitv of the intestine, and with the respiratory organs, is not yet determined with any certainty. The ITolothuriin, which also belong to the order of Echinodcrmata,are furnisiied with a complex apparatus of vessels, of which the exact functions are still unknown. In those species of Entozoa which ex- hibit a vascular structure, the canals appear rather to be ra- mifications of the intestinal tube, than proper vessels, for no distinct circulation can be traced in them: an organization of this kind has already been noticed in the Txnix:* It was, till very lately, the prevailing opinion among na- turalists that all true insects are nourished by imbibition, and that there exists in their system no real vascular circu- lation of juices. In all the animals belonging to this class and in every stage of their development, there is found a tubular organ, called the dorsal vessel, extending the whole length of the back, and nearly of uniform diameter, except where it tapers at the two ends. It contains a fluid, which appears to be undulated backwards and forwards, by means of contractions and dilatations, occurring in succession in different parts of the tube; and it is also connected with transverse ligamentary bands, apparently containing muscu- lar fibres, capable, by their action, of producing, or, at least, of influencing these pulsatory movements. An enlaro-ed re- presentation of the dorsal vessel of the Mclolontha vulgaris, or common cockchaffer, isolated from its attachments, is given in Fig. 333, showing the series of dilatations (v, v, v) which it usually presents in its course; and in Fig. 334, the same vessel is exhibited in connexion with the ligamentary * Page 64, in this volume, Tit^. 247. The family of rianarix present ex- ccptions to this genenxl rule: for many species possess a system of circu- lating vessels. Sec Duges, Amiales des Sciences Naturellesj xv. 161. 172 THE VITAL FUNCTIONS. and ninscular apparatus which surrounds it, seen from the lower side. In the hist of these figures, a is the tapering 336 ^ )i if f prolongation of the tube, proceeding towards the head of tlie insect; v, one of the dilated portions, or ventricles, as they have been called, of the dorsal part of the tube; f, one of the small tendinous folds, to which the ligamentary bands are attached; and l is one of these bands, having a triangular, or, if considered as continuous with that on the other side of the vessel, a rliomboidal shape, and attached at h, to the su- jierior segments of the abdomen. At i is seen a layer of the same fibres, wliich are partly ligamentous and partly muscu- lar, passihg underneath the dorsal vessel, and forming, in conjunction with the layer that passes above it, a sheath, which embraces and fixes that vessel in its place: these in- ferior layers have been removed from the other parts of the vessel, to allow the upper layers to be seen, as is the case at CIRCULATION IN INSECTS. 173 L. Fig. 335 gives a side view of the anterior extremity of the same vessel, showing tlie curve (a) which it dcscrihcs as it bends downwards in its course towards the head. The function performed by tlie dorsal vessel, which, judging from the universal presence of this organ in insects, must be one of great importance in their economy, was long a profound mystery. Its analogy in structure and position to the dorsal vessels of the Arachnida and the Annelida, r where it evidently communicates with channels of circu- lation, and exhibits movements of pulsation resembling those of insects, was a strong argument in favour of the opinion that it is the prime mover of a similar kind of circulation; but then, again, this hypothesis appeared to be overturned by the fact that no vessels of any kind could be seen extend- ing from it in any direction; nor could any channels for the transmission of a circulating fluid be detected in any part of the body. Those organs, which, in animals apparently of an inferior rank, are most vascular, such as the stomach, the intestinal tube, the eye, and other apparatus of the senses, seemed to be constructed, and to be nourished, by means to- tally different from those adopted in the former animals. Although extremely minute ramifications of air tubes are every where visible in the interior of insects, yet, neither Cuvier, nor any other anatomist, could succeed, by the closest scrutiny, in detecting the least trace of blood vessels; and the presumption, therefore, was, that none existed. But it still remained a question, if the dorsal vessel be not subservient to circulation, what is its real function? Marcel des Serres, who bestowed great pains in investi- gating this subject, came to the conclusion that its use is to secrete the fatty matter, which is generally found in great abundance in the abdominal cavity, and which is accumu- lated particularly around the dorsal vessel."^ A more at- tentive examination of the structure of the vessel itself brought to light a valvular apparatus, of which the only con- * See liis vai'ious papers in the Memolrcs clu Museum d'liist. Nat.; torn. iv. and v. 174 THE VITAL FUNCTIONS. ceivable purpose is that of determining the motion of the contained fluid in one constant course; a purpose necessarily- incompatible with its supposed alternate undulation in op- posite directions, from one end of the tube to the other. These valves are exhibited in Fig. 336, in a still more mag- nified view of a longitudinal section of the dorsal vessel, showing tlie semicircular folds (s, s) of its inner membrane, which perform the function of valves by closing the passage against any retrograde motion of the fluid. This discovery of valves in the dorsal vessel, again made the balance of pro- bability incline towards the opinion that it is the agent of some kind of circulation. All doubt as to the reality of a circulation in insects is now dispelled by the brilliant discoveries of Professor Ca- rus, who, in the year 1824, first observed this phenomenon in the larva of the Agrion puella. In the transparent parts of this insect, as w^ell as of many others, numerous streams of fluid, rendered manifest by the motions of the globules they contain, are seen meandering in the spaces which in- tervene between the layers of the integument, but without appearing to be confined within any regular vessels. The streams on the sides of the body all pass in a direction back- wards from the head, till they reach the neighbourhood of the posterior end of the dorsal vessel, towards which they all converge; they are then seen to enter that vessel, and to be propelled by its pulsations towards its anterior extremi- ty, where they again issue from it, and are subsequently di- vided into the scattered streams, which descend along the sides of the body, and which, after having thus completed their circuit, return into the pulsating dorsal vessel. This mixed kind of circulation, partly difl'used and partly vascular, is beautifully seen in the larva of the Ephemera marginata,'^' where, besides the main current, which, after • This insect is figured and described in Dr. Goring- and Mr. Pritchard's *' Microscopic Illustrations," and its circulation is very fully detailed, and il- lustrated by an engraving- on a large scale, by Mr. Bowerbank, in the Ento- mological Magazine, i. 239; plate ii. CIRCULATION IN INSECTS. 175 being discharged from the anterior extremity of the dorsal vessel, descends in a wide spreading stream on each side and beneath that vessel, another portion of the blood is con- veyed by two lateral trunks, which pass down each side of the body, in a serpentine course, and convey it into the lower extremity of the dorsal vessel, with which they are continuous. These are decidedly vessels, and not portions of the great abdominal cavity, for their boundaries arc clearly defined; yet they allow the blood contained in them to escape into that cavity, and mix with the portion previ- ously diffused. All these wandering streams sooner or later find their way into the dorsal vessel, being absorbed by it at various points of its course, where its membranous coat is reflected inwards to form the valves. In the legs, the tail, and the antennas, the circulation is carried on by means of vessels, which are continuous with the lateral vessels of the body, branching off from them in the form of loops, as- cending on one side, and then turning back to form the de- scending vessel, so that the currents in each move in con- trary directions. Fig. 337 represents the appearance of c.-.. - -^x ^ — * ^^^::":^>'''^•r■^r■•■.,^^•^^^'::i'■^^^^^^^ these parallel vessels in one of the antennae, of the Seynhlis viridis, magnified thirty times its natural size. The whole 176 THE VITAL PTTNCTIONS. system of circulating vessels in tliat insect, of which the for- mer is only a dctaclied part, is sliown in Fig. 33S, where the course of the Ijlood is indicated h}' arrows; A, repre- senting the currents in the antenna:; w, those in the rudi- mental wings; and t, those in the tail; in nil which parts the vessels form loops, derived from the main vessels of the trunk. In some larvre tlie vascular loops, conveying tliesc collateral streams, pass only for a certain distance into the legs; sometimes, indeed, they proceed no farther than the haunches. The currents of hlood in these vessels have not a uniform velocity, heing accelerated hy the impulsions they receive from the contractions of the dorsal vessel, which appears to he the prime agent in tlicir motion. As the insect advances to maturity, and passes through its metamorphoses, considerahle changes are observed to take place in the organization of the circulating system, and in the energy of the function it performs. The vessels in the extreme parts, as in the tail, are gradually obliterated, and the circulation in them, of course, ceases, the blood ap- pearing to retire into the more internal parts. In the wings, on the other hand, where the development proceeds rapidly, the circulation becomes more active; and even after tliey have attained their full size, and are yet in a soft state, the motion of the blood in the centre of all the nervures is dis- tinctly visible:^ but afterwards, as the wings become dry, it ceases there also, and is then confined to the vessels of the trunk. In proportion as the insect approaches to the com- pletion of its development, these latter vessels also, one after the other, shrink and disappear, till, at length, nothing which had once appertained to this system remains visible, except the dorsal vessel. But, as \vc observe this vessel still con- tinuing its pulsatory movements, we may fairly infer that they are designed to maintain some degree of obscure and imperfect circulation of the nutrient juices, through vessels, • These currents in the wing- of the Semblls hilineata have been described and delineated by Carus, in the Acta Acad. Cxs. Lcop. Carol. Nat. Cur. vol. XV. part ii. p. 9. CIRCULATION IN INSECTS. 177 which may, in their contracted state, corresponding to the diminished demands of the system, have generally escaped detection. In confirmation of these views, it may be stated, that several observers have, at length, succeeded in tracing minute branches, proceeding in diflcrent directions, from the dorsal vessel, and distributed to various organs. Tiie divi- sion of the anterior part of the dorsal vessel into descending branches was noticed by Comparetti. Dug^s has observed a similar division of this vessel in the corselet of several spe- cies of Phalcnx, and farther ramifications in that of the Gryllus lineola: and Audouin has traced them in many of the Hymenoptera.* • Annales des Sciences Natiirelles, xv. 308. The figures whicli follow (from 339 to 345) are representations, of the na- tural size, of the doi-sal vessel of the Sphinx ligustri, or Privet Hawk-moth, which has been dissected in its three different stages, with great care, by Mr. Newport, from whose drawings these figures have been engraved, and to whom I am indebted also for the description which follows: — The doreal vessel of this,insect is an elongated and gradually tapering ves- 339 sel extending from the hinder part of the abdomen, along the back, towards the head; and furnished with valves, which con-espond very nearly in their situation to the incisions of the body. During the changes of the insect from the lai-va to the imago state, it undergoes a slight modification of form. In every state it'may be distinguished into two portions, a dorsal and an aor- Vol. II. " 23 178 THE VITAL FUNCTIONS. The discovery of the circulation in insects, and of its va- rying energy at different periods of growth, has elucidated many obscure points in the physiology of this important ial. The dorsal portion, which is the ojie in which a pulsation is chiefly ob- servable, is furnished with distinct valves, is attached along the dorsal part of the body by lateral muscles, and has vessels which enter it laterally, pouring into it the circulating fluid, which is returning from the sides and inferior portions of tlie body. In the caterpillar, this portion of the dorsal vessel ex- tends from the twelfth to the anterior part of the fifth segment. It is fur- nished with eight double valves, which are formed, as Mr. Bowerbank has correctly described them in the Ephemera marginala — namely, the upper valve " by a reflecting inwards and upwards of the inner coat, or coats of the artery," (by which he means the dorsal vessel) "and the under one by a con- traction or projection of the like parts of a portion of the artery beneath, so as to come within the grasp of the lower part of the valve above it." The whole vessel is made up of three coats, the two innermost of which, the lining, or serous, and the muscular, or principal portion of the vessel, consti- tute the reflected portions, or valves; while the third, or outermost coat, which is exceedingly thin and delicate, is continued over the vessel nearly in a straight line, and does not appear at all to follow the reflexions of the other two. In the caterpillar, this portion of the vessel has eight pairs of small suspensory muscles, seen along the upper side of Fig. 339, which arise from the middle of the upper surface of each valve, and are continued back to be attached over the middle of the next valve: they seem to have considerable influence over the contractions of the valves. The Aortal, or anterior por- tion of the vessel, extends from the hinder part of the fourth segment to its termination and division into vessels, to be distributed to the head, which di- vision takes place after it has passed the oesophagus, and at a point immedi- ately beneath the supra-cesophageal ganglion, or brain of the insect. This portion of the vessel is much narrower than the dorsal, has no distinct valves or muscles; nor do any vessels enter it laterally; but it is very delicate and transparent, and gradually diminishes in size from its commencement to its anterior termination. Its course, in the caterpillar, is immediately beneath the integument, along the fourth and third segments, till it arrives at the hinder parts of the second segment; when it gradually descends upon the oesophagus, and, immediately behind the cerebral ganglion, gives off a pair of exceedingly minute vessels. It then passes beneath the ganglion, and, in the front part of the head, is divided into several branches, as noticed by Mr. Newport in the anatomical description he has given of the herves of this spe- cies of Sphinx: (l^hil. Trans. 1832, p. 385.) These branches are best ob- served in the chrysaUs (Fig. 339:) in all the stages they maybe divided into three sets; the first is given off immediately after the vessel has passed be- neath tlic ganglion; and consists of two lateral tnanks, the united capacity of CIRCULATION IN INSECTS. 179 class. It explains why insects, after they have attained their imago state, and the circulation, is nearly ohliterated, no longer increase in size, and require but little nourishment for the maintenance of life. This, however, Is a state not calculated for so long a duration as that in which the deve- lopment is advancing; and, accordingly, the period during which the insect remains in the imago condition is generally short, compared to that of the larva, where a large supply of nutriment, and a rapid circulation of the fluids, concur in maintaining the vital functions in full activity. Thus, the Ephemera, which lives for two or three years in the larva state, generally perishes in the course of a {gw hours after it has acquired wings, and reached its perfect state of ma- turity. which is equal to about one-third of that of the aorta; they descend, one on each side of the mouth, and are each divided into three branches. The se- cond set consists of two pairs of branches, one going- apparently to the tongue, the other to the antenna:. The third set is formed by two branches, which pass upwards, and are the continuations of the aorta; tliey divide into branches, and are lost in tlie integuments, and structures of the anterior part of the head. The pulsatory action of the dorsal vessel is continued along its whole course, and seems to terminate at the division of the vessel into branches. During the metamorphoses of the insect, this vessel becomes considerably shortened, but is stronger and more consolidated in its structure. Its course is likewise al- tered; from having, in the caterpillar, (Fig. 339,) passed along, nearly in a straight line, it begins in the chrysalis, (Fig. 340,) to descend in the fifth segment, and to pass under what is to become the division between the tiio- rax and abdomen in the perfect insect. It then ascends in the fourth seg- ment, and descends again in the second, so that when the insect has attained its perfect form, (Fig. 341,) its course is very tortuous. The vessels which enter it are situated in the abdomen, and pass in laterally among the muscles, chiefly at the anterior part of each segment or valve. Fig. 342 is a superior, or dorsal view of the same vessel, in the perfect state of tlie insect, which shows, still more distinctly, the vessels entering it laterally, intermixed witli the lateral muscles. Fig. 343 is a magnified lateral view of the anterior ex- tremity of the dorsal vessel, corresponding to Fig. 341; and Fig. 344, a simi- larly magnified view of the same portion of the vessel seen from above, cor- responding to Fig. 342. Fig. 345 shows the mode in which the valves ai'C formed by a duplicatare of the inner membrane in the perfect insect. ISO THE VITAL FUNCTIONS. In proportion as the cliangcs of form which the invSect un- dergoes arc less consitlcr;ihle, llie evidences of a circulation become more distinct. Such is the case in many of the Ap- terous Insects, comj)osing the family of Myriapoda: in the Scolopendra inorsitnns, (l^inn.,) for instance, Duges ob- served the dorsal vessel dividing into three large branches. INIost of tlic tribes belonging to the class of Arachnida liave, likewise, a dorsal vessel, very analogous in its struc- ture and situation to that of insects; and, as none of them undergo any metamorphosis, their vascular system admits of being considerably developed, and becomes a permanent part of the organization. Fig. 346 shows the dorsal vessel of the tJSrcniea domestica, or house spi- der, with some of the arterial trunks arising from it, lying embedded in a thick mass of substance, having a similar oily character to that which is contained, in large quantities, in the principal cavities of insects. It is, in general, difBcult to obtain a view of the circulation in the living spider, on account of the thick co- vering of hair which is spread over the body and the limbs; but if a species, which has no hair, be selected for examination, we can see very distinctly, through the microscope, the motion of the blood in the vessels, by means of the globules it contains, both in the legs and in other parts, where it presents appearances very similar to those already described in the limbs of the larva? of insects. A complete vascular circulation is established in all the animals which compose the class of Annelida; the vessels being continuous throughout, and having sufficient power to propel the blood through the whole of its circuit. Great va- rietv exists in the arrangement and distribution of these vessels, depending on the form of the animal, the compli- cation of its functions, and the extent of its powers. The first rudiment of a distinct system of circulating vessels, in- dependent of the ramified tubes proceeding from the inte^ CIRCULATION IN THE ANNELIDA. 181 tinal canal, occurs in the Plnncmre^ which are a trihc of flat vermiform animals^ in many respects allied to the more developed Entozoa, and ap])earing placed as an intermediate 3-16* link between them and the Annelida. In many species such as the Vhinaria nigra^ fitsca, and tremellaris, {Muller,) Duges ob- served two longitudinal trunks (Fig. 346*) running along tlie sides of the under surface of the animal, and joining together, both at their fore and hind extremities, so as to form a continuous channel of an oval form.f A great number of smaller vessels branch off from these main trunks in every direction, and ramify extensivel}^, often uniting with those from the opposite side, and establishing the freest communications between them. In the Annelida which have a more lengthened and cy- lindric form, the principal vessels have a longitudinal course, but are differently disposed in different species. There is, in all, a vascular trunk, extending along a middle line, the whole length of the back, and especially designated as the dorsal vessel: in general, there is also a corresponding trunk, occupying the middle line of the lower, or abdominal side of the body, and termed the abdominal vessel. This latter vessel is sometimes double; one being superficial, and ano- ther lying deeper; the principal nervous cord, and chain of ganglia being situated between them. Frequently, there are found, in addition to these, vessels which run alons: the sides of the body, and are therefore called the lateral ves- sels. In every case there are, as we have seen in the Plana- ria, numerous branches, and collateral communications be- tween the lateral, the abdominal, and dorsal vessels; more especially at the two extremities of the bod}-, where the great mass of blood, which has been flowing in one direction in one set of vessels, is transferred into others, which convey f De Blainvillc has described a structure similar to this In a Planaria from Brazil. Diet, dcs Sc. Nat t. xli. 216. 182 THE VITAL FUNCTIONS. it in the contrary direction, and complete the circuit of its course. The ramifications and kitcral connexions of the minuter branches are often so numerous as to compose a vascular net-work, covering a considerable extent of surface. This general description of the circulatory system is appli- cable to the tribes of Annelida possessing the simplest struc- ture, such as the Nais, the Nereis, and the Leech; genera which include a great variety of species of different shapes and sizes. Although the vessels themselves may be plainly discerned, it is not so easy to determine the real course which the blood takes while circulating within them; and we accordingly find very great discordance in the reports of different phy- siologists on this subject. De Blainville asserts that in all the Annelida, the blood in the dorsal vessel is carried back- wards, that is, from the head to the tail; a motion, which, of course, implies its return in the contrary direction, either in the lateral or the abdominal vessels. In the Nais, the Nereis, and the Leech, these last vessels are two in number, situated at the sides of the abdominal surface of the body. Carus adds his testimony in favour of this mode of considering the circulation in the Annelida. On the other hand, Spix, Bon- net, Sir Everard Home, and Duges, describe the course of the blood as quite the opposite of this, and maintain that it moves backwards, or towards the tail, in the abdominal ves- sels; and forwards, or towards the head, in the dorsal vessel. Morren, who is the latest authority on this subject, gives his testimony in favour of the latter view of the subject, as far as relates to the dorsal vessel of the Erpohdella vulga- ris* an animal allied to the Leech, and already noticed in the account of the mechanical functions of this tribe :t but he considers the abdominal vessel as performing also the same function of carrying the blood forwards towards the head, and the two lateral vessels as conveying it backwards, thus completing the circuit. This is illustrated by the diagram • Ilimdo vulgaris, (Linn.) Nephelis vulgaris. (Savigny.) I Vol. i. p. 195, where a delineation of this animal was given, Fig. 130. CIRCULATION IN THE ANNELIDA. 1S3 (Fig. 347;) where a is the anterior, and p the posterior ex- tremity of the animal, the dorsal vessel occupying the mid- dle straight line between the two lateral vessels, and the di- rection of the stream in each being indicated by the adjacent arrows. The blood in the abdominal vessel following the same course as that in the dorsal vessel, the same diagram represents also these vessels seen from below. Fig. 348 is an inferior view of the Erpobdella, showing the numerous ramifications of the abdominal vessel; the lesser branches encircling the nervous ganglia, and accompanying the prin- cipal nervous filaments which proceed from them: while the lateral vessels are seen pursuing a sliglitly serpentine course.* The tribe of Lumbrici, which includes the earth-worm, ♦ Duges represents the blood of this animal as moving- in different direc- tions in the rig-ht and in tlie left lateral vessels; g-enerally backwards in the former, and forwards in the latter: at the same time that it moves backwards in the dorsal, and forwards in the abdominal vessel. In the communicating branches which pass transversely from one lateral vessel to the other, the blood flows from left to right in those situated in the anterior half of the body, and from right to left in those of the posterior half: so that the plane in which its circuit is performed is horizontal, instead of vertical. It is curi- ous to find an example of a simiUu* transverse circulation, in the vegetable kingdom; this has recently been observed by Mr. Solly and Mr. Vai-ley, in a sprout of the Chara vulgaris, near the end of which the enclosed fluid re- volves continually on its own axis, instead of following the ordinary course of ascent and descent along the sides of tlic cylintkic cavity.— Sec Trans, of the Society of Arts, xlix. 180. 1S4 THE VITAL FUNCTIONS. is distinguished from the annclida ah'cady noticed, by being more highly organized, and possessing a more extensive cir- culation, and a more comj)licated apparatus for the perform- ance of this function. Tlic greater extent of vascular rami- fications appears to require increased powers for carrying the blood through the numerous and intricate passages it has to traverse; and these are obtained by means of muscular recc})taclcs, capable, by their successive contraction, of add- ing to the impulsive force witli which the blood is driven into the trunks that distribute it so extensively. These mus- cular appendages are globular or oval dilatations of some of the large vascular trunks, which bend round the sides of the anterior part of the body, and establish a free communication between the dorsal and the abdominal vessels. They are described by Duges as consisting, in the Linyibricus gigas, of seven vessels on each side, forming a series of rounded dilatations, about twelve in number, resembling a string of beads.* In the Lnmhricus ierresiris, or common earth-worm, there are only five pairs of these vessels; they have been de- scribed and figured by Sir E. Home if but the most full and accurate account of their structure has been given by Mor- ren, in his splendid work on the anatomy of that animal. J Fig. 349, which is reduced from his plates, represents these • They arc termed by Duges, Vaisseaux monlliformes, ou dorso-abdor^i- naux. — Annalcs dcs Sciences Natiirelles, xv. 299. f Pliilos. Transact, for 1817, p. 3: and Pi. iii. Fig. 4. t " De Lunibrici tcrrcstris Ilibtom naturalis, nccuon Anutomia Tractatus." Qto. Bruxelles, 1829. CIRCULATION IN THE CRUSTACEA. 185 Singular appendages to the vascular system of the cartli- worm, separated from their attachments, and viewed in con- nexion only with the dorsal and abdominal trunks in which they terminate. The abdominal vessel, (a, a,) on arriving near the oesophagus, is dilated, at the point r., into a globu- lar bulb (c,) which is followed, at equal intervals, by four others (c, c.) From each of these bulbs, or ventricles, as they are termed by Morren, a vessel (d) is sent off at right angles, on each side; this vessel also enlarges into several nearly globular dilatations (e,) followed by a still larger, and more elongated oval receptacle (f,) which completes the se- micircular sweep taken by the vessel in bending round the sides of the body, in order to join the dorsal vessel (g, g,) in which all the other four communicating vessels, presenting similar dilatations, terminate. Sir E. Home is of opinion that these dilated portions of the vessel are useful as reser- voirs of blood, for supplying it in greater quantity to the neighbouring organs, as occasion may require: but Morren ascribes to them the more important office of accelerating, by their muscular action, the current of circulating blood. If the latter of these views be correct, which the strong pul- sations, constantly visible in these bulbs, render extremely probable, this structure would offer the first rudiments of the organ which, in all the superior classes of animals, per- forms so important an office in the circulation of the blood, namely, the heart: and this name, indeed, is given by Cu- vier, Morren, and others, to these dilated portions of the vascular systems of the higher orders of Annelida."* Here, also, the statements of different anatomists are at variance, with regard to the direction taken by the blood while circulating in the vessels: Home and Duges represent it as proceeding forwards in the dorsal, and backwards in the abdominal vessels; a course which implies its descent • It is remai'kable tliat tlic blood in most of the Annelida has a bright scarlet colour, and resembles, in this respect, the blood of vcrtcbratcd ani- mals. Vol. H. 24 186 THE VITAL FUNCTIONS. alone; the lateral communicating vessels just clescril)ecl; while De jjlainvillc and Morrcn ascribe to it a course precisely the reverse. Amidst these conflicting testimonies, it is extreme- ly difficult to determine on which side the truth lies; and a suspicion will naturally arise, that tlie course of the blood in the vessels may not be at all times uniform, but may be lia- ble to ])artial oscillations, or be even completely reversed, by the operation of particular distiirbinii; causes. The larger Crustacea possess a circulatory apparatus still more extensive and complete, accompanied by a correspond- ing increase in the energy of the vital functions. As we follow this system in the more higlily organized tribes of this class, we find the powers of the dorsal vessel becoming more and more concentrated in its anterior extremity; till, in the Decapoda, a family which comj^rchends the Lobster and the Crab, we find this part dilated into an oval or globu- lar organ, with very muscular coats, capable of vigorous contractions, propelling its contents with considerable force into the vessels, and therefore clearly entitled to the appel- lation of heart. The distinction between arteries and veins, which can scarcely be made with any precision in the sys- tems of the inferior tribes, is here perfectly determined by the existence of this central organ of propulsion: for the ves- sels into which the blood is sent by its contractions, and which, ramifying extensively, distribute it to distant parts, are indisputably arteries; and, conversely, the vessels which collect the blood from all these parts, and bring it back to the heart, are as decidedly veins. The heart of the lobster is situated immediately under the carajDace, or shell of the dorsal region of the thorax, directly over the stomach; its pulsations are very distinct, and are performed with great regularity. The importance of the heart, as the prime agent in the circulation, increases as we advance to the higher classes of animals, whose more active and energetic functions require a continual and rapid renewal of nutrient fluid, and render necessary the introduction of farther refinements into its CIRCULATION IN THE VERTEBKATA. 1S7 structure. The supply of blood to the heart, beins; in a con- stant stream, produces a gradual dilatation of the cavity which receives it; and the muscular fibres of that cavity are not ex- cited to contraction, until tlicy are stretched to a certain point. But in order effectually to drive the blood into eve- ry part of the arterial system, where it has great resistances to overcome, a considerable impulsive force is required, im- plying a sudden as well as powerful muscular action. This object is attained, in all vertebrated animals, by providing a second muscular cavity, termed a ventricle, into which the first cavity, or auricle, throws the blood it has received from the veins, with a sudden impulse; and thus the ventricle, be- ing rapidly distended, is excited to a much more quick and forcible contraction than the auricle, and propels the blood it contains into the artery, with an impetus incomparably greater than could have resulted from the action of the au- ricle alone. Fig. 350 represents the heart with its two ca- vities; D being the auricle, and e the ventricle; together'witli the main trunks of the veins (c, c,) wliich convey the blood into the auricle; and those of the arteries (a,) which receive it from the ventricle for distribution over the whole system. The force of contraction in the principal cavity of i the heart being tlius increased, it becomes necessary to provide 188 THE VITAL FUNCTIONS. additional security against the retro2;rade motion of its fluid contents. Valves are accordingly interposed between the auricle and ventricle; and great refinement of mechanism is displayed in their construction. Fig. 351 represents their appearance (at v) when the cavities, both of the auricle (d,) and the ventricle (e) are laid open: c, c, as before, being the upper and lower venae cavre, and a, the main trunk of the aorta. These valves are composed of two loose membranes, the fixed edges of which are attached circularly to the aper- ture of communication between the cavities, and their loose edges project into the ventricle; so that they perform the office of flood-gates, allowing a free passage to the blood when it is impelled into the ventricle, and being pushed back the moment the ventricle contracts; in which latter case they concur in accurately closing the aperture, and preventing the return of a single drop into the auricle. These valves being attached to a wide circular aperture, it is necessary that they should be restrained from inverting themselves into the auricle, at each contraction of the ventricle. For this pur- pose thci-e are provided slender ligaments (which are seen in Fig. 351,) fixed by one end to the edge of the valve, and by the other to some part of the inner surface of the ventri- cle, so that the valve is ahvays kept within the cavity of the latter. In the auricle, the same purpose is answered by the oblique direction in which the veins enter it. CIRCULATION IN THE VERTEBRATA. 189 The arteries themselves, especially the main trunk of the aorta, as it issues from the heart, are muscular, and when suddenly distended, contract upon their contents. It was necessary, therefore, to provide means for preventing any reflux of blood into the ventricle during their contraction; and for this purpose a set of valves {v, Fig. 351,) is placed at the beginning of these tubes where they arise from the ventricle. These valve^consist usually of three membranes, which have the form of a crescent, and are capable of closing the passage so accurately, that not a drop of blood can pass 'between them.* In order to convey a more clear idea of the course of the blood in the circulatory sys- tem, I have drawn the diagram, Fig. 352, exhibiting the gene- ral arrangement of its compo- nent parts. The main arterial trunk, or Aorta (a,) while pro- ceeding in its course, gives off numerous branches, (b,) which divide and subdivide, till the ramifications (p) arrive at an extreme degree of minuteness; and they are finally distributed to every organ, and to the re- motest extremities of the body. They frequently, during their course, communicate with one another, or anastomose, as it is termed, by collateral branches, so as to provide against interruptions to the circu- lation, which might arise from accidental obstructions in any particular branches of this extended system of canals. The minutest vessels (p,) which, in incalculable numbers, pervade * In the arteiy of the shark, and other cartilag-lnous fishes, where the ac- tion of the vessel is very powerful, these valves are mucli more numerous, and arranged in rows, occupying' several parts of the artery. Additional valves are also met with in other fishes at tlie branching of largo arteries. 190 THE VITAL FUNCTIONS. every part of the frame, are named, from their behig finer than hciirs, cajnllary vessels. After the blood, thus transmitted to tlie dificrent parts of the body by the arteries, has supplied tlicm with the nourish- ment they require, it is conveyed back to the heart by the veins, which, commencing from the extreme ramifications of the arteries, bend back again in a course directed tow^ards the heart. Tlic smaller branches join in succession to form larger and larger trunks, till they are at length all united into one or two main pipes, called the Venx cavx, (c,) which pour their accumulated torrent of blood into the general re- servoir, the heart; entering first into the auricle (d,) and thence being carried forward into the ventricle (e,) which again propels it through the Aorta. The veins are larger and more numerous than the arteries, and may be compared to rivers, which, collecting all the water that is not imbibed by the soil, and reconveying it into its general receptacle, the ocean, perform an analogous office in the economy of the earth. The communications of the capillary arteries with the veins are beautifully seen, under the xnicroscope, in the trans- parent membranes of frogs or fishes. The splendid spectacle, thus brought within the cognizance of our senses,of unceasing activity in the minutest filaments of the animal frame, and of the rapid transit of streams of fluid, bearing along with them minute particles, which appear to be pressing forwards, like the passengers in the streets of a crowded city, through multitudes of narrow and winding passages, can never fail, when first beheld, to fill the mind with astonishment;'^ a feel- ing which must be exalted to the highest admiration, on re- flecting that what we there behold is at all times going on ♦ Lcwcnlioeck, speaking- of the dclig'ht he experienced on viewhig the circulation of the blood in tadpoles, uses the following expressions: " This pleasure has oftentimes been so recreating to me, that I do not believe that all the pleasure of fountains, or water-works, either natural or made by art, covdd have pk-ascd my sight so well, as the view of these creatures has given me."— rhil. Trans, xxii. 453. RESf»IRATOnY CIRCULATION. 191 within us, during the whole period of our lives, in every, even the minutest, portion oC our frame. How inadequate, then, must be any ideas we are capable of forming of the in- calculable number of movements and of actions, which are conducted in the living system; and liow infinite must be the prescience and the wisdom, by whicii these multifarious and complicated operations were so deeply planned, and so harmoniously adjusted! § 3. Respiratory Circulation. The object of the circulation is not merely to dislributc the blood through the general system of the body; it has, also, another and a very important ofTice to perform. The blood undergoes, in the course of its circulation, considera- ble changes, both in its colour and its chemical composition. The healthy l^lood transmitted by the arteries is of a bright scarlet hue; that brought back by the veins is of a dark pur- ple, from its containing an excess of carbon, and is conse- quently unfit to be again circulated. Whenever, from some derangement in the functions, this dark blood finds its way into the arteries, it acts as a poison on every organ which it reaches, and would soon, if it continued to circu- late, destroy life. Hence, it is ne- cessary that the blood f^hich returns by the veins should undergo purifi- cation, by exposure either to the air itself, or to a fluid containing air, for the purpose of restoring and pre- serving its salutary qualities. The heart and vascular system have, therefore, the additional task as- signed them of conveying the vi- tiated venous blood to certain or- gans, where it may have access to the air, and receive its 102 THE VITAL FUNCTIONS. vivifyino; influence; and to this oflicc a distinct set of arte- rics and veins is appropriated, constituting a distinct circu- lation. This I have endeavoured to illustrate by the dia- gram, Fig. 353, where d represents the auricle, and e the ventricle of the heart; and a and c, the main arterial and venous trunks; and where the two circulations are, for the sake of distinctness, supposed to be separated from one ano- ther, so that the two systems of vessels may occupy diffe- rent parts of the diagram. The vessels which pervade the body generally (li,) and are subservient to nutrition, belong to what is termed the greater^ or systemic circulation: those which circulate the blood through the respiratory organs, (r,) for the purpose of aeration, compose the system of the lesser, or respiratory circulation. Few subjects in Physiology present a field of greater in- terest than the comparison of the modes in which these two great functions are, in all the various classes of animals, ex- actly adjusted to each other. So intimately are the organs of circulation related to those which distribute the blood to the respiratory organs, that we never can form a clear idea of the first, without a close reference to the last of these sys- tems. While describing the several plans of circulation presented to us by the diflcrent classes, I shall be obliged to assume both the necessity of the function of respiration, and of a provision of certain organs for the reception of air, ei- ther in its gaipous form, or as it is contained in the water, where the blood may be subjected to its action. It is ne- cessary, also, to state that the organs for receiving atmosphe- ric air, in its gaseous state, are either lungs, or pulmonary cavities, while' those which are constructed for aquatic res- piration are termed gills, or hranchicn; the arteries and the veins which carry on this respiratory circulation, being termed pulfnoiiary, or branchial, according as they relate to the one or the other description of respiratory organs. In many animals it is only a part of the circulating blood which undergoes aeration; the pulmonary or branchial arte- ries and veins being merely branches of the general system RESPIRATORY CIRCULATION. 193 of blood vessels: so that in this case, which is that repre- sented in the preceding figure (353,) the lesser circulation is included as a part of the general circulation. But in all the higher classes the whole of the blood is, in some part of its circuit, subjected to the influence of the air; the pulmonary, being then distinct from the systemic circulation. In the An- nelida, for instance, the venas cav3e, which bring back the blood from the system, unite to form one or more vessels, which then assume the function of arteries, subdividing and ramifying upon the branchial organs; after this the blood is again collected by the branchial veins, which unite into one trunk to form the arteries of the systemic circulation. Most insects, especially when arrived at the advanced stages of their development, have too imperfect a circulation to ef- fect the thorough aeration of the blood: and indeed a greater part of that fluid is not contained within the vascular sys- tem, but permeates the cavities and cellular texture of the body. It will be seen, when I come to treat of respiration, that the same object is accomplished by means totally inde- pendent af the circulatory apparatus; namely, by a system of air-tubes, distributed over every part of the body. But an apparatus of this kind is not required in those Arachni- da, where the circulation is vigorous, and continues during the whole of life: here, then, we again meet with a pulmo- nary as well as a systemic circulation, in conjunction with internal cavities for the reception of air. In the Crustacea the circulation is conducted on the same general plan as in the Annelida; the blood from every part of the body being collected by the Venae Cava^, which arc exceedingly capacious, and extend, on eacli side, along the lower surface of the ab- domen. They send out branches, whicli distribute the blood to the gills; but these branches, at their ori- gin, suddenly dilate, so as to Vol. II. 25 194 THE VITAL FUNCTIONS. form large receptacles, which are called sinuses, where the blood is allowed to accumulate, and where, by the muscularity of the expanded coats of the vessels, it receives an additional force of propulsion. From the branchias the blood is re- turned by another set of veins to the elongated heart for- merly described, and propelled by that organ into the sys- temic arteries. Fig. 354 shows the relative situation of these vessels, when isolated and viewed from behind in the Muja squinado. c, c, are the venae cavse; e, e, the ve- nous sinuses above-mentioned; f, f, are the branchial ar- teries; G, the gills, or branchiae; and i, i, the branchial veins terminating in the heart l.* In the Mollusca, the heart acquires greater size, compared with the other organs, and exerts a proportionally greater influence as the prime mover in the circulation. In the de- velopment of its structure, in the different orders of this class, a beautiful gradation may be perceived: the Branchiopoda having two hearts, one placed upon each of the two lateral trunks of the branchial veins; the Gasteropoda having a single heart, furnished with an auricle; and the Jicephala being provided with a heart, which has a single ventricle, but two auricles, corresponding to the two trunks of the branchial veins.t The most remarkable variety of structure is that exhibited by the Cephalopoda. We have already seen, in the Crusta- cea, dilatations of the venae cavse, at the origin of the branch- ial arteries: but in the Nautilus the dilatations of the branch- ial veins are of such a size, as to be almost entitled to the appellation of auricles. The Sepia, in whose highly organ- ized system there is required great additional power to pro- pel the blood with sufficient force through the gills, is pro- vided with a large and complicated branchial apparatus; and • A minute account of the organs of circulation in the Crustacea is g-iven by Audouin and Milne Edwards, in the Annates des Sciences Naturelles, xi. 283 and 352, from which work the above figure is taken. f A great number of bivalve Mollusca exhibit the singular peculiarity of the lower portion of the intestinal tube traversing through the cavity of the heart. RESPIRATOnY CIRCULATION IN FISHES. 195 the requisite power is supplied by two additional hearts, situated on the venae cavae, of which they appear as if they were dilatations, immediately before the branchial arteries are sent ofT."^ They are shown at e, e, Fig. 355, which re- presents this part of the vascular system of the Loligo, de- tached from the surrounding parts; the course of the blood being indicated by arrows, c is one of the three trunks constituting the venae cavae, proceeding from above, dividing into two branches as it descends, and terminating, conjointly with the two venous trunks (d,) which are cominir from be- low, into the lateral or branchial hearts (e, e,) already men- tioned. Thence the blood is conveyed by the branchial arteries, (f, f,) on each side, to the gills (g,) and returned, by the branchial veins (i,) to the large central, or systemic heart (l,) which again distributes it, by means of the sys- temic arteries, to every part of the body. The cuttle-fish tribe is the only one thus furnished with three distinct hearts for carrying on a double circulation: none of these hearts are furnished with auricles. The remarkable distribution of the muscular powers which give an impulse to the circulating fluids, met with in * These veins arc surrounded by a great number of blintl pouches, wliich have the appearance of a fringe; the use of this singular structure is un- known. 196 THE VITAL FUNCTIOlSrS. the Sepia, constitutes a step in the transition from Mollusea to Fishes. In this latter class of animals, the two lateral hearts have united into a single central heart, while the aortic heart has entirely disappeared; and thus the position of the heart with respect to the two circulations is just the reverse of that which it has in the invertebrated classes. The plan in Fishes is shown in the diagram, Fig. 356, where the cen- tral organs are seen to consist of four cavities, c, d, e, f, opening successively the one into the other. The heart belongs exclusively to the gills; and there proceeds from it, not the aorta, but the trunk of those branchial arteries (f,) which convey the whole of the blood to the respiratory organs (g, h.) This blood, after being there aerated, is collected by the branchial veins (i,) which unite into a single trunk (a,) passing down the back, and performing, without any intermediate heart, the office of an aorta; that is, it divides into innumerable branches, and distributes the blood to every part of the system.* The blood is then reconveyed to the heart by the ordinary veins, which form a large vena cava (c.) This vein is generally considerably dilated at its termination, or just before it opens into the auricle, con- stituting what has been termed a venous sinus (s.) This, then, is followed by the auricle (d) and the ventricle (e;) but, besides these cavities, there is also a fourth (f,) formed by a dilatation of the beginning of the branchial artery, and termed the biilbus arterlosnSy contributing, doubtless, to • The caudal brancli of the aorta is protected by the roots of the inferior spinous processes, joining- to form arches through which it passes; and fre- quently the artery is contained in a bony channel, formed by the bodies of the vertebr?e, which effectually secures it from all external pressure- In the sturgeon even the abdominal aorta is thus protected, being- entirely concealed within this bony canal. 357 H M, '/,''' RESPIRATORY CIRCULATION IN REPTILES. 197 augment the impetus with which the blood is sent into the branchial arteries. The circulation in Reptiles is not double, like that of fishes; for only a part of the blood is brought under the in- fluence of the air in the pulmonary organs. All the animals belonging to this class are cold-blooded, sluggish, and inert; they subsist upon a scanty allowance of food, and arc as- tonishingly tenacious of life. The simplest form in which we meet with this mode of circuhition is in the Batrachia; it is .shown in the diagram, Fig. 357. The heart of the Frog, for example, may be considered as consisting of a single ventricle (e,) and a single auricle (d.*) From the former there proceeds one f|\N S^^^^ arterial trunk, which is proper- %\ ly the aorta. This aorta soon divides \iij into two trunks, which, after sending branches to the head and neck, bend downwards (as it is seen at o, p,) and unite to form a single trunk (a,) which is the descending aorta. From this vessel proceed all the arteries which are distributed to the trunk and to the limbs, and which are represented as situated at b: these arterial ramifica- tions are continued into the great ve- nous trunks, which, as usual, constitute the vena? cavce (c,) and terminate in the auricle (d.) From each of the trunks which arise from the primary division of the aorta, there proceed the small arteries (f,) which are distributed to the lungs (h,) and convey to those organs a part only of the mass of circulating blood. To * Dr. Davy has observed that although the auricle appeare sing-le, wlien viewed externally, its cavity is in reality divided into two compartments by a transparent membranous partition in which some muscular fibres are apparent: these communicate with the cavity of the ventricle by a common opening provided with three semilunai- valves. Edin. Phil. Joumal; xix. 161. 198 THE VITAL FUNCTIONS. these pulmonary arteries there correspond a set of veins, uniting in the trunks (i,) which bring back the aerated blood to the auricle of the heart (d,) where it is mixed with the blood which has returned by the venae cavae (c,) from the general circulation. Thus the blood is only partially aerated, in consequence of the lesser circulation being here only a branch of the greater. Nothing is more curious or beautiful than the mode in which, during the transformations of this animal, Nature conducts tbe gradual transition of the branchial circulation of the tadpole, into the pulmonary circulation of the frog. In the former, the respiratory organs are constructed on the model of those of fishes, and respiration is performed in the same manner as in that class of animals: the heart is conse- quently essentially branchial, sending the whole of its blood to the gills, the veins returning from which (describing the course marked by the dotted lines m, n, in the diagram,) unite, as in fishes, to form the descending aorta. As the lungs develope, small arterial branches, arising from the aor- ta, are distributed to those organs, and in proportion as these arteries enlarge, the branchial arteries diminish; until, on their becoming entirely obliterated, the course of the blood is wholly diverted from them, and flows through the en- larged lateral trunks (o, p,) of which the junction constitutes the descending aorta. This latter vessel now receives the whole of its blood directly from the heart; which, from be- ing originally a branchial, has become a systemic heart. The heart of the Chelonian reptiles, such as the ordinary species of Tortoises and Turtles, has two distinct auricles; the one, receiving the blood from the pulmonary veins; the other, from those of the body generally; so that the mixture of aerated and vitiated blood takes place, not in the auricle, but in the ventricle itself. When all the cavities are dis- tended with blood, the two auricles being nearly of the same size as the ventricle, the whole has the appearance of a union of three hearts. The circulatory system of the Ophidia is constructed on a plan very similar to that of the Chelonia. WARM-BLOODED CIRCULATION. 199 In the Saurian reptiles, the structure becomes again more complicated. In the Chameleon each auricle of the heart has a large venous sinus, appearing like two supplementary auricles.'* The heart of the Crocodile has not only two au- ricles, but its ventricle is divided by two partitions, into three chambers: each of the partitions is perforated to allow of a free communication between the chambers; and the pas- sages are so adjusted as to determine the current of aerated blood, returning from the lungs, into those arteries, more especially, which supply the head and the muscles of the limbs; while the vitiated blood is made again to circulate through the arteries of the viscera.t It is in warm-blooded animals that the two offices of the circulation are most efficiently performed; for the whole of the blood passes, alternately, through the greater and the lesser circulations, and a complete apparatus is provided for each. There are, in fact, two hearts, the one on the left side impelling the blood through the greater or systemic circu- lation ; the other, on the right side, appropriated to the lesser, or pulmonary circulation. The annexed diagram, (Fig. 35,9,) 359 .,-.^<:^ H illustrates the plan of the cir- ^^ culation in warm-blooded ani- mals. From the left ventri- cle (l) the blood is propelled into the aorta (a,) to be dif- fused through the arteries of the system (b) to every part, and penetrating into all the capillary vessels; thence it is returned by the veins, through the venae cavae (c,)to the right auricle (d,) which delivers it into the right ventricle (e.) This right ventricle impels the blood, thus received, through the pulmonary arteries * Houston; Trans. Roy. Irish Acad, xv. 189. f It would appear, from this arrangement of the vessels, that the brain, or 200 THE VITAL FUNCTIONS. (f,) into the lungs (at h,) where it is aerated, and whence it is reconveyed by the pulmonary veins (i,) into the left au- ricle (k,) which immediately pours it into the left ventri- cle (l,) the point from whence we had set out. Both the right and the left heart have their respective au- ricles and ventricles; but they are all united in one envelope, so as to compose, in appearance, but a single organ:* still, however, tlic riglit and left cavities are kept perfectly dis- tinct from one another, and are separated by thick partitions, allowing of no direct transmission of fluid from the one side to the other. These two hearts may, therefore, be com- pared to two sets of chambers under the same roof, having, each, their respective entrances and exits, with a party-wall of separation between them. This junction of the two hearts central organ of the nervous system, requires, more than any other part, a supply of oxyg-enated blood for the due performance of its functions. The cui-ious provision which is made for sending this partial supply of blood, of a particular quality, in the larger kinds of reptiles, such as the Crocodile, has been pointed out by many anatomists; but has been lately investigated more particularly by M. Martin St. Ange. (See the Report of G. St. Hilaire, Revue Medicale, for April, 1833.) It is found that in Uicse animals, as well as in the Chelonia, a partial respiratory system is provided for by the admission, through two canals opening externally, of aerated water into the cavity of the abdomen, where it may act upon the blood which is circulating in the vessels. Traces of canals, of this description, are also met with in some of the higher classes of vertebratcd animals, as, for instance, among the MammaUa, in the Monotremaia and the Marsupialia. * A remai'kable exception to this general law of consolidation occurs in the heart of the Bugong, repre- sented in Fig. 360, in which it may be seen that the two ventricles, e and L, are almost entirely detached from each other. In tliis figure, which is taken from the Philosopliical Trans- actions for 1820, T) is the systemic auricle, k the riglit or pulmonary ventricle, f the pulmonary arteiy, k the left or pulmonary auricle, l the left or systemic ventricle, and a the aorta. DISTRIBUTION OF BLOOD VESSELS. 201 is conducive to their mutual strength: for the fibres of each intermix and even co-operate in their actions, and both cir- culations are carried on at the same time; that is, both ven- tricles contract or close at the same instant; and the same applies to the auricles. The blood which has just returned from the body, and that from the lungs, the former by the venae cavic, the latter by the pulmonary veins, fdl their re- spective auricles at the same instant; and both auricles, con- tracting at the same moment, discharge their contents simul- taneously into their respective ventricles. In the like manner, at the moment when the left ventricle is propelling its aerated blood into the aorta, for the purposes of general nutrition, the right ventricle is likewise driving the vitiated blood into the pulmonary artery, in order that it may be pu- rified by the influence of the air. Thus, the same blood which, during the interval of one pulsation, was circulating through the lungs, is, in the next, circulating through tiie body; and thus do the contractions of the veins, auricles, ventricles, and arteries, all concur in the same general end, and establish the most beautiful and perfect harmony of ac- tion.* § 4. Distribution of Blood Vessels, In the distribution of the arteries in the animal system, we meet with numberless proofs of wise and j)rovident ar- rangement. The great trunks of both arteries and veins, which carry on the circulation in the limbs, are conducted always on the interior sides, and along the interior angles of the joints, and generally seek the protection of the adja- * Evidence is afforded of the human conformation being expressly adapted to tlie erect position of the bod}' by the position of the heart, as compared with quadrupeds; for, in the latter, the heart is placed directly in the middle of the chest, with the point towards the abdomen, and not occupying- any portion of the diaphragm; but, in man, the heart lies obliquely on the dia- phragm, with the apex turned towards the left side. Vol. II. 2r> 202 THE VITAL FUNCTIONS. cent bones. Grooves are formed in many of the bones, where arteries are lodged, with the evident intention of af- fording them a more secure passage. Thus, the principal arteries which supjily the muscles of the chest, proceed alonn- the lower ed";es of the ribs, in deep furrows formed for their protection. Arteries are often still more effectu- ally guarded against injury or obstruction by passing through complete tubes of solid bone. An instance occurs in the ar- teries supplying the teeth, which pass along a channel in the lower jaw, excavated througli tlie whole length of the bone. The aorta in fishes, after having supplied arteries to the vis- cera of the abdomen, is continued to the tail, and passes through a channel, formed by bony processes from the ver- tebra; and the same kind of protection is aflbrded to the corresponding artery in the Cetacea. In the fore leg of the Lion, which is employed in actions of prodigious strength, the artery, without some especial provision, would have been in danger of being compressed by the violent contrac- tions of the muscles: in order, therefore, to guard against this inconvenience, it is made to pass through a perforation in the bone itself, where it is completely secure from pres- sure.* The energy of every function is regulated in a great mea- sure by the quantity of blood which the organs exercising that function receive. The muscles employed in the most vigorous actions are always found to receive the largest share of blood. It is commonly observed that the right fore leg of quadrupeds, as well as the right arm in man, is stronger than the left. Much of this superior strength is, no doubt, the result of education; the right arm being habi- tually more used than the left. But still the different mode in which the arteries are distributed to the two arms consti- tutes a natural source of inequality. The artery supplying the risht arm willi blood is the first which arises from the • In like manner the coffin bone of the Horse is perforated for tlie safe conveyance of tlie arteries going to the foot. DISTRIBUTION OP BLOOD VESSELS. 203 aorta, and it proceeds in a more direct course from the heart than the artery of the left arm, whicli has its ori^ijin in com- mon with the artery of that side of the head. Hence it has heen inferred that the right arm is originally better supplied with nourishment than the left. It may be alleged, in con- firmation of this view, that in birds, where any inerpiality in the actions of the two wings would have disturbed the regularity of flight, the aorta, when it lias arrived at the centre of the chest, divides with perfect equality into two branches, so that both wings receive precisely the same quantity of blood; and the muscles, being thus equally nou- rished, preserve that equality of strength, which their func- tion rigidly demands. When a large quantity of blo^od is wanted in any particu- lar organ, and yet the force with which it would arrive, if sent immediately by' large arteries, might injure the texture of that organ, contrivances are adopted for diminishing its impetus, either by making the arteries pursue very winding and circuitous paths, or by subdividing them, before they reach their destination, into a great number of smaller arte- ries. The delicate texture of the brain, for instance, would be greatly injured by the blood being impelled with any considerable force against the sides of the vessels which are distributed to it; and yet a very large supply of blood is re- quired by that organ for the due performance of its func- tions. Accordingly we find that all the arteries which go to the brain are very tortuous in thcii- course; every flexure tending considerably to diminish the force of the current of blood. In animals that graze, and keep their heads for a long time in a dependent position, the danger from an excessive impetus in the blood flowing towards the head is much greater than in other animals; and we find that an extraor- dinary provision is made to obviate this danger. The arte- ries which supply the brain, on their entrance into the basis of the skull, suddenly divide into a great number of minute branches, forming a complicated net- work of vessels, an ar- 204 THE VITAL FUNCTIONS. rangemcnt wliich, on the well known principles of hydrau- lics, must greatly check the velocity of the hlood conducted through them. That such is the real purpose of this struc- ture is evident from the hranchcs afterwards uniting into larger trunks when they have entered the brain, through tiic substance of which they are then distributed exactly as in other animals, where no sucli previous subdivision takes place. In the Bradijpxis tridacii/lus^ or great American Sloth, an animal remarkable for the slowness of its movements, a plan somewhat analogous to the former is adopted in the structure of the arteries of the limbs. These arteries, at their entrance into both the upper and lower extremities, suddenly divide into a great number of cylindric vessels of equal size, communicating in various places by collateral branches. These curiously subdivided arteries are exclu- sively distributed to the muscles of the limbs; for all the other arteries of the body branch off in the usual manner. This structure, which was discovered by Sir A. Carlisle,* is not confined to the Sloth, but is met with in other animals, as the Lcinxir tardigradus, and the Lemur lo?'is, which re- semble the sloth in the extreme sluggishness of their move- ments. It is extremely probable, therefore, that this pecu- liarity in the muscular power results from, or is at least in some way connected with this remarkable structure in the arteries. In the Lion, and some other beasts of prey, a simi- lar construction is adopted in the arteries of the head, pro- bably with a view to confer a power of more permanent contraction in the muscles of the jaws for holding a strong animal, such as a buffalo, and carrying it to a distance. That we may form an adequate conception of the im- mense power of the ventricle, or prime mover in the circu- lation of the blood, we have but to reflect on the numerous obstacles impeding its passage through the arterial system. There is, first, the natural elasticity of the coats of the ar- • Pliil. Tr.ins. for 1800, p. 98, and for 1804, p. 17. FORCE OP THE HEART. 205 teries, which must be overcome before any blood can enter them. Secondly, the arteries are, in most places, so con- nected with many heavy parts of the body, that their dila- tation cannot be effected without, at the same time, commu- nicating motion to them. Thus, when we sit cross-legi^cd, the pulsation of the artery in the ham, which is pressed upon the knee of the other leg, is sufficiently strong to raise tlie whole leg and foot at each beat of the pulse. If we con- sider the great weight of the leg, and reflect upon the length of the lever by which that weight acts, we shall be convinced of the prodigious force which is actually exerted by the cur- rent of blood in the artery in thus raising the whole limb. Thirdly, the winding course, which the blood is forced to take, in following all the oblique and serpentine flexures of the arteries, must greatly impede its motion. But not- withstanding these numerous and powerful impediments, the force of the heart is so great, that, in defiance of all obstacles or causes of retardation, it drives the blood with immense velocity into the aorta. The ventricle of the human heart does not contain more than an ounce of blood, and it con- tracts at least seventy times in a minute; so that above three hundred pounds of blood are passing through this organ during every hour that we live. " Consider," says Paley, " what an affair this is when we come to very large animals. The aorta of a whale is larger in the bore than the main pipe of the water-works at London Bridge; and the water roaring in its passage through that pipe is inferior in impetus and ve- locity to the blood gushing through the whale's heart. An anatomist who understood the structure of the heart, might say before hand that it would play; but he would expect, from the complexity of its mechanism, and the delicacy of many of its parts, that it should always be liable to derange- ment, or that it would soon work itself out. Yet shall this wonderful machine go on night and day, for eighty years together, at the rate of a hundred thousand strokes every twenty-four hours, having at every stroke a great resistance to overcome, and shall continue this action for this length 206 THE VITAL FUNCTIONS. of time, witliout disorder and without weariness. To those who venture tlieir lives in a ship, it has often been said that there is only a plank between them and destruction; but in the body, and especially in the arterial system, there is in many parts only a membrane, a skin, a thread." Yet how well has every part been guarded from injury: how provi- dentiall}' have accidents been anticipated: how skilfully has danger been averted! The impulse which the heart, by its powerful contraction, gives to the blood, is nearly expended by the time it has reached the veins: nature has accordingly furnished them with numerous valves, all opening, of course, in a direction towards the heart; so that as long as the blood proceeds in its natural course, it meets with no impediment; while a retrograde motion is effectually prevented. Hence external pressure, occasionally ap- plied to the veins, assists in promoting the flow of blood towards the heart; and hence the effects of exercise in accelerating the cir- culation. Valves are more especially pro- vided in the veins which pass over the mus- cles of the extremities, or which run imme- diately beneath the skin; while they are not found in the more internal veins beloncrino; to the viscera, which are less exposed to une- qual pressure. These valves are delineated in Fig. 365, which represents the interior of one of the larger veins. The situation and structure of the valves belonjiino; to the hydraulic apparatus of the circulation furnish as une- quivocal proofs of design as any that can be adduced. It was the observation of these valves that first suc-o-ested to the mind of Harvey the train of reflections which led him to the discovery of the real course of the blood in the veins, the arteries and the heart. This great discovery was one of the earliest fruits of the active and rational spirit of in- quiry, which, at the era of Bacon^s writings, was beginning to awaken the human mind from its long night of slumber, VALVES OF THE VEINS. 20' and to dissipate the darkness which had, for so many ages, overshadowed the regions of philosophy and science. Wo cannot but feel a pride, as Englishmen, in the recollection, that a discovery of such vast importance as that of the cir- culation of the blood, which has led to all the modern im- provements in the medical art, was made by our own coun- tryman, whose name will for ever live in the annals of our race as one of its most distinguished benefactors. Tlic con- sideration, also, that it had its source in the study of com- parative anatomy and physiology, affords us a convincing proof of the great advantage that may result from the culti- vation of those sciences; to which Nature, indeed, seems, in this instance, expressly to have invited us, by displaying to our view, in the organs of the circulation, an endless di- versity of combinations, as if she had ])urposcly designed to \ elucidate their relations with the vital powers, and to assist our investigations of the laws of organized beings. ( 208 ) CHAPTER XL Respiration. * § 1. Respiration in General. The action of atmospheric air is equally necessary for the maintenance of animal, as of vegetable life; and as the ascending sap of the one cannot be perfected unless exposed to the chemical agency of air in the leaves, in like manner the blood of animals requires the perpetual renovation of its vital properties by the purifying influence of respiration. The great importance of this function is evinced by the con- stant provision which has been made by Nature, in every class of animals, for bringing each portion of their nutritive juices, in its turn, into contact with air. Even the circula- tion of these juices is an object of inferior importance, com- pared with their aeration; for we fmd that insects, which have but an imperfect and partial circulation of their blood, still require the free introduction of air into every part of their system. The necessity for air is more urgent than the demand for food; many animals being capable of sub- sisting for a considerable time without nourishment, but all speedily perishing when deprived of air. The influence of this clement is requisite as well for the production and de- velopment, as for the continuance of organized beings in a living state. No vegetable seed will germinate, nor will any egg, even of the smallest insect, give birth to a larva, if kept in a perfect vacuum. Experiments on this subject have been varied and multiplied without end by Spallanza- ni, who found that insects under an air pump, conflned in rarefied air, in general lived for shorter periods in propor- RESPIRATION. 209 tioii to the degree to which the exhaustion of air had been carried. Those species of infusoria, wliich are most tena- cious of life, lived in very rarefied air for above a month: others perished in fourteen, eleven, or eight days; and some in two days only. In this imperfect vacuum, tiicy were seen still to continue their accustomed evolutions, wheeling in circles, darting to the suriace, or diving to the bottom of the fluid, and producing vortices by the rapid vibration of their cilia, to catch the floating particles which serve as their food: in course of time, however, they invariably gave in- dications of uneasiness; their movements became languid, a general relaxation ensued, and they at lengtii expired. But when the vacuum was rendered perfect, none of the infu- sions of animal or vegetable substances, whicli, under ordi- nary circumstances, soon swarm with millions of these mi- croscopic beings, ever exhibited a single animalcule; although these soon made their appearance in great numbers, if the smallest quantity of air was admitted into the receiver. Animals which inhabit the waters, and remain constantly under its surface, such as fishes, and the greater number of mollusca, are necessarily precluded from receiving the di- rect action of atmospheric air in its gaseous state. But as all water exposed to the air soon absorbs it in large quanti- ties, it becomes the medium by which that agent is applied to the respiratory organs of aquatic animals; and the oxy- gen it contains may thus act upon the blood with considera- ble efi'ect; though not, perhaps, to the same extent as when directly applied in a gastious state. The air which is pre- sent in water is, accordingly, as necessary to these animals as the air of the atmosphere is to those which live on land: hence, in our inquiries into the respiration of aquatic ani- mals, it will be sufficient to trace the means by which the surrounding water is allowed to have access to the organs appropriated to this function; and in speaking of the action of the water upon them, it will always be understood that I refer to the action of the atmospheric air which that water contains. Vol. II. 27 210 THE VITAL FUNCTIONS. ResjDiration, in its dlfiferent modes, may be distinguished, according to the nature of the medium which is breathed, into aqualic or atmospheric; and in the former case, it is either cutaneous, or bi^anchial, according as the respiratory- organs are external or internal. Atmospheric respiration, again, is cither tracheal, or "puhnonary , according as the air is received by a system of air tubes, or trachex, or into pulmonary cavities, or lungs. § 2. •Aquatic Respiration, Zoophytes appear in general to be unprovided with any distinct channels for conveying aerated w^ater into the inte- rior of the bodies, so that it may act in succession on the nu- tritive juices, and after performing this office, may be ex- pelled, and exchanged for a fresh supply. It has according- ly been conjectured, on the presumption that this function is equally necessary to them as it is to all other animals, that the vivifying influence of the surrounding element is ex- erted through the medium of the surface of the body. Thus, it is very possible that in Polypi, while the interior surface of the sac digests the food, its external surface may perform the office of respiration : and no other mode of accomplishing this function has been distinctly traced in the Acalepha. Me- dusae, indeed, appear to have a farther object than mere pro- gression in the alternate expansions and contractions of the floating edges of their hemispherical bodies; for these move- ments are performed with great regularity under all circum- stances of rest or motion; and they continue even when the animal is taken out of the water and laid on the ground, as long as it retains its vitality. The specific name of the Medu- sa pulnio'* (the Pulmone Marino of the Italians,) is derived from the supposed resemblance of these movements to those of the lungs of breathing animals. The large cavities ad- jacent to the stomach, and which have been already pointed out in Fig. 249 and 252,t have been conje.cture,d to be res- • See the delineation of this animal in Fig. 135, vol. i. p. 198. •j- Pages 67" and 68 of this volume. AQUATIC RESPIRATION. 211 piratory organs, chiefly, I believe, because they are not known to serve any other purpose. The Enlozoa, m like manner, present no appearance of internal respiratory organs; so that they probal)ly receive the influence of oxygen only through the medium of tlie juices of the animals on which they subsist. Planar ix, which have a more independent existence, though endowed with a system of circulating vessels, have no internal respira- tory organs; and whatever respiration they perform must be wholly cutaneous. Such is also the condition of several of the simpler kinds of Annelida; but in those which are more highly organized, an apparatus is provided for respiration, which is wholly external to the body, and appears as an ap- pendage to it, consisting generally of tufts of projecting fibres, branching like a plume of feathers, and floating in the surrounding fluid. The Lumbricns marinus^ or lob-worm,* for example, has two rows of branchial organs of this de- scription, one on each side of the body; each row being com- posed of from fourteen to sixteen tufts. In the more sta- tionary Annelida, which inhabit calcareous tubes, as the Serpula and the Teredo^ these arborescent tufts are protected by a sheath which envelops their roots; and they are placed on the head, as being the only part w'hich comes in contact with the water. Most of the smaller Crustacea have branchiae in the form of feathery tufts, attached to the paddles near the tail, and kept in incessant vibratory motion, which gives an appear- ance of great liveliness to the animal, and is more especially striking in the microscopic species. The variety of shapes which these organs assume in different tribes is too great to allow of any specific description of them in this place: but amidst these varieties, it is sufficiently apparent that their construction has been, in all cases, designed to obtain a con- siderable extent of surface over which the minute subdivi- * Arenlcola piscatomm (Lam.) See a deHncation of this marine worm in Fig. 135, vol. i. p. 198. 212 THE VITAL FUNCTIONS. sions of the blood vessels might be spread, in order to ex- pose them fully to the action of aerated water. The Mollusca, also, present great diversity in the forms of their respiratory organs, although they are all, with but few exceptions, adapted to aquatic respiration. In many of the tribes which have no shell, as the Thelis, the Doris, and the Tritoiiia, there are arborescent gills projecting from different parts of the body, and floating in the water. In the Lepas, or barnacle, a curious family, constituting a connecting link between molluscous and articulated animals, these ofgans are attached to the bases of the cirrhi, or jointed tentacula, which are kept in constant motion, in order to obtain the full action of the water on the blood vessels they contain. We are next to consider the extensive series of aquatic animals in which respiration is carried on by organs situated in the interior of the body. The first example of internal aquatic respiration occurs in the Holothicria, where there is an organ composed of ramified tubes, situated in a cavity communicating with the intestine, and having an external opening for the admission of the aerated water, which is brought to act on a vascular net-work, containing the nutri- tive juices of the animal, and apparently performing a par- tial circulation of those juices. A still more complicated system of respiratory channels occurs, both in the Echinus and Ssterias^ where they open by separate, but very minute orifices, distinct from the larger apertures through which the feet protrude; and the water admitted through these tubes is allowed to permeate the general cavity of the body, and is thus brought into contact with all the orojans. The animals eomiposing the family of Jiscidix have a large respiratory cavity, receiving the water from without, and having its sides lined with a membrane, which is thrown into a great number of folds; thus considerably extending the surface on which the water is designed to act. The entrance into the oesophagus, or true mouth, is situated at the bottom of this cavity; that is, at the part most remote from the ex- ternal orifice; so that all the food has to pass through the AQUATIC RESPIRATION. 213 respiratory cavity, before it can be swallowed, and received into the stomach. In several of the Annelida, also, we find internal organs of respiration. The Lumhricus lerresiris, or common earth-worm, has a single row of apertures, al)out 120 in num- ber, placed along the back, and opening between the seg- ments of the body: they each lead into a respiratory vesicle, situated between the integument and the intestine.* The Leech has sixteen minute orifices of this kind on each side of the body, opening internally into the same number of oval cells, which are respiratory cavities; the water passing both in and out by the same orifices.t The *Bphrodita aculeata has thirty-two orifices on each side, placed in rows, opening into one large respiratory sac, which is situated immediately under the muscles of the back, but separated by a membrane from the abdominal cavity. Projecting into this sac, are seen several membranous vesi- cles, fifteen in number on each side, which have no external opening, but which receive, on the inner part, the ends of certain tubes, or caeca, sent off from the intestinal canal; so that the nutriment is aerated almost as soon as it is prepared by the digestive organs.^ In all the higher classes of aquatic animals, where the cir- culation is carried on by means of a muscular heart, and where the whole of the blood is subjected, during its circuit, to the action of the aerated water, the immediate organs of respiration consist of long, narrow filaments, in the form of a fringe, and the blood vessels belonging to the respiratory system are extensively distributed over the whole surface of ♦ A minute description of these organs is given by Morrcn, in pages S'2> and 148 of his work, ah-eady quoted. ■j- The blood, after being aci-atcd in these cells, is conveyed into the large lateral vessels, by means of canals, which pass transversely, forming loops, si- tuated between tlie c?eca of the stomach. These loops are studded with an immense number of small rounded bodies of a glandular appearance, resem- bling those which convey the vcnx cavae of the cephalopoda. % Home, Philos. Trans, for 1815, p. 259. 214 THE VITAL FUNCTIONS. these filaments. Organs of this description are denominated Branchice, or Gills; and the arteries which bring the blood to them are called the branchial arteries; the veins, which convey it back, being, of course, the branchial veins. The larger Crustacea have their branchiae situated on the under side of tiie body, not only in order to obtain protec- tion from the carapace, which is folded over them, but also for the sake of being attached to the haunches of the feet- jaws, and thoracic feet, and thus participating in the move- ments of those organs. They may be seen in the Lobster, or in the Crab, by raising the lower edge of the carapace. The form of each branchial lamina is shown at g, in Fig. 354:* they consist of assemblages of many thousands of mi- nute filaments, proceeding from their respective stems, like the fibres of a feather; and each group having a triangular, or a pyramidal figure. The number of these pyramidal bo- dies varies in the different genera; thus, the Lobster has twenty-two, disposed in rows on each side of the body; but in the Crab, there are only seven on each side. To these organs are attached short and flat paddles, which are moved by appropriate muscles, and arc kept in incessant motion, producing strong currents of water, evidently for the pur- pose of obtaining the full action of the element on every por- tion of the surface of the branchiae. In the greater number of INlollusca, these important or- gans, although external with respect to the viscera, are with- in the shell, and are generally situated near its outer margin. They are composed of parallel filaments, arranged like the teeth of a fine comb; and an opening exists in the mouth for admitting the water which is to act upon them.t In the * Page 193, of this volume. ■f- These filaments appear, in many instances, to have the power of pro- ducing* currents of water in their vicinity by the action of minute cilia, similar to those belong-ing" to the tentacula of many polypi, where the same pheno- menon is obseiTablc. Thus, if one of the branchial filaments of the fresh water muscle be cut across, the detached portion will be seen to advance in the fluid by a spontaneous motion, like tlic tentaculum of a polype, under AQUATIC RESPIRATION. 215 Gasteropoda, or inhabitants of univalve shells, this opening is usually wide. In the ^cephalu, or bivalve molhisca, the gills are spread out, in the form of laminae, round the mar- gin of the shell, as is exemplified in the oyster, where it is commonly known by the name of beard. The aerated wa- ter is admitted through a fissure in the mouth, and when it has performed its office in respiration, is usually expelled by a separate opening. The part of the mouth through which the water is admitted to the branchiae is sometimes prolonged, forming a tube, open at the extremity, and at all times al- lowing free ingress and egress to the water, even when the animal has withdrawn its body wholly within its shell. Sometimes one, and sometimes two tubes of this kind are met with; and they are often protected by a tubular portion of shell, as is seen in the Murex, Buccinuui, and Sirombus; in other instances, the situation of the tube is only marked by a deep notch in the edge of the shell. In those mollusca which burrow in the sand, this tube can be extended to a considerable length, so as to reach the water, which is alter- nately sucked in and ejected by the muscular action of the mouth. In those Acephala which are unprovided with any tube of this kind, the mechanism of respiration consists simply in the opening and shutting of the shell. By watch- ing them attentively, we may perceive that the surrounding water is moved in an eddy by these actions, and that the current is kept up without interruption. All the Sepia: have their gills enclosed in two lateral cavities, which communi- cate with a funnel-shaped opening in the middle of the neck, alternately receiving and expelling the water by the muscu- lar action of its sides. The forms assumed by the respira- tory organs in this class are almost infinitely diversified, while the general design of their arrangement is still the same. As we rise in the scale of animals, the respiratory func- the same circumstances. Similar currents of water, according- to the recent observations of Mr. Lister, and apparently determined by the same mechanism of vibratory cilia, take place in the branchial sac of Ascidix. 216 THE VITAL FUNCTIONS. tion assumes a higher importance. In fishes the gills form large organs, and the continuance of their action is more es- sential to life than it appears to be in any of the inferior classes: they are situated, as is well known, on each side of the throat in tlie immediate vicinity of the heart. Their usual form is shown at g g, Fig. 367, where they are repre- sented on one side only, but in their relative situations with respect to the auricle (d,) and ventricle (e,) of the heart; the bulbus arteriosus (b,) and the branchial artery (f.) They have the same fringed structure as in the mollusca, the fibres being set close to each other, like the barbs of a feather, or the teeth of a fine comb, and being attached on each side of the throat, in double rows, to the convex margins of four cartilaginous or osseous arches, which are themselves con- nected with the jaws by the bone called the os hyoides. The mode of their articulation is such as to allow each arch to have a small motion forwards, by which they are separated from one another; and by moving backwards they are again brought together, or collapsed. Each filament contains a slender plate of cartilage, giving it mechanical support, and enabling it to preserve its shape while moved by the streams of water which are perpetually rushing past. When their RESPIRATION IN FISHES. 217 surfaces are still more minutely examined, they are fountlto be covered with innumerable minute processes, crowded to- gether like the pile of velvet; and on these are distributed myriads of blood vessels, spread like a delicate net-work, over every part of the surface. The whole extent of this surface exposed to the action of the aerated water, by these thickly set filaments, must be exceedingly great.* A large flap termed the Operculum^ extends over the whole organ, defending it from injury, and leaving below a wide fissure for the escape of the water, whicli has per- formed its office in respiration. For this purpose the water is taken in by the mouth, and forced by the muscles of the throat through the apertures which lead to the branchial ca- vities: in this action the branchial arteries are brought for- wards and separated to a certain distance from each other; and the rush of w^ater through them unfolds and separates each of the thousand minute filaments of the branchiae, so that they all receive the full action of that fluid as it passes by them. Such appears to be the i)rincipal mechanical ob- ject of the act of respiration in this class of animals; and it is an oljject that requires the co-operation of a liquid such as water, capable of acting by its impulsive momentum in expanding every part of the apparatus on which the blood vessels are distributed. When a fish is taken out of the w^a- ter, this effect can no longer be produced; in vain the ani- mal reiterates its utmost eflforts to raise the branch ia2, and relieve the sense of suflbcation it experiences in consequence of the general collapse of the filaments of those organs, wliich adhere together in a mass, and can no longer receive the vi- vifying influence of oxygen. t Death is, in like manner, the consequence of a ligature passed round the fish, and prevent- ing the expansion of the branchiai and the motion of the . opercula. • Dr. Monro computed that in the state, the surface of the g-llls is, at the least, equal to the whole surface of the human body. •j- It lias been generally stated by physiolog-ists, even of the highest au- thority, such as Cuvier, that the principal reason why fibhcs cannot maintain Vol. II. 28 218 THE VITAL FUNCTIONS. In all osseous fishes the opening under the operculum for the exit of the respired water, is a simple fissure; hut in most of the cartilaginous trihes, there is no operculum, and the water escapes tiu'ough a series of apertures in the side of the throat. Sharks have live ohlong orifices of this descrip- tion, as may he seen in Fig. 367.* As the Lcni} ])7'cy cm\)\o\s its mouth more constantly than other fisli in laying hold of its prey, and adiiering to other bodies, the organs of respiration are so constructed as to be indc])endcnt of the mouth in receiving tho water. There are seven external openings on each side (Fig- 3GS,) lead- ing into the same number of separate oval pouches, situated horizontally, and the inner membrane of which has the same structure as gills: these pouches are seen on a larger scale than in the preceding figure, in Fig. 369. There is also an equal number of internal openings, seen in the lower part of this last figure, leading into a tube, the lower end of which is closed and the upper terminates by a fringed edge in the oesophagus. The water which is received by the seven la- teral openings, enters at one side, and after it has acted uj)on the gills, passes round the projecting membranes. The greater part makes its exit by the same orifices; but a por- tion escapes into the middle tube, and thence passes, either into the other cavities, or into the oesophagus.! life, when suiToundeJ by air instead of water, is tliat the branchix become diy, unJ lose the power of acting" when thus deprived of their natural mois- ture: for it might otherwise naturally be expected that the oxygen of atmo- spheric air woukl exert a more powerful action on the blood whicli circulates in the branchix, than that of merely aerated water. The rectification of this en'or is d\ie to Flourens, who pointed out the true cause of sulFocation, stated in the text, in a Memoir entitled " Experiences sur le Mcchanisme de la Res- piration des Poissons." — Annates des Sciences Naturelles, xx. 5. • They arc also visible in Fig. 293, (page 122,) which is that of the«S'^ua- lus prhtis, a species belonging to this tribe. ■j- It was commonly supposed that the respired water is ejected through the nostril; but this is certainly a mistake, for the nostril has no communi- cation through the mouth, as was pointed out by Sir E. Home. Phil. Trans, for 1815, p. 259. These oj^'ans have also been described by Bloch and Gaertncr. RESPIRATION IN PISHES. 219 In the Myxine, which feeds upon the internal parts of its prey, and huries its head and part of its body in the flesli, . the openinpjs of the respiratory organs are removed suffi- ciently far from the head to admit of respiration going on ■while the animal is so employed; and tlicre are only two external openings, and six lateral pouches on each side, with tubes similar to those in tlic lamprey. The Ferca scandens (DaldorlT, ' ) whicli is a fisli inhabiting the seas of India, has a very remarkable structure adapting it to the maintenance of respiration, and consequently to tlic support of life for a considerable time when out of the water: and hence it is said occasionally to travel on land to some distance from the coast.t The pharyngeal bones of this fish have a foliated and cellular structure, which gives them a capacity for retaining a sufficient quantity of water, not only to keep the gills moist, but also to enable them to per- form their proper office; while not a particle of water is suf- fered to escape from them, by the opercula being accurately closed. The same faculty, resulting from a similar structure, is possessed by the Ophicephalus, which is also met with in the lakes and rivers of India and China. Eels are enabled to carry on respiration when out of water, for a certain pe- riod, in consequence of the narrowness of the aperture for the exit of the water from the branchial cavity, wliich en- ables it to be closed, and the water to be retained in tliat cavity, t I have already stated that, in all aquatic animals, the water which is breathed is merely the vehicle by which the air it contains is brought into contact with the organs of respiration. This air is constantly vitiated by the respiration of these animals, and requires to be renewed by tlie absorption of a * Jlnthlas testudinus (Rloch:) Anahs (Cuv.) f This peculiar faculty has been ah'cady alluded to In volume i. p. oOl. t Dr, Hancock states that the Jhras cuslalus, (Siluriis costaltis, Linn.) or Hassar, in very dry seasons, is sometimes seen, in i^rcat nuniliers, making- long- marches over land, in search of water. Edin. Phil. Journal, xx. 396. 220 THE VITAL FUNCTIONS. fresh portion, which can only take place whpn the water freely communicates with the atmosphere: and if this re- newal be by any means prevented, the water is no longer capable of sustaining life. Fishes are killed in a very [ew hours, if confined in a limited portion of water, which has no access to fresh air. When many fishes arc enclosed in a narrow vessel, they all struggle for the uppermost place, (where the atmospheric air is first absorbed,) like the unfor- tunate men imprisoned in the black-hole at Calcutta. AVhen a small fisli pond is frozen over, the fishes soon perish, un- less holes be broken in the ice, in order to admit air: they may be seen flocking towards these holes, in order to re- ceive the benefit of the fresh air as it is absorbed by the water; and so great is their eagerness on these occasions, that they often allow themselves to be caught by the hand. Water from which all air has been extracted, either by the air-pump, or by boiling, is to fishes what a vacuum is to a breathing terrestrial animal. Humboldt' and Provencal made a series of experiments on the quantities of air which fishes require for their respiration. They found that river- water generally contains about one 3Gth of its bulk of air, of which quantity one-third consists of oxygen, being about one per cent., of the whole volume. A tench is able to brQ^the when the quantity of oxygen is reduced to the 5000th part of the bulk of the water, but soon becomes ex- ceedingly feeble by the privation of this necessary element. The fact, however, shows the admirable perfection of the or- gans of this fish, which can extract so minute a quantity of air from water to which that air adheres with great tenacity.* • The swimming bladder of fishes is regarded by many of the German naturalists as having some relations with the respiratoiy function, and as be- ing the rudiment of the pulmonary cavity of land animals; the passage of communication wit'i the oesophagus being conceived to represent the trachea. The air contained in the swimming bladder of fishes has been examined by many chemists, but although it is generally found to be a mixture of oxygen and nitrogen, the proportion in which these gases exist is observed to vary considerably, liiot concluded from his experiments, that in the air- ATMOSPHERIC RESPIRATION. 221 § 3. •^tmosjiheric licsjjiration. The next series of structures which are to come under our review, comprehends all those adapted to the respiration of atmospheric air in its gaseous form; and their physiology is no less diversified than that of the organs hy whicli water is respired. Air may be respired by irachese, or by pulmonary cavi- ties; the first mode is exemplified in insects; the second is that adopted in the larger terrestrial animals. The greater part of the blood of insects being diffused by transudation through every internal organ of their bodies, and a small portion only being enclosed in vessels, and cir- culating in them, the salutary influence of the air could not have been generally extended to that fluid by any of the or- dinary modes of respiration, where the function is carried on in an organ of limited extent. As the blood could not be brought to the air, it became necessary, therefore, that the air should be brought to the blood. For this purpose, there has been provided, in all insects, a system of continuous and ramified vessels, called trachex, distributing air to every part of the body. The external orifices, from which these air tubes commence, are called spiracles, or stiginaia, and bladder of fishes inliabiting" the greatest depths of the ocean, the quantity of oxyg-en is greater, while in those of fishes which come often to the surface, the nitrog-en is more abundant; and De k lloche came to the same concUision from his researches on the fishes of the Mediterranean. From the exjieriments of Humboldt and Provencal, on the other hand, we may conclude, that the quality of the air contained in the air-bladder is but remotely coniiected with respiration. (Memoires de la Socicte d'Arcueil, ii. 359.) According- to Ehrmann, the Colitis, or Loche, occasionally swallows air, which is decomposed in the alimentary canal, and effects a change in the blood vessels, with which it is brought into contact, exactly similar to that which occurs in ordinary respiration. It is also believed that in all fishes a partial aeration of the blood is the result of a similar action, taking place at the surface of the body under the scales of the integuments. Cuvicr, sur les Poissons, I. 383. 222 THE VITAL rUNCTIOWS. are usually situated in rows on each side of the bodj^, as is shown in Fig. 370, which represents the lower abdominal surface of the Dytisciis marginalis. They are seen very distinctly in the caterpillar, which has generally ten on each side, corresponding to the number of abdominal segments. In many insects we find them 2:uardcd by bristles, or tufts of hair, and sometimes by valves, placed at the orifice, to prevent the entrance of extraneous bodies. The spiracles are opened and closed by muscles provided for that purpose. Fig. 371 is a magnified view of spiracles of this description, from the Cerainhyx heros, (Fab.) They are the begin- nings of short tubes, which open into large trunks, (as shown in Fig. 372,) extending longitudinally on each side, and sending off radiating branches from the parts which are op- posite to the spiracles; and these branches are farther subdi- vided, in the same manner as the arteries of the larger ani- mals, so that their minute ramifications pervade every organ in the body. This ramified distribution has frequently oc- casioned their being mistaken for blood vessels. In the wings of insects, the nervures, which have the appearance of veins, are only large air-tubes. Jurlne asserts that it is by forcing air into these tubes that the insect is enabled sud- denly to expand the wings in preparing them for flight, RESPIRATION IN INSECTS. 003 giving them, by this means, greater buoyancy, as well as tension. The trachea} are kept continually pervious by a curious mechanism: they are formed of three coats, the external and internal of which are membranous; but the middle coat is ^constructed of an elastic thread coiled into a helix, or cylin- drical spiral, (as seen in Fig. 372;) and the elasticity of this thread keeps the tube constantly in a state of expansion, and therefore full of air. When examined under water, the tra- cheas have a shining silvery appearance, from the air they contain. This structure has a remarkable analogy with that of the air vessels of plants, which also bear the name of tra- chea; and in both similar variations are observed in the con- texture of the elastic membrane by which they are kept pervious.* i. The tracheae, in many parts of their course, present re- markable dilatations, wdiich apparently serve as reservoirs of air: they are very conspicuous in the Dytiscus margina- lis, which resides principally in water; but they also exist in many insects, as the Melolontha and the Cerambijx, which live wlK)lly in the air.t Those of the Scolia horto- rum (Fab.) are delineated in Fig. 373, considerably magni- fied. If an insect be immersed in water, air will be seen es- caping in minute bubbles at each spiracle; and in proportion as the water enters into the tubes, the sensibility is de- stroyed. If all the spiracles be closed by oil, or any other unctuous substance, the insect immediately dies of suflbca- tion; but if some of them be left open, respiration is kept up to a considerable extent, from the numerous communi- cations which exist among the air vessels. Insects soon • According' to the observation of Dr. Kidd these vessels are often annular in insects, as is also the case witli those of plants. He considers the long-i- tudinal trachex as connecting channels, by which the insect is enabled to direct the air to particular pai'ts for occasional purposes. Phil. Trans, for 1825, p. 234. t L^on Diifour, Annalcs dcs Sciences Naturellesj viii. 26. 024 THE VITAL FUNCTIONS. perish when placed in the receiver of an air-pump, and the air exhausted; but they are generally more tenacious of life under these circumstances than the larger animals, and often, after being apparently dead, revive on the readmission of air. Aquatic insects have tracheae, like those living in air, and are frequently provided with tubes, which are of sufficient length to reach the surface of the water, where they absorb air for respiration. In a few tribes a complicated mode of respiration is practised; aerated water is taken into the body, and introduced into cavities, when the air is extracted from it, and transmitted by the ordinary tracheai to the different parts of the system.* Such, then, is the extensive apparatus for aeration in ani- mals!^ which have either no circulation of their nutritious juices, or a very imperfect one; but no sooner do we arrive at the examination of animals possessing an enlarged sys- tem of blood vessels, than we find nature abandoning the system of tracheae, and employing more simple means of effecting the aeration of the blood. Advantage is taken of the facility afforded by the blood vessels of transmitting the blood to particular organs, where it may conveniently re- ceive the influence of the air. Thus, Scorpions are provided, on each side of the thorax, with four pulmonary cavities, seen at l, on the left side of Fig. 374, into each of which air is admitted by a separate external opening, a, b, is the dorsal vessel, which is connected with the pulmonary cavi- ties by means of two sets of muscles, the one set (m, m) be- ing longer than the other (m, m, m.) The branchial arte- ries (v) are seen ramifying over the inner surface of the • Mr. Dutrochct conceives that the principle on which this operation is conducted is the same with that by which g-ascs are reciprocally transmitted througli moistened membranes; as in the experiments of Humboldt and Gay Lussac, who, on enclosing- mixtures of oxygen, nitrogen, and carbonic acid gases, iTi any proportion, in a membranous bladder, which was then im- mersed in aerated water, found that there is a reciprocal transit of the g-ases; until at leng-th pure atmospheric air remains in the cavity of the bladder. RESPIRATION IN INSECTS. 225 pulmonary cavities (r) on tlic right side, whence the blood is conveyed by a corresponding set of branchial veins to the dorsal vessel: and other vessels, which are oi'dinary veins, are seen at o, proceeding from the abdominal cavity to join the dorsal vessel. The membrane which lines the pulmonary cavities is curiously plaited, presenting the ap- pearance of the teeth of a comb, and partaking of the struc- ture of gills; and on this account these organs are termed by Latreille pneumo-hranchix. Organs of a similar de- scription exist in Spiders, some species having eight, others four, and some only two: but there is one entire order of Arachnida which respire by means of tracheae, and in these the circulation is as imperfect as it is in insects. It may here be remarked that an essential diflcrence ex- ists in the structure of the respiratory organs, according to the nature of the medium which is to act upon them: for in Vol. II. 29 226 THE VITAL FUNCTIONS. aquatic respiration tlie air contained in water is made to act on the blood circulating in vessels which ramify on the external surface of the filaments of the gills; while in at- mospheric resj)iration the air in its gaseous state is always received into cavities, on the internal surface of which the blood vessels, intended to receive its influence, are distri- buted. It is not difllcult to assign the final cause of this change of jilan; for in each case the structure is accommo- dated to the mechanical properties of the medium respired. A liquid, being inelastic and ponderous, is adapted, by its momentum alone, to separate and surround the loose float- ing filaments composing the branchiae; but a light gaseous fluid, like air, is, on the contrary, better fitted to expand di- latable cavities into which it may be introduced. Occasionally, however, it is found that organs constructed like branchiae, and usually performing aquatic respiration, can be adapted to respire air. This is the case with some species of Crustacea, of the order Decapoda, such as the Crab, which, by means of a peculiar apparatus, discovered by Audouin, and Milne Edwards, retain a quantity of water in the branchial cavity so as to enable them to live a very long time out of the water. It is only in their mature state of development, however, that they are qualified for this amphibious existence, for at an early period of growth they can live only in water. There is an entire order of Gasteropodous Mollusca which breathe atmospheric air by means of pulmonary cavities. This is the case with the Limax, or slug, and also with the Helix, or snail, the Testucella, the Claiisilia, and many others, which, though partial to moist situations, are, from the conformation of their respiratory organs, essentially land animals. The air is received by a round aperture near the head, guarded by a sphincter muscle, which is seen to dilate or contract as occasion may require, but which is sometimes completely concealed from view by the mouth folding over it. The cavity, to which this opening leads, is lined by a membrane delicately folded, and overspread RESPIRATION BY LUNGS. 221 with a beautiful net-work of pulmonary vessels. Other mollusca of the same order, which arc more aquatic in their habits, have yet a similar structure, and are ol)liged at in- tervals to come to the surface of the water in order to breatiie atmospheric air: this is the case with the Onchidium, the Planorbis, the Lymnsea, &c. The structure of the pulmonary organs becomes gradually more refined and complicated as we ascend to the higher classes of animals. In all vertebrated terrestrial animals they are called lungs, and consist of an assemblage of vesi- cles, into which the air is admitted by a tube, called the trachea, or wind-pipe, extending downwards from the back of the mouth, parallel to the oesophagus. Great care is taken to guard the beginning of this passage from the intrusion of any solid or liquid that may be swallowed. A cartilaginous valve, termed the epiglottis, is generally provided for this purpose, which is made to descend by the action of the same muscles that perform deglutition, and which then closes ac- curately the entrance into the air-tube. It is an exceedingly beautiful contrivance, both as to the simplicity of the me- chanism, and the accuracy with which it accomplishes the purpose of its formation. At the upper part of the chest the trachea divides into two branches, called the bronchia, passing to the lungs on either side. Both the wind-pipe and the bronchia are prevented from closing by the inter- position of a series of firm cartilaginous ringlets, interposed between their inner and outer coats, and placed at small and equal distances from one another. The natural elasticity of these ringlets tends to keep the sides of the tube stretched, and causes it to remain open: it is a structure very analo- gous to that of the trachea of insects, or of the vessels of the same name in plants. The lungs of Reptiles consist of largo sacs, into the cavity of which the bronchia, proceeding from the bifurcation of the trachea, open at once, and without farther subdivision. Cells are formed within the sides of this great cavity, by fine membranous partitions, as thin and delicate as soap 228 THE VITAL FUNCTIONS. bubbles. The lungs of serpents have scarcely any of these partitions, but consist of one simple pulmonary sac, situated on the rijj^ht side, having the slender elongated form of all the other viscera, and extending nearly the whole length of the body. The lung on the left side is in general scarcely discernible, being very imperfectly developed. In the cha- meleon the lungs have numerous processes which project from them like ca^ca. In the Sauria, the lungs are more confined to the thoracic region, and are more completely cellular. The mechanism, by which, in these animals, the air is forced into the lungs, is exceedingly peculiar, and was for a long time a subject of controversy. If we take a frog as an example, and watch its respiration, wc cannot readily discover that it breathes at all, for it never opens its mouth to receive air, and there is no motion of the sides to indi- cate that it respires; and yet, on any sudden alarm, we see the animal blowing itself up, as if by some internal power, though its mouth all the while continues to be closed. We may perceive, however, that its throat is in frequent mo- tion, as if the frog were economizing its mouthful of air, and transferring it backwards and forwards between its mouth and lungs; but if we direct our attention to the nos- trils, we may observe in them a twirling motion, at each movement of the jaws; for it is, in fact, through the nostrils that the fros: receives all the air which it breathes. The jaws are never opened but for eating, and the sides of the mouth form a sort of bellows, of which the nostrils are the inlets; and by their alternate contraction and relaxation the air is swallowed, and forced into the trachea, so as to inflate the lungs. If the mouth of a frog be forcibly kept open, it can no longer breathe, because it is deprived of the power of swallowing the air required for that function; and if its nostrils be closed, it is, in like manner, suffocated. The respiration of most of the Reptile tribes is performed in a similar manner; and they may be said rather to swallow the air they breathe, than to draw it in by any expansive action RESPIRATION IN REPTILES. 229 of the parts which surround the cavity of lungs; for even the ribs of serpents contribute but little, by their motion, to this effect, being chiefly useful as organs of progressive motion. The Chelonia have lungs of great extent, passing back- wards under the carapace, and reaching to the posterior part of the abdomen. Turtles, which are aquatic, derive great advantages from this structure, which enables them to give buoyancy to their body, (encumbered as it is with a heavy shell,) by introducing into it a large volume of air; so that the lungs, in fact, serve the purposes of a large swim- ming bladder. That this use was contemplated in their structure is evident from the volume of air received into the lungs, being much greater than is required for the sole purpose of respiration. The section of the lungs of the tur- tle (Fig. 375,) shows their interior structure, composed of large cells, into which the trachea (t) opens. Few subjects in animal physiology are more deserving the attention of those whose object is to trace the operations of nature in the progressive development of the organs, than the changes which occur in the evolution of the tadpole trom the time it leaves the egg till it has attained the form of the 230 THE VITAL FUNCTIONS. perfect frog. We have already had occasion to notice seve- ral of these transformations in the organs of the mechanical functions, and also in those of digestion and circulation: but the most remarkable of all are the changes occurring in the respiratory apparatus, corresponding with the opposite na- ture of the elements which the same animal is destined to inhabit in the different stages of its existence. No less than three sets of organs are provided for respiration; the two first being branchiae, adapted to the fish-like condition of the tadpole, and the last being pulmonary cavities, for receiving air, to be employed when the animal exchanges its aquatic for its terrestrial life. It is exceedingly interesting to ob- serve that this animal at first breathes by gills, wdiich pro- ject in an arborescent form from the sides of the neck, and float in the water; but that these structures are merely tem- porary, being provided only to meet the immediate exigen- cy of the occasion, and being raised at a period when none of the internal organs are as yet perfected. As soon as ano- ther set of gills, situated internally, can be constructed, and are ready to admit the circulating blood, the external gills are superseded in their office; they now shrivel, and are re- moved, and the tadpole performs its respiration by means of branchiae, formed on the model of those of fishes, and acting by a similar mechanism. By the time that the system has undergone the changes necessary for its conversion into the frog, a new apparatus has become evolved for the respira- tion of air. These are the lungs, which gradually coming into play, direct the current of blood from the branchiae, and take upon themselves the whole of the office of respiration. The branchiae, in their turn, become useless, are soon obli- terated, and leave no other trace of their former existence than the orio;inal division of the arterial trunks, which had su])plied them with blood directly from the heart, but which, now uniting in the back, form the descending aorta.* , There is a small family called the Perenni-branchia, be- • See Fig. 357, p. 197. RESPIRATION IN REPTILES. 231 longing to this class, which, instead of undergoing all the changes I have been describing, present, during their whole lives, a great similitude to the first stage of the tadpole. This is the case with the */lxolotl, the Proteus angimius, the Siren lacertina, and the Menobranchiis lateralis, which permanently retain their external gills, while at the same time they possess imperfectly developed lungs. It would therefore seem as if, in these animals, the progress of deve- lopment had been arrested at an early stage, so that their adult state corresponds to the larva condition of the frog.* In all warm-blooded animals respiration becomes a func- tion of much greater importance, the continuance of life being essentially dependent on its vigorous and unceasing exercise. The whole class of Mammalia have lungs of an exceedingly developed structure, composed of an immense number of minute cells, crowded together as closely as pos- sible, and presenting a vast extent of internal surface. The thorax, or cavity in which the lungs, together with tlic heart and its great blood vessels, are enclosed, has somewhat the shape of a cone; and its sides are defended from compres- sion by the arches of the ribs, which extend from the spine to the sternum, or breast-bone, and produce mechanical sup- port on the same principle that a cask is strengthened by being girt with hoops, which, though composed of compara- tively weak materials, are yet capable, from their circular shape, of presenting great resistance to any compressive force. While Nature has thus guarded the chest, with such pe- culiar solicitude, against the efibrts of any external force, tending to diminish its capacity, she has made ample provi- sion for enlarging or contracting its diameter in the act of * GeofFroy St. Hilaire thinks there is g-vound for believing that Crocodiles and Turtles possess, in addition to the ordinary pulmonary respiration, a par- tial aquatic abdominal respiration, cficcted by means of the two channels of communication which have been found to exist between the cavity of the abdomen and the external surface of the body: and also that some analogy may l)e traced between this aquatic respiration in reptiles, by these pcrituncal canalSi and tlie supposed function of the swimming bladder of fishes, \\\ sub- serviency to a species of aerial rcspimtion. 232 THE VITAL FUNCTIONS. respiration. First, at tlie lower part, or that which cor- responds to the hasis of the cone, the only side, indeed, which is not defended by bone, there is extended a thin ex- pansion, partly muscular, and partly tendinous, forming a complete partition, and closing the cavity of the chest on the side next to the abdomen. This muscle is called the Dia- phragm: it is perforated, close to its origin from the spine, by four tubes, namely, the oesophagus, the aorta, the vena cava, and the thoracic duct. Its surface is not flat, but con- vex above, or towards the chest; and the direction of its fibres is such, that, when they contract, they bring down the middle part, which is tendinous, and render it more flat than before, (the passage of the four tubes already mentioned, not interfering with this action,) and thus, the cavity of the tho- rax may be considerably enlarged. It is obvious that if, upon the descent of the diaphragm, the lungs were to re- main in their original situation, an empty space would be left between them and the diaphragm. But no vacuum can take place in the body; the air cells of the lungs must al- ways contain, even in their most compressed state, a certain quantity of air: and this air will tend, by its elasticity, to ex- pand the cells; the lungs will, consequently, be dilated, and will continue to fill the chest; and the external air will rush in through the trachea in order to restore the equilibrium. This action is termed inspiration. The air is again thrown out when the diaphragm is relaxed, and pushed upwards, by the action of the large muscles of the trunk; the elasticity of the sides of the chest, concurring also in the same effect; and thus expiration is accomplished. The muscles which move the ribs conspire also to produce dilatations and contractions of the cavity of the chest. Each rib is capable of a small degree of motion on that extremity by which it is attached to the spine; and this motion, as- suming the chest to be in the erect position, as in man, is chiefly upwards and downwards. But, since the inclination of the ribs is such that their lower edges form acute angles with the spine, they bend downwards as" they proceed to- wards the breast; and the uppermost rib being a fixed point. RESPIRATION IN MAMMALIA. 233 the action of the intercostal muscles, which produces an ap- proximation of the ribs, tends to raise them, and to bring them more at riglit angles with the spine; the sternum, also, to which the other extremities of the ribs are articulated, is elevated by this motion, and, consequently, removed to a greater distance from the spine; the general result of all these actions being to increase the capacity of the chest. Thus, there are two ways in which the cavity of the thorax can be dilated; namely, by the action of the dia- phragm, and by the action of the intercostal muscles. It is only in peculiar exigencies that the whole power of this ap- paratus is called into action; for in ordinary respiration the diaphragm is the chief agent employed, and the princi])al effect of the action of the intercostal muscles is simply to fix the ribs, and thus give greater purchase to tlic diapliragm. The muscles of the ribs arc employed chiefly to give active Vol. II. 30 234 THE VITAL FUNCTIONS. assistance to the diaphragm, when, from any cause, a diffi- culty arises in dilating the chest. In Birds the mechanism of respiration proceeds upon a different plan, of which an idea may he derived from the view given of the lungs of the Ostrich, at l. l.. Fig. 377. The construction of the lungs of hirds is such as not to admit of any change in their dimensions; for they are very compact in their texture, and are so closely braced to the ribs, and up- per parts of the chest, by firm membranes, as to preclude all possibility of motion. They in part, indeed, project behind the intervals between the ribs, so that their whole mass is not altogether contained within the thoracic cavity. There is no large muscular diaphragm by which any change in the capacity of the chest could be effected, but merely a few narrow slips of muscles, arising from the inner sides of the ribs, and inserted into the thin transparent membrane which covers the lower surface of the lungs. They have the ef- fect of lessening the concavity of the lungs towards the abdo- men at the time of inspiration, and thereby assist in dilating the air-cells.* The bronchia, or divisions of the trachea (t,) after opening, as usual, into the pulmonary air-cells, do not terminate there, but pass on to the surface of the lungs, where they open by numerous apertures. The air is ad- mitted, through these apertures into several large air-cells (c c c,) which occupy a considerable portion of the body, and which enclose most of the large viscera contained in the abdomen, such as the liver, the stomach, and the intestines;! and there are, besides, many lateral cells in immediate com- munication with the lungs, and extending all down the sides of the body. Numerous air-cells also exist between the muscles, and underneath the skin; and the air penetrates even into the interior of the bones themselves, filling the spaces usually occupied by the marrow, and thus contributing ma- * Hunter in the Animal Economy, p. 78. -}■ It was asserted by the Parisian Academicians, that the air got admission into the cavity of the pericardium, in which the heart is lodged. This error was first pointed out by Dr. Macartney, in Rees's Cyclopaedia.— Art. Bird, RESPIRATION IN BIRDS. 235 terially to the litghtness of the fahric* All these cells are veryjarge and numerous in birds which perform the highest and most rapid flight, such as the eagle. The bill of the Toucan^ which is of a cellular structure, and also the cells between the plates of the skull in the Oi^;/, are,in like man- ner, filled with air, derived from the lungs: the barrels of the large quills of the tails and wings are also supplied with air from the same source. In birds, then, the air is not merely received into the lungs, but actually passes through them, bein|[ drawn for- wards by the muscles of the ribs when they elevate the chest, and produce an expansion of the subjacent air-cells. The chest is depressed, for the purpose of expiration, by another set of muscles, and the air driven back: this air, consequently, passes a second time through the lungs, and acts twice on the blood which circulates in those organs. It is evident that if the lungs of birds had been constructed on the plan of those of quadrupeds, they must have been twice as large to obtain the same amount of aeration in the blood; and consequently must have been twice as heavy, which would have been a serious inconvenience in an ani- mal formed for flying.t The diffusion of so large a quantity of air throughout the body of animals of this class presents an analogy with a similar purpose apparent in the confor- mation of insects, where the same object is effected by means of tracheae.:}: * In birds, not formed for extensive flight, as the gallinaceous tribes, tlie humerus is the only bone into which air is introduced. — Hunter on tlie Animal Economy, p. 81. \ I must mention, however, that the correctness of this view of the sub- ject is contested by Dr. Macartney, who thinks it probable that the air, on its return from the large air-cells, passes directly by the large air-holes into the bronchia, and is not brought a second time into contact with the blood. \ The peculiarities of structure in the rcspiratoiy system of birds have probably a relation to the capability we see them possess, of bearing with impunity, very quick and violent changes of atmospheric pressure. Thus, the Condor of the Andes is often seen to descend rapidly from a heiglit of above 20,000 feet, to the edge of the sea, where the air is more than twice 236 THE VITAL FUNCTIONS Thus, has the mechanism of respiration been varied in the diflferent classes of animals, and adapted to the particu- lar element, and mode of life designed for each. Combined with the peculiar mode of circulation, it affords a tolerably accurate criterion of the energy of the vital powers. In birds, the muscular activity is raised to the highest degree, in consequence of the double effect of the air upon the whole circulating blood in the pulmonary organs. The Mamma- lia rank next below birds, in the scale of vital energy; but they still p(^sess a double circulation, and breathe atmo- spheric air. The torpid and cold-blooded reptiles are sepa- rated from mammalia by a very wide interval, because, al- though they respire air, that air only influences a part of the blood; the pulmonary, being only a branch of the gene- ral circulation. In fishes, again, we have a similar result, because, although the w^hole blood is brought by a double circulation to the respiratory organs, yet it is acted upon only by that portion of air which is contained in the water respired, and which is less powerful in its action than the same element in its gaseous state. We may, in like man- ner, continue to trace the connexion between the extent of these functions and the degrees of vital energy throughout the successive classes of invertebrate animals. The vigour and activity of the functions of insects, in particular, have an evident relation to the effective manner in which the complete aeration of the blood is secured by the extensive distribution of trachea? through every part of their system. § 4. Chemical Changes effected by Resjnration. We have next to direct our attention to the chemical of- fices which respiration performs in the animal economy. It the density of that which the bird had been breathing-. We are, as yet, una- ble to trace the connexion which probably exists between the structure of the lung^, and this extraordinary power of* accommodation tq^such great and sudden variations of atmospheric pressure. CHEMICAL EFFECTS OF RESPIRATION. 237 is only of late years that we may be said to have obtained any accurate knowledge as to the real nature of this impor- tant function; and there is perliaps no brancli of physiology which exhibits in its history a more humiliating picture of the wide sea of error in which the human intellect is prone to lose itself, when the path of philosophical induction is abandoned, than the multitude of wild and visionary hypo- theses, devoid of all solid foundation, and perplexed by the most inconsistent reasonings, which formerly prevailed with regard to the objects and the processes of respiration. To give an account, or even a brief enumeration of these thco- ries, now sufficiently exploded, would be incompatible with the purpose to which I must confine myself in this treatise.* I shall content myself, therefore, with a concise statement of such of the leading facts relating to this function, as have now, by the labours of modern physiologists, been satisfac- torily established, and which serve to 'elucidate the benefi- cent intentions of nature in the economy of the animal sys- tem. Atmospheric air acts without difficulty upon the blood w^hile it is circulatins; through the vessels which are rami- fied over the membranes lining the air cells of the lungs; for neither these membranes, nor the thin coats of the ves- sels themselves, present any obstacle to the transmission of chemical elements from the one to the other. The blood being a highly compound fluid, it is exceedingly difficult to obtain an accurate analysis of it, and still more to ascertain with precision the different modifications which occur in its chemical condition at different times: on this account, it is scarcely possible to determine, by direct observation, what are the exact chemical changes, which that fluid undergoes * For an account of the history of the various chemical theories which have prevailed on this "interesting department of Physiolog-y, I must refer to the " Essay on Respiration," by Dr. Bostock, and also to the " Elementary System of Physiology," by the same author, which latter work comprises the most comprehensive and accurate compendium of the science which has yet appeared. 23S THE VITAL FUNCTIONS. during Its passage through the lungs; and we have only col- lateral evidence to guide us in the inquiry.* The most obvious effect resulting from the action of the air is a change of colour from the dark purple hue, which the blood has when it is brought to the lungs, to the bright vermilion colour, wlilch it is found to assume in those or- gans, and which accompanies its restoration to the qualities of arterial blood. In what the chemical difference between these two states consists may, in some measure, be collected from the changes which the air itself, by producing them, has experienced. The air of the atmosphere, which is taken into the lungs, is known to consist of about twenty per cent, of oxygen gas, seventy-nine of nitrogen gas, and one of carbonic acid gas. When it has acted upon the blood, and is returned from the lungs, it is found that a certain proportion of oxygen, which it had contained, has disappeared, and that the place of this oxygen is almost wholly supplied by an addition of carbonic acid gas, together with a quantity of watery vapour. It ap- pears also probable that a small portion of the nitrogen gas is consumed during respiration. For our knowledge of the fact of the disappearance of ox- ygen we are indebted to the labours of Dr. Priestley. It had, indeed, been long before suspected by Mayovv, that some portion of the air inspired is absorbed by the blood; but the merit of the discovery that it is the oxygenous part of the air which is thus consumed is unquestionably due to Dr. * Some experiments very recently made by Messrs. Macaire and Marcet, on the ultimate analysis of arterial and venous blood, taken from a rabbit, and dried, have shown that the former contains a lai'g-er proportion of oxy- g-en than the latter; and that the latter contains a larger proportion of carbon than the former: the proportions of nitrogen and hydrogen being the same in both. The following are the exact numbers expressive of these propor- tions: Carbon. Oxygen. JVitrogen. JJydrogen. Arterial blood 50.2 . . . 26.3 . . . 16.3 ... 6.6 Venous blood 55.7 . . . 21.7 . . . 16.2 ... 6.4 Memoires de la Society de Physique ei d'JIist. Naturelk de Geneve. T. v. p. 400. CHEMICAL EFFECTS OF RESPIRATION. 239 Priestley. The exact quantity of oxygen, which is lost in natural respiration, varies in different animals, and even in different conditions of the same animal. Birds, for in- stance, consume larger quantities of oxygen by their res- piration; and hence require, for the maintenance of life, a purer air than other vertebrated animals. Vauquelin, how- ever, found that many species of insects and worms pos- sess the power of abstracting oxygen from the atmosphere in a much greater degree than the larger animals. Even some of the terrestrial mollusca, such as snails, are capa- ble of living for a long time in the vitiated air in which a bird had perished. Some insects, which conceal them- selves in holes, or burrow under ground, have been known to deprive the air of every appreciable portion of its oxygen. It is observed by Spallanzani, that those animals, whose modes of life oblige them to remain for a great length of lime in these confined situations, possess this power in a greater degree than others, which enjoy more liberty of moving in the open air: so admirably have the faculties of animals been, in every instance, accommodated to their re- spective w^ants. Since carbonic acid consists of oxygen and carbon, it is evident that the portion of that gas which is exhaled from the lungs is the result of the combination of either the whole, or a part, of the oxygen gas, which has disappeared during the act of respiration, with the carbon contained in the dark venous blood, which is brought to the lungs. The blood having thus parted with its superabundant carbon, which escapes in the form of carbonic acid gas, regains its natural vermilion colour, and is now qualified to be again transmit- ted to the different parts of the body for their nourishment and growth. As the blood contains a greater proportion of carbon than the animal solids and fluids which are formed from it, this superabundant carbon gradually accumulates in proportion as its other principles, (namely, oxygen, hydro- gen, and nitrogen) are abstracted from it by the processes of secretion and nutrition. By the time it has returned to the 240 THE VITAL FUNCTIOJTS. heart, therefore, it is loaded with carbon, a principle, which, when in excess, becomes noxious, and requires to be re- moved from the blood, by combining it with a fresh quan- tity of oxygen obtained from the atmosphere. It is noty^t satisfactorily determined whether the whole of the oxygen, which disappears during respiration, is employed in the for- mation of carbonic acid gas: it appears, probable, however, from tlie concurring testimony of many experimentalists, that a small quantity is permanently absorbed by the blood, and enters into it as one of its constituents. A similar question arises with respect to nitrogen, of which, as I have already mentioned, it is probable that a small quantity disappears from the air when it is respired; although the accounts of experimentalists are not uniform on this point. The absorption of nitrogen during respiration was one of the results which Dr. Priestley had deduced from his experiments: and this fact, though often doubted, ap- pears, on the whole, to be tolerably well ascertained by the inquiries of Davy, Pfaflf, and Henderson. With regard to the respiration of cold-blooded animals, it has been satisfac- torily established by the researches of Spallanzani, and more especially by those of Humboldt and Provengal, on fishes, that nitrogen is actually absorbed. A confirmation of this result has recently been obtained by Messrs. Macaire and Marcet, who have found that the blood contains a larger proportion of nitrogen than the chyle, from which it is formed. We can discover no other source from which chyle could acquire this additional quantity of nitrogen, during its conversion into blood, than the air of the atmosphere, to which it is exposed in its passage through the pulmonary vessels.* According to these views of the chemical objects of res- piration, the process itself is analogous to those artificial operations which effect the combustion of charcoal. The food supplies the fuel, which is prepared for use by the di- * See the note at pag-e 238. CHEMICAL EFFECTS OF RESPIRATION. 241 gestive organs, and conveyed by the pulmonary arteries to the place where it is to undergo combustion: the diaphragm is the bellows, which feeds the furnace with air; and the tra- chea is the chimney, through which the carbonic acid, which is the product of the combustion, escapes. It becomes an interesting problem to determine whetlier this analogy may not be farther extended; and whether the combustion of carbon, which takes place in respiration, be not the exclusive source of the increased temperature, which all animals, but more especially those designated as warin- bloodedy usually maintain above the surrounding medium. The uniform and exact relation which may be observed to take place between the temperature of animals and the ener- gy of the respiratory function, or, rather, the amount of the chemical changes induced by that function, affords very strong pvidence in favour of this hypothesis. The coiiici- dence, indeed, is so strong, that, notwithstanding the objec- tions that have been raised against the theory founded upon this hypothesis, from some apparent anomalies which occa- sionally present themselves, we must, I think, admit that it affords the best explanation of the phenomena of any theory yet proposed, and that, therefore, it is probably the true one. The maintenance of a very elevated temperature appears to require the concurrence of two condi|ions; namely, first, that the whole of the blood should be subjected to the influ- ence of the air, and, secondly, that the air should be pre- sented to it in a gaseous state. These, then, are the circum- stances which establish the great distinction between warm and cold-blooded animals; a distinction which at once stamps the character of their whole constitution. It is the condition of a high temperature in the blood which raises the quadru- ped and the bird to a rank, in the scale of vitality, so far above that of the reptile: it is this which places an insupera- ble boundary between mammalia and fishes. However the warm-blooded Cetacea, who spend their lives in the ocean, may be found to approximate in their outward form, and in their external insti'umcnts of motion, to the other iniiabitants Vol II. 31 242 THE VITAL FUNCTIONS. of the deep, they are still, from the conformation of their respiratory organs, dependent on another element. If a seal, a porpoise, or a dolphin were confined, hut for a short time, under the surface of the water, it would perish with the same certainty as any otiicr of the mammalia, placed in the same situation. We ohserve them continually rising to the surface in order to hreathe, under every circumstance of pri- vation or of danger; and however eagerly they may pursue their prey, however closely they may be pressed by their enemies, a more urgent want compels them, from time to time, to respire air at the surface of the sea. Were it not for this imperious necessity, the Whale, whose enormous bulk is united with corresponding strength and swiftness, would live in undisturbed possession of the widely extended domains of the ocean, might view, without dismay, whole fleets sent out against him, and might defy all the efforts that man could practise for his capture or destruction. But the constitution of his blood, obliging him to breathe at the sur- face of the water, brings him within the reach of the fatal harpoon. In vain, on feeling himself wounded, does he plunge for refuge into the recesses of the deep; the same ne- cessity recurs, and compelling him again to present himself to his foes, exposes him to their renewed attacks, till he falls in the unequal strtMgle. His colossal form and gigantic strength are of little avail against the power of man, feeble though that power may seem, when physically considered, but which derives resistless might from its association with an immeasurably superior intellect. ( 243 ) CHAPTER XII. SECRETION. The capability of effecting certain chemical changes in the crude materials introduced into the body, is one of the powers which more especially characterize life; but although this power is exercised both by vegetable and by animal or- ganizations, we perceive a marked difference in the results of its operation in these two orders of beings. The food of plants consists, for the most part, of the simpler combina- tions of elementary bodies, which are elaborated in celhi- lar or vascular textures, and converted into various pro- ducts. The oak, for example, forms, by the powers of ve- getation, out of these elements, not only the green pulpy matter of its leaves, and the light tissue of its pith, but also the densest of its woody fibres. It is from, similar materials, again, that the olive prepares its oil, and the cocoa-nut its milk; and the very same elements in different states of com- bination, compose, in other instances, at one time the luscious sugar of the cane, at another the narcotic juice of the poppy, or the acrid principle of the euphorbium; and the same plant which furnishes in one part the bland farina of the potato, will produce in another the poisonous extract of the night- shade. Yet all these, and thousands of other vegetable pro- ducts, differing widely in their sensible qualities, agree very nearly in their ultimate chemical analysis, and owe their pe- culiar properties chiefly to the order in which their elements are arranged; an order dependent on the processes to which they have been subjected in the system of each particular vegetable. 244 THE VITAL FUNCTIONS. In the animal kingdom we observe these processes mul- tiplied to a still greater extent; and the resulting substances are even farther removed from the oriiirinal condition of un- organized matter. In the first place, the food of animals, instead of being simple, like that of plants, has always un- dergone previous preparation; for it has either constituted a portion of some otlier organized being, or it has been a product of organization; in each case, therefore, partaking of the complexity of composition which characterizes or- ganized bodies. Still, whatever may be its qualities when received into the stomach, it is soon converted by the pow- ers of digestion into a milky, or transparent fluid, having nearly the same uniform properties. We have seen that there is scarcely any animal or vegetable substance, how- ever dense its texture, or virulent its qualities, but is capa- ble of affording nourishment to various species of animals. Let us take as an example the elytra of cantharides, which are such active stimulants when applied in powder to the skin in the ordinary mode of blistering; we find that, not- withstanding their highly acrid qualities, they constitute the natural food of several species of insects, which devour them with great avidity; and yet the fluids of these insects, though derived from this pungent food, arc perfectly bland, and devoid of all acrimony. Cantharides are also, accord- ing to Pallas, the favourite food of the hedge-hog; although to other mammalia they are highly poisonous. It has also been found that even those animal secretions, (such as the venom of the rattle-snake,) which, when infused, even in the minutest quantity, into a wound, prove instantly fatal, may be taken into the stomach without producing any de- leterious effects. These, and a multitude of other well- known fiicts, fully prove how completely substances re- ceived as aliment may be modified, and their properties changed, or even reversed, by the powers of animal diges- tion. No less remarkable arc the transmutations, which the blood itself, the result of these previous processes, is subse- SECRETION. 245 quently made to undergo in the course of circulation, and when subjected to the action of the nutrient vessels and se- creting organs; being ultimately converted into the various textures and substances which compose all the parts of the frame. All the modifications of cellular substance, in its various states of condensation; the membranes, the liga- ments, the cartilages, the bones, the marrow; the muscles, with their tendons; the lubricating fluid of the joints; the medullary pulp of the brain; the transparent jelly of the eye; in a word, all the diversified textures of the various organs, which are calculated for such different ofiices, are derived from the same nutrient fluid, and may be considered as being merely modified arrangements of the same ultimate chemical elements. In what, then, we naturally ask, consists this subtle che- mistry of life, by which nature effects these multifarious changes; and in what secret recesses of the living frame has she constructed the refined laboratory in which she operates her marvellous transformations, far surpassing even those which the most visionary alchemist of former times had ever dreamed of achieving? Questions like these can only be fairly met by the confession of profound ignorance; for, although the subject of secretion has long excited the most ardent curiosity of physiologists, and has been prosecuted with extraordinary zeal and perseverance, scarcely any po- sitive information has resulted from their labours, and the real nature of the process remains involved nearly in the same degree of obscurity as at first. '" It was natural to ex- ♦ It is not yet precisely determined to what extent the organs of secretion are immediately instrumental in producing the sL'bstance which is secreted; and it has been even suggested that possibly their office is confined to the mere separation, or filtration from the blood, of cert:un animal products, which are always spontaneously forming in that fluid in the course of its circulation. This hypotliesis, in which the glands, and other secreting ap- paratus are regarded as only very fine strainers, is supported by a few facts, which seem to indicate the presence of these products in the blood, inde- pendently of the secreting processes by which they are usually supposed to be formed; but the evidence is as yet too scanty and equivocal to warrant the deduction of any general theory on the subject. 246 THE VITAL FUNCTIONS. pect that in this inquiry material assistance would be de- rived from an accurate anatomical examination of the or- gans by which the more remarkable secretions are formed; yet, notwithstanding the most minute and careful scrutiny of these organs, our knowledge of the mode in which they are instrumental in effecting the operations which are there conducted, has not in reality advanced a single step. To add to our perplexity, we often see, on the one hand, parts, to all appearance very differently organized, giving rise to secretions of a similar nature; and, on the other hand, sub- stances of very different properties produced by organs, which, even in their minutest details, appear to be identical in their structure. Secretions are often found to be poured out from smooth and membranous surfaces, such as those which line the cavities of the abdomen, the chest, and the head, and which are also reflected inwards, so as to invest the organs therein contained, as the heart, the lungs, the stomach, the intestines, the liver, and the brain.* In other instances, the secreting membrane is thickly set with mi- nute processes, like the pile of velvet: these processes are called villi, and their more obvious use, as far as we can perceive, is to increase the surface from which the secretion is prepared. At other times we see an opposite kind of structure employed; the secreting surface being the internal lining of sacs or cells, either opening at once into some larger cavity, or prolonged into a tube, or duct, for convcy- • Sometimes the secreting- organ appears to be entirely composed of a mass of vessels covered with a smooth membrane; in other cases, it appears to contain some additional material, or parenchyma, as it is termed. Verte- brated animals present us with numerous instances of glandular organs em- ployed for special purposes of secretion: thus, in the eyes of fishes there ex- ists a large vascular mass, which has been called the choroid gland, and which is supposed to be placed there for the purpose of replenishing some of the humours of the eye, in proportion as they are wasted. Within the air-bladder of several species of fishes there is found a vascular organ. Appa- rently sci-ving to secrete the air with which the bladder is filled; numerous ducts, filled with air, having been observed proceeding from the organ, and opening on the inner surface of the air-bladder. SECRETION. 247 ing the secreted fluid to a more distant point. These cells,- or follicles, as they are termed, are generally employed for the mucous secretions, and are often scattered throughout the surfaces of membranes:* at other times the secreting cavities are collected in great numbers into groups; and they then frequently consist of a series of lengthened tubes, like caeca, examples of which we have already seen in the hepatic and salivary glands of insects. A secretory organ, in its simplest form, consists of short, narrow and undivided tubes; we next find tubes which are elongated, tortuous or convoluted, occasionally presenting di- lated portions, or even having altogether the appearance of a collection of pouches, or sacs; while, in other cases, they are branched, and extend into minute ramifications. Sometimes they are detached, or isolated; at other times they are collected into tufts, or variously grouped into masses, where still the se- parate tubes admit of being unravelled. The secreting fila- ments of insects float in the general cavity, containing the mass of nutrient fluid, and thence imbibe the materials they require for the performance of their functions. It is only when they receive a firm investment of cellular membrane;, forming what is termed diCapsule, and assuming the appear- ance of a compact body, that they properly constitute a gland; and this form of a secreting organ is met with only among the higher animals.t Great variety is observable both in the form and struc- ture of difl'erent glands, and in the mode in which their blood vessels are distributed. In animals which are fur- nished with an extensive circulation, the vessels supplying the glands with blood are distributed in various modes; and it is evident that each plan has been designedly selected with reference to the nature of the particular secretion to * See p. 135 of this volume; and in particular Fig. 305. Sebaceous folli- cles are also noticed in Vol. i. p. 91. •j- Dr. Kidd, however, describes bodies apparently of a glandular charac- ter, disposed in rows on the inner surface of the intestinal canal of the Gryl' lotalpa, or mole-cricket. Phil. Tran. for 1825, p. 227. 248 THE VITAL FUXCTIOiNS. be performed, although we are here unable to follow the connexion between the means and the end. In some glands, for example, the minute arteries, on their arrival at the or- gan, suddenly divide into a great number of smaller branch- es, like the fibres of a camel-hair pencil: this is called the pencillaled structure. Sometimes the minute branches, in- stead of proceeding parallel to each other after their divi- sion, separate like rays from a centre, presenting a stel- lated, or star-like arrangement. In the greater number of instances, the smaller arteries take a tortuous course, and are sometimes coiled into spirals, but generally the convo- lutions are too intricate to admit of being unravelled. It is only by the aid of the microscope that these minute and delicate structures can be rendered visible; but the fallacy, to which all observations requiring the application of high inagnifying powers are liable, is a serious obstacle to the ad- vancement of our knowledge in this department of phy- siology. Almost the only result, therefore, which can be collected from these laborious researches in microscopic ana- tomy, is that nature has employed a great diversity of means for the accomplishment of secretion; but we still remain in ignorance as to the kind of adaptation, which must assuredly exist, of each structure to its respective object, and as to the nice adjustment of chemical affinities which has been pro- vided in order to accomplish the intended effects."* Elec- * The only instance in which we can perceive a correspondence between the chemical properties of the secretion, and the kind of blood from which it is prepared, is in the liver, which, imlike all the other glands, has venous, instead of arterial blood, sent to it for that purpose. The veins, which re- turn the blood that has circulated through the stomach, and other abdominal viscera, are collected into a large trunk, called the vena portas, which enters the liver, and is there again subdivided and ramified, as if it were an artery: its minuter branches here unite with those of the hepatic artery, and ramify through the minute lobules which compose the substance of the liver. After the bile is secreted, and carried off by hepatic ducts, the remaining blood is conducted, by means of minute hepatic veins, which occupy the centres of each lobule, into larger and larger trunks, till they all unite in the vena cava, going directly to the heart. (See Kiernan's Paper on the Anatomy and Phy- siology of the Liver, Phil. Trans, for 1833, p. 711.) A similar system of ve- • SECRETION. 249 tricity is, no doubt, an important agent in all these processes, but in the absence of all certain knowledge as to the mode in which it is excited and brought into play in the living body, the chasm can for the present be supplied only by remote conjecture. The process which constitutes the ultimate stage of nutri- tion, or the actual incorporation of the new material with the solid substance of the body, of which it is to form a part, is involved in equal obscurity with that of secretion. nous ramifications, though on a much smaller scale, has been discovered by Jacobson, in the kidneys of most fishes and reptiles, and even in some birds. Vol. II. 32 ( 250 ) . CHAPTER XIII. ABSORPTION. Absohption is another function, related to nutrition, which deserves special notice. The principal object of this function is the removal of such materials as have been al- ready deposited, and have become either useless or injurious, and their conveyance into the general mass of circulating fluids; purposes which arc accomplished by a peculiar set of vessels, called the Lymphatics. These vessels contain a fluid, which, being transparent and colourless like water, has been denominated the lymjyh. The lymphatics are perfect- ly similar in their structure, and probably, also, in their mode of action, to the lacteals, which absorb the chyle from the intestinal cavity: they are found in all the classes of verte- brated animals, and pervade extensively every part of the body. Exceedingly minute at their origin, they unite to- gether as they proceed, forming larger and larger trunks, generally following the course of the veins, till they finally discharge their contents either into the thoracic duct, or into some of the large veins in the vicinity of the heart. Throughout their whole course, they are, like the lacteals, provided with numerous valves, which, when the vessel is distended with lymph, give it a resemblance to a string of beads, Fig. 37S."* In the lower animals, it appears that the veins are occasionally en- dowed with a power of absorption, similar to that possessed • In warm-blooded animals, the lymphatics are made to traverse, in some part of their coui-se, certain bodies of a compact stmcture, resembling- glands, and termed, accordingly, tlic lymphatic glands. One of these is represented ABSORPTION. 251 by the lymphatics. None of the invcrtcbrata, indeed, pos- sess lymphatics, and absorption must consequently be per- formed by the veins, when these latter vessels exist. The addition of the system of lymphatic vessels, as auxiliaries to the veins, may therefore be regarded as a refinement in or- ganization, peculiar to the higher classes of animals.* Professor JVIuller, of Bonn, has lately discovered that the frog, and several other amphil)ious animals, are provided with large receptacles for the lymph, situated immediately under the skin, and exhibiting distinct and regular pulsa- tions, like the heart. The use of these lymphatic hearts, as they may be called, is evidently to propel the lymph in its proper course along the lymphatic vessels. In the frog four of these organs have been found; the two posterior hearts being situated behind the joint of the hip, and the two anterior ones on each side of the transverse process of the third vertebra, and under the posterior extremity of the scapula. The pulsations of these lymphatic hearts do not correspond with those of the sanguiferous heart; nor do those of the right and left sides take place at the same times, but they often alternate in an irregular manner. Professor Muller has discovered similar organs in the toad, the sala- mander, and the green lizard, and thinks it probable that they exist in all the amphibia. f in Fig. 378. They correspond in structure, and probably also in their func- tions, to the mesenteric ghuids, through which, in the nianimaha, the lacteals pass, before reaching- the thoracic duct. It is chiefly in the mammalia, in- deed, that these glands are met with? for they are rare among birds, and still more so among fishes and reptiles. * Fohmann, who has made extensive researches on the absorbent vessels throughout all the classes of vertebrated animals, has found that they termi- nate extensively in the veins. Sec his work, entitled " Anatomischc Untcr- suchungcn uber die Verbindung dcr Saugadcrn mit den Vencn." t Phil. Trans, for 1833, p. 89. ( 252 ) CHAPTER XIV. NERVOUS POWER. The organs which are appropriated to the performance of the various functions conducive to nutrition, are generally designated the vital ors;ans^ in order to distinguish them from those which are subservient to sensation, voluntary motion, and the other functions of animal life. The slight- est reflection on the variety and complication of actions comprised under the former class of functions in the higher animals, will convince us that they must be the result of the combined operation of several different agents; but the principal source of mechanical force required by the vital organs, is still, as in all other cases, the muscular power. The coats of the stomach and of the intestinal tube contain a large proportion of muscular fibres, the contractions of which effect the intermixture and propulsion of the con- tents of these cavities, in the manner best calculated to fa- vour the chemical operations to which they are to be sub- jected, and to extract from them all the nourishment they may contain. In like manner, all the tubular vessels, which transmit fluids, are endowed w^th muscular powers adapted to the performance of that office. The heart is a strong hol- low muscle, with power adequate to propel the blood, with immense force, through the arterial and venous systems. The blood vessels, also, especially the minute, or capillary arteries, besides being elastic, are likewise endowed with muscular power, which contributes its share in forwarding the motion of the blood, and completing its circulation. The quantity of blood circulating in each part, the velocity of its motion, and the heat which it evolves, are regulated NERVOUS POWER. 253 in a great measure by the particular mode of action of the • blood vessels of that part. The quantity, and sometimes even the quality of the secretions, are dependent, in like manner, on the conditions of the circulation; and the action of the ducts, which convey the secreted fluids to their re- spective destinations, is also resolvable into the cflfccts of a muscular power. The immediate cause which, in these organs, excites the muscular fibre to contraction, may frequently be traced to the forcible stretching of its parts. This is the case in all hollow and tubular muscles, such as the stomach, the heart, and the blood vessels, when they are mechanically distended, beyond a certain degree, by the presence of contained fluids, or other substances. At other times, the chemical quality of their contents appears to be the immediate stimulus in- citing them to contraction. But numerous instances occur, in the higher orders of animals, in which these causes alone are inadequate to explain the phenomena of the vital func- tions. No mechanical hypothesis will suffice to account for the infinite diversity in the modes of action of the organs which perform these functions, or aflford any clew to the means by which they are made to co-operate, with such nicety of adjustment, in the production of the ultimate ef- fect. Still less will any theory, comprising only the agency of the muscular power, and the ordinary chemical affinities, enable us to explain how an irritating cause, applied at one part, shall produce its visible efiects on a distant organ; or in what way remote and apparently unconnected parts shall, as if by an invisible sympathy, be brought, at the same mo- ment, to act in concert, in the production of a common ef- fect. Yet such co-operation must, in innumerable cases, be absolutely indispensable to the perfect accomplishment of the vital functions of animals. Nature has not neglected objects so important to the suc- cess of her measures, but has provided, for the accomplish- ment of these purposes, a controlling faculty, residing in the nervous system, and denominated the 7iervous power. Ex- 254 THE VITAL FUNCTIONS. pcriments have shown that the due performance of the vital functions of digestion, of circulation, and of secretion, re- quires the presence of an ai^ency, derived from different parts of tlie hrain and spinal marrow, and rci^ulatino; the or- der and combinations of the actions of the organs which are to perform those functions. The same influence, for exam- ple, which increases the power of secretion in any particu- lar gland, is found to increase, at the same time, the action of those blood vessels which supply that gland with the ma- terials for secretion; and conversely, the increased action of the blood vessels is accompanied by an increased activity of the secreting organ. Experience also shows that when the influence of the brain and spinal marrow is intercepted, although the afflux of blood may, for a time, continue, yet the secretion ceases, and all the functions dependent upon secretion, such as digestion, cease likewise. Thus, the ner- vous power combines together different operations, adjusts their respective degrees, and regulates their succession, so as to ensure that perfect harmony which is essential to the at- tainment of the objects of the vital functions; and thus, not only the muscular power which resides in the vital organs, but also the organic affinities which produce secretion, and all those unknown causes which effect the nutrition, deve- lopment, and growth of each part, are placed under the con- trol of the nervous povver.^ Although we are entirely ignorant of the nature of the nervous power, we know that, when employed in the vital functions, it acts through the medium of a particular set of fibres, which form part of the nervous system, and are classed, therefore, among the nerves. The principal fflaments of this class of nerves compose what is called the sympathetic nerve, from its being regarded as the medium of extensive * As the functions of plants arc sufficiently simple to admit of being con- ducted witliout the aid of muscidar power, still less do tiiey require the as- sistance of tlie ncn'ous encrg-y: both of which properties are the peculiar at- tributes of animal vitality. We accordingly find no traces citlier of nervous or of musctdar fibres in any of the vegetable structures. NERVOUS POWER. 255 sympathies among the organs; but the wliole assemblage of these nerves is more commonly known by the name of the ganglionic system^ from tlie circumstance of their being connected with small masses of nervous substance, termed ganglia, which are placed in difTcrent parts of their course. Fig. 379, represents a ganglion (g,) through which the nerve (n,) consisting at its origin of a number of separate filaments (f,) is seen to pass, before it subdivides into branches (b.) The numerous communications and inter- changes of filaments, which subsequently take place at vari- ous parts, forming what is called ^plexus, are shown in Fig. 380: where four trunks (t, t,) divide into branches, which are again separated, and variously reunited in their course, like a ravelled skein of thread, before they proceed to their respective destinations. The ganglia are connected by nervous filaments with every part of the brain and spinal marrow, the great central organs of the nervous system; and they also send out innu- merable branches, to be distributed all over the body. All the parts receiving blood vessels, and more especially the organs of digestion, are abundantly supplied with ganglionic nerves; so that, by their intervention, all these parts have extensive connexions with the brain and spinal marrow, and also with one another. The ganglia are more particularly the points of union between nervous fibres coming from 256 THE VITAL FUNCTIONS. many different parts: they may be considered, therefore, as performing, with regard to the vital functions, an office ana- logous to that which the brain and spinal marrow perform witli regard to tlic otlicr nerves, or as being secondary cen- tres of nervous power. Thus, there are two important ob- jects for whicli tlic nerves belonging to the ganglionic sys- tem have ])ccn provided; fil'st, to serve as the channels through which the affections of one organ miglit be enabled to influence a distant organ; and secondly, to be the medium through which the powers of several parts might be com- bined and concentrated for effecting particular purposes, re- quiring such co-operation. Hence it is by means of the gan- glionic nerves that all the organs and all the functions are rendered efficient in the production of a common object, and are brought into one comprehensive and harmonious system of operation. The nervous power, the effects of which we are here con- sidering, should be carefully distinguished from that power which is an attribute of another portion of the nervous sys- tem, and which, being connected with sensation, volition, and other intellectual operations, has been denominated sen- so7'ial poicer.^ The functions of digestion, circulation, ab- sorption, secretion, and all those included under the class of nutrient or vital functions, are carried on in secret, are not necessarily, or even usually attended with sensation, and are wholly removed from the control of volition. Nature has not permitted processes, which are so important to the preservation of life, to be in any way interfered with by the will of the animal. We know that in ourselves they go on as well during sleep as when we are awake, an4 whether our attention be directed to them or not; and though occa- sionally influenced by strong emotions, and other affections of mind, they arc in general quite independent of every in- tellectual process. In the natural and healthy condition of • TlVis distinction has been most clearly pointed out, and illustrated by Dr. A. P. W. Philip. Sec his <* Experiiiicntal Inquiry into tlic Laws of the Vi- tal Functions." NERVOUS POWER. 257 the system all its internal operations proceed quietly, stea- dily, and constantly, whether the mind be absorbed in thought or wholly vacant. The kind of existence resulting from these functions alone, and to which our attention has hither- to been confined, must be regarded as the result of mere vegetative, rather than of animal life. It is time that we turn our views to the higher objects, and more curious field of inquiry, belonging to the latter. Vol. II. 33 ( 258 ) PART III. THE SENSORIAL FUNCTIONS. CHAPTER I. SEiVSATION. The system of mechanical and chemical functions which we have been occupied in reviewing, has been established only as a foundation for the endowment of those higher fa- culties which constitute tlie great objects of animal exist- ence. It is in the study of these final purposes that the scheme of nature, in the formation of the animal world, opens and displays itself in all its grandeur. The whole of the phenomena we have hitherto considered concur in one essential object, the maintenance of a simply vital existence. Endowed with these properties alone, the organized system would possess all that is absolutely necessary for the conti- nuance and support of mere vegetative life. The machine- ry provided for this purpose is perfect and complete in all its parts. To raise it to this perfection, not only has the Divine Architect employed all the properties and powers of matter, which science has yet revealed to man, but has also brought into play the higher and more mysterious energies of nature, and has made them to concur in the great work that was to be performed. On the organized fabric there has been conferred a vital force; with the powers of mechanism have been conjoined those of chemistry; and to these have been SENSATION. 259 superadded the still more subtle and potent agencies of ca- loric and of electricity: every resource has been employed, every refinement studied, every combination exhausted that could ensure the stability, and prolong the duration of the system, amidst the multifarious causes which continually menace it with destruction. It has been supplied vvitii am- ple means of repairing the accidents to which it is ordinarily exposed; it has been protected from the injurious influence of the surrounding elements, and fitted to resist for a length- ened period the inroads of disease, and the progress of decay. But can this, which is mere physical existence, be the sole end of life? Is there no farther purpose to be answered by structures so exquisitel)'' contrived, and so bountifully pro- vided with the means of maintaining an active existence, than the mere accumulation and cohesion of inert materials, dif- fering from the stones of the earth only in the more arti- ficial arrangement of their particles, and the more varied configuration of their texture? Is the growth of an animal to be ranked in the same class of phenomena as the concre- tion of a pebble, or the crystallization of a salt? Must we not ever associate the power of feeling with the idea of animal life? Can we divest ourselves of the persuasion that the movements of animals directed like our own, to obvious ends, proceed from voluntary acts, and imply the operation of an intellect, not wholly dissimilar in its spiritual es- sence from our own? In vain may Descartes and his fol- lowers labour to sustain their paradox, that brutes are only automata, — mere pieces of artificial mechanism, insensible either to pleasure or to pain, and incapable of internal af- fections, analogous to those of which we arc conscious in our- selves. Their sophistry will avail but little against the plain dictates of the understanding. To those who refuse to admit that enjoyment, which implies the powers of sensation, and of voluntary motion, is the great end of animal existence, the object of its creation must for ever remain a dark and im- penetrable mystery; by such minds must all farther inquiry 260 THE SENSORIAL FUNCTIONS. into final causes be at once abandoned as utterly vain and hopeless. But it surely requires no laboured refutation to overturn a system that violates every analogy by which our reasonings on these subjects must necessarily be guided; and no artificial logic or scholastic syllogisms will long prevail over the natural sentiment, which must ever guide our con- duct, that animals possess powers of feeling, and of sponta- neous action, and faculties appertaining to those of intellect. The functions of sensation, perception, and voluntary mo- tion re(juire the presence of an animal substance, which we find to be organized in a peculiar manner, and endowed with very remarkable properties. It is called the medullary sub- stance; and it composes the greater part of the texture of the brain, spinal marrow, and nerves; organs, of which the assemblage is known by the general name of the nervous system. Certain affections of particular portions of this me- dullary substance, generally occupying some central situa- tion, are, in a way that is totally inexplicable, connected with affections of the sentient and intelligent principle; a princi- ple which we cannot any otherwise conceive than as being distinct from matter; although we know that it is capa- ble of being affected by matter operating through the me- dium of this nervous substance, and that it is capable of reacting upon matter through the same medium. Of the truth of these propositions there exist abundant proofs; but as the evidence which establishes them will more con- veniently come under our notice at a subsequent period of our inquiry, I shall postpone their consideration; and pro- ceeding upon the assumption that this connexion exists, shall next inquire into the nature of the intervening steps in the process, of which sensation and perception are the results. Designating, then, by the name oi brain this primary and essential organ of sensation, or the organ whose physical af- fections are immediately attended by that change in the percipient being which we term sensation; let us first in- quire what scheme has been devised for enabling the brain to receive impressions from such external objects, as it is NERVOUS SYSTEM. 261 intended that this sentient being shall be capable of per- ceiving. As these objects can, in the first instance, make impressions only on the organs situated at the surface of the body, it is evidently necessary that some medium of com- munication should be provided between the external organ and the brain. Such a medium is found in the nerves^ which are white cords, consisting of bundles of threads or fila- ments of medullary matter, enveloped in sheaths of mem- brane, and extending continuously from the external organ to the brain, where they all terminate. It is also indispen- sably requisite that these notices of the presence of objects should be transmitted instantly to the brain; for the slightest delay would be attended with serious evil, and might even lead to fatal consequences. The nervous power, of which, in our review of the vital functions, we noticed some of the operations, is the agent employed by nature for this import- ant office of a rapid communication of impressions. The ve- locity with which the nerves subservient to sensation trans- mit the impressions they receive at one extremity, along their whole course, to their termination in the brain, exceeds all measurement, and can be compared only to that of elec- tricity passing along a conducting wire. It is evident, therefore, that the brain requires to be fur- nished with a great number of these nerves, which perform the office of conductors of the subtle influence in question; and that these nerves must extend from all those parts of the body which are to be rendered sensible, and must unite at their other extremities in that central organ. It is of espe- cial importance that the surface of the body, in particular, should communicate all the impressions received from the contact of external bodies, and that these impressions should produce the most distinct perceptions of touch. Hence, we find that the skin, and all those parts of it more particularly intended to be the organs of a delicate touch, are most abun- dantly supplied with nerves; each nerve, however, commu- nicating a sensation distinguishable from that of every other, so as to enable the mind to discriminate between them, and 262 THE SENSORIAL FUNCTIONS. refer them to their respective origins in different parts of the surface. It is also expedient that the internal organs of the body should have some sensibility; but it is better that this should be very limited in degree, since the occasions are few in which its exercise would be useful, and many in which it would be positively injurious: hence, the nerves of sensation are distributed in less abundance to these organs. It is not sufficient that the nerves of touch should com- municate tlie perccj)tions of the simple pressure or resistance of the bodies in contact with the skin: they should also fur- nish indications of other qualities in those bodies, of which it is important that the mind be apprized; such, for example, as warmth, or coldness. Whether these different kinds of impressions are all conveyed by the same nervous fibres, it is difficult, and, perhaps, impossible to determine. When these nerves are acted upon in a way which threat- ens to be injurious to the part impressed, or to the system at large, it is also their province to give warning of the im- pending evil, and to rouse the animal to such exertions as may avert it; and this is effected by the sensation of pain, which the nerves are commissioned to excite on all these oc- casions. They act tlie part of sentinels, placed at the out- posts, to give signals of alarm on the approach of danger. Sensibility to pain must then enter as a necessary consti- tuent among the animal functions; for, had this property been omitted, the animal system would have been but of short duration, exposed, as it m.ust necessarily be, to perpe- tual casualties of every kind. Lest any imputation should be attempted to be tlirown on the benevolent intentions of • the great Author and Designer of this beautiful and wondrous fabric, so expressly formed for varied and prolonged enjoy- ment, it should always be borne in mind that the occasional suffering, to which an animal is subjected from this law of its organization, is far more than counterbalanced by the consc(iuences arising from the capacities for pleasure, with which it has been beneficently ordained that the healthy ex- ercise of the functions shall be accompanied. Enjoyment NERVOUS SYSTEM. 263 appears universally to be the main end, the rule, the ordi- nary and natural condition: while pain is but the casualty, the exception, the necessary remedy, which is ever tending to a remoter good, in subordination to a higher law of crea- tion. It is a wise and bountiful provision of nature that each of the internal parts of the body has been endowed with a par- ticular sensibility to those impressions which, in the ordina- ry course, have a tendency to injure its structure; while it has, at the same time, been rendered nearly, if not complete- ]}'', insensible to those which are not injurious, or to which it is not likely to be exposed. Tendons and ligaments, for example, are insensible to many causes of mechanical irrita- tion, such as cutting, pricking, and even burning: but the moment they are violently stretched, that being the mode in which they are most liable to be injured, they instantly com- municate a feeling of acute pain. The bones, in like man- ner, scarcely ever communicate pain in the healtliy state, except from the application of a mechanical force which tends to fracture them. The system of nerves, comprising those which are de- signed to convey the impressions of touch, is universally present in all classes of animals; and among the lowest or- ders, they appear to constitute the sole medium of commu- nication with the external world. As we rise in the scale of animals, we find the faculties of perception extending to a wider range, and many qualities, depending on the chemi- cal action of bodies, are rendered sensible, more especially those which belong to the substances employed as food. Hence arises the sense of taste, which may be regarded as a new and more refined species of touch. This difference in the nature of the impressions to be conveyed, renders it ne- cessary that the structure of the nerves, or, at least, of those parts of the nerves which are to receive the impression, should be modified and adapted to this particular mode of action. As the sphere of perception is enlarged, it is made to 264 THE SENSORIAL FUNCTIONS. comprehend, not merely those objects which are actually in contact with the body, but also those which are at a distance, and of the existence and properties of which it is highly im- portant that the animal, of whose sensitive faculties we are examining the successive endowment, should be apprized. It is more especially necessary that he should acquire an accurate knowledge of the distances, situations and motions of surrounding objects. Nature has accordingly provided suitable organizations for vision, for hearing, and for the perception of odours; all of which senses establish extensive relations between him and the external world, and give him the command of various objects which are necessary to sup- ply his wants, or procure him gratification; and which also apprize him of danger while it is yet remote, and may be avoided. Endowed with the power of combining all these perceptions, he commences his career of sensitive and intel- lectual existence; and though he soon learns that he is de- pendent for most of his sensations on the changes which take place in the external world, he is also conscious of an internal power, which gives him some kind of control over many of those changes, and that he moves his limbs by his own voluntary act; movements which originally, and of themselves, appear, in most animals, to be productive of great enjoyment. To a person unused to reflection, the phenomena of sen- sation and perception may appear to require no elaborate investigation. That he may behold external objects, nothing more seems necessary than directing his eyes towards them. He feels as if the sight of those objects were a necessary consequence of the motion of his eye-balls, and he dreams not that there can be any thing marvellous in the function of the eye, or that any other organ is concerned in this sim- ple act of vision. If he wishes to ascertain the solidity of an object within his reach, he knows that he has but to stretch forth his hand, and to feel in what degree it resists the pressure he gives to it. No exertion even of this kind is required for hearing the voices of his companions, or be- SENSATION. 265 ing apprized, by the increasing loudness of the sound of falling waters, as he advances in a particular direction, that he is coming nearer and nearer to the cataract. Yet how much is really implied in all these apparently simple phe- nomena! Science has taught us that these perceptions of external objects, far from being direct or intuitiv^e, are only the final results of a long series of operations, produced by agents of a most subtle nature, which act by curious and complicated laws, upon a refined organization, disposed in particular situations in our bodies, and adjusted with admi- rable art to receive their impressions, to modify and com- bine them in a certain order, and to convey them in regular succession, and without confusion, to the immediate seat of sensation. Yet this process, complicated as it may appear, constitutes but the first stage of the entire function o{ pei^ception: for ere the mind can arrive at a distinct knowledge of the pre- sence and peculiar qualities of the external object which gives rise to the sensation, a long series of mental changes must intervene, and many intellectual operations must be performed. All these take place in such rapid succession, that even when we include the movement of the limb, which is consequent upon the perception, and which we naturally consider as part of the same continuous action, the whole appears to occupy but a single instant. Upon a careful ana- lysis of the phenomena, however, as I shall afterwards at- tempt to show, we find that no less than twelve distinguish- able kinds of changes, or rather processes, some of which imply many changes, must always intervene, in regular succession, between the action of tiie external object on the organ of sense, and the voluntary movement of the limb which it excites. The external agents, which are capable of affecting the different parts of the nervous system, so as to produce sen- sation, are of different kinds, and are governed by laws pe- culiar to themselves. The structure of the organs must, accordingly, be adapted, in each particular case, to receive Vol. II. 34 266 THE SENSORIAL FUNCTIONS. the impressions made l)y these agents, and must be modi- fied in exact conformity with the physical laws they obey. Thus, the structure of that portion of the nervous system which receives visual impressions, and which is termed the retina, must Ix; adapted to the action of light; and the eye, throusili which the ravs are made to pass before reaching the retina, must be constructed with strict reference to the laws of optics. The ear must, in like manner, be formed to receive delicate imj)ressions from those vibrations of the air which occasion sound. The extremities of the nerves, in these and other organs of the senses, are spread out into a delicate expansion of surface, having a softer and more uniform texture than the rest of the nerve, whereby they acquire a susceptibility of being affected by their own ap- propriate agents, and by no other. The function of each nerve of sense is determinate, and can be executed by no other part of the nervous system. These functions are not interchangeable, as is the case with many others in the ani- mal system. No nerve, but the optic nerve, and no part of that nerve, except the retina, is capable, however im- pressed, of giving rise to the sensation of light: no part of the nervous system, but the auditory nerve, can convey that of sound; and so of the rest. The credulity of the public has sometimes been imposed upon by persons who pretend- ed to see by means of their fingers: thus, at Liverpool, the celebrated Miss M'Avoy contrived for a long time to per- suade a great number of persons that she really possessed this miraculous power. Equally unworthy of credit are all the stories of persons, under the influence of animal mag- netism, hearing sounds addressed to the pit of the stomach, and reading the pages of a book applied to the skin over that organ. In almost every case the impression made upon the sen- tient extremity of the nerve which is appropriated to sen- sation, is not the direct effect of the external body, but re- sults from the agency of some intervening medium. There is always a portion of the organ of sense interposed hetwcen SENSATION. 267 the object and the nerve on which the impression is to be made. The object is never allowed to come into direct con- tact with the nerves; not even in the case of touch, where the organ is defended by the cuticle, through which the im- pression is made, and by which that impression is modified so as to produce the proper effect on the subjacent nerves. This observation applies with equal force to the organs of taste and of smell, the nerves of which are not only sheathed with cuticle, but defended from too violent an action by a secretion expressly for that purpose. In the senses of hearing and of vision, the changes which take place in the organs interposed between the external impressions and the nerves, are still more remarkable and important, and will be re- spectively the subjects of separate inquiries. The objects of these senses, as well as those of smell, being situated at a dis- tance, produce their first impressions by the aid of some me- dium exterior to our bodies, through which their influence extends: thus, the air is the usual medium through which both light and sound are conveyed to our organs. Hence, in order to understand the whole series of phenemena be- longing to sensation, regard must be had to tlie physical laws which regulate the transmission of these agents. We are now to consider these intermediate processes in the case of each of the senses. ( 2GS ) CHAPTER IT. TOUCH. I HAVE already had occasion to point out the structure of the integuments, considered in their mechanical ofTice of protectinp; the general frame of the body;* but we are not to view them in their relation to the sense of touch, of which they are the immediate organ. It will be recollected that the corutm forms the principal portion of the skin; that the cuticle composes the outermost layer; and that between these there occurs a thin layer of a substance, termed there/e mu- cosum. The corium is constructed of an intertexture of dense and tough fibres, through which a multitude of blood vessels and nerves are interspersed; but its external sur- face is more vascular than any other part, exhibiting a fine and delicate net-work of vessels, and it is this portion of the skin, termed by anatomists the vascular plexus, which is the most acutely sensible in every point: hence we may- infer that it contains the terminations of all the nervous fila- ments distributed to this organ, and which are here found to divide to an extreme degree of minuteness. When examined with the microscope, this external sur- face presents a great number of minute projecting filaments. ISIalpighi first discovered this structure in the foot of a pig; and gave these prominences the name of papillae. It is pro- bable that each of these papilla? contains a separate branch of the nerves of touch, the ultimate ramifications of which are spread over the surfiice: so that we may consider these papiliaN of which the assemblage has been termed the cor- pus 2)upillare, as the principal and immediate organ of • Vol. I. p. 90. TOUCH. 269 touch. This structure is particularly conspicuous on those parts of the skin which are more especially appropriated to this sense, such as the tips of the fingers, the tongue, and the lips: in other parts of the surface, which are endowed with less sensibility, the papillae are scarcely visible, even with the aid of the microscope. The surface of the corium is exquisitely sensible to all ir- ritations, whether proceeding from the contact of foreign bodies, or from the impression of atmospheric air. This ex- treme sensibility of the corium would be a source of con- stant torment, were it not defended by the cuticle, which is unprovided with either blood vessels or nerves, and is, therefore, wholly insensible. For the same reason, also, it is little liable to change, and is thus, in both respects, admi- rably calculated to afford protection to the fmely organized corium. Although the cuticle exhibits no traces of vascularity, it is by no means to be regarded as a dead or inorganic sub- stance, like the shells of the mollusca. That it is still part of the living system is proved by the changes it frequently undergoes, both in the natural and the diseased conditions of the body. It is perpetually, though slowly, undergoing de- cay and renovation; its external surface drying off in mi- nute scales, and in some animals peeling off in large por- tions. When any part of the human skin is scraped with a knife, a gray dust is detached from it, which is found to con- sist of minute scales. By repeated friction, or pressure of any part of the skin, the cuticle soon acquires an increase of thickness and of hardness; this is observable in the soles of the feet, and palms of the hands, and in the fingers of those who make much use of them in laborious work. But this greater thick- ness in the parts designed by nature to suffer considerable pressure, is not entirely the eflect of education; for the cuti- cle, which exists before birth, is found even then to be much thicker on the soles of the feet, and palms of the hands, than on other parts. This example of provident care in origi- 270 THE SENSORIAL FUNCTIONS. nally adjusting the structures of parts to the circumstances in which they are to be placed at an after period, would of itself, were it a solitary instance, be well fitted to call forth our admiration. liut the proofs of design in the adaptation of organs to their respective purposes multiply upon us in such profusion, as we study in detail each department of the animal economy, that we are apt to overlook individual in- stances, unless they are particularly brought before our no- tice. IIow often have we witnessed and profited by the rapid renewal of the cuticle, when by any accident it has been destroyed, without adverting to the nature of the pro- cess which it implies; or reflected that the vessels of the skin must, on all these occasions, supply the materials, out of which the new cuticle is to be formed, must elfect their combination in the requisite proportions, and must deposite them in the precise situations in which they are wanted! Different animals present remarkable differences in the thickness and texture of the cuticle, according to the element they are destined to inhabit, and the situations in which they are most frequently placed. Provision is in many cases made for preserving the cuticle from the injury it would receive from the long continued action of the air or water; for it is apt to become rigid; and to peel off, from ex- posure to a very dry atmosphere; and the constant action of water, on the contrary, renders it too soft and spongy. In order to guard against both these effects, the skin has been furnished, in various parts of its surface, with a secreting apparatus, which pours out unctuous or mucilaginous fluids: the oily secretions being more particularly employed as a defence against the action of the air, and the mucilaginous fluids as a protection against that of water. The conditions on which the perfection of the sense of touch depends arc, first, an abundant provision of soft pa- pillae supplied with numerous nerves; secondly, a certain degree of fineness in the cuticle; thirdly, a soft cushion of cellular substance beneath the skin; fourthly, a hard resist- ing basis, such as that which is provided in the nails of the TOUCH. 271 human fingers; and lastly, it is requisite that the organ be so constructed as to be capable of being readily applied, in a variety of directions, to the unequal surfaces of bodies; for the closer the contact, the more accurate will be the percep- tions conveyed. In forming an estimate of the degree of perfection in which this sense is exercised in any particular animal, we must, accordingly, take into account the mobili- ty, the capability of flexion, and the figure of the parts em- ployed as organs of touch. As touch is the most important of all the senses, in^much as it is the foundation of all our knowledge of the material world, so its relative degrees of perfection establish marked differences in the intellectual sagacity of the several tribes, and have a considerable influence on the assignment of their proper station in the scale of animals. Although the power of receiving obscure impressions from the contact of external bodies, and of perceiving varia- tions of temperature, is probably possessed by all animals, a small number only are provided with organs specially ap- propriated for conveying the more delicate sensations of touch. The greater part of the surface of the body in the testaceous Mollusca is protected by a hard and insensible covering of shell. The integuments of Insects, especially those of the Coleoptera, are in general too rigid to receive any fine impressions from the bodies which may come in contact w^ith them; and the same observation applies, with even greater force, to the Crustacea. The scales of Fishes, and of Reptiles, the solid incasements of the Chelonia, the plumage of Birds, the dense coating of the Armadillo, the thick hides of the Rhinoceros, and other Pachydermata, arc evidently incompatible with any delicacy of touch. This nicer faculty of discrimination can be enjoyed only by ani- mals having a soft and flexible integument, such as all the naked Zoophytes, Worms, and Mollusca, among the lower orders, and Serpents, among the higher. The flexibility of the body or limbs is another condition which is extremely necessary towards procuring extensive and correct notions 272 THE SENSORIAL FUNCTIONS. of the relative positions of external objects. It is essential therefore that those instruments which are more particularly intended as organs of toucii. should possess this property. It will not be necessary to enter into a minute descri])tion of these organs, because they have, for the most part, been already noticed as instruments of prehension; for the sense of touch is in general exercised more particularly by the same ])arts which perform this latter function. Thus the tentacula of the various tribes of Polypi, of Actiniae, and of Annelida, are organs both of prehension and of touch. The tubular feet of the Asterias and Echinus arc, in like man- ner, subservient botii to tlie sense of touch, and to the fa- culty of progressive motion. Tl^e feet of Insects and of Crustacea are well calculated, indeed, by their jointed struc- ture, for being applied to the surfaces, and to different sides of bodies; but they are scarcely ever employed in this capa- city; being superseded by the palpi, which are situated near the mouth. When insects are walking, the palpi are inces- santly applied to the surface on which they advance, as if these organs were especially employed to feel their way. There can be little doubt, however, that, in most insects, TOUCH. 273. the principal organs of touch are the sflnlcnnx^ also deno- minated, from their supposed oilice, \\\q. feelers.^ Some idea of the great variety in the forms of the anten- nae of insects may be obtained from the specimens deh'ne- ated in Fig. 3S1, which shows a few of the most remarka- ble.f The universality of these organs among every species of this extensive class of animals, their great flexibility, arising from their jointed structure,! their incessant motion when the insect is walking, and their constant employment in exa- mining the surfaces of all the bodies with which they come in contact, sufficiently point them out as instruments of a very delicate sense of touch. Organs of this kind were par- ticularly necessary to insects, since the horny nature of the • The German name for ^\em, fiihlh'dmer, or the feeling horns, is founded on the same notion. i f In this figure, A represents the form of antennae, technically denomi- nated Antenna capitulo uncinato, as exemplified in the Pausus. B . is the A . piloso-verticillata, as in the Psychoda ocellaris. C . . A . biclavata, (Clavlger longicornis.) D . . A . triang-ularis, (Lophosia.) £ . . A . clavata, (Masaris.) F . . A . capit. lamellato, (Melolontha mas.) G . . A . capit. fissile, (Jphodlusfossor.) H . . A . fusiformis, {Zygaena.) I . . A . capitata, (Ascalaphus.) K . . A . furcata, (Nepa.) L. . A . bipectinata, {Bombyx.) M . . A . irregularis, {Agaon paradoxum.) N . . A . cordata, (Diaperis boleti.) O . . A . bipectinata, (Ctenophora.) P . . A . palmataj (Nepa cinerea.) Q . . A . ensiformis, (TVuxalis.) R . . A . setacea, (Cerambyx.) i The number of segments into which these organs are divided is often very great. In the Grylloialpa, or mole cricket, it amounts to above 100. (Kidd, Phil. Trans, for 1825, p. 211.) This insect has, besides the antennae on the head, two posterior or caudal antennae, which are not jointed, except- ing at their very commencement. These are extremely sensible, and sei-ve, probably, to give the animal notice of the approach of any annoyance from behind, lb. p. 216. Vol. II. 35 274 THE SENSORIAL FUNCTIONS. intcoruments of the greater number, precludes them from im- parting any accurate perceptions of touch. It has been conjectured that the antennae of insects are the organs of other senses besides that of touch. If an insect be deprived of its antennas, it either remains motionless, or if it attempt to walk or fly, appears bewildered, and moves with- out any apparent object. Ilubcr found that bees are ena- bled, by feeling with their antennae, to execute their various works in the interior of the hive, where, of course, they can have no assistance from light. They employ these organs perpetually while building the combs, pouring honey into the magazines, ascertaining the presence of the queen, and feeding and tending the larvae. The same naturalist ob- serves, also, that it is principally by means of the antennae that these social insects communicate to one another their impressions and their wants. The different modes in which ants, when they happen to meet during their excursions, mutually touch one another with their antennae, appears to constitute a kind of natural language understood by the whole tribe. This contact of the antennse evidently admits of a great variety of modifica- tions, and seems capable of supplying all the kinds of in- formation which these insects have occasion to impart. It would seem impossible, indeed, for all the individuals com- posing these extensive societies to co-operate effectually in the execution of many works, calculated for the general be- nefit of the community, unless some such means of commu- nication existed. There is no evidence that sound is the medium of this intercourse; for none, audible to us at least, was ever known to be emitted by these insects. Their mode of conversing together appears to be simply by touching one another in different ways with the antennae. Iluber's ob- servations on this subject are exceedingly curious.* He re- marks that the signal denoting the apprehension of danger, is made by the ant striking its head against the corselet of • See his '*Rcchcrches sur les mccursdcs foiirmis indigenes." TOUCH. 275 every ant which it chances to meet. Each ant, on receiving this intimation, immediately sets about repeating the same signal to the next ant which comes in its way; and the alarm is thus disseminated with astonishing rapidity tliroughoutthc whole society. Sentinels are at all times stationed on the outside of the nests, for the purpose of apprizing the inha- bitants of any danger that may be at hand. On the attack of an enemy, these guardians quickly enter into the nest, and spread the intelligence on every side: the whole swarm is soon in motion, and while the greater number of ants rush forwards with desperate fury to repel the attack, others who are intrusted with the office of guarding the eggs and the larvae, hasten to remove their charge to places of greater se- curity. When the queen bee is forcibly taken away from the hive, the bees which are near her at the time do not soon appear sensible of her absence, and the labours of the hive are car- ried on as usual. It is seldom before the lapse of an hour, .that the working-bees begin to manifest any symptoms of uneasiness: they are then observed to quit the larvae which they had been feeding, and to run about in great agitation, to and fro, near the cell which the queen had occupied be- fore her abduction. They then move over a wider circle, and on meeting with such of their companions as are not aware of the disaster, communicate the intelligence by cross- ing their antennae, and striking lightly with them. The bees which receive the news become, in their turn, agitated, and conveying this feeling wherever they go, the alarm is soon participated by all the inhabitants of the hive. All rush forwards with tumultuous precipitation, eagerly seek- ing their lost queen; but after continuing the search for some hours, and finding it to be fruitless, they appear resigned to their misfortune; the noisy hubbub subsides, and the bees quietly resume their labours. A bee, deprived of its antennae, immediately becomes dull and listless: it desists from its usual labours, remains at tlie bottom of the hive, seems attracted only by the light, and 276 THE SENSORIAL FUNCTIONS. takes the first opportunity of quitting the hive, never more to return. A queen bee, thus mutilated, ran about, without apparent object, as if in a state of delirium, and was incapa- ble of directing her trunk with precision to the food which was offered to her. Latreille relates that, having deprived some labouring ants of their antennae, he replaced them near the nest; but they wandered in all directions, as if bewil- dered, and unconscious of what they were doing. Some of their companions were seen to notice their distress; and, ap- proaching them with apparent compassion, applied their tongues to the wounds of the sufferers, and anointed them with their saliva. This trait of sensibility was repeatedly witnessed by Latreille, while watching their movements with a magnifying glass. The Arachnida, from the mobility of their limbs, and the thinness of their cutaneous investment, have a very delicate sense of touch. Among the Mollusca, it is only the higher orders of Cephalopoda that enjoy this sense in any con- siderable degree, and they are enabled to exercise it by means of their long and flexible tentacula. Many bivalve mollusca have, indeed, a set of tentacula placed near the mouth, but they are short, and of little power. It is pro- bable that the foot may also be employed by these animals as an organ of touch. Fishes are, in general, very ill-constructed for the exer- cise of this sense; and their fins are used for no other pur- poses than those of progressive motion. That part of the surface which possesses the most acute feeling is the under- side, where the integuments are the thinnest. The chief seat of the sense of touch, however, is the lip, or end of the snout, which is largely supplied with nerves; and perhaps the cirrhi^ or little vermiform processes called bm^bcls, which in some species are appended to the mouth, may be subser- vient to this sense. * These processes in the Silurus glanis are moved by particular muscles. ♦ These kind of tentacula are remarkable for their length and mobility in the Lnphhvf piscatorius, or Angler; and it is said that they are employed by TOUCH. 277 Serpents, from the great flexibility of their spine, are ca- pable of grasping and twining round objects of almost any shape, and of taking, as it were, their exact measure. This conformation must be exceedingly favourable to the acqui- sition of correct perceptions of touch. As it is these per- ceptions, which, as we shall afterwards find, lay the founda- tion of the most perfect acquaintance with the tangible pro- perties of surrounding bodies, we may presume that this power contributes much to the sagacity possessed by these animals. It has been said of Serpents, that their whole body is a hand, conferring some of the advantages of that instru- ment. Hellman has shown that the slender bifurcated tongue of these animals is used for the purposes of touch.* In those species of Lizards which are enabled by the structure of their feet to clasp the branches of trees, as the Gecko and the Chameleon, and whose tails also are prehen- sile, we must, for the same reason, presume that the sense of touch exists in a more considerable degree than in other saurian reptiles, which do not possess this advantage The toes of Birds are also well calculated to perform the office of organs of touch, from the number of their articulations and their divergent position, and from the papillae with which their skin abounds, accompanied as they are with a large supply of nerves. Those birds, which, like the Parrot, em- ploy the feet as organs of prehension, probably enjoy a greater development of this sense. The skin which covers the bills of aquatic birds is supplied by very large nerves, and consequently possesses great sensibility. This struc- ture enables them to find their food, which is concealed in the mud, by the exercise of the sense of touch residing in that organ. A similar structure, probably serving a similar purpose, is found in the Ornithorhyncus. Among Mammalia, we find the seat of this sense frequent- ly transferred to the lips, and extremity of the nostrils, and the fish, while lurking" in ambush, as a decoy to other fishes, which they en- tice by their resemblance to worms. * tiuoted by Blumenbach. 278 THE SENSORIAL FUNCTIONS. many have the nose prolonged and flexible, apparently with this view. This is the case with the Shrew and the Mole, which are burrowing animals, and still more remarkably with the Pachydermata, where this greater sensibility of the parts about the face seems to have been bestowed as some compensation for the general obtuseness of feeling resulting from the tliickness of the hide which covers the rest of the body. Thus, the Rhinoceros has a soft, hook-shaped exten- sion of the upper lip, which is always kept moist, in order to preserve its sensibility as an organ of touch. The Hog has the end of the nose also constructed for feeling; though it is not so well calculated for distinguishing the form of ob- jects, as where the organ is prolonged in the form of a snout, which it is in the Tapir, and in a still higher degree in the admirably constructed proboscis of the Elephant, which, as an organ, both of prehension and of touch, forms the nearest approach to the perfect structure of the human hand. The Lion, Tiger, Cat, and other animals of the genus Fe- lls, have whiskers, endowed at their roots with a particular sensibility, from being largely supplied with nerves. The same is the case with the whiskers of the Seal. The prehensile tails of the American monkeys are doubt- less fitted to convey accurate perceptions of touch, as well as the feet and hands: as may be inferred from the great size of the nervous papillae, and the thinness of the cuticle of those parts. The sense of touch attains its greatest degree of excellence in the human hand, in which it is associated with the most perfect of all instruments of prehension. But as the struc- ture and functions of this organ are the exclusive subjects of another of these treatises, I shall refrain from any farther remarks respecting them. ( 279 ) CHAPTER III. TASTE. The senses of taste and smell are intended to convey im- pressions resulting from the chemical qualities of bodies, the one in the fluid, the other in the gaseous state.* There is a considerable analogy between the sensations derived from these two senses. The organ of taste is the surface of the tongue, the skin of which is furnished with a large propor- tion of blood vessels and nerves. The vascular plexus im- mediately covering the corium is here very visible, and forms a distinct layer, through which a great number of papillse pass, and project from the surface, covered with a thin cuti- cle, like the pile of velvet. In the fore part of the human tongue these papillae are visible even to the naked eye, and especially in certain morbid conditions of the organ.t They are of different kinds; but it is only those which are of a co- nical shape that are the seat of taste. If these papilla be touched with a fluid, which has a strong taste, such as vine- gar, applied by means of a camel-hair pencil, they will be seen to become elongated by the action of the stimulus, an effect which probably always accompanies the perception of taste. • Bellini contended that the different flavors of saline bodies were owing- to the peculiar figures of their crystalline particles. It is strang-e tliat Dumas should have thouglit it worth while seriously to combat this exti-avag-ant liy- pothesis, by a laboured refutation. j- This is particularly the case in scarlatina, In the early stag-e of wliich dis- ease they are elongated, and become of a bright red colour, from their mi- nute blood vessels being- distended with blood. As the fever subsides, tbe points of the papilla; collapse, and acquire a brown hue, giving- rise to tlie appearance known by the name of the strawberry tongue. 280 THE SENSORIAL FUNCTIONS. The primary use of this sense, the organ of which is plaped at the entrance of the alimentary canal, is evidently to guide animals in the choice of their food, and to warn them of the introduction of a noxious substance into the sto- mach. With respect to the human species, this use has been, in the present state of society, superseded by many acquired tastes, which have supplanted those originally given to us by nature: but in tlie inferior animals it still retains its pri- mitive oflice, and is a sense of great importance to the safety and welfare of the individual, from its operation being coin- cident with those of natural instincts. If, as it is said these instincts are still met with among men in a savage state, they are soon weakened or effaced by civilization. The tongue, in all the inferior classes of vertebrated ani- mals, namely, Fishes, Reptiles, and Birds, is scarcely ever constructed with a view to the reception of delicate impres- sions of taste; being generally covered with a thick, and often horny cuticle; and being, besides, scarcely ever employed in mastication. This is the case, also, with a large propor- tion of quadrupeds, which swallow their food entire, and which cannot, therefore, be supposed to have the sense of taste much developed. Insects which are provided with a tongue or a proboscis may be conceived to exercise the sense of taste by means of these organs. But many insects possess, besides these, a pair of short feelers, placed behind the true antennae; and it has been observed that, while the insect is taking food, these organs are in incessant motion, and are continually employed in touching and examining the food, before it is introduced into the mouth: hence, some entomologists have concluded that they are organs of taste. But it must be obvious that in this, as in every other instance in which our researches extend to beings of such minute dimensions, and which oc- cupy a station, in the order of sensitive existence, so remote from ourselves, we are wandering into regions where the only light that is afforded us must be borrowed from vague and fanciful analogies, or created by the force of a vivid and deceptive imagination. ( 281 ) CHAPTER IV. SMELL. Animal life being equally dependent upon the salubrious qualities of the air respired, as of the food received, a sense has been provided for discriminating the nature of the for- mer, as well as of the latter. As the organs of taste are placed at the entrance of the alimentary canal, so those of smell usually occupy the beginning of the passages for res- piration, where a distinct nerve, named the olfactory, ap- propriated to this office, is distributed. The sense of smell is generally of greater importance to the lower animals than that of taste; and the sphere of its perceptions is in them vastly more extended than in man. The agents, which give rise to the sensations of smell, are certain effluvia, or particles of extreme tenuity, which are disseminated very quickl}'- through a great extent of atmo- spheric air. It is exceedingly difficult to conceive how mat- ter so extremely rare and subtle as that which composes these odorous effluvia can retain the power of producing any sensible impression on the animal organs: for its tenuity is so extraordinary as to exceed all human comprehension. The most copious exhalations from a variety of odoriferous substances, such as musk, valerian, or asafoetida, will be continually emanating for years, without any perceptible loss of weight in the body which supplies them. It is well known that if a small quantity of musk be enclosed for a few hours in a gold box, and then taken out, and the box cleaned as carefully as possible with soap and water, that box will retain the odour of musk for many years; and yet Vol. II. 36 282 THE SENSORIAL FUNCTIONS. the nicest balance will not show the smallest increase of its wei*'ht from this impregrnation. No facts in natural philo- sophy afford more striking illustrations of the astonishing^ and indeed inconceivable divisibility of matter, than those relatinc; to odorous effluvia. It would apj)ear that most animal and vegetable bodies are continually emitting these subtle effluvia, of which our own organs are not sufficiently delicate to apprize us, unless wlicn they arc much concentrated, but which are readily perceived and distinguished by the lower animals; as may be inferred from their actions. A dog is known to follow its master by the scent alone, through the avenues and turn- ings of a crowded city, accurately distinguishing his track amidst thousands of others. The utility of the sense of smell is not confined to that of being a check upon the respiration of noxious gases; for it is also a powerful auxiliary to the sense of taste, which, of itself, and without the aid of smell, would be very vague in its indications and limited in its range. What may have been its extent and delicacy in man, while he existed in a savage state, we have scarcely any means of determining; but in the present artificial condition of the race, resulting from civilization and the habitual cultivation of other sources of knowledge, there is less necessity for attending to its per- ceptions, and our sensibility to odours may perhaps have di- minished in the same proportion. It is asserted both by Soemmerring and Blumenbach that the organ of sm.ell is smaller in Europeans, and other civilized races of mankind, than in those nations of Africa or America, which are but little removed from a savage state: it is certainly much less developed in man than in most quadrupeds. To the carni- vorous tribes, especially, it is highly useful in enabling them to discover their natural food at great distances. The cavity of the nostrils, in all terrestrial vertebrated animals is divided into two by a vertical partition; and the whole of its internal surface is lined by a soft membrane, SMELL. 283 called the Schneiderian membrane,* which is constantly- kept moist, is supplied with numerous hlood vessels, and upon which are spread the ultimate ramifications of the ol- factory nerves. The relative magnitude of these nerves is much greater in carnivorous quadrupeds than in those which subsist on vegetable ^ood. In quadrupeds as well as in man, these nerves are not collected into a single trunk in tlicir course towards the brain, but compose a great number of fila- ments, which pass separately through minute perforations in a plate of bone, (called the ethm,oid boiie) before they en- ter into the cavity of the skull, and join that part of the ce- rebral substance with which they are ultimately connected. The surface of the membrane which receives the impres- sions from odorous effluvia, is considerably increased by several thin plates of bone, which project into the cavity of the nostrils, and arc called the turbinated hones. These are delineated at t, t, in Fig. 3S2, as they appear in a vertical and longitudinal section of the cavity of the human nostril, where they are seen covered by the Schneiderian mem- * It has been so named in lionour of Schneider, the first anatomist who gave an accurate description of this membrane. 284 THE SENSORIAL FUNCTIONS. brane."* A transverse and vertical section of these parts is given in Fig. 3S3.t The turbinated bones are curionsly folded, and often convoluted in a spiral form, with the evi- dent design of obtaining as great an extent of surface as pos- sible within the confined space of the nasal cavity. This tur- binated, or spiral shape, chiefly cliaracterizes these bones among herbivorous quadrupeds: in the horse, for example, the turbinated bones are of a large diameter, and extend the whole length of the prolonged nostrils. Their structure is exceedingly intricate; for while they retain, externally, the general shape of an oblong spiral shell, they arc pierced on all their internal sides with numerous perforations, through • This fig-ure sliows the branches of the olfactory nerve (o,) passing through tlie thin cribriform plate of the ethmoid bone, and distributed over that membrane. Several of the cells, which open into the cavity, are also vSecn; such as the large sphenoidal sinus (s,) the frontal sinus (f,) and one of the ethmoidal cells (c.) s, is the nasal bone; p, the palate; and e, the mouth of the Eustachian tube, which leads to the ear. ■|- In this figure, s, is the septum, or partition of the nostrils, on each side of which are seen the sections of the turbinated bones projecting into the cavity; the ethmoid cells (c,) situated between the orbits (o;) and the An- trum maxillare (a,) which is another large cavity communicating with the < iiostrils. SMELL. 285 which the membrane, together with the fine branches of the nerves, passes freely from one side to the other. The ca- vities resulting from the convolutions are inlersccted by un- perforated partitions of extraordinary tenuity, serving both to support the arches of bone, and to furnish a still greater surface for the extension of the olfactory membrane. In the Sheep, the Goat, and the Deer, the structure is very si- milar to that just described; but the convolutions are double, with an intermediate partition, so as to resemble in its trans- verse section the capital of an Ionic column.* They are shown at (t) in the transverse section of the nostrils of a sheep in Fig. 384. In carnivorous quadrupeds the structure of these bones is still more intricate, and is calculated to afford a far more ex- tensive surface for the distribution of the olfactory nerve. In the Seal this conformation is most fully developed, and the bony plates are here not turbinated, but ramified, as * In a species of Antelope described by Mr. Hodgson, cavities exist, si- tuated immediately behind the ordinary nostrils, and communicating- with them. The accessory nostrils are conjectured to be useful to this excceding-- ly fleet animal by facilitating its breathing, while it is exerting its utmost speed; for the expansion of the nostrils opens also these posterior cavities, the sides of which, being elastic, remain dilated. Journal of the Asiatic So- ciety, Feb. 1832, p. 59. 286 THE SENSORIAL FUNCTIONS. shown at t in Fig. 385. Eight or more principal branches arise from the main trunk; and each of these is afterwards divided- and subdivided to an extreme degree of minuteness, so as to form, in all, many luindred plates. The olfactory- membrane, with all its nerves, is closely applied to every plate in this vast assemblage, as well as to the main trunk, and to the internal surface of the surrounding cavity: so that its extent cannot be less than 120 square inches in each nos- tril. An organ of such exquisite sensibility requires an ex- traordinary provision for securing it against injury, by the power of voluntarily excluding noxious vapours; and nature has supplied a mechanism for this express purpose, enabling the animal to close, at pleasure, the orifice of the nostril. The boo;, which, in its natural state, subsists wholly on ve- getable food, resembles herbivorous tribes in the external form and relative magnitude of the turbinated bones; but they are more simple in their structure, being formed of sin- gle, and slightly convoluted plates, without partitions or per- forations. In this respect, they approach to the human structure, which is even less complicated, and indicates a greater affinity with vegetable than with animal feeders. Man, indeed, distinguishes more accurately vegetable odours than those proceeding from animal substances; while the reverse is observed with regard to quadrupeds whose habits are decidedly carnivorous. A dog, for instance, is regard- less of the odour of a rose or violet; and, probably, as he derives from them no pleasure, is unable to discriminate the one from the other. Predacious animals, as Sir B. Harwood observes, require both larger olfactory nerves, and a more extensive surface for their distribution, than the vegetable eaters. The food of the latter is generally near at hand; and as they have occasion only to select the wholesome from the noxious plants, their olfactory organs are constructed for the purpose of arresting the effluvia of odorous substances immediately as they arise. The former are often under the necessity of discovering the lurking places of their prey at a considerable distance, and are, therefore, more sensible to SMELL. 287 the weak impressions of particles widely diffused through thl5 surrounding medium, or slightly adhering to those bo- dies, with which the ohjcct of their pursuit may have come into contact. The olfactory bones of birds are constructed very much on the model of the spiral bones of herbivorous quadrupeds, and vary but little in the different species. Fig. 386 cxhi- 386 bits their appearance in the Turkey: but the size of the ol- factory nerves of birds of prey greatly exceeds that of the •same nerves in granivorous birds. In the latter, indeed, they are exceedingly small^ and as the natural food of that tribe has but little odour, we find that they are easily de- ceived by any thing which bears a resemblance to it. Sir Busick Harwood relates that some poultry, which were usually fed with a mixture of barley meal and water, were found to have swallowed, by mistake, nearly the whole con- tents of a pot of white paint. Two of the fowls died, and two others became paralytic. The crops of the latter were opened, and considerably more than a pound of the poison- ous composition taken from each; and the crops, either na- turally, or from the sedative effects of the paint, appeared to have so little sensibility that, after the wounds were sewed up, both the fowls eventually recovered. The olfactory nerves are conspicuous in the Duck, both from their size and mode of distribution. They are seen 288 THE SENSORIAL FUNCTIONS. in Fig. 387, passing out through the orbit of the eye (o) in two large branches, an upper one (u,) and a lower one (l,) the ramifications of which are spread over the mandibles, both wathin and without. For the protection of the highly sensible extremity of the beak against the injurious impres- sions of hard bodies, a horny process (p,) similar, both in form and office, to the human nail, is attached to it, and its edtJ-es guarded by a narrow border of the same horny mate- rial; these receive a first, and fainter impression, and admo- nish the animal of approaching danger; if none occur, the matter is then submitted to the immediate scrutiny of the n.erves themselves, and is swallowed or rejected according to their indication.* It has been generally asserted that Vultures, and other birds of prey, are gifted with a highly acute sense of smell; and that they can discover by means of it the carcass of a dead animal at great distances: but it appears to be now suf- ficiently established by the observations and experiments of Mr. Audubon, that these birds in reality possess the sense of smell in a degree very inferior to carnivorous quadru- peds; and that so far from guiding them to their prey from a distance, it affords them no indication of its presence, even when close at hand. The following experiments appear to be perfectly conclusive on this subject. Having procured the skin of a deer, Mr. Audubon stuffed it full of hay; and after the whole had become perfectly dry and hard, he placed it in the middle of an open field, laying it down on * Such is tlie account given by Sir Busick Harwood, in his *' System of Comparative Anatomy and Physiology," p. 26. SMELL. 289 its back, in the attitude of a dead animal. In the course of a few minutes afterwards, he observed a vulture flying to- wards it, and alighting near it. Quite unsuspicious of the deception, the bird immediately proceeded to attack it, as usu^l, in the most vulnerable points. Failing in his object, he next, with much exertion, tore open the scams of the skin, where it had been stitched together, and appeared earnestly intent on getting at the flesh, which he expected to find within, and of the absence of which, not one of his senses was able to inform him. Finding that his efibrts, which were long reiterated, led to no other result than the pulling out large quantities of hay, he at length, though with evident reluctance, gave up the attempt, and took flight in pursuit of other game to which he was led by the sight alone, and which he was not long in discovering and securing. Another experiment, the converse of the first, was next tried. A large dead hog was concealed in a narrow and winding ravine, about twenty feet deeper than the surface of the earth around it, and filled with briers and high cane. This was done in the month of July, in a tropical climate, where putrefaction takes place with great rapidity. Yet, although many vultures were seen, from time to time, sail- ing in all directions over the spot where the putrid carcass was lying, covered only with twigs of cane, none ever dis- covered it; but in the mean while, several dogs had found their way to it, and had devoured large quantities of the flesh. In another set of experiments, it was found that young vultures, enclosed in a cage, never exhibited any tokens of their perceiving food, when it could not be seen by them, however near to them it was brought.* It has been doubted whether fishes, and other aquatic ani- mals, possess the sense of smell; in some of the whale tribe, ♦Edinburgh New Journal of Science, ii. 172. The accuracy of these re- sults, which had been contested by Mr. Waterton, is fully established by the recent observations and experiments of M. Bachman, which are detailed in Loudon's Magazine of Nat. Hist. vii. 167. Vol. II. 37 290 THE SENSORIAL FUNCTIONS. indeed, neither the organ of smell nor the olfactory nerves are found.* Some physiologists have gone the length of de- nvin«j; the capability of water to serve as the vehicle of odo- rous effluvia. But as water is known to contain a large quantity of air, which acts upon the organs of respiration, it is easy to conceive that it may also convey to the nostrils the peculiar agents which are calculated to excite perceptions of smell. Fishes arc, in fact, observed to be attracted from great distances by the effluvia of substances thrown into the water; and they are well known to have a strong predilec- tion for all highly odoriferous substances. Baits used by ansjlers are rendered more attractive by being impregnated with volatile oils, or other substances having a powerful scent, such as asafoetida, camphor, and musk. Mr. T. Bellt has discovered in the Crocodile and Alligator, a gland, which secretes an unctuous matter, of a strong, musky odour, si- tuated beneath the lower jaw, on each side. The external orifice of this gland is a small slit, a little within the lower edo-e of the jaw; and the sac, or cavity containing the odo- riferous substance, is surrounded by two delicate bands of muscular fibres, apparently provided for the purpose of first bringing the gland into a proper position, and then, by com- pressing it, discharging its contents. Mr. Bell conceives that the use of this secretion is to act as a bait for attracting fish towards the sides of the mouth, where they can be rea- dily seized in the mode usual to the alligator, which is that of snapping sideways at the objects he aims at devouring. The organs of smell in Fishes are situated in cavities, placed one on each side, in front of the head: they are mere- ly blind sacs, having no communication with the mouth or throat, and, indeed, no other outlet but the external open- ings, which are generally two to each sac. The principal entrance is furnished with a valve, formed by a moveable membrane, appearing like a partition dividing each nostril * Home; Lectures on Comparative Anatomy, i. 17. j IMiil. Trans, for 1827, p. 132. SMELL. 291 into two cavities, and serving the purpose of preventing the introduction of any foreign body. The organ itself is si- tuated behind this valve, and consists either of a membrane, curiously plaited into numerous semicircular folds, or of tufted or .arborescent filaments. Fig. 3SS shows this cavity (s,) with its plaited membrane in the Perch; and Fig. 3S9, in the Skate; the laminae in the former being radiated, and in the latter, foliated, or parallel to each other. On the sur- face of these organs, whatever be their. shape, the olfactory nerves (n,) arising from the anterior lobes (o) of the brain, are distributed; and the great size of these nerves would lead us to infer considerable acuteness in the sense which they supply. When the fish is swimming, their situation in front of the snout exposes them to the forcible impulse of the water which strikes against them. According to Gcof- froy St. Hilaire, the water enters the cavity by the upper orifice, and escapes by the lower. Scarpa alleges that fishes exercise this sense by compressing the water against the membrane. On the other hand, it is contended by Dumerll, that the perceptions communicated by this organ, being the result of the action of a liquid instead of a gas, should be classed under the head of taste rather than of smell. This seems, however, to be a mere verbal criticism, in making which it appears to have been forgotten that the impressions of odorous effluvia, even in animals breathing atmospheric air, always act upon the nerve through the intermedium of the fluid which lubricates the membrane of the nostril. That the nasal cavities of fishes are rudimental forms of 292 THE SENSORIAL FUNCTIONS. o those of the mammalia, although they do not, as in the latter class, open into the the respiratory organs, is shown by the curious transformation of the one into the other during the development of the tadpole, both of the frog and of the sa- lamander. During the first periods of their existence, these animals arc perfectly aquatic, breathing water by means of gills, and having all tlicir organs formed on the model of the fish. Their nasal cavities are not employed for respiration at this early period, nor even for some time after they have begun to take in air, which they do by the mouth, swallow- ing it in small portions at a time, and afterwards throwing it out in bubbles by the same channel. But when they quit the water, and become land animals with pulmonary respi- ration, the nostrils are the channels through which the air is received and expelled ; and it is here also that the sense of smell continues to be exercised. We know very little respecting the seat of the sense of smell in any of the invertebrated animals, though it is very evident that insects, in particular, enjoy this faculty in a very high degree. Analogy would suggest the spiracles as the most probable seat of this sense, being the entrances to the respiratory passages. This office has, however, been as- signed by many to the antennae; while other entomologists have supposed that the palpi are the real organs of smell.* Experiments on this subject are attended with great diffi- culty, and their results must generally be vague and incon- clusive. Those which Mr. P. Huber made on bees, seem, however, to establish, with tolerable certainty, that the spira- cles are insensible to strong odours, such as that of oil of turpentine, which is exceedingly offensive to all insects. It was only when a fine camel-hair pencil containing this pun- gent fluid was presented near the cavity of the mouth, above the insection of the proboscis, that any visible effect was pro- duced upon the insect, which then gave decisive indications • On the subject of this sense in insects. See Kirby and Spence's Introduc- tion to Entomology, vol. iv. p. 249. SMELL. 293 of strong aversion. Mr. Kirby has discovered in the ante- rior part of the nose of the Necrophorus vespillo^ or bu- rying beetle, which is an insect remarkable for the acute- ness of its smell, a pair of circular pulpy cushions, covered with a membrane, beautifully marked with fine transverse furrows. These he considers as the organs of smell; and he has found similar structures in several other insects.* No distinct organs of smell have been discovered in any of the Mollusca; but as there is evidence that some of the animals belonging to that class possess this sense, it has been conjectured that it resides either in the whole mucous surface of the mantle, or in the respiratory organs. Swam- merdam observed, long ago, that snails are evidently af- fected by odours; and cuttle-fish are said to show a decided aversion to strongly scented plants. * Kirby and Spence's Introduction to Entomology, vol. iii. 481? and iv. 254. ( 294 ) CHAPTER V. HEARING. § 1. Acoustic Principles. The knowledge acquired by animals of the presence and movements of distant objects is derived almost wholly from the senses of hearing and of sight; and the apparatus, ne- cessary for the exercise of these senses, being more elabo- rate and refined than any of the organs we have yet exa- mined, exhibit still more irrefragable evidence of those pro- found designs, and that infinite intelligence, which have guided the construction of every part of the animal frame. Sound results from certain tremulous or vibratory motions of the particles of an elastic medium, such as air or water, excited by any sudden impulse or concussion given to those particles by the movements of the sounding body. These sonorous vibrations are transmitted with great velocity through these fluids, till they strike upon the external ear; and, then, after being concentrated in the internal passages of the organ, they are made to act on the filaments of a par- ticular nerve called the acoustic, or auditory nerve, of which the structure is adapted to receive these peculiar im- pressions, and to communicate them to the brain, where they produce changes, which are immediately followed by the sensation of sound. Sound cannot traverse a void space, as light does; but always requires a ponderable material ve- hicle for its transmission; and, accordingly, a bell suspended in the vacuum of an air-pump, gives, when struck, no audi- HEARING. 295 ble sound, although its parts are visibly thrown into the usual vibratory motions. In proportion as air is admitted into the receiver, the sound becomes more and more dis- tinct; and if, on the other hand, the air be condensed, the sound is louder than when the bell is surrounded by air of the ordinary density.* The impulses given by the sounding body to the contigu- ous particles of the elastic medium, are propagated in every direction, from particle to particle, each, in its turn, striking against the next, and communicating to it the whole of its own motion, which is destroyed by the reaction of the particle against which it strikes. Hence, after moving a certain de- finite distance, a distance, indeed, which is incalculably small, each particle returns back to its former situation, and is again ready to receive a second impulse. Each particle, being elastic within a certain range,t suffers a momentary com- pression, and immediately afterwards resumes its former shape: the next particle is, in the mean time, impelled, and undergoes the same succession of changes; and so on, throughout the whole series of particles. Thus, the sono- rous undulations have an analogy with waves, which spread in circles on the surface of water, around any body, which, by its motion, ruffles that surface; only that, instead of merely extending in a horizontal plane, as waves do, the so- norous undulations spread out in all directions, forming, not circles in one plane, but spherical shells; and, whatever be the intensity of the sounds, the velocity with which the un- dulations advance is uniform, as long as they continue in a medium of uniform density. This velocity in air, is, on an average, about 1100 feet in a second, or twelve and a half * These facts were first ascertained by Dr, Ilauksbee. See Philosophical Transactions for 1705, vol. xxiv., p. 1902, 1904. ■|- The particles of water are as elastic, within a limited distance, as those of the most solid body, although, in consequence of their imperfect cohe- sion, or, rather, theii* perfect mobility in all directions, this property cannot be so easily recog^iised in masses of fluids, as it can in solids. 296 THE SENSORIAL FUNCTIONS. miles in a minute: it is greater in dense, and smaller in rare- fied air; being, in the same medium, exactly proportioned to the elasticity of that medium. Water is the medium of sound to aquatic animals, as the air is to terrestrial animals. Sounds are, indeed, conveyed more quickly, and to greater distances, in water than in air, on account of the greater elasticity of the constituent parti- cles of water, within the minute distance required for their action in propagating sound. Stones, struck together under water, are heard at great distances by a person whose head is under water. Franklin found, by experiment, that sound, after travelling above a mile through water, loses but little of its intensity. According to Chladni, the velocity of sound in water is about 4900 feet in a second, or between four and five times as great as it is in air. Solid bodies, especially such as are hard and elastic, and of uniform substance, are also excellent conductors of sound. Of this we may easily convince ourselves by applying the ear to the end of a log of wood, or a long iron rod, in which situation we shall hear very distinctly the smallest scratch made with a pin at the other end; a sound, which, had it passed through the air only, would not have been heard at all. In like manner, a poker suspended by two strings, the ends of which are applied to the two ears, communicates to the organ, when struck, vibrations which would never have been heard under ordinary circumstances. It is said that the hunters in North America, when desirous of hearing the sounds of distant footsteps, which would be quite inaudible in any other way, apply their ears close to the earth, and then readily distinguish them. Ice is known to convey sounds, even better than water: for if cannon be fired from a distant fort, where a frozen river intervenes, each flash of light is followed by two distinct reports, the first being con- veyed by the ice, and the second by the air. In like man- ner, if the upper part of the wall of a high building be struck with a hammer, a person standing close to it on the ground, HEARING. 297 will hear two sounds after each hlow, the first descending through the wall, and the second through the air. As sounds are weakened by diflfusion over a larger sphere of particles, so they are capable of having tlieir intensity in- creased by concentration into a smaller space; an cflect which may be produced by their being reflected from the solid walls of cavities, shaped so as to bring the undulations to unite into a focus; it is on this principle that the ear- trumpet, for assisting persons dull of hearing, is construct- ed: and the same effect sometimes takes place in echoes, which occasionally reflect a sound of greater loudness than the original sound which was directed towards them. If the impulses given to the nerves of the ear be repeated at equal intervals of time, provided these intervals be very small, the impressions become so blended together as not to be distinguishable from one another, and the sensation of a uniform continued sound, or musical note, is excited in the mind. If the intervals between the vibrations be long, the note is grave; if short, that is, if the number of vibrations in a given time be great, the note is, in the same proportion, acute. The former is called a low, the latter a high note; designations which in all probability were originally derived from the visible motions of the throat of a person who is singing these diff'erent notes; for, independently of this cir- cumstance, the terms of high and low are quite arbitrary; and it is well known that they were applied by the ancients in a sense exactly the reverse of that in which we now use them. The difierent degrees of tension given to the cord or wire of a stringed musical instrument, as well as its diflferent lengths, determine the frequency of its vibrations; a greater tension, or a shorter length, rendering them more frequent, and consequently producing a higher note; and on the con- trary, the note is rendered more grave by either lessening the tension, or lengthening the cord or wire. In a wind instrument, the tone depends altogether upon the length of the tube producing the sound. Vol. II. 3S 298 THE SENSORIAL FUNCTIONS. Tlicre are, therefore, two qualities in sound recognisable by the ear, namely, loudness, or intensity, and quality, or tone; the former depending on tlie force of the vibrations; the latter, on tlieir frequency. These acoustic principles arc to l)e borne in mind in studying the comparative physi- ology of hearing; and since the functionsof the different parts of the organ of this sense are, as yet, but imperfectly under- stood, 1 sliall, in treating of this subject, deviate from the plan I liave hitherto followed, and premise an account of the structure of the ear in its most perfectly developed state, wluch it appears to be in Man. § 2. Physiology of Hearing in Man. That part of the organ of hearing, which, above all others is essential to the performance of this function, is the acous- tic nerve, of which the fibres are expanded, and spread over the surface of a fine membrane, placed in a situation adapt- ed to receive the full impression of the sonorous undulations, which are conveyed to them. This membrane, then, with its nervous filaments, may be regarded as the immediate or- gan of the sense; all the other parts being merely accessory apparatus, designed to collect and to condense the vibrations of the surrounding medium, and to direct their concentrated action on the auditory membrane. I have endeavoured, in Fig. 390, to exhibit, in one view^, the principal parts of this complicated organ, as they exist in man, in their relative situations, and of their natural size: thereby affording a scale by which the real dimensions of those portions, which I shall afterwards have occasion to explain by magnified representations, may be properly ap- preciated.* The Coiicha, or external ear (c,) is formed of an elastic plate of cartilage, covered by integument, and presenting va- • In lliis and uU the following' figures, the parts of the right car are shown, and similar parts arc always indicated by the same letters. HEARING. 299 rious elevations and depressions, which form a series of pa- rabolic curves, apparently for the purpose of collecting the sonorous undulations of the air, and of directing them into a funnel-shaped canal (m,) termed the meatus auditoriiis, which leads to the internal ear. This canal is composed partly of cartilage, and partly of bone; and the integument lining it is furnished with numerous small glands, which supply a thick oily fluid, of an acrid quality, apparently de- signed to prevent the intrusion of insects: the passage is also guarded by hairs, which appear intended for a similar pur- pose. The meatus is closed at the bottom by a membrane (d,) which is stretched across it like the skin of a drum, and has been termed, from this resemblance, the membrane of the tympanum, or the ear-drum.'^ It performs, indeed, an of- fice corresponding to its name; for the sonorous undulations of the air, which have been collected, and directed inwards by the grooves of the concha, strike upon the ear-drum, and throw it into a similar state of vibration. The ear-drum is * The Inner surface of the ear-drum is shown in this figure, the cavity of the tympanum, which is behind it, being- laid open. 300 THE SENSORIAL FUNCTIONS. composed of an external membrane, derived from the cuti- cle whicli lines the meatus; an internal layer, which is con- tinuous witli that of the cavity beyond it; and a middle layer, wiiich consists of radiating muscular fibres, proceed- ing from the circumference towards the centre, where they are inserted into the extremity of a minute bony process (h,) presently to be described.* This muscular structure apj)ears designed to vary the degree of tension in the ear- drum, and tlius adapt the rate of its vibrations to those com- municated to it by the air. There is, also, a slender muscle, situated internally, which, by acting on this delicate process of bone, as on a lever, puts the whole membrane on the stretch, and enables its radiating fibres to effect the nicer adjustments required for tuning, as it may be called, this part of the organ. t Immediately behind the membrane of the ear-drum, there is a hollow space (t,) called the cavity of the tympanum^ of an irregular shape, scooped out of the most solid part of the temporal bone, which is here of great density and hard- ness. This cavity is always filled with air; but it would obviously defeat the purpose of the organ if the air were confined in this space; because unless it were allowed occa- sionally to expand or contract, it could not long remain in equilibrium with the pressure exerted by the atmosphere on the external surface of the ear-drum; a pressure which, as is w^ell known, is subject to great variations, indicated by the rise and fall of the barometer. These variations would ex- pose the membrane of the ear-drum to great inequalities of pressure at its outer and inner surfaces, and endanger its being forced, according to the state of the weather, either outwards or inwards, which would completely interfere with the delicacy of its vibrations. Nature has guarded against • In many quadrupeds their insertion into this process is at some distance from the centre of the membrane. These muscular fibres arc dcUncated in Fig. 45, vol. i. p. 105. f Home, Lectures, &c., iii. 268. HEARING. 301 these evils by establishing a passage of communication be- tween the tympanum and the external air, by means of a tube (e,) termed the Eustachian tube, which begins by a small orifice from the inner side of the cavity of the tympa- num, and opens by a wide mouth at the back of the nos- trils.* This tube performs the same office in the ear, as the hole which it is found necessary to make in the side of a drum, for the purpose of opening a communication with the external air; a communication which is as necessary for the functions of the ear, as it is for the proper sounding of the drum. We find accordingly that a degree of deafness is induced whenever the Eustachian tube is obstructed, which may happen either from the swelling of the membrane lining it, during a cold, or from the accumulation of secre- tion in the passage. It is also occasionally useful as a chan- nel through which sounds may gain admittance to the inter- nal ear; and it is perhaps for this reason that we instinct- ively open the mouth when we are intent on hearing a very faint or distant sound. On the side of the cavity of the tympanum, which is op- posite to the opening of the Eustachian tube, is situated the beginning of another passage, leading into numerous cells, contained in the mastoid process of the temporal bone, and therefore termed the mastoid cells: these cells are likewise ;393 1 a filled with air. The innermost side of the same cavity, that is, the side opposite to the ear-drum, and which is shown in • This opening is seen at e, in Fig". 283, p. 383, representing a vertical and longitudinal section of the right nostril. 302 THE SENSORIAL FUNCTIONS. Fie;. 391, is occupied by a rounded eminence (p,) of a tri- ane;ular shape, termed the pro77iojitor}j ; on each side of which there is an opening in the bone, closed, however, by' the membrane lining the whole internal surface of the cavi- ty. The opening (o,) which is situated at the upper edge of the promontory, is called the fenestra ovalis, or oval window; and that near the under edge (r,) is the fenestra rotunda, or round window. Connected with the membrane of the ear-drum, at one end, and with the fenestra ovalis at the other, there extends a chain of very minute moveable bones, seen at (b,) in Fig. 390; but more distinctly in Fig. 392, which is drawn on a somewhat larger scale, and in which as before (d) is the ear-drum; (p) the promontory, (o,) the fenestra ovalis; and (r) the fenestra rotunda. These bones, which may be called the tymjjanic ossicula, are four in number, and are repre- sented, enlarged to twice the natural size, in Fig. 393.. The names they have received are more descriptive of their shape than of their office. The first is the malleus, or ham- mer (m;) and its long handle (h) is affixed to the centre of the ear-drum: the second is the incus, or anvil (i;) the third, which is the smallest in the body, being about the size of a millet seed, is the orbicular bone (o;)* and the last is the stapes, or stirrup (s,) the base of which is applied to the membrane of the fenestra ovalis. These bones are regularly articulated together, with all the ordinary appa- ratus of joints, and are moved by small muscles provided for that purpose. Their office is apparently to transmit the vibrations of the ear-drum to the membrane of the fenestra ovalis, and probably, at the same time, to increase their force. The more internal parts of the ear compose what is de- • lilumenbach, and other anatomists, consider this as not being a separate bone, but only a process of the incus,- a view of the subject which is sup- ported by the observations of Mr. Shrapnell, detailed in the Medical Ga- zette, xii., 172. HEARING. 303 signaled, from the intricacy of its winding passages, the la- byrinth. It is seen at (s v k) in Fig. 390, in connexion with the tympanum; but in Fig. 394, it is represented, on a very large scale, detached from every other part, and separated from the solid bone in which it lies embedded. It consists of a middle portion, termed the vestibule (v,) from which, on its upper and posterior side, proceed the three tubes (x, T, z,) called, from their shape, the semicircular canals; while to the lower anterior side of the vestibule there is attached a spiral canal, resembling in ap- pearance the shell of a snail, and on that account denomi- nated the Cochlea (k.) All these bony cavities are lined with a very delicate membrane, or periosteum, and are fdled with a transparent watery, or thin gelatinous fluid, which is termed by Breschet, Wiq perilym,ph.^ Within the cavity of the osseous labyrinth,, now de- scribed, are contained membranes having nearly the shape of the vestibule and semicircular canals, but not extending into the cochlea. These membranes, which compose what has been termed, for the sake of distinction, the membranous labyrinth, form pne continuous, but closed sac, containing a fluidjf perfectly similar in appearance to the perilymph, which surrounds it on the outer side, and intervenes be- tween it and the sides of the osseous labyrinth, preventing any contact with those sides. In Fig. 395, which is on a still larger scale than the preceding figure, the osseous laby- rinth is laid open, so as to show the parts it encloses, and * Annales des Sciences Naturelles, xxix. 97. It has also been called the Aqua lahyrinnd, and the Jliiid of Cotunnius, from the name of the Anato- mist who first distinctly described it. f De Blainville has termed tiiis fluid " la vitrine auditive," from its sup- posed analogy with the vitreous humour of the eye. 304 THE SENSORIAL FUNCTIONS. mor§ especially the membranous labyrinth, floating in the perilymph (p.) The form of this latter part is still more distinctly seen, in Fig. 39G, where it is represented in a po- sition exactly corresponding to the former figure, but whol- ly detached from the bony labyrinth, and connected only with the nervous fdaments which arc proceeding to be dis- tributed to its diflerent parts. A simple inspection of these figures, in both of which the corresponding parts are marked by the same letters, will show at once the form and the connexions of the three semi- circular canals, (x, y, z,) each of which present, at their ori- gin from the vestibule, a considerable dilatation, termed an cnnjmlla (a, a, a,) while, at their other extremities, where they terminate in the vestibule, there is no enlargement of their diameter: and it will also be seen that two of these ca- nals (x and y) unite into one before their termination. The HEARING. 305 same description applies in all respects botli to the osseous and to the membranous canals contained witliin them; the space (p) which intervenes between the two, being filled with the perilymph. But the form of the membranous vestibule demands more particular notice, as it is not so exact an imi- tation of that of the osseous cavity; being composed of two distinct sacs, opening into each other: one of these (u) is termed the utricle;'^ and the other (s,) the sacculus. Each sac contains in its interior a small mass of white calcareous matter, (o, o,) resembling powdered chalk, which seems to be suspended in the fluid contained in the sacs by the interme- dium of a number of nervous filaments proceeding from the acoustic nerves (g and n,) as seen in Fig. 396. From the universal presence of these cretaceous substances in the la- byrinth of all the mammalia, and from their much greater size and hardness in aquatic animals, there can he little doubt that they perform some office of great importance in the physiology of hearing.t Their size and appearance in the Dog is shown in Fig. 397: and in the Hare, in Fig. 398. The Cochlea, again, is an exceedingly curious structure, being formed of the spiral convolutions of a double tube, or rather of one tube, separated into two compartments by a partition (l,) called the lamina spiralis, which extends its whole length, except at the very apex of the cone, where it suddenly terminates in a curved point, or hook (h,) leaving an aperture by which the two portions of the tube commu- nicate together. In Fig. 395, a bristle (b, b) is passed through this aperture. The central pillar, round which these tubes take two and a half circular turns, is termed the modiolus. Its apex is seen at (m.) One of these passages is distin- guished by the name of the vestibular tiibe,X in consequence • Scarpa and Weber term it the sinus or alveus titrlculosus ,- it is called by others the sacculus vestihuU. Brcschct ^ives it tlie name of Ic sinus median. See the Memoir already quoted, p. 98. I These cretaceous bodies are termed by IJrcschct otolithcs, and otoconicSy according as they are of a hard or soft consistence. Ibid. p. 99. \ Sea la vestibuli. Vol. II. 39 306 THE SENSORIAL FUNCTIONS. of its arising from the cavity of the vestibule; and the other by that of the tympanic tube,'^ because it begins from the inner side of the membrane which closes the fenestra ro- tunda, and forms the only separation between the interior of that tube, and the cavity of the tympanum. The trunk of the auditory nerve occupies a hollow space immediately be- hind the ventricle, and its branches pass through minute holes in the bony plate which forms the wall of that cavity, being finally expanded on the different parts of the mem- branous lal)yrinth.t Great uncertainty prevails with regard to the real func- tions performed by the several parts of this very complex apparatus. It is most probable, however, that the sonorous viljrations of the air which reach the external ear, are di- rected down the meatus, and striking against the ear-drum which closes the passage, throw that membrane into vibra- tions of the same frequency; to which the action of its mus- cles, which appear intended to regulate its tension, may also contribute. The vibrations of the ear-drum, no doubt, ex- cite corresponding motions in the air contained in the cavity of the tympanum; which, again, communicates them to the membrane of the fenestra rotunda; while, on the other hand, the membrane closing the fenestra ovalis, receives similar impressions from the stapes, conveyed through the chain of tympanic ossicula, which appear to serve as solid conductors of the same vibrations. Thus, the perilymph, or fluid con- tained in the labyrinth, is affected by each external sound, both tlirough the medium of the air in the tympanum, and by means of the ossicula: the undulations thus excited pro- • Scala tympani. f In Fig. 396, the anterior tmnk of the auditory nerve is seen (at g) dis- tributing branches to the ampullse (a, a,) the utricle (u,) and the calcareous body it contains; while the posterior trunk (u) divides into a branch, which supplies the sacculus (s) and its calcareous body (o) and a second branch ( K ) which is distributed over the cochlea, (d) is the nei*ve called the por- iio duruy which merely accomi)anies the auditoiy nerve, but has no relation to the sense of hearing. In Fig. 390, the auditory nerve (n) is seen enter- ing at the back of the vestibule. HEARING. 307 duce impressions on the extremities of the nervous filaments, which are spread over the membranous labyrinth; and these impressions being conveyed to the brain, are immediately followed by the sensation of sound. With regard to the purposes which are answered by the winding passages of the semicircular canals, and cochlea, hardly any plausible conjecture has been oflered; yet no doubt can be entertained that the uses of all these parts are of considerable importance, both as to delicacy and correct- -ness of hearing. There is an obvious correspondence be- tween the positions of the three semicircular canals, (two of which are vertical, and one horizontal, and of which the planes are reciprocally perpendicular to one another,) and the three dimensions by which the geometrical relations of space are estimated; and it might hence be conjectured that the ob- ject of this arrangement is to allow of the transmission of vi- brations of every kind, in whatever direction they may ar- rive. It is not an improbable supposition tliat tiie return into the vestibule, of undulations which have passed through these canals, has the effect of at once putting a stop to all farther motion of the fluid, and preventing the continuance of the impression which has been already made on the nerves. The same use may be assigned to the double spiral convolutions of the tubes of the cochlea: for the undulations of the fluid in the tympanic tube, received from tlie mem- brane of the fenestra rotunda, will meet those proceeding along the vestibular tube, derived from the membrane of the fenestra ovalis, and like two opposing waves, will tend to destroy one another. Thus each external sound will pro- duce but a single momentary impression; the prolongation of the undulations of the fluid of the labyrinth being pre- vented by their mutual collision and neutralization.*' * The preliminary steps in the process above described are not absolutely essential to hearing-, for many instances have occuiTed in which the power of bearing has been perfectly retained after the membrane of the ear-drum, and also the ossicula had been destroyed by disease. A small aperture in the membrane does not interfere with its power of vibration; but if the whole 808 THE SENSORIAL FUNCTIONS. § 3. Comjjarative Physiology of Heariiig. The structure of the organs of hearing in the lower ani- mals presents a regular gradation from the simple vestibule, with its membranous sac, supplied with nervous filaments, which may be regarded as the only essential part of this or- gan, through the successive additions of semicircular canals, fenestra ovalis, tympanic cavity, ossicula, ear-drum, meatus auditorius, cochlea, and concha, till we arrive at the combi- nation of all these parts in the higher orders of the Mam- malia. The simpler forms are generally met with in aqua- tic animals, probably because the sonorous undulations of water are communicated more readily, and with greater force, than tiiose of air, and require no accessory apparatus for their concentration. The loftster, for instance, has a ves- tibular cavity (seen at v, in Fig. 399,) containing a membra- nous sac, with a striated groove (g,)* and receiving the fila- ments of the auditory nerve. This vestibule is protected by tho shell on all sides, except at one part, where it is closed only by a membrane (e,) which may therefore be considered as corresponding to the fenestra ovalis. The outer side of this membrane in the Jlstaciis Jluviatilis, or cray-fish, is seen at f in Fig. 401; while Fig. 402, shows an ear-drum be deslroyed, and the ossicula lost, an almost total deafness g-ene- rally ensues. After a time, however, the hearing- may be in a great measure recovered, with an undiminished power of distinguishing musical tones. See two papers by Sir Astlcy Cooper, in the Phil. Trans, for 1800, p. 151; and for 1801, p. 437. ♦ This g-roove is represented magnified in Fig. 400. HEARING. 309 interior view of the same membrane (f,) with the vestibule (v) laid open, and the auditory nerve (n) passing through the shell to be distributed on the sacculus. It appears from a variety of observations that Insects, both in their larva and their perfect state, possess the faculty of hearing; but no certain knowledge has been obtained of tbe parts which exercise this sense. Tlic prevailing opinion among entomologists is that it resides in some part of the antennae; organs, which are supposed to have a peculiar sen- sibility to aerial undulations. This hypothesis is founded principally on the analogy of the Crustacea, whose antennae contain the vestibular cavity already described; but on the other hand it is opposed by the fact that Spiders, which hear very acutely, have no antennae; and it is also reported that insects, when deprived of their antennae, still retain the power of hearing.* None of the Mollusca appear to possess, even in the small- est degree, the sense of hearing, if we except the highly or- ganized Cephalopoda; for in them we find, at the lower part of the cartilaginous ring, which has been supposed to exhi- bit the first rudiment of a cranium, a tubercle, containing in its interior two membranous vesicles, contiguous to each other, and surrounded by a fluid. They evidently corre- spond to the vestibular sacs, and contain each a small calca- reous body, suspended from the vesicles by slender nervous filaments, like the clapper of a bell, and probably performing an office analogous to that instrument; for, being thrown into a tremulous motion by every undulation of the sur- rounding fluid, they will strike against the membrane, and communicate similar and still stronger impulses to the nerves by which they are suspended, thus increasing the impression made on those nerves. The mechanical effect of an apparatus of this kind is shown by the simple experiment, • Camparetti has described structures in a great number of insects, which he imagined were organs of hearing; but liis observations have not been con- firmed by subsequent inquirers, and their accuracy is therefore doubtful. See De Blainville " De I'Organisation des Animaux," i. 565. 310 THE SENSORIAL FUNCTIONS. mentioned by Camper, of enclosing a marble in a bladder full of water, and held in the hand; when the slightest shaking of the bladder will be found instantly to communicate mo- tion to tiie marble, the reaction of which on the bladder gives an unexpected concussion to the hand. The ear of Fishes contains, in addition to the vestibule, the three semicircular canals, which arc, in general, greatly developed.* An enlarged view of the membranous laby- rinth of the Lophiiis piscatoruis is given in Fig. 403, show- ing the form and com})lication of its parts, which are repre- sented of twice the natural size, x, y, z, are tlie semicircular canals, with their respective ampullae (a, a, a.) m is the Sinus rtiedianus^ or principal vestibular sac, with its ante- rior expansion, termed the Utricle (u.) The Saccidus (s) has, in like manner, a posterior appendage (c) termed the Cysticule. The hard calcareous bodies (o, o, o) are three in number; and tlie branches of nerves (i, i, i) by which they are suspended in the fluid contained in the membranes, are seen passing into them; while the ampullae are supplied by other branches (n, n, n.) In all the osseous fishes, the labyrinth is not enclosed in the bmies of the cranium, but projects into its cavity; but in the larger cartilaginous fishes, ♦ In the lamprey, these canals exist only in a rudimental state, appearing" as folds of tlie membrane of the vestibule; and there are also no cretaceous bodies in the vestibular sac. HEARING. 311 as the ray and shark tribes, it is surrounded by solid bone, and is not visible within the cranium. In these latter fishes, we first meet with a rudiment of the meatus, in a passage extending from the inner side of the vestibule, to the upper and back part of the skull, where it is closed by a mem- brane, which is covered by the skin. Aquatic reptiles have ears constructed nearly on the same plan as those of fishes: thus, the Triton or Newt has a vesti- bule containing only one cretaceous body, and three semi- circular canals, unprotected by any surrounding bone. In the Frog, however, we first perceive the addition of a dis- tinct cavity, closed by a membrane, which is on a level with the integuments, on each side of the head. From this cavity, which corresponds to that of the tympanum, there proceeds an Eustachian tube; and within it, extending from the external membrane, which must here be regarded as an ear-drum, to the membrane of the vestibule, or fenestra ovalis, is found a bone, shaped like a trumpet, and termed unci the Columeim. This bone is seen at c in Fig. 404, attached by its base (b) to the fenestra ovalis of the vestibule (v,) which contains the cretaceous body (o.) There is also a small bone (i) attached in front to the columella. In the Chelonia, the structure of the ear is essentially the same as in the Frog, but the tympanum and columella are of greater length. In the saurian reptiles the cavity of the tympanum is still more capacious, and the ear-drum very distinctly marked, and these animals possess great delicacy of hearing. The labyrinth of tlie Crocodile is enclosed in bone, and con- tains three calcareous bodies: it presents also an appendage 312 THE SENSORIAL FUNCTIONS. which has been regarded as tlie earliest rudiment of a coch- lea; and there are two folds of the skin, resembling; eye-lids, at the external orifice of the organ, which appear like the first step towards the development of an external ear. The structure of the car in the Crocodile is but an ap- proximation to that wliich we fmd prevailing in Birds, where the organ is of large size compared with that of the head. The rudimcntal cochlea, as seen at k in Fig. 405, which represents these organs in the Turkey, is of large size, and slightly curved. In the cavity of the tympanum (t) is seen the columella, which extends to the fenestra ovalis; and beyond it, the semicircular canals (s,) the bony cells (b) which communicate with the tympanum, the os quadratum (q,) the zygomatic process (z,) and the lower jaw (j.) The ear-drum is now no longer met with at the surface, but lies concealed at the bottom of a short meatus, the orifice of which is surrounded with feathers arranged so as to serve as a kind of imperfect concha, or external ear. In Owls these feathers are a prominent ancjfcharacteristic feature; and in these birds there is, besides, a membranous flap, acting as a valve to guard the passage. The chief peculiarity observable in the internal ears of ]\Iammalia is the great development of the cochlea, the tubes of wliich are convoluted, turning in a spiral, and as- suming the figure of a turbinated shell. From an extensive comparison of the relative size of the cochlea in different tribes of quadrupeds, it has been inferred that it bears a to- leraldy constant proportion to the degree of acuteness of hearing, and that, consequently, it contributes essentially to the perfection of that faculty: bats, for instance, which are known to possess exquisite delicacy of hearing, have a coch- lea of extraordinary size, compared with the other parts of the ear. The tympanic ossicula are completely developed only in the JMammalia.* It is also in this class alone that • TIjcsc tympanic ossicula are reg-arded by GcofFroy St. liilaire as cor- responding to the opercular hones of fishes, where, according to his theory, they have attained the highest degree of development. HEARING. 313 we meet with a concha, or external ear, distinctly marked; and the utility of this part, in catching and coliectinj^ the sonorous undulations of the air, may be inferred from the circumstance, that a large and very moveable concha is ge- nerally attended with great- acuteness of hearing. This is more particularly the case with feeble and timid quadrupeds, as the hare and rabbit, which are ever on the watch to catch the most distant sounds of danger, and whose ears are turned backwards, or in the direction of their pursuers; while, on the contrary, the ears of predaceous animals are directed forwards, that is, towards the objects of their pursuit. This difference in direction is not confined to the external ear, but is observable also in the bony passage leading to the tympanum. The Cetacea, being strictly inhabitants of the water, have no external ear; and the passage leading to the tympanum is a narrow and winding tube, formed of cartilage instead of bone, and having a very small external aperture. In the Dolphin tribe the orifice will barely admit the entrance of a pin; it is also exceedingly small in the Dugong; these structures being evidently intended for preventing the en- trance of any quantity of water.* It is apparently with the same design that in the Seal the passage makes a circular turn; and that, in the Ornithorhyncus paradoxus, it winds round the temporal bone, and has its external orifice at a great distance from the vestibule. The internal parts of the organ of hearing in the Whale, and other cetacea, are en- closed in a bone of extraordinary hardness, which, instead of forming a continuous portion of the skull, is connected to it only by ligaments, and suspended in a kind of osseous ca- vity, formed by the adjacent bones. The cochlea is less de- veloped than in quadrupeds, for it only takes one turn and a half, instead of two and a half. The existence of the se- * It is probable that in these animals the principal channel by which sounds reach the internal organ is the Eustachian tube. Vol. II. 40 314 THE SENSORIAL FUNCTIONS. micircular canals in the cetacea was denied by Camper; but they have since been discovered by Cuvier. Several quadrupeds which are in the habit of burrowing, or of diving, as the Soy^ex fodiens, or water-shrew, are fur- nished with a valve, composed of a double membrane, capa- ble of accurately closing the external opening of the meatus, and protecting it from the introduction of water, earth, or other extraneous bodies.* In like manner the external ear of the Hippopotamus, which feeds at the bottom of rivers, is guarded by an apparatus which has the effect of a valve. We fmd, indeed, the same provident care displayed in this as in every other department of the animal economy: every part, however minute, of the organ of this important sense, being expressly adapted, in every species, to the par- ticular circumstances of their situation, and to that degree of acuteness of perception, which is best suited to their respec- tive wants and powers of gratification.! • GeofTroy St. Hllaire; Memoires du Museum, i. 305. f The Comparative Physiology of Voice, a function of which the object, in animals as well as in man, is to produce sounds, addressed to the ear, and expressive of their ideas, feeling's, desires and passions, forms a natural se- quel to that of Hearing-; but Sir Charles Bell having- announced his intention of inti'oducing- it in his Treatise on the Hand, I have abstained from enter- ing into tliis extensive subject. ( 315 ) CHAPTER VI. VISION, § 1. Object of the Sense of Vision. To those who study nature with a view to the discovery of final causes, no subject can be more interesting or instruc- tive than the physiology of Vision, the most refined and most admirable of all our senses. However well we may be acquainted with the construction of any particular part of the animal frame, it is evident that we can never form a correct estimate of the excellence of its mechanism, unless we have also a knowledge of the purposes to be answered by it, and of the means by which those purposes can be accom- plished. Innumerable are the works of creation, the art and contrivance of which we are incompetent to understand, because we perceive only the ultimate effects, and remain ig- norant of the operations by which those effects are produced. In attempting to investigate these obscure functions of the animal or vegetable economy, we might fancy ourselves en- gaged in the perusal of a volume, written in some unknown language, where we have penetrated the meaning of a few words and sentences, sufficient to show us that the whole is pregnant with the deepest thought, and conveys a tale of sur- passing interest and wonder, but where we are left to gather the sense of connecting passages by the guidance of remote analogies or vague conjecture. Wherever we fortunately succeed in deciphering any continued portion of the dis- course, we find it characterized by a perfection of style, and grandeur of conception, whicli at once reveal a master's hand., 316 THE SENSORIAL FUNCTIONS. and which kindle in us the most ardent desire of supplying the wide chasms perpetually intervening in the mysterious and inspiring narrative. But in the suhject which now claims our attention we have hccn permitted to trace, for a consi- derable extent, the continuity of the design, and the length- ened series of means employed for carrying that design into execution; and the view which is thus unfolded of the mag- nificent scheme of creation is calculated to give us the most sublime ideas of the wisdom, the power, and the bene- volence OF God. On none of the works of the Creator, which we are per- mitted to behold, have the characters of intention been more deeply and legibly engraved than in the organ of vision, where the relation of every part to the efifect intended to be produced is too evident to be mistaken, and the mode in which they operate is at once placed within the range of our comprehen- sion. Of all the animal structures, this is, perhaps, the one which most admits of being brought into close comparison with the works of human art; for the eye is, in truth, a re- fined optical instrument, the perfection of which can never be fully appreciated until we have instituted such a comparison; and the most profound scientific investigations of the anato- my and physiology of the eye concur in showing that the whole of its structure is most accurately and skilfully adapt- ed to the physical laws of light, and that all its parts are finished with that mathematical exactness which the preci- sion of the effect requires, and which no human effort can ever hope to approach, — far less to attain. To the prosecution of this inquiry we are farther invited by the consciousness of the incalculable advantages we derive from the sense of sight, the choicest and most enchanting of our corporeal endowments. The value of this sense must, indeed, appear inestimable, when we consider of how large a portion of our sensitive and intellectual existence it is the intermediate source. Not only has it given us extensive command over the objects which surround us, and enabled us to traverse and explore the most distant regions of the VISION. 317 globe, but it has introduced us to the knowledge of the bo- dies which compose the solar system, and of the countless hosts of stars which are scattered through the firmament, thus expanding our views to the remotest confines of crea- tion. As the perceptions supplied by this sense are at once the quickest, the most extensive, and the most varied, so they become the fittest vehicles for the introduction of other ideas. Visual impressions are those which in infancy, fur- nish the principal means of developing the powers of the un- derstanding: it is to this class of perceptions that the philoso- pher resorts for the most apt and perspicuous illustrations of his reasonings; and it is also from the same inexhaustible fountain that the poet draws his most pleasing and graceful, as well as his sublimest imagery. The sense of Vision is intended to convey to its posses- sor a knowledge of the presence, situation, and colour of external and distant objects, by means of the light which those objects are continually sending off, either spontane- ously, or by reflection from other bodies. It would appear that there is only one part of the nervous system so pecu- liarly organized as to be capable of being affected by lumi- nous rays, and conveying to the mind the sensation of light, and this part is the Retina^ so named from the thin and delicate membranous net-work, on which the pulpy extre- mities of the optic nerves^ establishing an immediate com- munication between that part and the brain, are expanded. If the eye were so constructed as to allow the rays of light, which reach it from surrounding objects, simply to impinge on the retina as they are received, the only per- ception which they could excite in the mind, would be a general sensation of light, proportionate to the total quantity which is sent to the organ from the whole of the opposite hemisphere. This, however, does not properly constitute Vision; for in order that the presence of a particular object in its real direction and position with respect to us, may be recognised, it is necessary that the light, which comes from it, and that light alone, produce its impression exclusively 318 THE SENSORIAL FUNCTIONS. on some particular part of the retina; it being evident that if the light, coming from any other object, were allowed to act, together with the former, on the same part, the two ac- tions would interfere with one another, and only a confused impression would result. The objects in a room, for exam- ple, are all throwing light on a sheet of paper laid on the floor; but this light, being spread equally over every part of the surface of the paper, furnishes no means of distinguish- ing the sources from which each portion of the light has proceeded; or, in other words, of recognising the respective figures, situations, and colours of the objects themselves. We shall now proceed to consider the modifications to be introduced into the structure of the organ, in order to attain tlicse objects. § 2. Modes of accomjjlishing the Objects of Vision. Let us suppose that it were proposed to us, as a problem, to invent an apparatus, by which, availing ourselves of the known properties of light, we might procure the concentra- tion of all the rays, proceeding from the respective points of the object to be viewed, on separate points of the retina, and obtain likewise the exclusion of all other rays; and also to contrive that the points of the retina, so illuminated, shall have the same relative situations among one another, which the corresponding points of the surrounding objects have in nature. In other words, let us suppose ourselves called upon to devise a method of forming on the retina a faithful deli- neation, in miniature, of the external scene. As it is a fundamental law in optics that the rays of light, while tliey arc transmitted through the same medium, pro- ceed in straiglit lines, the simplest mode of accomplishing the proposed end would be to admit into the eye, and con- vey to each particular point of the retina, only a single ray proceeding directly from that part of the object which is to be depicted on it, and to exclude all other rays. For car- VISION. 319 rying this design into effect we have the choice of two me- thods, both of which we find resorted to by nature under different circumstances. The first method consists in providing for each of these single rays a separate tube, with darkened sides, allowing the ray which traverses it, and no other, to fall on its re- spective point of the retina, which is to be applied at the opposite end of the tube. The most convenient form to be given to the surface of the retina, which is to be spread out to receive the rays from all these tubes,- appears to be that of a convex hemi- sphere; and the most eligible distribution of the tubes is the placing them so as to constitute diverging radii, perpendicular, in every part, to the surface of the retina. This arrangement will be understood by reference to Fig. 406, which represents a section of the whole organ: (t, t,) being the tubes disposed in radii every where perpendicular to the convex hemispheri- cal surface of the retina (r.) Thus will an image be formed, composed of the direct rays from each respective point of the objects, to which the tubes are directed; and these points of the image will have, among themselves, the same rela- tive situation as the external objects, from which they ori- ginally proceeded, and which they will accordingly faith- fully represent. The second method, which is nearly the inverse of the first, consists in admitting the rays through a small aperture into a cavity, on the opposite and internal side of which the retina is expanded, forming a concave, instead of a convex hemispherical surface. The mode in which this arrange- ment is calculated to answer the intended purpose will be easily understood by conceiving a chamber (as represented in Fig. 407,) into which no light is allowed to enter, except what is admitted through a small hole in a shutter, so as to fall on the opposite side of the room. It is evident that 320 THE SENSORIAL FUNCTIONS. each ray will, in that case, illuminate a different part of the wall, and that the whole external scene will be there faith- fully represented; for the several illuminated points, which constitute these images, preserve among themselves the same relative situation whicli the objects they represent do in nature, although with reference to the actual objects they have an inverted position. This inversion of the image is a necessary consequence of the crossing of all the rays at the same point; namely, the small aperture in the shutter, througli which they are admitted. JB ^^ \. One inconvenience attending the limiting of the illumina- tion of each point of the wall to that of a single ray, in the mode last pointed out, is that the image produced must ne- cessarily be very faint. If, with a view of remedying this defect, the aperture were enlarged, the image would, indeed, become brighter, but would at the same time be rendered more indistinct, from the intermixture and mutual interfe- rence of adjacent rays; for all the lines would be spread out, the outlines shaded off, and the whole picture confused. The only mode by which distinctness of image can be ob- tained with increased ilhimination, is to collect into one poin^ a great number of rays proceeding from the corre- sponding point of the object to be represented. Such a col- lection of rays proceeding from any point, is termed, in the language of optics, a pencil of rays; and the point into VISION. 321 niilch Ihey arc collected is called n focus. For the purpose of collecting a pencil of rays into a focus, it is evident that all of them, except the one which proceeds in a straight line from the ohject to that focus, must be dejlected, or bent from their rectilineal course. This effect may be produced b\- refractio7i, which takes place according to another optical law; a law of which the following is the exposition. It is only when the medium which the rays are traversing is of uniform density tiiat their course is constantly recti- lineal. If the density change, or U the rnys pass obliquely from one medium into another of a difierent density, they are refracted; each ray being deflected towards a line situ- ated in the medium of greatest density, and drawn from the point where the ray meets the new medium, perpendicular to the refracting surface. Thus, the ray, r, Fig. 40S, striking obliquely on the surface of a denser medium, at the point s, instead of pursuing its original course along the line s o, is refracted, or turned in the direction s t, which is a line situated between s o, and s p; this latter line being drawn £^^^^ B ^ 1 nmsssmm 40S|IH||j| r 1 iS ■Hj ^^HnHSrail fjfp ■ 1 : ' lUgQutJMHliM Y 'J perpendicularly to the surface of the medium, at the point s, and within that medium. When the ray arrives at t, and meets the posterior surface of the dense medium, passing thence into one that is less dense, it is again refracted ac- cording to the same law; that is, it inclines towards the per- pendicular line T I, drawn from t, within the denser me- dium, and describes the new course t u instead of t v. The amount of the deflection corresponds to the degree of ob- VoL. II. 41 THE SENSORIAL FUXCTIO^fS. liquity of the ray to the surface which refracts it; and is mathematically expressed, by the law that the sines of the two angles formed with the perpendicular by the incident and the refracted rays retain, amidst all the variations of those angles, the same constant proportion to one another. We may hence derive a simple rule for placing the plane of the refracting surface so as to produce the particular refrac- tion we wish to obtain. When a ray is to be deflected from its original course to a particular side, we have only to turn the surface of the medium in such a manner as that the per- pendicular line to that surface, contained within the denser medium, shall lie still farther on the same side. Thus, in Fig. 408, if we wish to turn the ray r s, from s o to s t, we must place the dense medium so that the perpendicular s p, which is within it, shall be still farther from s o,than s xis; that is, shall lie on the other side of s t. The same rule ap- plies to the contrary refraction of the ray s t from t v to t u, when it passes out of a dense, into a rare medium; for the perpendicular t i must still be placed on the same side of T V as T u is situated. Let us now aj)ply these principles to the case before us; that is, to the determination of the form to be given to a dense medium, in order to collect a pencil of rays, proceed- ing from a distant object, accurately to a focus. We shall 409 •B suppose the object in question to be very remote, so that the rays composing the pencil may be considered as being parallel to each other; for at great distances their actual de- VISION. 32-3 viation from strict parallelism is wholly insensible; and let a, B, c, D, E, (Fig. 409,) represent these rays. There must evidently be one of these rays (c,) and only one, which by continuing its rectilineal course, would arrive at the point (r)' intended to be the focus of the rays. This ray, then, may be suffered to pass on, without being subjected to any refraction; the surface of the medium should, therefore, be presented to the ray (at i) perpendicularly to its course, so that it may pass through at right angles to that surface. Those rays (b and d) wliich are situated very near to this direct or central ray (c,) will require but a small degree of refraction in order to reach the focus (r:) this small refrac- tion w^ill be effected by a slight degree of obliquity in the medium at the points (h and k) where those rays meet it. In proportion as the rays (such as those at a and e) are more distant from the central ray, a greater amount of refraction, and consequently a greater obliquity of the surfaces (g and l) will be required, in order to bring them to the same focus. The convergence of these rays, after they have passed this first surface, may be farther increased by interposing new surfaces of other media at the proper angles. If the new medium be still denser than the last, the inclination of its surface must be similar to that already described; if rarer, they must be in an opposite direction. This last case is il- lustrated in the figure, where m, n, o, p, q, represent the in- clinations of the surfaces of a rarer medium, calculated to increase the convergence of the rays, that is to bring them , to a nearer focus (f.) The result of the continued change of direction in the refracting surface, is a regular curvilineal surface, which, in the present case, approaches very nearly to that of a sphere. Hence by giving these refractive me- dia spherical surfaces, we adapt them, with tolerable exact- ness, to produce the convergence of parallel rays to a focus, and by making the denser medium convex on both sides (as shown in Fig. 410,) both surfaces will conspire in pro- ducing the desired efiects. Such an instrument is termed 324 THE SENSORIAL FUNCTIONS. a double convex lens; anil it has the property of collecting into a focus rays proccedinj; from distant points."^ 410 Ilavinii: obtained this instniincMil, wc mav now venture to enlarge the a])ertui'e tlii-oni;h wlilch the light was admit- ted into our dark chamber, and fit into the apertivrc a dou- ble convex lens. Vv'e hnve thus constructed the well known optical instrument called the Camera Ohscura, in which the imaj];es of external objects are formed upon a white sur- face of paper, or a 5cnii-transj)arent ])latc of glass; and these images must evidently bo in an inverted position with re- •pect to the actual objects which they represent. Such is precisely the construction of the eye, which is, to • The refraction by spherical surfiiccs docs not, strictly speakint^, unite x pencil of pnrallel or divergent rays into a muthematicul point, or focus; for in reality the rays wiiich are near the central line are made to converg-e to a point a little more distant than that to which tiie remoter rays converg-e: an •ffect which 1 have endeavoured to ilUistrate by the diagram Fig-. 411} where, in order to render it obvious to tlie eye, the disparity is much exaj^fgerated. But, on ordiuMiy occiisions, where great nicety is not required, this differ- ence in the degree of convcrgiMicc l)et\vcen the central rays and tliose near the circumference of titc lens, giving r'.sc to \vi;at is termed the Aberration ef Sphericity, is too small to attract notice. VISION. 325 all intents, a camera obscura: for in both these instrumenls, the objects, the principles of consti^ction, and the mode of operation are exactly the same; and tlie only difference is, that the former is an infinitely more perfect instrument thart the latter can ever be rendered by the utmost efforts of hu- man art. With a view of simplifying the subject, I have assumed, in the account given in the text, that the rays which arrive at the eye are parallel, which in mathematical strictness they never are. The focus of the rays refracted by a convex lens is more remote in proportion as the rays are more divergent or, in other words, proceed from nearer objects. This is illustrated by Fi- gures 412, 413, and 414j to which I shall again have occasion to refer in the sequel. § 3. Structure of the Eye, One of the many points of superiority which the eye pos- sesses over the ordinary camera obscura is derived from its spherical shape, adapting the retina to receive every portion of the images produced by refraction, which are themselves curved: whereas, had they been received on a plane surface, as they usually are in the camera obscura, a considerable portion of the image would have been indistinct. This sphe- rical form is preserved by means of the firm membranes which protect the eye, and which are termed its Coats; and the transparent media which they enclose, and which effect 326 THE SENSORIAL FUNCTIONS. the convergence of the rays, are termed the Humours of the Eye. There are in tliis organ three principal coats, and three humours, composing altogether what is called the Globe of the Eye. Fig. 415, which gives an enlarged view of a liorizontal section of the right eye, exhibits distinctly all these parts. Tiie outermost coat (s,) which is termed the Sclerotica, is exceedingly firm and dense, and gives to the globe of the eye the mechanical support it requires for the performance of its delicate functions. It is perforated behind by the op- tic nerve (o,) which passes onwards to be expanded into the retina (ii.) The sclerotica docs not extend farther than about four-fifths of the globe of the eye; its place in front being supplied by a transparent convex membrane (c,) called the Cornea, which is more prominent than the rest of the eye- ball. A line passing through the centre of the cornea and the centre of the globe of the eye, is called the axis of the eye. The Sclerotica is lined internally by the Choroid coat VISION. 327 (x,) which is chiefly made up of a tissue of blood vessels, for supplying nourishment to the eye. It has on its inner surface a layer of a dark coloured viscid secretion, known by the name of the Pigrnenliim nigritvi, or black pigment. Its use is to absorb all the light which may happen to be ir- regularly scattered througli the eye, in consequence of re- flection from different quarters; and it serves, theiefore, the same purpose as the black paint with which the inside of optical instruments, sucli as telescopes, microscopes, and ca- meraB obscurae, is darkened. Within the pigmentum nigrum, and almost in immediate contact with it,* the Retina (r) is expanded, forming an exceedingly thin and delicate layer of nervous matter, supported by a fine membrane. More than three-fourths of the globe of the eye are filled with the vilreous humour (v,) which has the appearance of a pellucid and elastic jelly, contained in an exceedingly de- licate texture of cellular substance. The Crystalline hu- motcr, (l,) which has the shape of a double convex lens, is formed of a denser material than any of the other humours, and occupies the fore part of the globe of the eye, immedi- ately in front of the vitreous humour, which is there hol- low^ed to receive it. The space which intervenes between the lens and the cornea is filled with a watery secretion (a,) called the tdqueous /tumour. This space is divided into an anterior and a posterior chamber by a flat circular partition (i,) termed the Iris. The iris has a central perforation (p,) called the Pupil, and it is fixed to the edge of the choroid coat, by a white elastic ring (q,) called the Ciliary Ligament. The poste- rior surface of the iris is called the Uvea, and is lined with a dark brown pigment. The structure of the iris is very peculiar, being composed of two layers of contractile fibres; the one, forming concentric circles; the other, disposed like radii between the outer and inner margin.! When the * Between the pigmentum and the retina there is found a very fine mem- brane, discovered by Dr. Jacobson: its use has not been ascertained, .f See Fig. 47, vol. i.p. 105. 323 THE SENSORIAL FUNCTIONS. former act, the pupil is contracted: when the latter act, the breadth of tlie iris is diminished, and the pu])il is, of course, dilated. By varying the size of the pupil the quantity of li^ht admitted into tlie interior of the eye is regidated, and accommodated to tlie sensibility of the retina. When the intensity of the light would be injurious to that highly deli- cate organ, the pupil is instantly contracted, so as to exclude the greater portion; and, on the contrary, when the light is too leeblc, it is dihUed, in order to admit as large a quantity as possible. The iris also serves to intercept such rays as woukl have fallen on parts of the crystalline lens less fitted to produce their regular refraction, the object of which will be better understood when we have examined the functions of this latter part. But, before engaging in this inquiry, it will be proper to complete this sketch of the Anatomy of the Eye by describing the principal parts of the apparatus belonging to that organ, which are exterior to the eye-ball, and may be considered as its appendages. The purposes answered by the parts exterior to the eye- ball are chiefly those of motion, of lubrication, and of pro- tection. As it is the central part of the retina which is endowed with tlic greatest share of sensibility, it is necessary that the images of the objects to be viewed should be made to fail on this part; and, consequently, that the eye should be ca- pable of having its axis instantly directed to those objects, wherever they may be situated. Hence, muscles are pro- vided within the orbits, for effecting the motions of the eye- ball. A view of these muscles, with their attachments to the ball of the eye, but separated from the other parts, is given in Fig. 416. Four of these proceed in a straight course from the bottom of the orbit, arising from the margin of the aperture through which the optic nerve passes, and being inserted by a broad tendinous expansion into the VISION. 329 fore part of the sclerotic coat. Three of these arc marked A, B, and c, in the figure: and the edge of the fourth is seen behind and above b. These straight muscles, as they are called, surround the optic nerve and the eye-hall, forming four longitudinal bands; one (a) being situated above for the purpose of turning the eye upwards; a second (c,) situated below, for turning it downwards; and the two others, on either side, for performing its lateral motions to the riglit or left. The cavity of the orbits being considera])ly larger tlian the eye-ball, the intervening space, especially at the back part, is filled up by fat, which serves as a soft cushion for its protection, and for enabling it to roll freely in all direc- tions. J3esides these straight muscles, there are also two others (s and i) termed the ohlique muscles, which give the eye- ball a certain degree of rotation on its axis. When these act in conjunction, they draw the eye forwards, and serve as an- tagonists to the combined power of the straight muscles. The upper oblique muscle (s) is remarkable for the artificial manner in which its tendon passes through a cartilaginous pulley (p) in the margin of the orbit, and then turns back again to be inserted into the eye-ball, so that the effect pro- duced by ^he action of the muscle is a motion in a direction exactly the reverse of that in which its fibres contract. This mechanism, simple as it is, afibrds one of the most palpable instances that can be adduced of express contrivance; for in no other situation could the muscle have been so conve- niently lodged as within the eye-ball; and in no other way could its tendon have been made to pull in a direction con- trary to that of the muscle, than by the interposition of a pulley, turning the tendon completely round. The fore-part of the globe of the eye, which is of a white colour, is connected with the surrounding integuments by a membrane, termed the Conjunctiva* This membrane, on * An abundant supply of nerves has been bestowed on this membrane for the purpose of conferring upon it that exquisite degree of sensibility which Vol. II. 42 330 THE SEKSOllIAL FUNCTIONS. arriving at the base of the eye-lids, is folded forwards so as to line their inner surfaces, and to be continuous with the skin whicli covers their outer sides. The surfaces of the conjunctiva and of tiie cornea are kept constantly moist by the tears, which are as constantly secreted by the Lacry- mal inlands. Each i»;hind, (as shown at l, Fig. 417,) is si- tuated above the eye, in a hollow of the orbit, and the ducts (d) proceeding from it open upon the inner side of the up- per eye-lid (e.) This fluid, the uses of which ar(f obviously to wash away dust, or other irritating substances which may happen to get introduced, is distributed over the outer sur- face of the eye by means of the eye-lids. Each lid is sup- ported by an clastic plate of cartilage, shaped like a cres- cent, and covered by integuments. An orbicular muscle, the fibres of which run in a circular direction, immediately underneath the skin, all round the eye,* is provided for closing them. The upper eye-lid is raised by a separate muscle, contained within the orbit, immediately above the was necessary to give immediate warning of the slightest clanger to so im- portant an organ as the eye from the intrusion of foreign bodies. That this is the intention is apparent from the fact that the internal parts of the eye possess but little sensibility compared with the external surface. • Sec Fig. 46, vol. i. p. 105. VISION. * 331 upper straight muscle of the eye-ball. The eye-lashes are curved in opposite directions, so as not to interfere with each other when the eye-lids are closed. Their utility in guarding the eye against the entrance of various substances, such as hairs, dust, or perspiration, and also in shading the eye from too strong impressions of light, is sufiiciently ap- parent. The eye-lids, in closing, meet first at the outer corner of the eye; and their junction proceeds along the line of their edges, towards the inner angles, till the contact is complete: by this means the tears are carried onwards in that direction and accumulated at the inner corner of the eye, an effect which is promoted by the bevelling of the margins of the eye-lids, which, when they meet, form a channel for the fluid to pass in that manner. When they arrive at the inner corner of the eye, the tears arc conveyed away by two slender ducts, the orifices of which, called the puncta lacrymalia (p, p,) are seen at the inner corner of each eye-lid, and are separated by a round projecting body (c,) connected with a fold of the conjunctiva, and termed the lacryrnal caruncle. The two ducts soon unite to form one passage, which opens into a sac (s,) situated at the upper part of the sides of the nose, and terminating be- low (at n) in the cavity of the nostrils, into which the tears are ultimately conducted. When the secretion of the tears is too abundant to be carried off by this channel, they over- flow upon the cheeks; but when the quantity is not exces- sive, the tendency to flow over the eye-lid is checked by an oily secretion proceeding from a row of minute glands, si- tuated at the edge of the eye-lids, and termed the Meibo- mian glands. The eye-brows arc a fiU'ther protection to the eyes, the direction of the hairs being such as to turn away from them any drops of rain or of perspiration which may ciiancc to fall from above. Excepting in front, where tlie eyes are covered and pro- tected jjy the eye-lids, these important organs are on all 332 THE SENSORIAL FUNCTIONS. sides effectually guarded from injury by Ijeing contained in a hollow bony socket, termed the orbit, and composed of seven portions of bone. Tliese seven elements may be re- cognised in the skulls of all the mammalia, and perhaps also in those of all other vertebrated animals, affording a remark- able illustration of the unity of the plans of nature in the construction of the animal fabric. § 4. Physiologi/ of perfect Vision. The rays of light, proceeding from a distant object, strike upon the convex surface of the cornea, which being of great- er density than the air, refracts them, and makes them con- verge towards a distant focus. This effect, however, is in part counteracted on their emergence from the concave pos- terior surface of the cornea, when the rays enter into the aqueous humour. On the whole, however, they are refract- ed, and made to converge to a degree equal to that which they would have undergone if they had at once impinged against the convex surface of the aqueous humour, supposing the cornea not to have been interposed. A considerable portion of the light which has thus en- tered the aqueous humour is arrested in its course by the iris; so that it is only those rays which are admitted through the pupil that are subservient to vision. These next arrive at the crystalline lens, where they undergo two refractions, the one at the anterior, the other at the posterior surface of that body. Both these surfaces being convex outwardly, and the lens being a denser substance than either the aque- ous or the vitreous humours, the effect of both these refrac- tions is to increase the convergence of the rays, and to bring them to unite in a focus on the retina at the bottom of the eye. The most considerable of these refractions is the first; because the difference of density between the air and the cornea, or rather the aqueous humour, is greater than that of any of the humours of the eye compared with one ano- ther. VISION. 333 The accurate converp;cnce of all the rays of light, which enter through the pupil, to their respective foci on the reti- na, is necessary for the perfection of the images there formed; but, for the complete attainment of this end, various nice ad- justments are still requisite. In the first place, the Aberration of Sphericity * which is a consequence of the geometrical law of refraction, intro- duces a degree of confusion in the image; which is scarcely perceptible, indeed, on a small scale, but which becomes sensible in instruments of much power; being one of the greatest difficulties which the optician has to overcome in the construction of the telescope and the microscope. Na- ture, in framing the human eye, has solved this difficulty by the simplest, yet most effectual means, and in a manner quite inimitable by human art. She has, in the first place, given to the surfaces of the crystalline lens, instead of the spheri- cal form, curvatures more or less hyperbolical or elliptical; and has, in the next place, constructed the lens of an infinite number of concentric layers, which increase in their densi- ty, as they succeed one another from the surface to the cen- tre. The refracting power, being proportional to the density, is thus greatest at the centre, and diminishes as we recede from that centre. This admirable adjustment exact- ly corrects the deficiency of refraction, which always takes place in the central portions of a lens composed of a mate- rial of uniform density, as compared with the refraction of the parts more remote from the centre.! The second adjustment for perfect vision has reference to the variations in the distance of the focus which take place according as the rays arrive at the eye from objects at diffe- rent distances, and which may be called the Merrations of * See Fig. 411, and the note referring to it, p. 324. ■j- Sir David Brewster has ascertained that the variations of density pro- ducing- the doubly refracting structure, in the crystalline lens of fishes, arc related, not to the centre of the lens, but to the diameter which forms the axis of vision: an arrangement peculiarly adapted for correcting the splicri- cal aberrations. Philos. Trans, for 1810, p. 317. 334 THE SENSORIAL FUNCTIONS. Parallax. When the distance of the object is very great, the rays proceeding from each point arrive at the eye with so little divergence, that each pencil may be considered as composed of rays which arc parallel to each other; the ac- tual deviation from parallelism being quite insensible. But if the same object be brought nearer to the eye, the diver- gence of the rays becomes more perceptible; and the effect of the same degree of refraction is to collect them into a focus more remote than before.* For every distance of the object there is a corresponding focal distance; and when the eye is in a state adapted for distinct vision at one distance, it will have confused images of objects at another distance; because the exact foci of the rays will be situated either be- fore or behind the retina. It is evident that if the retina be not placed exactly at the point where the focus is situated, it will either intercept the pencil of rays before they are united into a point, or receive them after they have crossed one another in passing through the focus: in either of which cases, each pencil will throw upon the retina a small circle of light, brighter at the middle and fainter at the edges, which will mix itself with the adjacent pencils, and create confusion in thd image. It is found, however, that the eye has a power of accommo- dating itself to the distinct vision of objects at a great variety of distances, according as the attention of the mind is di- rected to the particular object to be viewed. The mode in which this change in the state of the eye is effected has been the subject of much controversy. The increase of the re- fracting power of the eye necessary to adapt it to the vision of near objects is evidently the result of a muscular eifort, of which we are distinctly conscious when we accurately at- tend to the accompanying sensations. The researches of • This is illustrated by Fig. 412, 413, and 414; the first of which shows the rapid convergence of rays proceeding from a veiy distant object, and which may be considered as parallel. The second shows that divergent i-ays unite at a more distint focus; and the third, that the focus is more distant the greater the divergence. VISION. 335 Dr. Young have rendered it probable that some change takes place in the figure of the lens, whereby its convexity, and perhaps, also, its distance from the retina, are increased. He has shown, by a very decisive experiment, that any change which may take place in the convexity of the cornea has but little share in the production of the effect; for the eye retains its power of adaptation when immersed in water, in which the form of the cornea can in no respect influence the refraction. But the rays of light are of different kinds; some exciting the sensation of red, others of yellow, and others again of blue; and these different species of light are refracted, under similar circumstances, in different degrees. Hence, the more refrangible rays, namely, the violet and the blue, are brought to a nearer focus, than those which are less refran- gible, which are the orange and-the red rays: and this want of coincidence in the points of convergence of these different rays, (all of which enter into the composition of white light,) necessarily impairs the distinctness of all the images pro- duced by refraction, shading off their outlines with various colours, even when the object itself is colourless. This de- fect, which is incident to the power of a simple lens, and which is termed the Chromatic Aberration, is remedied almost perfectly in the eye, by the nice adjustment of the powers of the different refracting media, which the rays of light have to traverse before they arrive at the retina, pro- ducing what is called an achromatic combination;^' and it is found that the eye, though not an absolutely achromatic in- strument, as was asserted by Euler,t is yet sufficiently so for all the ordinary practical purposes of life. The object, then, of the whole apparatus appended to the optic nerve, is to form inverted images of external objects on the retina, which, as we have seen, is the expanded ex- * For the exposition of the principles on which these achromatic combi- nations of lenses correct this source of aberration, I must refer to works which treat professedly on Optics. f For the rectification of this error we are indebted to Ur. Young. 336 THE SENSORIAL FUNCTIONS. tremity of that nerve. That this cflcct is actually produced, may be easily shown by direct observation; for if the scle- rotic and choroid coats be carefully dissected off from the posterior part of the eye of an ox, or any other large quad- ruped, leaving ordy the retina, and tlic eye so prepared be j)laced in a hole in a window-shutter, in a darkened room, with the cornea on the outside, alL the illuminated objects of the external scene will be beautifully depicted, in an in- verted j)osition, on the retina. Few spectacles are more calculated to raise our admira- tion than this delicate picture, which nature has, with such exquisite art, and with the finest touches of her pencil, spread over the smooth canvass of this subtle nerve; a pic- ture, which, though scarcely occupying a space of half an inch in diameter, contains the delineation of a boundless scene of earth and sky, full of all kinds of objects, some at rest, and others in motion, yet all accurately represented as to their forms, colours and positions, and followed in all their changes, without the least interference, irregularity, or confusion. Every one of those countless and stupendous orbs of fire, whose light, after traversing immeasurable re- gions of space, at length reaches our eye, is collected on its narrow curtain into a luminous focus of inconceivable mi- nuteness; and yet this almost infinitesimal point shall be sufllcient to convey to the mind, through the medium of the optic nerve and brain, a knowledge of the existence and po- sition of the far distant luminary, from which that light has emanated. How infinitely surpassing all the limits of our conception must be the intelligence, and the power of that Being, who planned and executed an instrument comprising, within such limited dimensions, such vast powers as the eye, of which the perceptions comprehend alike the nearest and most distant objects, and take cognizance at once of the most minute portions of matter, and of bodies of the largest magnitude ! VISION. 337 § 5. Comparative Physiology of Vision. In the formation of every part of the animal machinery we may generally discern the predominance of the law of gradation; but this law is more especially observed in those organs which exhibit, in-their most perfect state, the great- est complication and refinement of structure: for on follow- ing all their varieties in the ascending series, we always find them advancing by slow gradations of improvement, before they attain their highest degree of excellence. Thus, the organ of vision presents, amidst an infinite variety of con- structions, successive degrees of refinement, accompanied by corresponding extensions of power. So gradual is the pro- gress of this development, that it is not easy to determine the point where the faculty of vision, properly so called, be- gins to be exercised, or where the first rudiment of its or- gan begins to appear. Indications of a certain degree of sensibility to light are afibrded by many of the lower tribes of Zoophytes, while no visible organ appropriated to receive its impressions can be traced. This is the case with many microscopic animalcules; and still more remarkably with the Hydra, and the Actinia, which show by their movements that they feel the influ- ence of this agent; for, when confined in a vessel, they al- ways place themselves, by preference, on the side where there is the strongest light.* The Veretillum cynomo- rium, on the other hand, seeks the darkest places, and con- tracts itself the moment it is exposed to light.t In a per- fectly calm sea, the Medusae which are rising towards the surface, are seen to change their course, and to descend again, as soon as they reach those parts of the water which receive the full influence of the sun's rays, and before any part of * Such is the uniform report of Trembley, Baker, Bonnet, Goeze, Ha- now, RcEsel, and Schseffer. f Rapp; Nov. Act. Acad. Nat. Cur. of Bonn, xlv. 645. Vol. II. 43 33S THE SENSORIAL FUNCTIONS. their bodies has come into contact with the atmosphere.* But, in all these instances a doubt may arise whether the observed actions be not prompted by the mere sensation of warmth excited by calorific rays which accompany those of light; in which case they would be evidence only of the operation of a finer kind of touch. The first unequivocal appearance of visual organs is met with in the class of Annelida; although the researches of Ehrenberg would induce us to believe that they may be traced among animals yet lower in the scale; for he has no- ticed them in several of the more highly organized Infuso- ria, belonging to the order Rotifera, and particularly in the Hydatina senta, where he has found the small black points observable in other species, united into a single spot of larger size. Nitsch, also, states that the Cercaria viridis, possesses three organs of this kind. Planarix present two or three spots, which have been regarded as visual organs; and these have been found by Baer to be composed, in the Planaria torva, of clusters of black grains, situated underneath the white or transparent integu- ment.! The eyes of the Nais pi^oboscidea are composed, ac- cording to Gruithuisen, simply of a small mass of black pig- ment, attached to the extremity of the optic nerve ;| and or- gans apparently similar to these are met with in many of the inferior tribes of Annelida. In all these cases it is a matter of considerable doubt whether the visual organs are constructed with any other intention than merely to con- vey general sensations of light, without exciting distinct per- ceptions of the objects themselves from which the light pro- ceeds; this latter purpose requiring, as we have seen, a spe- cial optical apparatus of some degree of complexity. An ap- proach to the formation of a crystalline lens takes place in the genus Eiuiice of Cuvier, {Lycoris, Sav.,) which, from the • Grant; Edin. Journal of Science: No. 20. f Nov. Act Acad. Nat. Cur. of Bonn, xiii. 712. See also the Memoir of Dugt'S, entitled *' Recherches sur I'Organisation etles Mocui'sdes Planaires," in the Annales dcs Sc. Nat. xv. 148. i Nov. Act. Acad. N.it. Cur. of Bonn, xi. 242. VISION. 339 account given by Professor MuUer,* has four eyes, situated on the hinder part of the head, and covered with the epidermis, but containing in their interior a spherule, composed of an opaque white substance, surrounded by a black pigment, and penetrated by an optic nerve, which is continued to the brain. On the other hand. Professor Weber found in the Hirudo medicinalis, or common leech, no less than ten minute eyes, arranged in a semicircle, in front of the head, and project- ing a little from the surface of the integument: they present externally a convex, and perfectly transparent cornea; while internally, they are prolonged into cylindrical tubes, con- taining a black pigment;! structures, apparently subservient to a species of vision of a higher order than that which con- sists in the simple recognition of the presence of light. No organs having the most distant relation to the sense of vision, have ever been observed in any of the Acephalous, or bivalve MoUusca; but various species of Gasteropoda have 418 organs which appear to exercise this sense, situated sometimes at the base, sometimes at the middle, and frequently at the ex- tremity of the tentacula. Of the latter we have examples in the common slug and snail, where these tentacula, or horns, are four in number, and are capable of being protruded and again retracted, by folding inwards like the finger of a glove, at the pleasure of the animal. According to Muller,J the eye of the Helix pomatia, represented at e, (Fig. 41S;) is situ- ated a little to one side of the rounded extremity, or papilla (p,) of the tentaculum, and is attached to an oval bulb of a • Annales des Sciences Naturelles, xxii. 23, f Meckel, Archiv. fiir Anatomic und Physiologic; 1824, p. 301. t Annales des Sciences Naturelles; xxii. 12. 340 THE SENSORIAL FUNCTIONS. black colour. It receives only a slender branch (o) from a large nerve (n n) which is distributed to the papilla of the tentaculum, and appears to be appropriated exclusively to the sense of touch. The bulb, with the eye attached to it, is represented, in this figure, as half retracted within the tu- bular sheath of the tentaculum (s s;) but it can exercise its proper function only when fully exposed, by the complete unfolding and protrusion of the tentaculum. This eye con- tains, within its choroid coat, a semi-fluid and perfectly transparent substance, filling the whole of the globe; and JNIuUer also discovered at the anterior part, another transpa- rent body, having the shape of a lens.* A structure very similar to this was found to exist in the eye of the Murex iritonis, with the addition of a distinct iris, perforated so as to form a pupil; a part which had also been observed, to- gether with a crystalline lens of very large size, in the Va- luta cymbium, by De Blainville.t Thus, the visual organs of these Gasteropoda appear to possess every requisite for dis- tinct vision, properly so called. Experiments are said to have been recently made, both by Leuchs, and by Steifen- sand,t in which a snail was repeatedly observed to avoid a small object presented near the tentaculum; thus affording evidence of its possessing this sense. The accurate investigation of the anatomy of the eyes of insects presents considerable difficulty, both from "the mi- nuteness of their parts and from the complication of their structure; so that notwithstanding the light which has re- cently been thrown on this interesting subject by the patient and laborious researches of entomologists, great obscurity still prevails with regard to the mode in which these dimi- • MtiUer thus confirms the accuracy of Swammerdam's account of the anatomy of the eye of the snail, which had been contested by Sir E. Home (Phil. Trans. 1824, p. 4) and other writers. ■j- Principes d'Anatomic Comparce, i. 445. % Quoted by MuUer; ibid. p. 16. These results also corroborate the testi- mony of Swammerdam, who states that he had obtained proofs that the snail could see by means of these organs. VISION. 341 nutive beings exercise the sense of vision. Four descrip- tions of visual organs are met with in the class of Articu- lated Animals; the first are the simple eyes, or siemmata, as they are termed, which appear as lucid spots, resembling those we have noticed in the higher orders of Annelida; the second, are the conglomcrale eyes, which consist of clus- ters or aggregations of simple eyes; the third, are the com- "pound eyes, which are formed of a vast assemblage of small tubes, each having its respective apparatus of humours, and of retina, and terminating externally in a separate cornea, slightly elevated above the general surface of the organ: the fourth kind of eyes, which have not yet been distinguished by any particular appellation, are constituted by a number of separate lenses, and subjacent retinae, but the whole co- vered by a single cornea common to them all. Few insects are wholly destitute of visual organs, either in their larva or perfect states.^ The larvae of those insects which undergo a complete metamorphosis have only stem- mata; but those which are subjected only to a partial change of form, as the Orthoptera, the Hemiptera, and the aquatic Neuroptera, have compound as well as simple eyes. Perfect insects, with the few exceptions above noticed, have always compound eyes, generally two in number, placed on the sides of the head: and they are often accompanied by stem- mata situated between, or behind them, on the upper part of the head. These stemmata, when met with, are generally three in number, and are either placed in a row, or form a triangle. Their structure has been minutely examined by Professor Muller, who found them to contain a hard and spherical crystalline lens, a vitreous hum.our, and a choroid coat, with its accompanying black pigment; the whole being covered externally by a convex cornea. The stemmata of * This is the case, however, with the genus Claviger, among the Coleoptera Braula (Nltzch) among Diptera, and also some of the species of Pupipara, Nyderibia, and 31elophagus, which are all parasitic insects: there are also five species of ants, whose neuters have no eyes. (Muller, Annales dcs Sc. Nat. xvii. 366,) 342 THE SENSORIAL FUNCTIONS. a caterpillar, which has eight of these eyes, are shown in Fig. 419, connected together by a circular choroid membrane o o o e o o o o o o o o e o o O o o e o o O o o o o O o (x x) common to the whole: together with the separate branches (o o) of the optic nerve (n) belonging to each. All the Arachnida possess eyes of this latter description; and from their greater size afford facilities for dissection, which are not met with among proper insects. Their num- ber in Spiders is generally eight, and they are disposed with great symmetry on the upper side of the head. Fig. 420 represents, on a magnified scale, one of the large stemmata, on the head of the Scorpio tunensis, dissected so as to dis- play its internal parts; in which are seen the cornea (c,) de- rived from an extension of the integument (i;) the dense spherical crystalline lens (l;) the choroid coat with its pig- ment (x,*) forming a wide opening, or pupil; the vitreous hu- mour (v,) covered behind the retina (r,) which is closely ap- plied to it; and the optic nerve (o,) with which the retina is continuous. Examples of the conglomerate eye occur in the Myriapo- da: in the Scolopendra, for instance, they consist of about twenty contiguous circular pellucid lenses, arranged in five lines, with one larger eye behind the rest, which Kirby com- pares to a sentinel, or scout, placed at some little distance from the main body. In the Julus terrestriSi or common Millepede, these eyes, amounting to 28, form a triangle, be- • Marcel dc Serres states, that some of the stemmata of the insects which he examined contain a thin choroid, having a silvery lustre, as if intended as a reflector of the light which falls on it. VISION. 343 ing disposed in seven rows, the number in each regularly diminishing from the base to the apex; an arrangement which is shown in Fig. 421.* The compound eyes of insects arc formed of a vast num- ber of separate cylinders or elongated cones,t closely packed together on the surface of a central bulb, which may be con- sidered as a part of the optic nerve; while their united bases or outer extremities constitute the surface of a hemispherical convexity, which often occupies a considerable space on each side of the head. The usual shape of each of these bases is that of a hexagon, a form which admits of their uni- form arrangement with the greatest economy of space, like the cells of a honey-comb; and the hexagonal divisions of the surface are very plainly discernible on viewing the sur- face of these eyes with a microscope, especially as there is a thin layer of black pigment intervening between each, like mortar between the layers of brick. The appearance they present in the Melolontha, when highly magnified, is shown in Fig. 422.% The internal structure of these eyes will be best understood from the section of that of the Lihelhda vulgata, or gray Dragon-fly, shown in Fig. 424, aided by the highly magnified views of smaller portions given in the succeeding figures, in all of which the same letters of refe- rence are used to indicate the same objects.^ The whole outer layer (c c) of the compound eye may be considered • Kirby and Spence's Introduction, Sec, ill. 494. f The number of these cones or cylinders which compose the entire org^n differs much in different species. In the ant, there are only 50; in a Scara- baeus, 3180; in the Bomhyxmori^ 6236; in the house-fly (Musca domestica,) 8000; in the Melolontha vulgaris, 8820; in the Phalena cossus, 11,300; in the Libellula, 12,544; in the Papilio, 17,325; and in the Mordella, 25,088. ^ In the Fhaknae, and other tribes, they are arranged in squares (as shown in Fig-. 423,) instead of hexagons, and frequently much less regularly; as must necessarily happen, in many parts, from the curvature of the spherical surface. § These figures, as well as the account of the anatomy of the eye of the Libellula, are taken from the memoir of Duges, in the Annales des Science* Naturelles, xx. 341. 344 THE SENSORIAL FUNCTIONS. as corresponding to the cornea: each separate division of which has heen termed a Corneicle, heing composed of a horn}^ and perfectly transparent material. Each corneule 422 « • # .# 1i 4 |# # # • # # # • r 423 424 ^^A J^O ^ 1 ^ «y-^MMJW» (c) has the form of a truncated pyramid, the length of which (l) is between two and three times the diameter of the base (b.) The outer surface (b) is very convex; but the internal, or truncated end (d) is concave; and the concavity of the latter being smaller than the con- vexity of the former, its optical effect is that of a menis- cus, or concavo-convex lens, with power of converging to a distant focus the rays of light which traverse it. With- in these corneules there is extended a layer of an opaque VISION. 345 black pigment (x,) probably connected with a choroid coat, which, from the delicacy of its texture, has hitherto escaped observation. There exists opposite to the centre, or axis of each corneule, a circular perforation (p,) which performs the functions of a pupil.* Duges states, in- deed, that he has witnessed in this part movements of con- traction and dilatation, like those of the iris in vertebrated animals. He has likewise found that tliere is a small space (a) intervening between the extremity of each corneule and the iris, and filled with an aqueous humour. The compart- ments formed by the substance of the choroid (x) are conti- nued inwards towards the centre of the general hemisphere, the cylindrical spaces which they enclose being occupied each by a transparent cylinder (v,) consisting of an outer membrane, filled with a viscid substance analogous to the vi- treous humour. Their general form and situation, as they lie embedded in the pigment, may be seen from the magnified sections; each cylinder commencing by a rounded convex base, immediately behind its respective pupil, and slightly tapering to its extremities, where it is met by a filament (n) of the optic nerve; and all these filaments, after passing for a certain distance through a thick mass of pigment, are united to the large central nervous bulb (g, Fig. 427,) which is termed the optic ganglion.^ * This pupillary aperture was discovered by Muller, and had eluded all the efforts of former observers to detect it; and it was accordingly the pre- vailing notion that the black pigment lined the whole surface of the cornea, and interposed an insuperable barrier to the passage of light beyond the cornea. It was evidently impossible, while such an opinion was entertained, that any intelligible theory of vision, with eyes so constructed, could be formed. + Numberless modifications of the forms of e^h of these constituent parts occur in different species of insects. Very frequently the vitreous humour (v,) instead of forming an elongated cylinder, has the shape of a short cone, terminating in a fine point, as shown in Fig. 426. Straus Durckheim ap- pears to have mistaken this part for an enlarged termination of the optic nerve, believing it to be opaque, and to form a retina applied to the back of the corneule, which latter part he considered as properly the crystalline lens. In his elaborate work on the anatomy of the Melolontha, he describes the Vol. II. 44 346 THE SENSORIAL FUNCTIONS. It thus appears that each of the constituent eyes, which compose this vast agji;regate, consists of a simple tube, fur- nished with all the elements requisite for distinct vision, and capable of receivinsj; impressions from objects situated in the direction of the axis of the tube. The rays tra- versing adjacent corneules are prevented from mixing them- selves with those which arc proper to each tube by the in- terposition of the black pigment, which completely surrounds the transparent cylinders, and intercepts all lateral or scat- tered light. Thus has nature supplied the want of mobility in Ihc eyes of insects, by the vast multiplication of their number, and by providing, as it were, a separate eye for each separate point which was to he viewed; and thus has she realized the hypothetical arrangement, which suggested itself in the outset of our inquiries, while examining all the possible modes of eficcting this object. This mode of vision is probably assisted by the converging powers of each corneule, although in parts which are so mi- nute it is hardly possible to form an accurate estimate of these powers by direct experiment. In corroboration of this view I am fortunately enabled to cite a valuable observation of the late Dr. Wollaston, relative to the eye of the Aslacus Jhiviatilis, or cray-fish, where the length of each compo- nent tube is short, compared with that of the Libellula. On measuring accurately the focal distance of one of the cor- neules. Dr. Wollaston ascertained that it corresponds with filaments (f) of the optic nerve, in tlieir progress inwards, as passing through a second membrane (k, Fig. 428,) which he denominates the common cho- roid, and afterwards uniting to form an expanded layer, or more general re- tina (h,) whence proceed a small number of short but thick nervous co- lumns (>',) still converging towards the large central ganglion (g,) in which they terminate. The use he ascribes to this second clioroid is to intercept the light, which, in so diminutive an organ, might otherwise penetrate to the general retina and produce confusion, or injurious irritation. The co- lour of the pigment is not always black, but often has a bluish tint: in the common fly, it is of a bright scarlet hue, resembling blood. In nocturnal insects the transvci-sc layer of pigment between the corneule and the vitre- ous humour is absent. VISION. 347 great exactness to the length of the tul)e attached to it; so that an image of an external ohject is formed precisely at the point where the retina is placed to receive it* Little is known of the respective functions of these two kinds of eyes, the simple and the compound, both of which are generally possessed by the higher orders of winged in- sects. From the circumstance that the compound eyes are not developed before the insect acquires the power of flight, it has been inferred that they are more particularly adapted to the vision of distant objects; but it must be confessed that the experiments made on this subject have not, hitherto, led to any conclusive results. Duges found, in his trials, that after the stemmata had been covered, vision remained appa- rently as perfect as before, while, on the other hand, when insects were deprived of the use of the compound eyes, and saw only with the stemmata, they seemed to be capable of distinguishing nothing but the mere presence or absence of light. Others have reported, that if the stemmata be co- vered with an opaque varnish, the insect loses the power of guiding its flight, and strikes against walls or other obsta- cles: whereas, if the compound eyes be covered while the stemmata remain free, the insect generally flies away, rising perpendicularly in the air, and continuing its vertical ascent as long as it can be followed by the observer. If all tho eyes of an insect be covered, it will seldom make any at- tempt whatsoever to fly. The eyes of insects, whether simple or compound, are immoveably fixed in their situations; but the compound eyes of the higher orders of the class of Crustacea, arc placed at the ends of moveable pedicles, so as to admit of being turned at pleasure towards the objects to be viewed.f This, how- * This interesting fact was communicated to me by Captain Kater, who, together with Mr. Children, assisted Dr. Wollaston in this examination. f Latreille describes a species of Crab, found on the shores of the Medi- terranean, having its eyes supported on a long jointed tube, consisting of two articulations, which enables the animal to move them in various direc- tions, like the arms of a telegraph. 348 THE SENSORIAL FUNCTIONS. ever, Is not the case with the Entomostraca^ comprising the various species of Monocidi, in which the two eyes are brought so close to one another, as apparently to constitute a sini!;lc organ, corresponding in its structure to the fourth class of eyes already enumerated; that is, the separate lenses it contains have a general envelope of a transparent mem- brane, or cornea. Muscles are provided for moviiTg the eye in its socket; so that we have here indications of an approach to the structure of the eye which prevails in the higher classes of animals. There is, however, a still nearer approximation to tlie latter in the eye of the Cephalopoda; for Sepix differ from all the tribes belonging to the inferior orders of mollusca in having large and efficient eyes, con- taining a hemispherical vitreous humour, placed immediate- ly before a concave retina, and receiving in front a large and higlily convex crystalline lens, which is soft at its exte- rior, but rapidly increases in density, and contains a nucleus of great hardness: there is also a pigmentum nigrum, and a distinct iris, with a kidney-shaped pupil. This eye is re- markable for the total absence of a cornea; the integuments of the head being continued over tlic iris, and reflected over the edges of the pupil, giving a covering to the external sur- face of the lens: there is, of course, no chamber for contain- ing an aqueous humour. The globe of the eye is nearly spherical, but the sclerotica is double, leaving, at the poste- rior part, between its two portions, a considerable space, oc- cupied by the large ganglion of the optic nerve, with its nu- merous filaments, which are embedded in a soft glandular substance.* The eyes of Fishes differ from those of sepiae principally in the addition of a distinct cornea, exterior to the lens and iris, but having only a slight degree of convexity. This, indeed, is the case with all aquatic animals; for, since the difference of density between the cornea and the external medium is but small, the refractive power of any cornea, • See Cuvier, sur lea Mollusques; Memoir sur Ic Poulpe, p. 37. In the Octopus there are folds of the skin, which appear to be nutiments of eye-lids. ^ VISION. 349 however convex, would be inconsiderable; and the chief agent for performing the requisite refraction of the rays is the crystalline lens. We, accordingly, in general, find the cornea nearly flat, and the globe of the eye approaching in shape to a hemisphere; while the lens itself is nearly sphe- rical, and of great density. The circumstances are shown in the section of the eye of the Ferch, Fig. 430.''' The flatness of the cornea leaves scarcely any space for aqueous humour, and but little for the motions of the iris. The surface of the eye in fishes, being continually washed by the water in which it is immersed, requires no provision 43Q of a secreted fluid for that purpose; and there are consequently neither lacrymal apparatus, nor proper eye-lids; the integu- ments supplying only a thin transparent membrane, which passes over and protects the cornea, serving the oilice of a conjunc- tiva. The eye retains its form by the support it receives from the sclerotic coat, which is of extraordinary thickness and density. In the Shay^k and the Skate the eye is sup- ported from the bottom of the orbit, by a cartilaginous pe- dicle, which enables it to turn as on a pivot, or lever. Sir David Brewster has recently made an interesting ana- lysis of the structure of the crystalline lens of the Cod, to which he was led by noticing some remarkable optical ap- pearances presented by thin layers of this substance when transmitting polarized light. He found that the hard cen- tral portion is composed of a succession of concentric, and perfectly transparent, spheroidal lamina}, the surfaces of which, though apparently smooth, have the same kind of iridescence as mother-of-pearl, and arising from the same cause; namely, the occurrence of regularly arranged lines, • In this figure, as in the others, c is tlic cornea; l, the lens; v, the vitreous humour; n, the retina; o, the optic nerve; and s, the sclerotica. There is also found in the eyes of most fishes an organ, lodged in the space k, termed the Choroid gland, which envelops the optic nerve, is shaped like a horse- shoe, is of a deep red colour, and highly vascular; its use is quite unknown. 350 THE SENSORIAL FUNCTIONS. or strixJ* These lines, which mark the edges of the sepa- rate fibres, composing each lamina, converge like meridians from the equator to the two poles of the spheroid, as is shown in Fig. 431. The fibres themselves arc not cylindri- cal, but flat; and they taper at each end as they approach the points of convergence. The breadth of the fibres in the most external layer, at the equator, is about the 5,500th part of an inch. The observation of another optical phenomenon, of a still more delicate kind, led Sir David Brewster to the farther discovery of the curious mode in which, (as is re- presented in Fig. 432,) the fibres are locked together at their edges by a series of teeth, resembling those of rack- work. He found the number of teeth in each fibre to be 12,500; and, as the whole lens contains about 5,000,000 fibres, the total number of these minute teeth amounts to 02,500,000,000. t Some fishes, which frequent the depths of the ocean, be- ing found at between three and four hundred fathoms below the surface, to which it is impossible that any sensible quan- tity of the light of day can penetrate, have, like nocturnal quadrupeds, very large eyes. J In a few species, which ♦ Sec vol. i. p. 169. f As far as his observations have extended, this denticulated structure ex- ists in the lenses of all kinds of fishes, and likewise in those of birds. He has also met with it in two species of Lizards, and in the Ornithorhyncus; but he has not been able to find it in any of the Mammalia, not even in the Ce- iacea. (Phil. Trms. for 1833, p. 323.) i See *' ObsciTations sur les Poissons recuciUis dans un Voyage aux lies Baleares ct Pythiuses. Par M. Delaroche." VISION. 351 dwell in the muddy banks of rivers, as the Csecilia, and Muraena cxca, or blind eel, the eyes are quite rudimental, and often nearly imperceptible; and in the Gasi7'ohranchus, De Blainvillc states that it is impossible, even by the most careful dissection, to discover the least trace of eyes. Reptiles, being destined to reside in air as well as in wa- ter, have eyes accommodated to these variable circumstances. By the protrusion of the cornea, and the addition of an aque- ous humour, they approach nearer to the spherical form than the eyes of fishes; and the lens has a smaller refractive pow- er, because the principal refraction is now performed by the cornea and aqueous humour. Rudiments of eye-lids are met with in the Salamander, but they are not of suffi- cient extent to cover the whole surface of the eyes. In some serpents, the integuments pass over the globe of the eye, forming a transparent conjunctiva, or external cornea, be- hind which the eye-ball has free motion. This membrane is shed, along with the cuticle, every time that the serpent is moulting; and at these epochs, while the cornea is pre- paring to detach itself, air insinuates itself underneath the external membrane and renders it opaque: so that until this operation is completed and an entire separation effected, the serpent is rendered blind. Serpents have no proper eye- lids; but the cornea is covered by a transparent integument, which does not adhere to it.* Lizards have usually a sin- gle perforated eye-lid, which, when closed by its orbicular muscle, exhibits merely a horizontal slit. There is also a small internal fold, forming the rudiment of a third eye-lid. The Chameleon has remarkably projecting eyes, to which the light is admitted through a very minute perforation in * It was the general opinion, until very lately, that serpents are unpro- vided with any lacrymal apparatus; but a small lacrymal passage has been recently discovered by Cloquet, leading from the space in the inner corner of the eye, between the transparent integument and the cornea. This la- crymal canal opens into the nasal cavity in venomous snakes, and into the moutli in those that are not venomous. 352 THE SENSORIAL FUNCTIONS. the skin constituting the outer eye-lid. This animal has the power of turning each eye, independently of the other, in a great variety of directions. The eyes of Tortoises exhibit an approach to those of birds: they are furnished with large lacrymal glands,. and with a very moveable memhrana nicitans or third oye-lid. Birds present a still fartlier development of all these parts: tlieir eyes are of great size compared with the head, as may be seen from the large portion of the skull wdiich is occupied on each side by the orbits. The chief peculiari- ties of the' internal structure of these organs are apparently designed to accommodate them to vision through a very rare medium, and to procure their ready adjustment to ob- jects situated at very dillerent distances. The form of the eye appears calculated to serve both these purposes; for the great prominence of its anterior portion, which has often the shape of a short cone, or cylinder, prefixed to the front of a hemispherical globe, and which is terminated by a very convex cornea, aflbrds space for a larger quantity of aqueous humour, and also for the removal of the lens to a greater distance from the retina, whereby the vision of near ol)jccts is facilitated, while at the same time, the refracting powers are susceptible of great variation. For the purpose of preserving the hemispherical form of the sclerotica, this membrane in birds is strengthened by a circle of bony plates, which occupy the fore-part, and are lodged between the two layers of which it consists. These plates vary in number from fifteen to twenty, and they lie close together, their edges successively overlapping each other. There is manifest design in this arrangement: for it is clear that a ring formed of a number of separate plates is better fitted to resist fracture than an entire bony circle of the same thickness. There is a dark-coloured membrane, called the Marsu- pium^ situated in the vitreous humour, the use of which is unknown, though it appears to be of some importance, as it VISION. 353 is found in almost every bird having extensive powers of vision.* The comparative anatomy of the eye offers, in- deed, a great number of special structures of which we do not understand the design, and which I have therefore pur- posely omitted to notice, as being foreign to the object of this treatise. In most birds the memhrana nictiians, or third eye-lid, is of considerable size, and consists of a semi-transparent fold of the conjunctiva, lying, when not used, in the inner cor- ner of the eye, with its loose edge nearly vertical: it is re- presented at N, Fig. 434, covering half riie surface of the eye: its motion, like that of a curtain, is horizontal, and is effected by two muscles: the first of which, seen at q, in Fig. 435, is called from its shape the quadratus^ and arises from the upper and back part of the sclerotica: its fibres de- scending in a parallel course towards the optic nerve, where 433 434 ^Zb they terminate, by a semicircular edge, in a tubular tendon. This tendon has no particular attachment, but is employed for the purpose of serving as a loop for the passage of the long tendon of the second muscle (p,) which is called the jjyramidalis, and which arises from the lower and back part of the sclerotica. Its tendon (t,) after passing through the • It is shown at m, Fig. 433, wliich is a mag-nified section of the eye of a Goose, c is the cornea; i, the iris; p, the ciliary processes, s, the sclerotic coat, and o, the optic nerve. Vol. II. 45 354 THE SENSORIAL FUNCTIONS. channel above described, which has the effect of a pulley, is conducted throuf^h a circular sheath, furnished by the scle- rotica to the under part of the eye, and is inserted into the lower portion of the loose edge of the nictitating membrane. lly the united action of these two muscles, the former of which serves merely to guide tlie tendon of the latter, and increase the velocity of its action, the membrane is rapidly drawn over the front of the globe. Its return to its former position is effected simply by its own elasticity, which is sufficient to bring it back to the inner corner of the eye.' If the membrane itself had been furnished with muscular fibres for effecting this motion, they would have interfered with its use by obstructing the transmission of light. Tlie eyes of quadrupeds agree in their general structure with those of man. In almost all the inferior tribes they are placed laterally in the head, each having independent fields of vision, and the two together commanding an extensive por- tion of the whole sphere. This is the case very generally among fislics, reptiles, and birds. Some exceptions, indeed, occur in particular tribes of the first of these classes, as in the Uranoscopus^ where the eyes are directed immediately upwards; in the /?f/y and the Caliionymus, where their di- rection is oblique; and in the Pleuroncctes, where there is a remarkable want of symmetry between the right and left sides of the bod}-, and where both eyes, as well as the mouth, are apparently situated on one side. Among birds, it is only in the tribe of Owls, which are nocturnal and predacious, that we find both eyes placed in front of the head. In the lower quadrupeds, the eyes are situated laterally, so that the optic axes form a very obtuse angle with each other. As we ascend towards the quadrumana we find this angle be- coming smaller, till at length the approximation of the ffelds of view of the two eyes is such as to admit of their being both directed to the same object at the same time. In the human species the axes of the two orbits approach nearer to parallelism than in any of the other mammalia; and the fields , • of vision of both eyes coincide nearly in their whole extent. VISION. 355. This is probably a circumstance of considerable importance with regard to our acquisition of correct perceptions by this sense. In the magnitude of the organ compared with that of the body, we may occasionally observe some relation to the cha- racter of the animal and the nature of its pursuits. Herbi- vorous animals, and especially those whose bulk is great, as the Elephant^ the Rhinoceros ^\^i\ tlie Hippopotamus^ have comparatively small eyes; for that of the elephant docs not exceed two inches in diameter. The eye of the Whale is not much more than the 200th part of the length of tlic body. When the natural food of an animal is stationary and re- quires no effort of pursuit, the eye is generally small, and the sight less keen; while in the purely carnivorous tribes, which are actively engaged in the chase of living prey, the organ of vision is large and occupies a considerable portion of the head; the orbit is much developed, and encroaches on the bones of the face; while, at the same time, the bony par- tition separating at the globe of the eye from the temporal muscle is supplied by ligament alone: so that when that muscle is in strong action, the eye is pressed outwards, giving to the expression of the countenance a peculiar fe- rocity. While nature has thus bestowed great acuteness of sight on pursuing animals, she has, on the other hand, been no less careful to arm those which are the objects of pursuit, with powers of vision, enabling them to perceive their enemies from afar, and avoid the impending danger. Thus, large eyes are bestowed on the Rodentia and the Ruminantia. Those tribes which pursue their prey by night, or in the dusk of the evening, as for example the Lemur and the Cat, are furnished with large eyes. Bats, however, form an ex- ception to this rule, their eyes being comparatively small; but a compensation has been afforded them in the superior acuteness of their other senses. In many quadrupeds a por- tion of the choroid coat is highly glistening, and reflects a 356 THE SENSORIAL FUNCTIONS. great quantity of coloured light: the object of this structure, which is termed the Tapeiiim, is not very apparent. Among the lesser quadrupeds which burrow in the ground, we find many whose eyes are extremely minute, so much so, indeed, as to be scarcely serviceable as visual organs. The eye of the Sorex, or shrew mouse, is very small, and surrounded by thick liair, which completely obstructs vi- sion, and requires to be removed by the action of the subcu- taneous muscles, in oi;der to enable the animal to derive any advantage from its eyes. These organs in the Mole are still more remarkably deficient in their development, not being larger than the head of a pin, and consequently not easily discovered.* It is therefore probable that this animal trusts chiefly to its sense of hearing, which is remarkably acute, for intimations of the approach of danger, especially as, in its subterranean retreats, the vibrations of the solid earth are readily transmitted to its ears. The Mus typhlus,ov blind rat of Linnaeus; (the Zemni of Pallas,) which is an inhabitant of the western parts of Asia, cannot be supposed to possess even the small degree of vision of the mole: for no external organ of this sense has been detected in any part of that ani- mal. The whole side of the head is covered with a conti- nuous integument of uniform thickness, and equally over- spread with a thick velvetty hair. It is only after removing the skin that a black spot is discovered on each side, of ex- ceedingly small size, and apparently the mere imperfect ru- diment of an eye, and totally incapable of exercising any of the functions of vision. Those mammalia, whose habits are.aquatic, having the eye frequently immersed in a dense medium, require a special provision for accommodating the refractive power of that organ to this variation of circumstances. Accordingly, it is found that in the Seal, and other amphibious tribes, the • Magcndie asserts that the mole has no optic nei*ve; but G. St. Hilaire and Carus recognise the existence of a very slender nci*vous filament, arising from tlic brain, and distributed to the eye of that animal. VISION. 357 structure of the eye approaches to that of fishes, the lens be- ing denser and more convex than usual, the cornea thin and yielding, and both the anterior and posterior segments of the sclerotic thick and firm; but the middle circle is very thin and flexible, admitting of the ready separation or ap- proximating of the other portions, so as to elongate or con- tract the axis of the eye;" just as a telescope can be drawn out or shortened, in order to adapt it to the distance of the object to be viewed. The whole eye-ball is surrounded by strong muscles which are capable of effecting these requisite changes of distance between the cornea and the retina. The Dolphin^ which lives more constantly in the water, has an eye still more nearly approaching in its structure to that of fishes; the crystalline lens being nearly spherical, and the globe of the eye furnished with strong and numerous mus- cles. In birds which frequently plunge their heads under water the crystalline lens is more convex than in other tribes; and the same is true, also, of aquatic reptiles. ( 35S ) CHAPTER VII. PERCEPTION. The object of nature in establishing the organizations we have been reviewing is to produce certain modified impres- sions on the extremities of particular nervous filaments pro- vided to receive them; but these impressions constitute only the commencement of the series of corporeal changes which terminate in sensation; for they have to be conveyed along the course of the nerves to the brain, or central organ of the nervous system,* where, again, some physical change must take place, before the resulting affection of the mind can be produced. The particular part of the brain, where this last physical change, immediately preceding the mental change, takes place, is termed the Sensorium.. Abundant proofs exist that all the physical changes here referred to really oc- cur, and, also, that they occur in this order of succession: for they are invariably found to be dependent on the healthy state, not only of the nerve, but, also, of the brain; thus, the destruction, or even compression of the nerve, in any part of its course between the external organ and the sensorium, totally prevents sensation; and the like result ensues from even the slightest pressure made on the sensorium itself. Although the corporeal or physical change taking place in the sensorium, and the mental affection we term sensation, arc linked together by some inscrutable bond of connexion, • It is usual to designate the end of the nerve which is next to the senso- rium, as the origin of tliat nerve; whereas, it should more properly be re- garded as its termination; for the scries of changes which end in sensation commence at the organ of sense, and arc thence propagated along the nerve to the sensorium. PERCEPTION. 359 they are, in their nature, as perfectly distinct as the subjects in which they occur; that is, as mi?id is distinct from mat- ter; and they cannot, therefore, be conceived by us as having the slightest resemblance the one to the other. Yet sensations invariably suggest to the mind ideas, not only of the existence of an external agent as producing them, but also of various qualities and attributes belonging to these agents; and the belief, or rather the irresistible conviction, thus forced upon us, of the reality of these external agents, which we conceive as constituting the material world, is termed P accept ion. Various questions here present themselves concerning the origin, the formation, and the laws of our perceptions. This vast field of curious but difficult inquiry, situated on the confines of the two great departments of human knowledge, (of which the one relates to the phenomena of matter, and the other to those of mind,) requires for its successful culti- vation the combined effi^rts of the physiologist and the me- taphysician. For although our sensations are purely men- tal affections, yet inasmuch as they are immediately depend- ent on physical causes, they are regulated by the physical laws of the living frame; whereas the perceptions derived from these sensations, being the results of intellectual pro- cesses, are amenable rather to the laws which regulate men- tal than physical phenomena. It is certain, from innume- rable facts, that in the present state of our existence, the operations of the mind are. conducted by the instrumentality of our bodily organs; and that unless the brain be in a healthy condition, these operations become disordered, or altogether cease. As the eye and the ear are the instruments by which we see and hear, so the brain is the material in- strument by which we retrace and combine ideas, and by which we remember, we reason, we invent. Sudden pres- sure on this organ, as in a stroke of apoplexy, puts a total stop to all these operations of the mind. If the pressure be of a nature to admit of remedy, and has not injured the tex- ture of the brain, recovery may take place; and immediately 360 THE SENSORIAL FUNCTIONS. on the return of consciousness, the person awakes as from a dream, having no sense of the time which has elapsed since the moment, of the attack. All causes which disturb the healthy condition of the brain, such as alcohol, opium, and other narcotic drugs, or wliich disorder more especially the circulation in that organ, such as those inducing fever, or inflammation, produce corresponding derangements of the intellectual powers; modifying the laws of the association of ideas, introducing confusion in the perceptions, irregularity in the trains of thought, and incapacity of reasoning, and leading to the infinitely diversified forms of mental halluci- nation, delirium, or insanity. ICven the strongest minds are subject to vicissitudes arising from slighter causes, which aff'ect the general tone of the nervous system. Vain, in- deed, was the boast of the ancient Stoics that the human mind is independent of the body, and impenetrable to ex- ternal influences. No mortal man, whatever may be the vi- gour of his intellect, or the energy of his application, can withstand the influence of impressions on his external senses; for, if sufiiciently reiterated or intense, they will al- ways have power, if not to engross his whole attention, at least to interrupt the current of his thoughts, and direct them into other channels. Nor is it necessary for producing this efiect that cannon should thunder in his ears; the mere rat- tling of a window, or the creaking of a hinge will often be sufficient to disturb his philosophical meditations, and disse- ver the whole chain of his ideas. . "Marvel not," says Pas- cal, " that this profound statesman is just now incapable of reasoning justly; for behold, a fly is buzzing round his head. If you wish to restore to him the power of correct thinking, and of distinguishing truth from falsehood, you must first chase away the insect, holding in thraldom that exalted rea- son, and that gigantic intellect, which govern empires and decide the destinies of mankind." Although we must necessarily infer, from the evidence furnished by experience, that some physical changes in the brain accompany the mental processes of thought, we are in PERCEPTION. 3i)l utter ignorance of the nature of those actions; and all our knowledge on this suhject is limited to the changes which we are conscious are going on in the mind. It is to these mental changes, therefore, that our attention is now to be directed. In experiencing mere sensations, whatever be their as- semblage or order of succession, the mind is wholly passive: on the other hand, the mind is active on all occasions when we combine into one idea sensations of different kinds, (such as those w^hich are derived from each separate sense,) when we compare sensations or ideas with one another, when we analyze a compound idea, and unite its elements in an order or mode of combination different from that in which they were originally presented. Many of these active operations of mind are implied in the process of perception; for al- though it might be supposed that the diversity in the nature of our sensations would sufficiently indicate to us a corre- sponding variety in the qualities of the material agents, which produce their impressions on our senses, yet these very qualities, nay, even the existence of the objects them- selves, are merely inferences deduced by our reasoning powers, and not the immediate effects of those impressions on the mind. We talk, for instance, of seeing a distant body; yet the immediate object of our perception can only be the light, which has produced that particular impression on our retina; whence we infer, by a mental process, the existence, the position, and the magnitude of that body. When we hear a distant sound, the immediate object of our perception is neither the sounding body whence it emanates, nor the successive undulations of the medium conveying the effect to our ear; but it is the peculiar impression made by the vibrating particles of the fluid, which are in direct con- tact with the auditory nerve. It is not difficult to prove that the objects of perception are mere creations of the mind, suggested, probably instinctively, by the accompanying sen- sations, but having no real resemblance or correspondence either with the impressions themselves, or with the agencies Vol. II. 46 362 THE SENSORIAL FUNCTIONS. which produce them; for many are the instances in which our actual perceptions are widely different from the truth, and have no external prototype in nature. In the absence of light, any mechanical pressure, suddenly applied to the eye, excites, by its cifcct on the retina, the sensation of vivid light. That tiiis sensation is present in the mind we are certain, because we are conscious of its existence: here there can be no fallacy. But the perception of light, as a cause of this sensation, being inseparably associated with such sen- sation, and wholly dependent on it, and corresponding in all respects, both as to its duration and intensity, with the same circumstances in the sensation, we cannot avoid having the perception as well as the sensation of light: yet it is cer- tain that no light has acted. The error, then, attaches to the perception; and its source is to be traced to the mental process by which perception is derived from sensation. INIany other examples might be given of fallacious per- ceptions, arising from impressions made in an unusual man- ner on the nerves of the senses. One of the most remark- able is the appearance of a flash of light from the transmis- sion of the galvanic influence through the facial nerves. If a piece of silver, or of gold, be passed as high as possible between the upper lip and the gums, while at the same time a plate of zinc is laid on the tongue, or applied to the inside of the cheeks; and if a communication be then made be- tween the two metals, either by bringing them into direct contact, or by means of a wire touching both of them at the same time, a flash of light is seen by the person who is the subject of the experiment. This appearance is the effect of an impression made either on the retina, or on the optic nerve, and is analogous to that occasioned by a mechanical impulse, such as a blow directed to the same part of the ner- vous system, both being phenomena totally independent of the presence of light. A similar fallacy occurs in the per- ception of taste, which arises in the well known experiment of placing a piece of zinc and another of silver, the one on the upper and the other on the under surface of the tongue, PERCEPTION. 363^ and making them communicate, when a pungent and disa- greeable metallic taste is instantly perceived: this happens because the nerves of the tongue, being acted upon by the galvanism thus excited, communicate the same sensation as that w^hich would be occasioned by the actual application of sapid bodies to that organ. Thus, it appears that causes which are very different in their nature, may, by acting on the same nerves, produce the very same sensation; and it follows, therefore, that our sensations cannot be depended upon as being always exactly correspondent with the quali- ties of the external agent which excites them. Evidence to the same effect may also be gathered from the consideration of the narrowness of those limits within which all our senses are restricted. It requires a certain in- tensity in the agent, whether it be light, or sound, or che- mical substances applied to the senses of smell or taste, in order to produce the very lowest degree of sensation. On the other hand, when their intensity exceeds a certain limit, the nature of the sensation changes, and becomes one of pain. Of the sensations commonly referred to the sense of touch, there are many which convey no perception of the cause producing them. Thus, a slighter impression than that which gives the feeling of resistance produces the sensa- tion of itching, which is totally different in its kind. The sensation of cold is equally positive with that of warmth, and differs from it, not in degree merely, but in species; al- though we know that it is only in its degree that the exter- nal cause of each of these sensations differs. The only distinct notions we are capable of forming re- specting Matter, are that it consists of certain powers of attraction and repulsion, occupying certain portions of space, and capable of moving in space; and that its parts thereby assume different relative positions or configurations. But of mind, our knowledge is more extensive and more precise, because we are conscious of its existence, and of many of its operations, which arc comprised in the general term thought. To assert that thought can be a property of mat 364 THE SENSORIAL FUNCTIONS. ter, is to extend the meaning of the term matter to that with which we cannot perceive it has any relation. All that we know of matter has regard to space: nothing that we know of the properties and affections of mind has any relation whatsoever to space. A similar incongruity is contained In the proposition that thought is :{ function of the hrain. It is not the brain which thinks, any more than it is the eye which sees, though each of these material organs is necessary for the production of these respective effects. That which sees and which thinks is exclusively the mind; although it is by the instrumentali- ty of its bodily organs that these changes take place. At- tention to this fundamental distinction, which, although ob- vious when explicitly pointed out, is often lost sight of in ordinary discourse, will furnish a key to the solution of many questions relating to perception, which have been considered as difficult and embarrassing. The sensations derived from the different senses have no resemblance to one another, and have, indeed, no property in common, except that they are felt by the same percipient being:. A colour has no sort of resemblance to a sound; nor have either of these any similarity to an odour, or a taste, or to the sensations of heat, or cold. But the mind, which receives these incongruous elements, has the power of giving them, as it were, cohesion, of comparing them with one another, of uniting them into combinations, and of form- ing them into ideas of external objects. All that nature presents is an infinite number of particles, scattered in diffe- rent parts of space; but out of these the mind forms indivi- dual groups, to which she gives a unity of her own crea- tion. All our notions of material bodies involve that of space; and we derive this fundamental idea from the peculiar sensa- tions which attend the actions of our voluntary muscles. These actions first give us the idea of our own bodies, of its various parts, and of their figure and movements; and next teach us the position, distances, magnitudes, and figures of PERCEPTION. 365 adjacent objects. Combined with tbese ideas are the more immediate perceptions of touch, arising from contact with the skin, and especially with the fingers. All these percep- tions, variously modified, make us acquainted with those mechanical properties of bodies, which have been regarded by many as primary or essential qualities. The perceptions derived from the other senses can only add to the former the ideas of partial, or secondary qualities, such as tempera- ture, the peculiar actions which produce taste and smell, the sounds conveyed from certain bodies, and, lastly, their visi- ble appearances. The picture formed on the retina by the refracting power of the humours of the eye, is the source of all the perceptions which belong to the sense of vision: but the visible appear- ances which these pictures immediately suggest, when taken by themselves, could have given us no notion of the situa- tion, distances, or magnitudes of the objects they represent; and it is altogether from the experience acquired by the ex- ercise of other senses that we learn the relation which these appearances have with those objects. In process of time the former become the signs and symbols of the latter; while ab- stractedly, and without such reference, they have no meaning. The knowledge of these relations is acquired by a process exactly analogous to that by which we learn a new language. On hearing a certain sound in constant conjunction with a certain idea, the two become inseparably associated together in our minds; so that on hearing the name, the correspond- ing idea immediately presents itself. In like manner, the visible appearance of an object is the sign, which instantly impresses us with ideas of the presence, distance, situation, form, and dimensions of the body, that gave rise to it. This association is, in man at least, not original, but acquired. The objects of sight and touch, as Bishop Berkeley has justly ob- served, constitute two worlds, which although they have a very important correspondence and connexion, yet bear no sort of resemblance to one another. The tangible world has three dimensions, namely, length, breadth, and thickness; the 366 THE SENSORIAL FUNCTIONS. visible world only two, namely, length and breadth. The objects of sight constitute a kind of language, which Nature addresses to our eyes, and by wliich she conveys informa- tion most important to our welfare. As, in any language, the words or sounds bear no resemblance to the things they denote, so in this particular language the visible ob- jects bear no sort of resemblance to the tangible objects they represent. The theory of Berkeley received complete confirmation by the circumstances attending the well known case, described by Cheselden, of a boy, who, from being blind from birth, suddenly acquired, at the age of twelve, the power of see- ing, by the removal of a cataract. He at first imagined that all the objects he saw touched his eyes, as what he felt did his skin; and he was unable either to estimate distances by the sight alone, or even to distinguish one object from another, until he had compared the visual with what has been called the tact ualim-pression. This theory also affords a satisfactory solution of a ques- tion which has frequently been supposed to involve consi- derable difficulty; namely, how it happens that we see ob- jects in their true situation, when their images on the retina, by which we see them, are inverted. To expect that the impression from an inverted image on the retina should pro- duce the perception of a similar position in the object viewed, is to commit the error of mistaking these images for the real objects of perception, whereas they are only the means which suggest the true perceptions. It is not the eye which sees; it is the mind. The analogy which the optical part of the €ye bears to a camera obscura has perhaps contributed to the fallacy in question; for, in using that instrument, we really contemplate the image which is received on the paper, and re- flected from it to our eyes. But in our own vision nothing of this kind takes place. Far from there being any contem- plation by the mind of the image on the retina, we are ut- terly unconscious that such an image exists, and still less can we be sensible of the position of the image with respect VISUAL PERCEPTIONS. 367. to the object. All that we can distinguish as to the locality of the visual appearance which an object produces, is that this appearance occupies a certain place in the field of vi- sion; and we are taught, by the experience of our other senses, that this is a sign of the existence of the external object in a particular direction with reference to our own body. It is not until long after this association has been established, that we learn, by deduction from scientific principles, that the part of the retina, on which the impression causing this appearance is made, is on the side opposite to that of the object itself; and also that the image of a straight object is curved as well as inverted. But this subsequent information can never inter- fere with our habitual, and perhaps instinctive reference of the appearance resulting from an impression made upon the upper part of the retina, to an object situated below us, and vice versa. Hence we at once refer impressions made on any particular part of the retina to a cause proceeding from the opposite side. Thus, if we press the eye-ball with the finger applied at the outer corner of the orbit, the lumi- nous appearance excited by the pressure is immediately re- ferred to the opposite or inner side of the eye. If we place a card perpendicularly between the two eyes, and close to the face, the card will appear double, because, although each surface is seen by the eye which is adjacent to it, in the direction in which it really is with regard to that eye, yet, being out of the limits of distinct vision, it is referred to a much greater distance than its real situation ; and consequently, the two sides of the object appear separated by a wide interval, and as if they belonged to two different objects. Many other examples might be given of similar fallacies in our visual perceptions. All impressions made on the nerves of sensation have a definite duration, and continue for a certain interval of time after the action of the external agent has ceased. The ope- ration of this law is most conspicuous in those cases where the presence or absence of the agent can readily be deter- mined. Thus, we retain the sensation of a sound for some 353 THE SENSORIAL FUNCTIONS. time after the vibrations of the external medium have ceased; as is shown by the sensation of a musical note being the re- sult of the rco-ular succession of aerial undulations, when the impression made by each continues during the whole inter- val between two consecutive vibrations. The impulses of lio-ht on the retina are unquestionably consecutive, like those of sound, but being repeated at still shorter intervals, give rise to a continuous impression. A familiar instance of the same principle occurs in the appearance of an entire lumi- nous circle, from the rapid whirling round of a piece of liglUed charcoal; for the part of the retina which receives the brilliant image of the burning charcoal, retains the im- pression with nearly the same intensity during the entire revolution of the light, when the same impression is re- newed. For the same reason a rocket, or a fiery meteor, shooting across the sky in the night, appears to leave behind it a long luminous train. The exact time, during which these impressions continue, after the exciting cause has been withdrawn, has been variously estimated by different experi- mentalists, and is very much influenced, indeed, by the in- tensity of the impression.* * Many curious visual illusions may be traced to the operation of this principle. One of the most remarkable is the curved appearance of the spokes of a carriage wheel rolling- on the ground, when viewed through the intervals between vertical parallel bare, such as those of a palisade, or Vene- tian window-blind. On studying the circumstances of this phenomenon, t found that it was the necessary result of the traces left on the retina by the parts of each spoke which became in succession visible through the apertures, and assumed the curved appearances in question. A paper, in which I gave an account of the details of these observations, and of the theory by which I explained them, was presented to the Koyal Society, and published in the Philosophical Transactions, for 1825, p. 131. About three years ago, Mr. Faraday prosecuted the subject with tlie usual success which attends all his philosophical researches, and devised a great number of interesting experi- ments on the appearances resulting from combinations of revolving wlipels; the details of which are given in a paper contained in the first volume of the Journal of tlie Koyal Institution of Great Britain, p. 205. Tliis again direct- ed my attention to the subject, and led me to the invention of the instrument which has since been introduced into notice under the name of the Phantas- VISUAL PERCEPTIONS. 369 When the impressions are very vivid, another phenome- non often takes place; namely, their suhsequent recurrence, after a certain interval, during whicli tliey are not felt, and quite independently of any renewed aj)plicatioii of the cause which had originally excited them. If, for example, we look steadfastly at the sun for a second or two, and then im- mediately close our eyes, the image or spectrum of the sun remains for a long time present to the mind, as if its light were still acting on the retina. It then gradually fades and disappears; hut if we continue to keep the eyes shut, the same impression will, after a certain time, recur, and again vanish; and this phenomenon will be repeated at intervals, the sensation becoming fainter at each renewal. It is pro- bable that these reappearances of the image, after the light which produced the original impressioji has been withdrawn, are occasioned by spontaneous affections of the retina itself, which are conveyed to the sensorium. In other cases, where the impressions are less strong, the physical changes producing these spectra are perhaps confined to the senso- rium. These spectral appearances generally undergo vari- ous changes of colour, assuming first a yellow tint, passing then to a green, and lastly becoming blue, before they finally disappear. Another general law of sensation is, that all impressions made on the nerves of sense tend to exliaust their sensibility, so that the continued or renewed action of the same exter- nal cause produces a less effect than at first: while, on tiie other hand, the absence or diminution of the usual excite- ment leads to a gradual increase of sensibility, so that the subsequent application of an exciting cause produces more than the usual effect. One of the most obvious exemplifica- tions of this law presents itself in tlic case of the sensations of temperature. The very same body may appear warm to mascope or Pheniiklsticope. I constructed several of these at that period, (in the spring- of 1 831 ) wiiich I showed to many of my friends; but in conse- quence of occupations and cares of a more serious kind, I did not publish any account of this invention, which was last year reproduced on Uic continent. Vol. II. 47 370 THE SENSORIAL FUNCTIONS. the touch at one time, and cold at another, (although its real temperature has not varied,) according to the state of the or«'an induced hy previous impressions: and a very different judgment will be formed of its temperature, when felt by each hand in succession, if the one has immediately before been exposed to cold, while the other has retained its natu- ral warmth. Similar phenomena may be observed with regard to all the other senses: thus, the flavour of odorous, as well as sapid bodies, depends much on the previous state of the organ by which they are perceived; any strong im- pression of taste made on the nerves of the tongue, render- ing them, for some time, nearly insensible to weaker tastes. Sounds, wliich make a powerful impression on the auditory nerves, will, in like manner, occasion temporary deafness with rcc;ard to faint sounds. The converse of this is ob- served when hearing has been suddenly restored in deaf persons, by the operation of perforating the ear-drum.* The sensibility of the auditory nerves, which had not been ac- cessible to impressions of sound, is found to be increased to a morbid degree. This was remarkably exemplified in the case of a gentleman, who, for several years, had been very deaf, in consequence of the obliteration of the Eustachian tube, so that he could scarcely hear a person speaking in a loud voice close to his ear. As soon as the instrument which had made the perforation was withdrawm, the by-standers began to address him in a very low tone of voice, and were surprised at receiving no answer, and at his remaining im- moveable in his chair, as if stunned by a violent blow. At length, he burst out into the exclamation, " For God's sake, gentlemen, refrain from crying out so terribly loud! you are giving me excessive pain by speaking to me." The sur- geon,t upon this, retired across the room; unfortunately, however, the creaking of his boots caused the gentleman to start up in an agony from his chair, at the same time apply- ing his hand instinctively to cover his ear; but in doing this, • Sec the note in p. 307 of this volume. f M. Maunoir, of Geneva, on whose authority I have given this account. VARIATIONS OF SENSIBILITY. 371 the sound of his fingers coming in contact with his head was a fresh source of pain, producing an efTcct similar to that of a pistol suddenly fired close to him. For a long time after, when spoken to, even in the lowest whisper, he complained of the distressing loudness of the sounds; and it was several weeks before this excessive sensibility of the auditory nerves wore off: by degrees, however, tlicy accommodated them- selves to their proper function, and became adapted to the ordinary impressions of sound. Some time afterwards, this gentleman had a similar operation performed on the other ear, and with precisely the same results; the same degree of excessive sensibility to sounds was manifested on the resto- ration of hearing in this ear as had occurred in the first; and an equal time elapsed before it was brought into its natural state. The most striking illustrations of the extent of this law are furnished by the sense of vision. On entering a dark chamber, after having been for some time exposed to the glare of a bright sunshine, we feel as if we were blind; for the retina, having been exhausted by the action of a strong light, is insensible to the weaker impressions which it then receives. It might be supposed that the contraction of the pupil, which takes place on exposure to a strong light, and, of course, greatly reduces the quantity admitted to the re- tina, is a cause adequate to account for this phenomenon: hut careful observation will show that the pupil very rapid- ly enlarges to its full expansion when not acted upon by liffht; while the insensibilitv of the retina continues for a much longer time. It regains its usual sensibility, indeed, only by slow degrees. ]5y remaining in the dark its sensi- bility is still farther increased, and a faint light will excite impressions equal to those produced in Ihc ordinary state of the eye by a much stronger light; and while it is in this state, the sudden exposure to the light of day produces a dazzling and painful sensation. This law of vision was usefully applied by vSir William Herschel in training his eye to the acquisition of extraordi- 372 THE SENSORIAL FUNCTIONS. nary sensibility, for the purpose of observing very faint ce- lestial objects. It often happened to him, when, in a fine winter's night, and in the absence of the moon, he was oc- cupied during four, five, or six hours in taking sweeps of the heavens with his telescope, that, by excluding from the eye the liglit of surrounding objects, by means of a black hood, the sensibility of the retina was so much increased, that when a star of the third magnitude approached the field of view, he found it necessary immediately to with- draw his eye, in order to preserve its powers. He relates that on one occasion the appearance of Sirius announced itself in the field of the telescope like the dawn of the morn- ing, increasing by degrees in brightness, till the star at last presented itself with all the splendour of the rising sun, obliging him quickly to retreat from the beautiful but over- powering spectacle. The peculiar construction of the organ of vision allows of our distinguishing the effects of impressions made on parti- cular parts of the retina from those made on the rest, and from their general effect on the whole surface. These par- tial variations of sensibility in the retina give rise to the phe- nomena of ocular spectra^ as they are called, which were first noticed by Buffon, and afterwards more fully investi- gated by Dr. Robert Darwin. A white object on a dark ground, after being viewed steadfastly till the eye has be- come fiitigued, produces, when the eye is immediately di- rected to another field of view, a spec^um of a darker co- lour than the surrounding space, in consequence of the ex- haustion of that portion of the retina on which its image had been impressed. The converse takes place, when the eye, after having been steadfastly directed to a black object on a light ground, is transferred to another part of the same field; and in this case a bright spectrum, of the object is seen. It is a still more curious fact that the sensibility of the re- tina to any particular kind of light, may, in like m.anner, be increased or diminished, without any change taking place in its sensibility to other kinds of light. Plence the spectrum OCULAR SPECTRA. 373 of a red object appears green; because the sensibility of that portion of the retina, on which the red image has been im- pressed, is impaired with regard to the red rays, while the yellow and the blue rays still continue to produce their usual effect; and these, by combining their influence, produce the impression of green. For a similar reason, the spectrum of a green object is red; the rays of that colour being those which alone retain their power of fully impressing the re- tina, previously rendered less sensible to the yellow and the blue rays composing the green liglit it had received from the object viewed. The judgments we form of the colours of bodies are in- fluenced, in a considerable degree, by the vicinity of other coloured objects, which modify the general sensibility of the retina. When a white or gray object of small dimensions, for instance, is viewed on a coloured ground, it generally appears to assume a tint of the colour which is complemen- tary to that of the ground itself* It is the etiquette among the Chinese, in all their epistles of ceremony, to employ paper of a bright scarlet hue: and I am informed by Sir George Staunton, that for a long time after his arrival in China, the characters written on this kind of paper appeared to him to be green; and that he was afterwards much sur- prised at discovering that the ink employed was a pure black, without any tinge of colour, and on closer examina- tion he found that the marks were also black. The green appearance of the letters, in this case, was an optical illu- sion, arising from the tendency of the retina, which had been strongly impressed with red light, to receive impres- sions corresponding to the complementary colour, which is green. A philosophical history of the illusions of the senses would afford ample evidence that limits have been intentionally as- signed to our powers of perception; but the subject is much • Any two colours which, when combined tog-cther, produce white li^ht, are said to be compkmentary to one another. 374 THE SENSORIAL FUNCTIONS. too extensive to be treated at len^jth in the present work.* I must content myself with remarking, that these illusions are the direct consequences of the very same laws, which, in ordinary circumstances, direct our judgment correctly, but are then acting under unusual or irregular combinations of circumstances. These illusions may be arranged under three classes, according as they are dependent on causes of a physical, physiological, or mental kind. The first chiss includes those illusions in which an impres- sion is really made on tlic organ of sense by an external cause, but in away to wiiicli we have not been accustomed. To this class belong the acoustic deceptions arising from echoes, and from the art of ventriloquism; the deceptive appear- ances of the mirage of the desert, the looming of the horizon at sea, the Fata Morgana of the coast of Calabria, the gi- gantic spectre of the Brockcn in the Hartz, the suspended images of concave mirrors, the visions of the phantasmago- ria, the symmetrical reduplications of objects in the field of the kaleidoscope, and a multitude of other results of the simple combinations of the laws of optics. The second class comprehends those in which the cause of deception is more internal, and consists in the peculiar condition of the nervous surface receiving the impressions. Ocular spectra of various kinds, impressions on the tongue and the eye from galvanism, and those which occasion sing- ing in the ears, arising generally from an excited circulation, are among the many perceptions which rank under this head. The third class of fallacies comprehends those which are essentially niental in their origin, and are the consequences of errors in our reasoning powers. Some of these have al- ready been pointed out with regard to the perceptions of vision and of hearing, the formation of which is regulated • In the Gulstonian Lectures, whicli I was appointed to read to the Royal College of Physicians, in May, 1832, 1 took occasion to enlarge on this sub- ject. A summary of these lectures was given in the London Medical Ga- zette, vol. X. p. 273, Illusions of the senses. 375 by the laws of the association of ideas. But even the sense of touch, which has been generally regarded as the least lia- ble to fallacy, is not exempt from this sourceof error, as is proved by the well known experiment of feeling a single ball, of about the size of a pea, between two fingers which are crossed; for there is then a distinct perception of the presence of two balls instead of one. But limited as our senses arc in their range of perception, and liable to occasional error, we cannot but perceive, that, both in ourselves, and also in every class of animals, they have been studiously adjusted, not only to the properties and the constitution of the material world, but, also, to the re- spective wants and necessities of each species, in the situa- tions and circumstances where it has been placed by the gra- cious and beneficent Author of its being. , If the sensorial functions had been limited to mere sensa- tion and perception, conjoined with the capacity of passive enjoyment and of suffering, the purposes of animal existence w^ould have been but imperfectly accomplished; for, in or- der that the sentient being m^ay secure the possession of those objects which are agreeable and salutary, and avoid or reject those which are painful or injurious, it is necessary that he possess the power of spontaneous action. Hence, the faculty of Voluntary Motion is superadded to the other sensorial functions. The muscles which move the limbs, the trunk, the head, and organs of sense, — all those parts, in a word, which establish relations ^vith the external world, are, through the intermedium of a separate set of nervous filaments, totally distinct from those which are subservient to sensation,^' made to communicate directly with the senso- rium, and are thereby placed under the direct control and guidance of the will. The mental act of volition is doubt- less accompanied by some corresponding physical change in that part of the sensorium, whence the viotor nerves^ or * On this subject I must refer the reader to the researches of Sir Charles Bell, and Magcndie, who have completely cstablisiicil the disitniction be- tween these two classes of nerves. 376 THE SENSORIAL FUNCTIONS. those distributed to the muscles of voluntary motion, arise. Here, then, we pass from mental phenomena to such as are purely physical; and the impression, whatever may be its nature, originating in the sensorium, is propagated along the course of the nerve to those muscles, whose contraction is required for the production of the intended action. Of the function of voluntary motion, as far as concerns the moving powers and the mechanism of the instruments employed,* I have already treated at suITicient length in the first part of this work. Every excitement of the sensorial powers is, sooner or later, followed by a proportional degree of exhaustion; and when this has reached a certain point, a suspension of the exercise of these faculties takes place, constituting the state of sleej:), during which, by the continued renovating action of the vital functions, these powers are recruited, and ren- dered again adequate to the purposes for which they were bestowed. In the ordinary state of sleep, however, the ex- haustion of the sensorium is seldom so complete as to pre- clude its being excited by internal causes of irritation, which would be scarcely sensible during our waking hours: and hence arise dreams, which are trains of ideas, suggested by internal irritations, and which the mind is bereft of the power to control, in consequence of the absence of all im- ♦ A voluntary action, occurnng' as the immediate consequence of the ap- plication of an external agent to an organ of the senses, thougli apparently a simple phenomenon, implies the occurrence of no less than twelve succes- sive processes, as may be seen by the following enumeration. First, there is the modifying action of the organ of tlie sense, the refractions of the rays, for instance, in the case of the eye: secondly, the impression made on the extremity of the nerve: thirdly, the propagation of this impression along the nerve: fourthly, the impression or physical change in the sensorium. Next follow four kinds of mental processes, namely, sensation, perception, associa- tion, and volition. Then, again, there is another physical change taking place in the sensorium, immediately consequent on the mental act of voli- tion: this is followed by the propagation of the impression downwards along tlie motor nei've; then an impression is made on the muscle; and, lastly, we obtain the contraction of the muscle, which is the object of the whole series of operations. VOLUNTARY MOTION. 377 pressions from the external senses.* In many animals, a much more general suspension of the actions of life, extend- ing even to the vital functions of respiration and circulation, takes place during the winter months, constituting what is termed Hybernation. * The only indications of dreaming- given by the lower animals occur in those possessed of the gi-eatest intellectual powers, such as the I)o^, amonir quadrupeds, and the Parrot, among birds. Vol. II. 48 ( 378 ) CHAPTER VIII. COMPARATIVE PHYSIOLOGY OF THE NERVOUS SYSTEM. § 1. Nervous Systems of Invertehrated Animals. Our knowledge of the exact uses and functions of the various parts which compose the nervous system, and espe- cially of its central masses, is unfortunately too scanty to enable us to discern the correspondence, which undoubtedly exists, between the variations in the functions and the di- versities in the organization. The rapid review which I propose to take of the different plans, according to which the nervous system is constructed in the several classes of animals, will show that these central masses are multiplied and developed in proportion as the faculties of the animal embrace a wider range of objects, and arc carried to higher degrees of excellence. In none of the lowest tribes of Zoophytes, such as Spoiiges, Polypi, and MedusXy have any traces of organs, bearing the least analogy to a nervous system, been discovered; not even in the largest specimens of the last named tribe, some of 'which are nearly two feet in diameter. All these ani- mals give but very obscure indications of sensibility; for the contractions they exhibit, when stimulated, appear to be rather the efifect of a vital property of irritability than the result of any sensorial faculty. Analogy, however, would lead us to the belief that many of their actions are really prompted by sensations and volitions, though in a de- gree very inferior to those of animals higher in the scale of being: but whatever may be their extent, it is probable that the sensorial operations in these animals take place without NERVOUS SYSTEM OP INVERTEBRATA. 37$) the Intervention of any common scnsorlnm, or centre of ac- tion. It is at the same time remarkable that their move- ments are not effected by means of muscular fibres, as they are in all other animals, the granular flesh, of which their whole body is composed, appearing to have a generally dif- fused irritability, and perhaps also some degree of sensibi- lity; so that each isolated granule may be supjiosed to be endowed with these combined properties, performing, inde- pendently of the other granules, the functions both of nerve and muscle. Such a mode of existence exhibits apparently the lowest and most rudimental condition of the animal functions. Yet the actions of the Hydra, of which I have given an account, are indicative of distinct volitions; as are also, in a still more decided manner, those of the Lifusoria. In the way in which the latter avoid obstacles while swim- ming in the fluid, and turn aside when they encounter one another, and in the eagerness with which they pursue their prey, we can hardly fail to recognise the evidence of volun- tary action. To seek for an elucidation of these mysteries in the struc- ture of animals whose minuteness precludes all accurate ex- amination, would be a hopeless inquiry. Yet the indefati- gable Ehrenberg has recently discovered, in some of the larger species of animalcules belonging to the orticr Roii- ferUy an organization, which he believes to be a nervous sys- tem. He observed, in the Hydatina senta, a series of six or seven gray bodies, enveloping the upper or dorsal part of the oesophagus, closely connected together, and perfectly distinguishable, by their peculiar tint, from the viscera and the surrounding parts. The uppermost of these bodies, which he considers as a ganglion, is much larger than the others, and gives off slender nerves, which, by joining another ganglion, situated under the integuments at the back of the neck, form a circle of nerves, analogous to that which surrounds the oisophagus in the mollusca: from this circle two slender nervous filaments are sent off to the head, and a larger branch to the abdominal surface of 380 THE SENSORIAL FUNCTIONS. the body. The discovery of a regular structure of muscu- lar bands of fibres, in these animalcules, is a farther evi- dence of the connexion which exists between nerves and muscles. We again meet with traces of nervous filaments, accom- panied also witli muscular bands of fibres, in some of the more highly organized Enlozoci. In the Ascaris^ or long round worm, a slender and apparently single filament is seen passing forwards, along the lower side of the abdomen, till it reaches the oesophagus, where it splits into two branches, one passing on each side of that tube, but without exhibit- ing any ganglionic enlargement. This may be considered as the first step towards the particular form of the nervous system of the higher classes of articulated animals, where the principal nervous cord is obviously double throughout its whole length, or, if partially united at different points, it is always readily divisible into two, by careful manipula- tion. In addition to this characteristic feature, these cords present, in their course, a series of enlargements, appearing like knots; one pair of these generally corresponding to each of the segments of the body, and sending off, as from a cen- tre, branches in various directions. It is probable that these knots, or ganglia, perform, in each segment of the worm, an office analogous to that of the brain and spinal marrow of vertcbrated animals, serving as centres of nervous, and per- haps, also, of sensorial powers. Many facts, indeed, tend to show that each segment of the body of articulated animals, of an annular structure and cylindric form, such as the long w^orms and the myriapoda, has in many respects an inde- pendent sensitive existence, so that when the body is di- vided into two or more parts, each portion retains both the faculty of sensation, and the power of voluntary motion. As far as we can judge, however, the only external sense which is capable of being exercised by this simple form of nervous system, is that of touch; all the higher senses evidently re- quiring a much more developed and concentrated organiza- tion of nervous ganglia. NERVOUS SYSTEM OF ARTICULATA. 3S'l In this division of the animal kingdom, the primary ner- vous cords always pass along the middle of the lower sur- face of the body, this being the situation which, in the ab- sence of a vertebral bony column, aflbrds them the best pro- tection. They may be considered as analogous to the spi- nal marrow, and as serving to unite the series of ganglia, through which they pass, into one connected system. On arriving at the oesophagus, they form round it a circle, or collar, studded with ganglia, of which the uppermost, or that nearest the head, is generally of greater size than the rest, and is termed the oesophogcal, cephalic, or cerebral gan- glion, being usually regarded as analogous to the brain of larger animals. Perhaps a more correct view of its func- tions would be conveyed by calling it the principal brain, and considering; the other 2;ani2;lia as subordinate bi-ains. This large ganglion, whicli supplies an abundance of ner- . vous filaments to every part of the heacl, seems to be the chief organ of the higher senses of vision, of hearing, of taste, and of smell, and to be instrumental in combining th.eir impressions, so as to constitute an individual percipient animal, endowed with those active powers which are suited to its rank in the scale of being. Such is the general form of the nervous system in all the */innelida: but in the higher orders of Jirticulata we find it exhibiting various degrees of concentration. The pro- gress of this concentration is most distinctly traced in the Crustacea* One of the simplest forms of these organs oc- curs in a little animal of this class, which is often found in immense numbers, spread over tracts of sand on the sea shore, and which is called the Talitrus locust a, or Sand- ^oo hopper, (Fig. 43S.) The central parts y^^M^j^fe^ of its nervous system are seen in Fig. f f^^S|^^ 439, wiiich represents the abdominal side of this animal laid open, and mag- nified to twice the natural size. The two primary nervous * See the account of the researches of Victor Audouin, and II. M. Ed- wards, on this subject, given in the Ann. dcs Sc. Nat. xix. 181. 382 THE SENSORIAL FUxVCTIONS. cords, which run in a longitudinal direction, are here per- fectly distinct from one another, and even separated by a small interval: they present a series of ganglia, which are nearly of equal size, and equidistant from one another, one pair corresponding to each segment of the body,* and united by transverse threads: and other filaments, diverging late- rally, proceed from each ganglion. During the progress of growth, the longitudinal cords approach somewhat nearer to each other, but still remain perfectly distinct. The first 43.9 440 pair of ganglia, or the cephalic, have been considered, though improperly, as the brain of the animal. The next step in the gradation occurs in the Fhyllosoma (Leach,) where the ganglia composing each pair in the ab- domen and in the head, are united into single masses, while those in the thoracic region are still double. In the Cymo- ihoa, (Fab.,) which belongs to the family of Onisciis, there is the appearance of a single chain of ganglia, those on the one side having coalesced with those on the other; each pair composing a single ganglion, situated in the middle line; while the longitudinal cords which connect them still re- • These segments arc numbered in this and the following- figure in their proper order, beginning witli that near the head, a is the external antenna? a, tlic internal antenna; and e, the eye. NERVOUS SYSTEM OP CRUSTACEA. 383 main double, as is shown in Fig. 440, which represents the interior of this crustaceous animal, nearly of the natural size. But in the higher orders of Crustacea, as in the Lobster, these longitudinal cords are themselves united in the abdo- minal region, though still distinct in the thorax. , In following the ascending series of crustaceous animals, we observe also an approximation of the remoter ganglia to- wards those near the centre of the body: this tendency al- ready shows itself in the shortening of the hinder part of the nervous system of the Cymolhoa, as compared with the Talitrus; and the concentration proceeds farther in other tribes. In the Palemon, for example, most of the thoracic ganglia, and in the Palinuriis (Fab.,) all of them, have co- alesced into one large oval mass, perforated in the mid- dle, and occupying the centre of the thorax; and, lastly, in the Maia squinado, or Spider Crab (Fig. 441,"^) this mass • In this figure are seen the great thoracic g-anglion (n,) from which pro- ceed the superior thoracic nerves (t,) those to the fore feet (f,) to the liiiulcr 3S4 THE SENSORIAL FUNCTIONS. acquires still greater compactness, assumes a more globular form, and has no central perforation. These different forms of structure are also exemplified in the progress of the development of the liigher Crustacea: thus, in the Lobster, the early condition of the nervous sys- tem is that of two separate j^arallcl cords, each having a dis- tinct chain of ganglia, as is the case in the Talitrus: then the cords are observed gradually to approximate, and the gan- glia on each side to coalesce, as represented in the Cymo- thoa: and at the period when the limbs begin to be deve- loped, the thoracic ganglia approach one another, unite in clusters, and acquire a rapid enlargement, preparatory to the growth of the extremities from that division of the body, the abdominal ganglia remaining of the same size as before. The cephalic ganglion, which was originall)^ double, and has coalesced into one, is also greatly developed, in corre- spondence with the growth of the organs of sense. The next remarkable change is that taking place in the hinder por- tions of the nervous cords, which are shortened, at the same time that their ganglia are collected into larger masses, pre- paratory to the growth of the tail and hinder feet; so that throughout the whole extent of the system the number of ganglia diminishes in the progress of development, while their size is augmented. All Insects have the nervous system constructed on the same general model as in the last mentioned classes; and it assumes, as in the Crustacea, various degrees of concentra- tion in the different stages of development. As an example we may take the nervous system of the Sphmx ligusiri, of which representations are given in the larva, pupa, and ima- feet (f.) and the abdominal nervous trunk (n;) the cephalic ganglion (c,) communicating by means of two nervous cords (o,) which surround the cEsophagus and entrance into the stomach (s,) with the thoracic ganglion (b;) and sending ofT the optic nerve (e) to the eyes (e,) and the motor ner\'es (m,) to the muscles of those organs; and also the nerves (a) to the internal antennse, and the nerves (x) to the external antennsc (a.) NERVOUS SYSTEM OF INSECTS. 385 go States, wholly detached from the body, and of their na- tural size, in Figures 442, 443, and 444.* This system in the larva (Fig. 442) has the same simple form as in the Annelida, or in the Talitrus, for it consists of * These figures were drawn by Mr. Newport, from orlj^inal preparations made by himself. The same numbers in each refer to the same parts; so that by comparing the figures with one another, a judgment may be formed of the changes of size and situation which occur in the progress of the prin- cipal transformations of the insect. Numbers 1 to 11 indicate the scries of ganglia which are situated along the under side of the body, and beneath the alimentary canal. Of these the first five are the thoracic, and the last six the abdominal ganglia; while the cephalic, or cerebral ganglion (17) is situated above the oesophagus and dorsal vessel, and communicates by two nervous chords with the first of tlie series, or sub-asopliageal ganglion (1,) which is, in eveiy stage of the insect, contained witliin the head, and distri- butes nerves to the parts about tiie mouth. The next ganglion (2) becomes obhterated at a late period of the change from the pupa to the imago state: the third (3) remains, but the two next (4, 5) coalesce to form, in the ima- go, the large thoracic ganglion; while the two which follow (6 and 7,) be- come wholly obliterated before the insect attains the imago stale, the inter- vening cords becoming shorter, and being, with the nerves they send out, carried forwards. The last four (8, 9, 10, 11) of the abdominal ganglia re- main, with but little alteration, in all the stages of metamorphosis: in the larva, they supply nerves to the false feet. The nerves (12, 13) which sup- ply the wings of the imago, are very small in the larva; and tliey arise by two roots, one derived from the cord, and one from the ganglion. The nerves sent to the three pairs of anterior, or true legs, are marked 14, 15, 16. The nervous system of the larva is exhibited in Fig. 442, that of tlie pupa in Fig. 443, and that of the imago in Fig. 444. It will be seen that in the pupa the abdominal ganglia are but little changed; but those situated more forward (6, 7) are brought closer together by the shortening of the inter- vening cord, preparatory to their final obliteration in the imago; a cluuige which those in front of them (4, 5) have already undergone. The pro- gressive development of the optic (18) and antennal (19) nerves may also be traced. Mr. Newport has also traced a set of nerves (20) which arise from distinct roots, and which he found to be constantly distributed to the organs of respiration. A detailed account of the anatomy of the nervous system of the Sphinx ligusiri, and of the changes it endergoes up to a certain period, is given by Mr. Newport in a paper in the Phil. Trans, for 1832, p. 383. lie has since completed the inquiry to the last transformation of this and other insects, and has lately presented to the Uoyal Society an account of his researches. Vol. II. 49 386 THE SENSORIAL FUNCTIONS. a longitudinal series of ganglia, usually twelve or thirteen in number, connected in their whole length by a double fila- ment. By degrees the difi'erent parts of which it consists approach each other, the thoracic ganglia, in particular, coa- lescing into larger masses, and becoming less numerous, some being apparently obliterated; the whole cord becomes in con- 444 443 442 sequence shorter, and the abdominal ganglia are carried for- wards. The optic nerves are greatly enlarged during the latter stages of transformation, and are often each of greater magnitude than the brain itself. A set of nerves has also been discovered, the course of which is peculiar, and appears to correspond with the sympathetic or ganglionic system of nerves in vertcbrated animals, while another nerve resem- bles in its mode of distribution, the pneumo-gasiric nerve, or par vagum. Very recently Mr. Newport has distinctly traced a separate nervous tract, which he conceives gives NERVOUS SYSTEM OF INSECTS. 387 origin to the motor nerves, while the subjacent column sends out the nerves of sensation. In the next great division of the animal kingdom, which includes all molluscous animals, the nervous ganglia have a circular, instead of a longitudinal arrangement. The first example of this type occurs in the JJsterias, where the ner- vous system (Fig. 445) is composed of small ganglia, equal 446 445 448 in number to the rays of the animal, and disposed in a cir- cle round the central aperture or mouth, but occupying si- tuations intermediate between each of the rays. A nerve is sent off from both sides of each ganglion, and passes along the side of the rays, each ray receiving a pair of these nerves. In the Holothuria there is a similar chain of ganglia, encircling the oesophagus; and the same mode of arrangement prevails in all the bivalve Mollusca, except that, besides the oesophageal ganglia, others are met with in different parts of the body, distributing branches to the viscera, and connected with one another and with the oesophageal ganglia, by filaments, so as to form with them one continuous nervous system. In the Gasteropoda, which are furnished with a distinct head and organs of the higher 388 THE SENSORIAL FUNCTIONS. senses, (such as the *^plysia, of which the nervous system is exhihited in Fig. 446,) there is generally a special cephalic ganglion (c,) which may be supposed to serve the office of brain.* In others, again, as in the Patella (Fig. 447,) the cephalic ganglion is scarcely discernible, and its place is sup- plied by two lateral ganglia (l, l;) and there is besides a a transverse ganglion (t,) below the oesophagus. The ce- phalic ganglion, on the other hand, attains a considerable size in the Cephalopoda (c, Fig. 448,) where it has extensive connexions with all the parts of the head: the optic ganglia (o, o,) in particular, are of very great size, each of them, singly, being larger than the brain itself.t § 2. Nervous System of Vertebrated Jinimals. The characteristic type of the nervous system of verte- brated animals is that of an elongated cylinder of nervous matter, (m, z. Fig. 449,) extending down the back, and lodged in the canal formed by the grooves and arches of the vertebrce. It has received the name of spinal marrow^ or, more properly, spinal cord: and, (as is seen in the transverse section. Fig. 450,) is composed of six parallel columns, two posterior, two middle, and two anterior, closely joined to- gether, but leaving frequently a central canal, which is filled with fluid. On each side of the spinal cord, and between all the adjacent vertebras, there proceed two sets of nervous filaments, those which are continuous with the posterior co- lumns (p,) being appropriated to the function of sensation; and those arising from the anterior columns (a,) being sub- * This figure also shows ag-ang-lion (a,) which is placed higher, and com- municates by lateral filaments with the cephalic gang-lion (c;) two lateral gunglia (l, l,) of great size; and a large abdominal ganghon (g.) f Some peculiarities in the structure of the cephalic ganghon of the Sepia have been supposed to indicate an approach to the vertebrated structure; for this ganglion, together with the labyrinth of the ear, is enclosed in a cartila- ginous ring, perforated at the centre to allow of the passage of the CESopha- giis, and imagined to be analogous to a cranium. NERVOUS SYSTEM OP VERTEBRATA. 389 servient to voluntary motion. The former, soon after their exit from the spine, pass through a small ganglion (g,) and 451 then unite with the nerves from the anterior column, com- posing, by the intermixture of their fibres, a single nervous 390 THE SENSORIAL FUNCTIONS. trunk (n,) which is afterwards divided and subdivided in the course of its farther distribution, both to the muscular and the sentient organs of the body. Each of these spinal nerves also sends branches to the ganglia of the sympathetic nerve, which, as was formerly described, passes down on each side, parallel and near to the spine. Enlargements of the spinal marrow are observed in those parts, (w and l. Fig. 449,) which supply the nerves of the extremities, the increase of diameter being proportional to the size of the limbs requiring these nerves. In Serpents, which are wholly destitute of limbs, the spinal marrow is not enlarged in any part, but is a cylindrical column of uni- form diameter. In Fishes, these enlargements are in pro- portion to the relative size and muscularity of the lateral fins, and correspond to them in their situation. The Piper Gur- nard ( Tri'^la lyra,) which is a species of flying fish, having very large pectoral fins, that portion of the spinal marrow supplying their muscles with nerves (as seen in the space between m and s. Fig. 451,) has numerous enlargements, presenting a double row of tubercles. Fishes which possess electrical organs have a considerable dilatation of the spinal marrow, answering to the large nerves w^hich are distributed to those organs. Birds which fly but imperfectly, as the Gallinaceous tribe and the Scansoi^es, have the posterior enlargement much greater than the anterior; a disproportion which is particularly remarkable in the Ostrich. On the contrary, the anterior enlargement is much more considera- ble than the posterior in birds which have great power of flight. In the Dove, of which the brain and whole extent of the spinal marrow are shown in Fig. 449, the enlargements (w and l) corresponding to the wings and legs respectively, are nearly of equal size. In Quadrupeds, we likewise find the relative size of these enlargements corresponding to that of fore and hind extremities. When the latter are absent, as in the Cetacea, the posterior dilatation does not exist. The brain (b) may be regarded as an expansion of the an- terior or upper end of the spinal marrow; and its magnitude, NERVOUS SYSTEM OF VERTEBRATA. 391 as well as the relative size of its several parts, vary much in the different classes and families of vertebrated animals. This will appear from the inspection of the figures I have given of this organ in various species, selected as specimens from each class, viewed from above; and in all of which I have, indicated corresponding parts by the same letters of reference. The portion (m) of the brain, which appears as the im- mediate continuation of the spinal marrow (s,) is termed the medulla oblongata. The single tubercle (c,) arising from the expansion of the posterior columns of the spinal mar- row, is termed the cerehellum, or little brain. Next follow the pair (t) which are termed the oj^lic tubercles, or lobes,* and appear to be productions from the middle columns of the spinal marrow. These are succeeded by another pair of tubercles (h,) which are called the cerebral hemispheres, and the origin of which may be traced to the anterior columns of the spinal marrow. There is also generally found, in front of the hemispheres, another pair of tubercles (o,) which, being connected with the nerves of smelling, have been called the olfactory lobes, or tuberclesA These are the principal parts of the cerebral mass to be here noticed, for I purposely omit the mention of the minuter divisions, which, though they have been objects of much attention to anato- mists, unfortunately furnish no assistance in understanding the physiology of this complicated and wonderful organ. On comparing the relative proportions of the brain and of the spinal marrow in the four classes of vertebrated ani- mals, a progressive increase in the size of the former will be observed as we ascend from Fishes to Reptiles, Birds, and Mammalia. This increase in the magnitude of the brain arises chiefly from the enlargement of the cerebral hemi- spheres (h,) which, in the inferior orders of fishes, as in the * In the Mammalia, and in Man, tliey have been often desig-nated by the very inappropriate name of Corpora quadriiremina. ■\ Several cavities, termed Ventricles, are occasionally found in the inte- rior of the principal tubercles of the brain; but their use is unknow"- 392 THE SENSORIAL FUNCTIONS. « Trigla lyra, or Piper Gurnard, (Fig. 451,) and in the Mu- rxna conger, or Conger Eel, (Fig. 452,) are scarcely dis- cernible. They are very small in the Percct Jluviatilis, or common Perch (Fig. 453;) but more developed in Reptiles, as in the Tcstudo inydas, or Green Turtle, (Fig. 454,) and in the Crocodile, (Fig. 455;) and still more so in Birds, as is seen in the brain of the Dove, (Fig. 449;) but, most of all, in the JNIammalia, as is exemplified in the brain of the Lion, (Fig. 456.) On the other hand, the optic tubercles (t) are largest, compared with the rest of the brain, in Fishes; and their relative size diminishes as we ascend to Mammalia: and the same observation applies also to the olfactory lobes, (o.) The relative positions of the parts of the brain are much influenced by their proportional development. This will be rendered manifest by the lateral views of the brains of the Perch, the Turtle, the Dove, and the Lion, presented in Figures 457, 458, 459, and 460, respectively, where the same letters are employed to designate the same parts as in the preceding figures. In Fishes, all the tubercles which compose this organ, are disposed nearly in a straight line, continuous with the spinal marrow, of which, as they scarce- ly exceed it in diameter, they appear to be mere enlarge- ments. As the skull expands more considerably than the brain, this organ does not fill its cavity, but leaves a large space filled with fluid. Some degree of shortening, how- ever, may be perceived in the brain of the Perch (Fig. 457;) for the medulla oblongata (m) is doubled underneath the ce- rebellum (c,) pushing it upwards, and rendering it more prominent than the other tubercles. This folding inwards, and shortening of the whole mass, proceeds to a greater extent as we trace the structure upwards, as may be seen in the brain of tlie Green Turtle (Fig. 458.) In that of Birds, of which Fig. 459 presents a vertical section, the optic tubercles have descended from their former place, and assumed a lateral po- sition, near the lower surface of the brain, lying on each side of the medulla oblongata, at the part indicated by the letter NERVOUS SYSTEM OF VERTEBRATA. 393 T. In the Mammalia, as the Lion (Fig. 4G0,) they are lodged quite in the interior of the organ, and concealed hy the expanded hemispheres (h;) their position only being marked by the same letter (t.) These changes are conse- quences of the increasing development of the brain, com- pared with that of the cavity in which it is contained, re- quiring every part to be more closely packed; thus, the lay- ers of the hemispheres in INIammalia are obliged, from their great extent, to be plaited and folded on one another, pre- senting at the surface curious windings, or convolutions , as they are called (seen in Fig. 456,) which do not take place in the hemispheres of the inferior classes. The foldings of the substance of the cerebellum produce, likewise, even in birds, transverse furrows on the surface; and from the in- terposition of a substance of a gray colour between the la- minae of the white medullary matter, a section of the ce- rebellum presents the curious appearance (seen in Fig. 459,) denominated, from its fancied resemblance to a tree, the Jirhor Vitx, 461 .' Thus far we have followed an obvious gradation in the development and conccutration of the dilierent parts of the Vol. II. 50 394 THE SENSORIAL FUNCTIONS. brain: but on arriving at Man the continuity of the series is suddenly disturbed by the great expansion of the hemi- spheres, (Fig. 4f)l,) whicli, compared with those of quadru- peds, bear no sort of proportion to the rest of the nervous system. Both Aristotle and Pliny have asserted that the absolute, as well as the comparative size of the human brain is "-rcater than in any other known animal: exceptions, how- ever, occur in the case of the Elephant, and also in that of the Whale, wliosc brains arc certainly of greater absolute bulk than that of man. But all the large animals, with which we are familiarly acquainted, have brains considera- bly smaller; as will readily appear from an examination of their skulls, which are narrow and compressed at the part occupied by the brain; the greater part of the head being taken up by the development of the face and jaws. In Man, on the other hand, the bones of the skull rise perpendicu- larly from the forehead, and are extended on each side, so as to form a capacious globular cavity for the reception and defence of this most important organ. It is chiefly from the expansion of the hemispheres, and the development of its convolutions, that the human brain derives this great aug- mentation of size.* • This will be apparent from the vertical section of the human brain, Fig-. 461; where, as before, s is the spinal marrow; ji, the medulla oblongata; c, the cerebellum, with the arbor vitae,- t, the optic tubercles, or corpora quad- rigemina, dwindled to a very small size, compared with their bulk in fishes: p, the pineal gland, supposed by Des Cartes to be the seat of the soul; v, one of the lateral ventricles; a, the corpus callosum; and h, ii, h, the hemi- spheres. Several expedients have been proposed for estimating the relative size of the brain in different tribes of animals, with a view of deducing conclusions as to the constancy of the relation which is presumed to exist between its greater magnitude and the possession of higiier intellectual faculties. The most celebrated is that devised by Camper, and which he ternied the facial angle, composed of two lines, one drawn in the direction of the basis of the skull, from the car to the roots of the upper incisor teeth, and the other from the latter point, touching the most projecting l^ai-t of the forehead. Cam- per conceived that th.e magnitude of this angle would correctly indicate the size of the brain, as compared with the organs of the principal senses which FUNCTIONS OF THE BRAIN. 395 § 3. Funciions of the Brain. Physiologists have m all ages sought for an elucidation of the functions of the brain by the accurate examination of its structure, which evidently consists of a congeries of me- dullary fibres, arranged in the most intricate manner. Great pains have been bestowed in unravelling the tissue of these fibres, in the hope of discovering some clew to the perplex- ing labyrinth of its organization: but nearly all that has been learned from the laborious inquiry, is that the fibres of the brain are continuous with those which compose the columns of the spinal marrow; that they pass, in their course, through masses of nervous matter, which appear to be analogous to ganglia; and that their remote extremities extend to the sur- face of the convolutions of the brain and cerebellum, which are composed of a softer and more transparent gray matter, termed the cortical or cineritioiis substance of the brain. It is a remarkable fact, that in vertebrated animals all the organs which are subservient to the sensorial functions are double, those on one side being exactly similar to those on the other. We see this in the eyes, the ears, the limbs, and all the other instruments of voluntary motion; and in like manner the parts of the nervous system which arc connect- ed with these functions are all double, and arranged sym- metrically on the two sides of the body. The same law of symmetry extends to the brain: every part of that organ which is found on one side is repeated on the other; so that, strictly speaking, we have two brains, as well as two optic nerves and two eyes. But in order that the two sets of fibres may co-operate, and constitute a single organ of sen- sation, corresponding with our consciousness of individual- ity, it was necessary that a free communication should be compose the face: but the Hdlacy of this criterion of animal sag'acity has been shown in a great many cases. 396 THE SENSORIAL FUNCTIONS. cstablislicd between llie parts on both sides. For this pur- pose there is provided a set of medullary fibres, passing di- rectly across from one side of the brain to the other; these constitute what are called the Cominissiires of the brain.* The question, however, still recurs: — What relation does all this artificial intertexture and accumulation of fibres bear to the mental operations of which we are conscious, such as memory, abstraction, judgment, imagination, volition? Are there localities set apart for our different ideas in the store- house of the cerebral hemispheres, and are they associated by the material channels of communicating fibres? Are the mental phenomena the effects, as was formerly supposed, of a subtle fluid, or anivial spirits, circulating with great ve- locity along invisible canals in the nervous substance? or shall we, with Hartley, suppose them to be the results of vibrations and vibratiuncles, agitating in succession the finer threads of which this mystic web has been construct- ed? But a little reflection will suffice to convince us that these, and all other mechanical hypotheses, which the most fanciful imagination can devise, make not the smallest ap- proach to a solution of the difficulty; for they, in fact, do not touch the real subject to be explained, namely, how the affections of a material substance can influence and be influ- enced by an immaterial agent. All that we have been able to accomplish has been to trace the impressions from the organ of sense along the communicating nerve to the senso- rium: beyond this the clew is lost, and we can follow the process no farther. • The principal commissure of the human brain, called tlie corpus callosum, is seen at a, Fig. 461. Dr. Macartney, in a paper which he read at the late meeting at Cambridge of the Britisli Association for the Advancement of Science, described the structure of the human brain, as discovered by his pe- culiar mode of dissection, to be much more complicated than is generally supposed. He obsei-ved that its fibres are interlaced in the most intricate manner, resembling the plexuses met with among the nerves, and establish- ing the most extensive and general communications between every part of the cerebral mass. FUNCTIONS OP THE BRAIN. 397 The exact locality of the sensorium has ])ccn eagerly sought for by }3hysiologists in every age. It would appear, from the results of the most recent inquiries, that it cer- tainly does not extend to the whole mass of the brain, but has its seat more especially in the lower part, or basis of that organ. It differs, however, in its locality, in different classes of animals. In man, and tlie mammalia wliich ap- proach the nearest to him in their stiucture, it occupies some part of the region of the medulla oblongata, probably the spot where most of the nerves of sense are observed to terminate. In the lower animals it is not confined to this region, but extends to the upper part of the spinal marrow. As we descend to the inferior orders of the animal kingdom, we find it more and more extensively diffused over the spi- nal marrow; and in the Invertebrata the several ganglia ap- pear to be endowed with this sensorial property: but, be- coming less and less concentrated in single masses, the cha- racter of individuality ceases to attach to the sensorial phe- nomena; until, in Zoophytes, vv^e lose all traces of ganglia and of nervous filaments, and every part appears to possess an inherent power of exciting sensation, as well as perform- ing muscular contractions. Beyond this point we can derive no farther aid from Ana- tomy, since the intellectual operations of which we arc con- scious bear no conceivable analogy with any of the configu- rations or actions of a material substance. Although the brain is constructed with evident design, and composed of a number of curiously wrought parts, we are utterly unable to penetrate the intention with which they are formed, or to perceive the slightest correspondence which their configu- ration can have with the functions they respectively per- form. The map of regions which modern Phrenologists have traced on the surface of the head, and which they sup- pose to have a relation to different faculties and propensities, does not agree either with the natural divisions of the brain ^or with the metaphysical classification of mental phenome- 39R THE SENSORIAL FUNCTIONS. na.* Experiments and pathological observations, however, seem to show that the hemispheres of the brain are the chief instruments by whicli the intellectual operations are carried on; that the central parts, such as the optic lobes and the medulla oblongata, are those principally concerned in sen- sation; and that the cerebellum is the chief sensorial agent in voluntary motion. § 4. Comparative Physiology of Perception. Of the perceptions of the lower animals, and of the laws which they obey, our knowledge must, of necessity, be ex- tremely imperfect, since it must be derived from a compari- son with the results of our own sensitive powers, which may differ very essentially from those of the subjects of our observation. The same kind of organ which, in ourselves, conveys certain definite feelings, may, when modified in other animals, be the source of very different kinds of sen- sations and perceptions, of which our minds have not the power to form any adequate conception. INIany of the qualities of surrounding bodies, which escape our more ob- tuse senses, may be distinctly perceived, in all their grada- tions, by particular tribes of animals, furnished with more delicate organs. Many quadrupeds and birds possess pow- ers of vision incomparably more extensive than our own; in acuteness of hearing, we are excelled by a great number of animals, and in delicacy of taste and smell, there are few quadrupeds which do not far surpass us. The organ of smell, in particular, is often spread over a vast extent of sur- face, in a cavity occupying the greatest part of the head; so that the perceptions of this sense must be infinitely diver- sified. • For a summary of the doctrines of EJrs, Gall and Spurzheim, I beg leave to refer the reader to an account which I drew up, many years ago, for the Encyclopedia liritannica, and which composed the article "CnASfioscoPY" in the last supplement to that work, edited by Mr. Napier. FUNCTIONS OF THE BRAIN. 399. Bats have been supposed to possess a peculiar, or sixth sense, enabling them to perceive the situations of external ob- jects without the aid either of vision or of touch. The prin- cipal facts upon which this opinion has been founded were discovered by Spallanzani, who observed that these aniniala would fly about rapidly in the darkest chambers, although various obstacles were purposely placed in their way, with- out striking against or even touching tlicm. They continued their flight with the same precision as before, threading their way through the most intricate passages, when their eyes were completely covered, or even destroyed. Mr. Jurine, who made many experiments on these animals, concludes that neither the senses of touch, of hearing or of smell, were the media through which bats obtain perceptions of the pre- sence and situation of suri'ounding bodies; but he ascribes this extraordinary faculty to the great sensibility of the skin of the upper jaw, mouth, and external ear, which are fur- nished with very large nerves. '■ The wonderful acuteness and power of discrimination which many animals exercise in the discoverv and selection of their food, has often suggested the existence of new senses, different from those which we possess, and Anveying pecu- liar and unknown powers of perception. An organ, which appears to perform some sensitive function of this kind, has been discovered in a great number of quadrupeds by Jacob- son. t In the human skeleton there exists a small perfora- tion in the roof of the mouth, just behind the sockets of the incisor teeth, forming a communication with the under and fore part of the nostrils. This canal is perceptible only in the dried bones; for, in the living body, it is completely closed by the membrane lining the mouth, which sends a prolongation into it: but in quadrupeds, this passage is per- vious even during life, and is sometimes of considerable width. Jacobson found, on examinina; this structure with * Sir Anthony Carlisle attributes this power to the extreme delicacy of hearing- in this animal. f See Annales du Miiscej xviii. 412. 400 THE SENSORIAL FUNCTIONS. uttcnllon, that the canal led to two glandular organs of an oblong shape, and enclosed in cartilaginous tubes: each gland has in its centre a cavity which communicates above with the general cavity of the nostrils. These organs lie can- cealed in a hollow groove within the bone, where they are carefully protected from injury: and they receive a great number of nerves and blood vessels, resembling in this re- spect the organs of the senses. Their structure is the same in all quadrupeds in which they have been examined; but they are largest in the family of the Rodentia, and next in that of the Riiminantia; in the Horse, they are still very large, but the duct is not pervious; while, in carnivorous quadru- peds, they are on a smaller scale. In Monkeys, they may still be traced, although extremely small, appearing to form a link in the chain of gradation connecting this tribe with the human race, in whom every vestige of these organs has disappeared, excepting the aperture in the bones already no- ticed. Any use that can be attributed to these singularly constructed organs must evidently be quite conjectural. The ample supply of nerves which they receive would indi- cate their performing some sensitive function; and their si- tuation would Aoint them out as fitting them for the ap- preciation of objects presented to the mouth to be used as food; hence it is probable that the perceptions they convey have a close affinity with those of smell and taste. The larger cartilaginous fishes, as Shm^ks and Rays, have been supposed by Treviranus to be endowed with a peculiar sense, from their having an organ of a tubular structure on the top of the head, and immediately under the skin; Roux con- siders it as conveying sensations intermediate between those of touch and hearing; while De Blainville and Jacobson re- gard it merely as the organ of a finer touch. The perceptive powers of Insects must embrace a very different, and, iii many respects, more extended sphere than our own. These animals manifest by their actions that they perceive and anticipate atmospheric changes, of which our senses give us no information. It is evident; indeed, that PERCEPTIONS OP ANIMALS. 401 the impressions made by external objects on their sentient organs must be of a nature widely difl'erent from those which the same objects communicate to ourselves. While with re- gard to distance and mas2;nitudc our perceptions take the widest range, and appear infinitely extended when com- pared with those of insects, yet they may, in other respects, be greatly inferior. The delicate discrimination of the more subtle affections of matter is, perhaps, compatible only with a minute scale of organization. Thus, the varying degrees of moisture or dryness of the atmosphere, the continual changes in its pressure, the fluctuations in its electrical state, and va- rious other physical conditions, may be objects of distinct perception to these minuteanimals. Organsmayexistinthem, appropriated to receive impressions, of which we can have no idea; and opening avenues to various kinds of knowledge, to which we must ever remain utter strangers. Art, it is true, has supplied us with instruments for discovering and measuring many of the properties of matter, which our un- assisted senses are inadequate to observe. But neither our thermometers, nor our electroscopes, our hygrometers, nor our galvanometers, how^ever skilfully devised or elaborately constructed, can vie in delicacy and perfection with that re- fined apparatus of the senses which nature has bestowed on the minutest insect. There is reason to believe, as Dr. Wol- laston has showm, that the hearing of insects comprehends a range of perceptions very different from that of the same sense in the larger animals; and that a class of vibrations too rapid to excite our auditory nerves, is perfectly audible to them. Sir John Herschel has also very clearly proved that, if we admit the truth of the undulatory theory of lighl, it is easy to conceive how the limits of visible colour may be established; for if there be no nervous fibres in unison with vibrations, more or less frequent than certain limits, such vibrations, though they reach the retina, will produce no sensation. Thus, it is perfectly possible that insects, and other animals, may be incapable of being affected by any of the colours which we perceive; while they may be suscepti- VoL. II. , 51 402 THE SENSORIAL FUNCTIONS. ble of receiving; distinct luminous impressions from a class of vibrations which, applied to our visual organs, excite no sensation.* The functions of the antennae, which, though of various forms, are organs universally met with in this class of animals, must be of great importance, though obscurely known; for insects when deprived of them appear to be quite lost and bewildered. The Torpedo, tlie Gymnotiis, and several other fishes, arc furnislied with an electrical apparatus, resembling the Voltaic battery, wnich they liave the power of charging and discharging at pleasure. An immense profusion of nerves is distributed upon this organ; and we can hardly doubt that they comniunicate perceptions, with regard to electricity, very different from any that we can feel. In general, in- deed, it may be remarked, that the more an organ of sense differs in its structure from those which we ourselves pos- sess, the more uncertain must be our knowledge of its func- tions. We may, without any great stretch of fancy, conceive ourselves placed in the situation of the beasts of the forest, and comprehend what are the feelings and motives which animate the quadruped and the bird. But how can we transport ourselves, even in imagination, into the dark re- cesses of the ocean, which we know are tenanted by multi- tudinous tribes of fishes, zoophytes, and mollusca? How can we figure to ourselves the sensitive existence of the worm or the insect, organized in so different a manner to ourselves, and occupying so remote a region in the expanse of creation? How can we venture to speculate on the perceptions of the animalcule, whose world is a drop of fluid, and whose fleet- ing existence, chec^uered, perhaps, by various transforma- tions, is destined to run its course in a few hours? Confining our inquiries, then, to the more intelligible in- tellectual phenomena displayed by the higher animals, we readily trace a gradation which corresponds with the de- velopment of the central nervous organ, or brain. That the • Encyclopjcdiu McUupolilanu, Article *' Ligut." PERCEPTIONS OF ANIMALS. 403 comparison may be fairly made, however, it is necessary to distinguisli those actions which are the result of the exer- cise of the intellectual faculties, from those which are called instinctive, and are referrible to other sources. Innumera- ble are the occasions in which the actions of animals appear to be guided by a degree of sagacity not derivable from ex- perience,and apparently implying a foreknowledge of events, v^rhich neither experience nor reflection could have led them to anticipate. We cannot sufliciently admire the provident care displayed by nature in the preservation both of the in- dividual and of the species, which she has intrusted, not to the slow and uncertain calculations of prudence, but to in- nate faculties, prompting, by an unerring impulse, to the performance of the actions required for those ends. We see animals providing against the approach of winter, the effects of which they have never experienced, and employ- ing various means of defence against enemies they have ne- ver seen. The parent consults the welfare of the offspring she is destined never to behold; and the young discovers and pursues without a guide that species of food which is best adapted to its nature. All these unexplained, and, perhaps, inexplicable facts, we must content ourselves with classing under the head o^ instinct, a name which is, in fact, but the expression of our ignorance of the nature of that agency, of which we cannot but admire the ultimate cflects, while we search in vain for the eflicient cause. In all the inferior orders of the animal creation, wliere in- stincts are multiplied, while tiie indications of intellect are feeble, the organ which performs the office of the brain is comparatively small. The sensitive existence of these ani- mals appears to be circumscribed within the perceptions of the moment, and their voluntary actions have reference chiefly to objects which arc present to tlie sense. In i)ro- portion as the intellectual faculties of animals are multiplied, and embrace a wider sphere, additional magnitude and com- plication of structure are given to the nervous substance which is the organ of those faculties. The greater the power 404 THE SENSORIAL FUNCTIONS. of combining; ideas, and of retaining them in the memory, the greater do we find the development of the cerebral he- mispheres. These parts of ihe brain are comparatively small, as we have seen, in fishes, reptiles, and the greater number of birds; but in the mammalia they are expanded in a degree nearly proportional to the extent of memory, sa- gacity and docility. In man, in whom all the faculties of sense and intellect are so harmoniously combined, the brain is not only the largest in size, but beyond all comparison the most complicated in its structure.* A large brain has been bestowed on man, evidently with the design that he should exercise superior powers of intel- lect; the great distinguishing features of which are the ca- pacity for retaining an immense variety of impressions, and the strength, the extent, and vast range of the associating principle, which combines them into groups, and forms them into abstract ideas. Yet the lower animals also possess their share of memory, and of reason: they are capable of ac- quiring knowledge from experience; and, on some rare oc- casions, of devising expedients for accom])lishing particular ends. But still this knowdcdoe and these efforts of intellect are confined within very narrow^ limits; for nature has as- signed boundaries to the advancement of the low^er animals, which they can never pass. If one favoured individual be selected for a special education, some additional share of in- telligence may, perhaps, with infinite pains, be infused; but the improvement perishes with that individual, and is w'holly lost to the race. By far the greater portion of that know- ledge which it imports them to possess is the gift of nature, who has wisely implanted such instinctive impulses as are necessary for their preservation. Man, also, is born with instincts, but they are few in number, compared with those • All Ihc parts nut witli m the brain of animals exist also in the brain of man; while sevcnil of those foiiiid in man arc either extremely small, or alto- gether absent in the brains of the lower animals. Soemmerring- has enu- njerated no less than fifteen material anatomical diflerences between the hu- man brain and tliat of the ape. PERCEPTIONS OF ANIMALS. 405 of the lower animals; and, unless cultivated and improved by reason and education, would, of themselves, produce but inconsiderable results. That of which the effects are most conspicuous, and which is the foundation of all that is noble and exalted in our nature, is the instinct of Sympathy. The affections of the lower animals, even between individuals of the same species, are observable only in a few instances: for in general they are indifferent to each other's joys or suffer- ings, and regardless of the treatment experienced by their companions. The attachment, indeed, of the mother to her offspring, as long as its wants and feebleness require her aid and protection, is as powerful in the lower animals, as in the human species: but its duration, in the former case, is con- fined, even in the most social tribes, to the period of help- lessness; and the animal instinct is not succeeded, as in man, by the continued intercourse of affection and kind offices, and those endearing relations of kindred, which are the sources of the purest happiness of human life. While Nature has, apparently, frowned on the birth of man, and brought him into the world weak, naked, and de- fenceless, unprovided with the means of subsistence, and exposed on every side to destruction, she has, in reality, im- planted in him the germ of future greatness. The helpless- ness of the infant calls forth the fostering care and ten- derest affections of the mother, and lays the deep founda- tions of the social union. The latent eneruries of his mind and body are successively, though slowly developed. AVhile the vital organs are actively engaged in the execution of their different offices, while the digestive apparatus is exer- cising its powerful chemistry, while myriads of minute ar- teries, veins, and absorbents are indefatigably at work in building and modelling this complex frame, the sentient principle is no less assiduously and no less incessantly em- ployed. From the earliest dawn of sensation it is ever busy in arranging, in combining, and in strengthening the im- pressions it receives. Wonderful as is the formation of the bodily fabric, and dilhcult as it is to collect its history, still 406 THE SENSORIAL FUNCTIONS. more marvellous is the progressive construction of the hu- man mind, and still more arduous the task of tracing the finer threads which connect the delicate web of its ideas, which fix its fleeting perceptions, and which establish the vast system of its associations, and of following the long se- ries of gradations by which its affections are expanded, pu- rified, and exalted, and the soul prepared for its higher des- tination in a future stage of existence. Here, indeed, we perceive a remarkable interruption to that regular gradation, which we have traced in all other parts of the animal series; for between man and the most sagacious of the brutes there intervenes an immense chasm, of which we can hardlv estimate the magnitude. The func- tions which are purely vital, and are necessary for even the lowest degree of sensitive existence, are possessed equally by all animals: in the distribution of the faculties of mere sensation a greater inequality may be perceived: the intel- lectual faculties, again, are of a more refined and nobler cha- racter, and being less essential to animal life, are dealt out by nature with a more sparing and partial hand. Between the two extremities of the scale we find an infinite number of intermediate degrees. The more exalted faculties are possessed exclusively by man, and constitute the source of the immense superiority he enjoys over the brute creation, which so frequently excels him in the perfection of subor- dinate powers. In strength and swiftness he is surpassed by many quadrupeds. In vain may he wish for the power of flight possessed by the numerous inhabitants of air. He may envy that range of sight which enables the bird to dis- cern from a height at which it is itself invisible to our eyes, the minutest objects on the surface of the earth. He may regret the dulness of his own senses, w^hen he adverts to the exquisite scent of the hound, or the acute hearing of the bat. While the delicate perceptions of the lower animals teach them to seek the food which is salutary, and avoid that which is injurious, man alone seems stinted in his pow- ers of discrimination, and is compelled to gather instruction INTELLECTUAL FACULTIES OF MAN. 407 from a painful and hazardous experience. Bui if nature has created him thus apparently helpless, and denied him those instincts with which she has so liberally furnished the rest of her offspring, it was only to confer upon him gifts of in- finitely higher value. While in acuteness of sense he is surpassed by inferior animals, in the powers of intellect he stands unrivalled. In the fidelity and tenacity with which impressions are retained in his memory, in the facility and strength with which they are associated, in grasp of compre- hension, in extent of reasoning, in capacity of progressive improvement, he leaves all other animals at an- immeasura- ble distance behind. He alone enjoys in perfection the gift of utterance; he alone is able to clothe his thoughts in words; in him alone do we find implanted the desire of examining every department of nature, and the power of extending his views beyond the confines of this globe. On him alone have the high privileges been bestowed of recognising and of adoring the Power, the Wisdom, and the Goodness of the Author of the Universe, from whom his being has ema- nated, to whom he owes all the blessino-s which attend it, and by whom he has been taught to look forward to bright- er skies and to purer and more exalted conditions of exist- ence. Heir to this high destination, Man discards all alli- ance with the beasts that perish; confiding in the assurance that the dissolution of his earthly frame destroys not the germ of immortality which has been implanted within him, and by the development of which the great scheme of Pro- vidence here commenced, will be carried on, in a future state of being, to its final and perfect consummation. ( 408 ) PART IV. THE REPRODUCTIVE FUNCTIONS. CHAPTER I. REPRODUCTION. Limits have been assigned to tlic duration of all living beings. The same power to vvbom they owe their creation, their organization, and their endowments, has also subjected them to the inexorable Law of Mortality; and has ordained that the series of actions which characterize the state of life, shall continue for a definite period only, and shall then ter- minate. The very same causes which, at the earlier stages of their existence, promoted their development and growth, and which, at a maturer age, sustained the vigour and ener- gies of the system, produce, by their continued and silent operation, gradual changes in the balance of the functions, and, at a later period, effect the slow demolition of the fabric they had raised, and the successive destruction of the faculties they had originally nurtured and upheld.* With the germs of life, in all organized structures, are conjoined the seeds of decay and of death; and however great may be the powers of their vitality, we know that those powers are finite, and that a time must come when they will be ex- • See the article '• Age," in the Cydopxdia of Practical Mcdiciney where I have enlarged on this subject. EEPRODUCTIO.V. 409 pended, and when their renewal, in that individual, is no longer possible. But although the individual perishes. Nature has taken special care that the race shall be constantly preserved, by providing for the production of new individuals, each spring- ing from its predecessor in endless perpetuity. The pro- cess by which this formation, or rather this apparent crea- tion, of a living being is effected, surpasses the utmost powers of the human comprehension. No conceivable com- binations of mechanical, or of chemical powers, bear ihe slightest resemblance, or the most remote analogy, to or- ganic reproduction, or can afford the least clew to the solu- tion of this dark and hopeless enigma. We must be con- tent to observe and generalize the phenomena, in silent wonder at the marvellous manifestation of express con- trivance and design, exhibited in this department of the economy of created beings. Throughout the whole, both of the vegetable and animal world. Nature has shown the utmost solicitude to secure not only the multiplication of the species, but also the dis- semination of their numbers over every habitable and acces- sible region of the globe, and has pursued various plans for the accomplishment of these important objects. The simplest of all the modes of multiplication consists in the spontaneous division of the body of the parent into two or more parts; each part, when separated, becoming a distinct individual, and soon acquiring the size and shape of the parent. We meet with frequent examples of this pro- cess of fissiparous generation, as it is termed among the infusory animalcules. Many species of Monads, for in- stance, which are naturally of a globular shape, exhibit at a certain period of their development a slight circular groove round the middle of their bodies, which by degrees be- coming deeper, changes their form to that of an hour-glass; and the middle part becoming still more contracted, they present the appearance of two balls, united by a mere point. The monads in this state are seen swimming irregularly in Vol. II. 52 410 THE REPRODUCTIVE FUNCTIOXS. the fluid, as if animated by two different volitions; and, ap- parently for the purpose of tearing asunder the last connect- ing fibres, darting through the thickest of the crowd of sur- rounding animalcules; and the moment this slender ligament is broken, each is seei; moving away from the other, and beginning its independent existence. This mode of sepa- ration is illustrated by Fig. 462, representing the successive changes of form during this progress. In this animalcule the division is transverse, but in others, for example in the 462 463' 0 8 3 8? o Vorticella, (as shown in Fig. 463,) and in most of the larger species, the line of separation is longitudinal. Each animal- cule, thus formed by the subdivision of its predecessor, soon grows to the size which again determines a farther spon- taneous subdivision into two other animalcules; these, in course of time, themselves undergo the same process, and so on, to an indefinite extent. The most singular circum- stance attending this mode of multiplication is that it is im- possible to pronounce which of the new individuals thus formed out of a single one should be regarded as the parent, and which as the offspring, for they are both of equal size. Unless, therefore, we consider the separation of the parts of the parent animal to constitute the close of its individual ex- istence, we must recognise an unbroken continuity in the vitality of the animal, thus transmitted in perpetuity from the original stem, throughout all succeeding generations. This, however, is one of those metaphysical subtleties for which the subject of reproduction aflfords abundant scope, but which it would be foreign to the object of this work to discuss. REPRODUCTION. 411 It is in the animal kingdom only that we meet with in- stances of this spontaneous division of an organic being into parts, where each reproduces an individual of the same spe- cies. All plants, however, are capable of being multiplied by artificial divisions of this kind; thus, a tree may be di- vided longitudinally into a great number of portions, or slips, as they are called, any one of which, if planted sepa- rately and supplied with nourishment, may continue to grow, and may, in time, reproduce a tree similar in all re- spects to the one from which it had originated. This inhe- rent power of reproduction exists even in smaller fragments of a plant; for, when all circumstances are favourable, a stem will shoot from the upper end of the fragment, and roots will be sent forth from its lower end; and, ultimately, a com- plete plant will be formed.* These facts, which are well known to agriculturists, exhibit only the capal)ilities of ve- getative power under circumstances which do not occur in the natural course of things, but have been the effect of hu- man interference. Reproductive powers of a similar kind are exhibited very extensively in the lower departments of the animal king- dom. The Hydra, or fresh water polype, is capable of in- definite multiplication by simple division: thus, if it be cut asunder transversely, the part containing the head soon sup- plies itself with a tail; and the detached tail soon shoots forth a new head, with a new set of tentacula. If any of the ten- tacula, or any portion of one of them, be cut off, the mutila- • Among the conditions necessary for these evolutions of organs .ire, first, the previous accumulation of a store of nourishment in the detached frag- ment, adequate to supply the growth of the new parts; and, secondly, the presence of a sufficient quantity of circulating sap, as a vehicle fbr the trans- mission of that nourishment. It has been found that when these conditions are present, even the leaf of an orange tree, when planted in a favourable soil, sends down roots, and is capable of giving origin to an entire tree. Ac- cording to the observations of Mirandola, the leaf of the BryophyUum, when simply laid on moist groimd, strikes out roots, which quickly penetrate into the soil. (De CandoUe, Physiologic Vcgctale, ii. 677.) The leaves of the, monocotyledonous plants often present the same phenomenon. 412 THE REPRODUCTIVE FUNCTIONS. tion is soon repaired; and if the whole animal be divided into a great number of pieces, each fragment acquires, in a short time, all the parts wliich are wanting to render it a complete individual. The same phenomena are observed, and nearly to the same extent, in the Flanaria. The *^5- terias, the %/lciinia, and some of the lower species of Anne- lida, as the Kais, arc also capable of being multiplied by ar- tificial divisions, each segment having the power of supply- ing others, and containing within itself a kind of separate and individual vitality. A power of more partial regeneration of mutilated parts by new growths, which is very analogous to that of com- plete reproduction, exists in the higher orders of animals, though it does not extend to the entire formation of two in- dividuals out of one. The claws, the feet, and the antennae of the Crustacea, and the limbs of the Arachnida, are re- stored, when lost, by a fresh growth of these organs. If the head of a Snail be amputated, the whole of that part of the animal, including the telescopic eyes, and other organs of sense, will be reproduced. Even among the Vertebrata we find instances of these renovations of mutilated parts; as hap- pens with respect to the fins of fishes: for Broussonet found that in whatever direction they are cut, the edges easily unite; and the rays themselves are reproduced, provided the smallest part of their base has been left. The tails of Newts, and of some species of Lizards, will grow again, if lost: and, what is more remarkable, the eyes themselves, with all their complex apparatus of coats and humours, will, if removed, be replaced by the growth of new eyes as perfect as the for- mer. We have seen that the teeth of Sharks and other fishes are renewed with the utmost facility, when by acci- cident they have been lost. Among Mammalia, similar powers exist, although they are restricted within much nar- rower limits; as is exemplified in the formation of new bones, replacing those which have perished. When we advert to the numberless instances of the reparation of injuries hap- pening to various partJ' of our own frame, we have abun- REPRODUCTION. 413 dant reason to admire and be grateful for the wise and boun- tiful provisions which nature has made for meeting these contingencies. The multiplication of the species by buds, or Cernmipa- rous reproduction, \s exemplified on the largest scale in the vegetable creation. ■ Almost every point of the surface of a plant appears to be capable of giving rise to a new shoot, which, when fully developed, exactly resembles the parent stock, and may, therefore, be regarded as a separate organic being. The origin of buds is wholly beyond the sphere of our observation; for they arise from portions of matter too minute to be cognizable to our organs, with every assistance which the most powerful microscopes can supply. These imperceptible atoms, from which organic beings take their rise, are called gerins. Vegetable germs are of two kinds; those which produce stems, and those which produce roots: and although both may be evolved from every part of the plant, the former are usually developed at the axillse of the leaves; that is, at the angles of their junction with the stem; and also at the ex- tremities of the fibres of the stems; their development being determined by the accumulation of nourishment around them. They first produce buds, which expanding, and put- ting forth roots, assume the form of shoots; and the succes- sive accumulation of shoots, which remain attached to the parent plant,* and to each other, 'is what constitutes a tree. What are called knots in wood are the result of germs, which, in consequence of the accumulation of nourishment around them, are developed to a certain extent, and then * In some rare instances the shoots are removed to a distance from the pa- rent plant, by a natural process: this occurs in some creeping' plants, which propagate themselves by tlie horizontal extension of their branches on the ground wliere thay dip, and strike out new roots, g'iving- rise to stems inde- pendent of the orig-inal plant. This also sometimes happens in the case of tuberous roots, as the potato, which contain a number of germs, surround- ed by nutritive matter, ready to be developed when circumstances are fa- vourable. These portions are called eyes; and each of them, when planted separately, are readily evolved, and give rise to an individual plant. 41 t THE REPRODUCTIVE FUNCTIONS. cease to grow. The Lemna^ or common Duckweed, which, consists of a small circular leaf, floating on the surface of sta«-nant pools, presents a singular instance of the develop- ment of germs from the edges of the leaves, and the suhse- quent separation of the new plant thus formed. In this re- spect the process is analogous to the natural mode of multi- plication met with in the lower orders of Zoophytes, such as the Hydra. At the earliest period at which the young of this animal is visihle, it appears like a small tuhercle, or bud, rising from the surface of the parent hydra: it grows in this situation, and remains attached for a considerable period; at first deriving its nourishment, as well as its mechanical sup- ])ort, from the parent; then occasionally stretching forth its tentacula, and learning the art of catching and of swal- lowing its natural prey. The tube, which constitutes its sto- mach, at first communicates by a distinct opening with that of its parent: but this opening afterwards closes; and the fila- ments by which it is connected with the parent becoming more and more slender, at length break, and the detached hydra immediately moves away, and commences its career of independent existence. This mode of multiplication, in its first period, corresponds exactly with the production of a vegetable by buds; and may therefore be classed among the instances of gemmiparous reproduction; although at a la- ter stage, it differs from it in the complete detachment of the offspring from the parent. Another plan of reproduction is that in which the germs are developed in the interior of the animal, assuming, at the earliest period when they become animated, the form of the parent. In this case they are termed gemmules instead of buds. This mode of reproduction is exemplified in the Vol- vox, which, as we have already seen, is an infusorial animal- cule of a spherical form, exhibiting incessant revolving move- ments.* The germs of this animal are developed, in great numbers, in its interior, having a globular shape, and visible^ • Vol. i. p. 159. This animal is delineated in Fig. 79. REPRODUCTION. 415 by the aid of the microscope, through the transparent co- vering; and while yet retained within the hody of the pa- rent, other still minuter glohulcs are developed within these, constituting a third generation of these animals. After a certain period, the young, which have thus been formed, escape by the bursting of the parent volvox, which, in con- sequence, perishes. Similar phenomena are presented by many of the Infusoria. In some of the Entozoa, likewise, as in the Hydatid, the young are developed within the pa- rent; and this proceeds successively for an indefinite number of generations.* In most cases of the spontaneous evolution * The mode in which infiisory animalcules are produced and multiplied is involved in much obscurity. Many distinguished naturalists, adopting- the views of Buffon, have regarded them as the product of an inherent power belonging- to a certain class of material particles, which, in circumstances fa- vourable to its operation, tends to form these minute organizations, and in this manner they explain how the same organic matter which had composed I former living aggregates, on the dissolution of their union, reappears under new forms of life, and gives rise to the phenomenon of innumerable animal- cules, starting into being, and commencing a new, but fleeting career of ex- istence. Yet the analogy of every other department of the animal and ve- getable kingdoms is directly opposed to the supposition that any living being can arise without its having been originally derived from an individual of the same species as itself, and of which it once formed a part. The difficulty which the hypothesis of the spontaneous production of infusory animalcules professes to remove, consists in our inability to trace the pre-existence of the germs in the fluid, where these animalcules are found to arise; and to follow the operations of nature in these regions of infinite minuteness. Tiie disco- veries of Ehrenberg relative to the organization of the Rotifera go far to- wards placing these diminutive beings more on a level, both in structure and in functions, with the larger animals, of whose history and economy we have a more familiar and certain knowledge, and in superseding the hypothesis above referred to, by showing that the bold assumption on which it rests, is not required for the explanation of the obsen'cd phenomena. In many of these animalcules, he has seen the ova excluded in the form of extremely minute globules, the 12,000th of an inch in diameter. AVhcn tlicse had gi'own to the size of the 1700th of an inch, or seven times their original dia- meter, they were distinctly seen to excite cuirents, and to swallow food. The same diligent observer detected the young of the Rotifer vulgaris, per- fectly formed, moving in the interior of the parent animalcule, and excluded in a living state, thus constituting them viviparous animals, as the fonncr were 416 THE REPRODUCTIVE FUNCTIONS. of gemmules within the parent, channels are provided for their exit: but the gemmules of the Actinia force their way through the sides of the body, which readily open to give them passage; after which, the lacerated part soon heals. In the instances which have now passed under our review, the progeny is, at first, in direct communication with its pa- rent, and docs not receive the special protection of mem- branous envelopes, containing a store of nourishment for its subsequent growth. But in all the more perfect structures, both of animals and vegetables, the germ is provided with auxiliary coverings of this kind, the whole together com- posing what is called a seed, or an ovum: the former term being usually applied to vegetable, and the latter to animal productions; and, in both cases, the organ which originally contained them is termed the ovary. The formation and evolution of vegetable seeds take place, not indiscriminately, at every point, as we have seen is the case with simple germs, but only in particular parts of the plant. The Filices, or fern tribes, may be taken as exam- ples of this mode of reproduction, the seeds being formed at the under surface of the leaves, apparently by a simple pro- cess of evolution; and when detached and scattered on the ground, being farther developed into a plant similar to the parent. The Linnean class of CryptogaJiiia includes all the plants coming under this description. In Animals, likewise, it is only in the particular organs termed ovaries, that ova are formed, and they are generally divided into compart- ments, the whole being enclosed in a membranous covering, bearing a great resemblance to the seed-capsules of plants. The propagation of living beings by means of ova or seeds, is a process of a totally different class from their multipli- cation by mere slips or buds; and the products of the former oviparous. Other species, ag'ain, imitate the hydra, in being what is termed gemmiparousy tliat is, producing- gemmules (like the budding of a plant,) which shoot forth from the side of the parent, and are soon provided with cilia, enabling them, when separated, to provide for their own subsistence, although they are of a very diminutive size when thus cast off. REPRODUCTION. 417- retain less of the peculiar characters of the individual from which they spring, than those of the latter. This is re- markably exemplified in the case of orchard trees, such as apples and pears; for all the trees which derive their ori«rin from shoots, or grafts from the same individual, j)artake of the same properties, and prochice a fruit of the same flavour and qualities; whereas, trees of the same sj)ccies, which e Dipluzoon paradoxum, from its having- the semblance of two distinct animals of u lengthened shape, each bent at an obtuse angle, aiul joined together in the form of the letter X. The right and left halves of this cross are perfectly similar in their organization, leaving each a complete and independent system of vital organs, excepting that the two alimentary canals join at the centre of the cross to form a single •cavity, or stomacli, (Aimales des Sciences NatureHes, xxx. o7o.) ORGANIC DEVELOPMENT. 427 by the obliteration of the parts which are in contact, into single tubes, throughout a considerable portion of their length.* Nature, ever vigilant in her anticipations of the wants of the system, has accumulated round the embryo amj)le stores of nutritive matter, sufficient for maintaining the life of tlic chick, and for the building of its frame, while it continues in the egg, and is, consequently, unable to obtain supplies from without: yet, with the same foresight of future circum- stances, she delays not, longer than is necessary for the complete establishment of the circulation, to construct the apparatus for digestion, on which the animal is to rely for the means of support in after life. The alimentary canal, of which no trace exists at an earlier period, is constructed by the formation of two laminae, arising from folds of the in- nermost of the pellicles which invest the embryo; that is, on the surface opposite to the one which has produced the spi- nal marrow. These laminae, which are originally separate, and apart from one another, are brought together, and by the junction or soldering of their opposite edges, formed into a tube,t which, from being, at first, uniform in diameter, af- terwards expands into several dilated portions, correspond- ing with the cavities of the stomach, crop, gizzard, &c., into which they are to be converted, when the time shall come for their active employment. These new organs are, however, even in this, their rudimental state, trained to the perform- ance of their proper offices, receiving into their cavities, through a tube temporarily provided for that purpose, the fluid of the yelk, and preparing nourishment from it. In the mean time, early provision is mado for the aera- tion of the fluids by an extensive but tcm])orary system of * These facts were fii-st observed by Scn-cs (Annates dcs So. Nut. xxi. 8,) and their accuracy has been confirmed by the observations ofDr. Allen 'I'honi. son. In Reptiles this union of the tw( onstituent trunks of the aorta is ef- fected only at the posterior part, wh' the anterior portion rcmuiijs perma- nently double. (Sec Fig. '357, vol. li. p. 197.) f Wolf! is the author of tliis discovery. 428 THE REPRODUCTIVE FUNCTIONS. vessels, spread over the membrane of the egg, and receiving tlie influence of atmospheric oxygen tlirough the substance of tlie shell, which is sufllcicntly porous to transmit it; and these vessels, being brought into communication with the circulatory system of the chick, convey to its blood this vi- vifying agent. As the lungs cannot come into use till after the bird is emancipated from its prison, and as it was suffi- cient that they^ be in readiness at that cj^och, these organs are amonji the last which are constructed: and as the me- chanism of respiration in this class of animals does not re- quire the play of the diaphragm, this muscular partition, though begun, is not completed, and there is no separation between the cavities of the thorax and the abdomen. The succession of organic metamorphoses is ecjually re- markable in the formation of the diversified apparatus for aeration, which is required to be greatly modified, at differ- ent periods, in order to adapt it to different elements: of this we have already seen examples in those insects which, after being aquatic in their larva state, emerge from the water when they have acquired wings; and also in the steps of transition from the tadpole to the frog. ]3ut similar, though less conspicuous changes occur in the higher vertebrated animals, during the early periods of their formation, corre- sponding to the differences in the modes of aeration em- ployed at different stages of development. In the primeval conditions this function is always analogous to that of aqua- tic animals, and requires for its performance only the sim- pler form of heart already described, consisting of a single set of cavities: but the system being ultimately designed to exercise atmospheric respiration, requires to be gi'adually adapted to this altered condition; and the heart of the Bird and the Quadruped must be separated into two compart- ments, corresponding to the double function it will have to perform. For this purpose a partition wall must, be built in its cavity; and this wall is accordingly begun around the interior circumference of the ventricle, and is gradually car- ried on towards the centre, there being, for a time, an aper- ORGANIC DEVELOPMENT. 429 ture of communication between the right and left cavities; but this aperture is soon closed, and the ventricle is now effectually divided into two. Next the auricle, which at first was single, becomes double; not, however, by the growth of a partition, but by the folding in of its sides, along a middle line, as if it were encompassed by a cord, which was gradu- ally tightened. In the mean while the partition, which had divided the ventricle, extends itself into the trunk of the main artery, which it divides into two channels; and these afterwards become two separate vessels; that which issues from the left ventricle being the aorta; and the other, which proceeds from the right ventricle, being the pulmonary ar- tery; and each being now prepared to exercise its appropri- ate function in the double circulation which is soon to be established.* A mode of subdivision of blood vessels, very similar to that just described, takes place in those which are sent to the first set of organs provided for aeration, and which re- semble branchiae. These changes may be very distinctly followed in the Batrachia;\ for we see, in those animals, the trunk of the aorta undergoing successive subdivisions, by branches sent off from it, and forming loops, which ex- tend in length and are again subdivided, in a manner not unlike the unravelling of the strands of a rope; each subdi- vision, however, being preceded by the formation of a dou- ble partition in the cavity of the tube; so that at length the whole forms an extensive ramified system of branchial arte- ries and veins. Still all these are merely temporary struc- tures; for when the period of change approaches, and the branchiae are to be superseded in their office, every vessel, one after another, becomes obliterated, and there remain only the two original aorta, which unite into a single trunk lower down, and from which proceed the pulmonary arte- ries, conveying either the whole, or a portion of the blood, to the newly developed respiratory organs, the lungs. * The principal authorities for the facts here stated are Bacr and Rolando. See the paper of Dr. Thomson already quoted. t See the investigations of Kusconi, and of Bacr, on this subject. 430 THE KEPRODUCTIVE FUNCTIONS. By a similar process of continued bifurcation, or the de- tachment of branches in the form of loops, new vessels are developed in other parts of the body, as has been particu- larly observed in the finny tail, and the external gills of the frog, and the newt, parts which easily admit of microscopi- cal examination.* Progress is in the mean while making in the building of the skeleton, the forms of the principal bones being modelled in a gelatinous substance, which is converted into cartilage, beginning at the surface, and gradually advancing towards ' the centre of each portion or element of the future bone; and thus a temporary solid and elastic scaffolding is raised, suit- ed to the yielding texture of the nascent organs: lastly, the whole fabric is surrounded by an outer wall, the building of which is begun from the dorsal region, and conducted round the sides of the body, till the two portions come to meet in the middle abdominal line, where they are finally united into one general and continuous integument. The eyes, which were hitherto unprotected, receive special means of defence, by the addition of eyelids, which are formed by a farther extension and folding of these integuments; and the greater part of the surface of the body gives rise to a growth of temporary down, which, as we have seen, is provided as a covering to the bird at the time it is ready to quit the shell. But this hard shell, which had hitherto afforded it protection, is now opposed to its emancipation; and the chick, in order to obtain its freedom, must, by main force, break through the walls of its prison; its beak is, however, as yet too tender to apply the force requisite for that pur- pose. Here, again, we find Nature expressly interposing her assistance; for she has caused a pointed horny projection to grow at the end of the beak, for the special object of giving the chick the power of battering its shell, and making a practicable breach, through which it shall be able to creep out, and begin its new career of life. That this horn is pro- • Sucli is the result of the concurring observations of Spallanzani, Fonta- na, and DoUinger. ORGANIC DEVELOPMENT. 431 vided only for this temporary use appears from the circum- stance of its falling off spontaneously in the course of three or four days after it has been so employed. But though tlic bird has now gained its liberty, it is still unable to provide for its own maintenance, and requires to be fed by its parent till it can use its wings, and has learned the art of obtaining food. The pigeon is furnislied by na- ture with a secretion from the crop, with which it feeds its young. In the Mammalia the same object is provided for still more expressly, by means of glands, whose oflice it is to prepare milk, a fluid which, from its chemical qualities, is admirably adapted to the powers of the digestive organs, when they first exercise their functions. The Cetacea have also mammary glands; bwt as the structure of the mouth and throat of the young in that class does not appear adapted to the act of sucking, there has always been great difliculty in understanding how they obtain the nourishment so pro- vided. A recent discovery of Geoffroy St. Ililaire appears to have resolved the mystery with respect to ihc De/p/u?ius globiceps; for he found that the mammary glands of that ani- mal contain each a large reservoir, in which milk is accu- mulated, and which the dolphin is capable, by the action of the surrounding muscles, of emptying at once into the mouth of its young, without requiring from the latter any effort of suction.* The rapid sketch which I have attempted to draw of the more remarkable steps of the early stages of organic deve- lopment in the higher animals, taken in conjunction with the facts already adverted to in various parts of this Treatise, and particularly those relating to ossification, dentition, the formation of hair, of the quills of the porcupine, of the an- tlers of the stag, and of the feathers of birds, will suffice to show that they are regulated by laws which arc definite, and preordained according to the most enlarged and profound • The account of this discovery is contained in a memoir wliich was read at the "Institut." March 24, 1834. 432 THE REPRODUCTIVE FUNCTIONS. views of the future circumstances and wants of the system. The double origin of all the parts of the frame, even those which appear as single organs, and the order of their forma- tion, wliich, in each system, commences with the parts most remote from the centre, and proceeds inwards, or towards the mesial plane, are among the most singular and unex- pected results of this train of inquiries.* We cannot but be forcibly struck with the numerous forms of transition through which every organ has to pass before arriving at its ultimate and comparatively permanent condition: we cannot but won- der at the vast apparatus which is provided and put in action for eflecting all these changes; nor can we overlook the in- stances of express contrivance in the formation of so many temporary structures, which are set up, like the scaffold of an edifice, in order to afford the means of transporting the materials of the building in proportion as they are wanted; nor refuse to recognise the evidence of provident design in the regular order in which the w^ork proceeds, every organ growing at its appointed time, by the addition of fresh par- ticles brought to it by the arteries, while others are carried away by the absorbents, and gradually acquiring the form which is to qualify it for the performance of its proper ofTice in this vast and complicated system. • The first of these two laws is termed by Serres, who has zealously pro- secuted these investigations, " la hi dc symmHrit;'" and the second, "^f hi de conjugaison.*' He maintains that they are strictly applicable to all the parts of the body having- a tubular form, such as the trachea, the Eustachian tube, the canals, and perforations of bones, &,c. See the preliminary dis- course to his *' Anatomie comparee du cerveau," p. 25; and also his several memoirs in the " Annales des Sciences Naturelles," vols. xi. xii. xvi. and xxu An excellent summary of the principal facts relating to the development of the embryo is given by Mr. Herbert Mayo, in the third edition of his ^* Outlines of Human Physiology.'* ( 433 ) CHAPTER III. DECLINE OF THE SYSTEM. To follow minutely the various steps by which Nature conducts the individual to its state of maturity, would en- gage us in details incompatible with the limits of the pre- sent work. I shall only remark, in general, that during the period when the body is intended to increase in size, the powers of assimilation are exerted to prepare a greater abun- dance of nourishment, so that the average supply of mate- rials rather exceeds the consumption: but when tlie fabric has attained its prescribed dimensions, the total quantities furnished and expended being nearly balanced, the vital powers are no longer exerted in extending the fabric, but are employed in consolidating and perfecting it, and in qua- lifying the organs for the continued exercise of their re- spective functions, during a long succession of years. Yet, while every function is thus maintained in a state of healthy equilibrium, certain changes are in progress, which, at the appointed season, will inevitably bring on the decline and ultimate destruction of the system.* The process of * It would appear, from the researches of De CandoUe, that the vegetab?e system is not, like the animal, subject to the destructive operation of internal causes; for the agents which destroy vegetable life are always extitmeous to its economy. Each individual tree is composed of an accumulation of the shoots of every successive year since the commencement of its growth; and although, from the continued deposition of lignin, and the consequent obli- teration of many of its cells and vessels, the vitaUty of the interior wood may be destroyed, and it then becomes liable to decay by the action of foreign agents, yet the exterior layers of the liber still vegetate with umiiininishcd vigour; and unless injured by causes extraneous to its own system, the life of the tree will continue to be sustained for an indefinite period. If, on the Vol. II. 55 434 DECLINE OF THE SYSTEM. consolidation, begun from the earliest period of development, is still advancing, and is producing in the fluids greater thick- ness, and a reduction of their total (juantity; and in the so- lids, a diminution in the proportion of gelatin, and the con- version of this element into albumen. Hence, all the tex- tures acquire increasing solidity, the cellular substance becomes firmer and more condensed, and the solid structures more rigid and inelastic: hence, the tendons and ligamentous fibres growing less flexible, the joints lose their suppleness, and the contractile power being also impaired, the muscles act more tardily as well as more feebly, and the limbs no longer retain the elastic spring of youth. The bones them- selves grow harder and more brittle; and the cartilages, the tendons, the serous membranes, and the coats of the blood vessels, acquire incrustations of ossilic matter, which inter- fere with their uses. Thus are all the progressive modifica- tions of structure tending, slowly but inevitably, to disqua- lify the organs for the due performance of their functions. Among the most important of the internal changes con- sequent on the progress of age are those which take place in the vascular system. A large proportion of the numerous arteries, which were in full activity during the building of the fabric, being now no longer wanted, are thrown, as it were, out of employment; they, in consequence, contract, and becoming impervious, gra(hially disappear. The parts of the body, no longer yielding to the power applied to ex- tend them, oppose a gradually increasing resistance to the propelling force of the heart: w^hile, at the same time, this force, in common with all the others, is slowly diminishing. Thus do the vital powers become less equal to the demands made upon them; the waste of the body exceeds the supply, other hand, we were to regard each separate shoot as an individual organic body, and every layer us constituting a distinct generation of shoots, the older being covered and enclosed in succession by the younger, the great longevity of a tree would, on this hypothesis, indicate only the permanence of the species, not the indefinitely protracted duration of the individual plant. DECLINE OF THE SYSTEM. 435 and a cllmhiution of energy becomes apparent in every func- tion. Such are the insensible gradations by whicli, while gliding down the stream of time, we lapse into old age, which in- sidiously steals on us before we are aware of its approach. But the same provident power which presided at our birth, which superintended the growth of all the organs, which infused animation into each as they arose, and which has conducted the system unimpaired to its maturity, is still ex- erted in adjusting the conditions under which it is placed in its season of decline. New arrangements are made, new energies are called forth, and new resources arc em})loyed, to accommodate it to its altered circumstances, to pro}) the shattered fabric, and retard tiie progress of its decay. In proportion as the supply of nutritive materials has become less abundant, a more strict economy is practised with re- gard to their disposal; the substance of the body is husband- ed with greater care; the absorbent vessels are employed to remove such parts as are no longer useful; and when all these adjustments have been made, the functions still go on for a considerable len2;th of time without material altera- tion. The period prescribed for its duration being at length completed, and the ends of its existence accomplished, the fabric can no longer be sustained, and preparation must be made for its inevitable fall. In order to form a correct judgment of the real intentions of nature, with regard to this last stage of life, its phenomena must be observed in cases where the systeni has been wholly intrusted to the operation of her laws. When death is the simple consequence of age, we find that the extinction of the powers of life observes an order the reverse of that which was followed in their evolu- tion. The sensorial functions, wliicli were llu; last perfect- ed, are the first which decav: and their decline is found to commence with those mental faculties more inimediately dependent on tiic j)hysical conditions of tlu^ sensorium, and more especially with tlie memory, which is often much im- 436 DECLINE OF THE SYSTEM. paired, while the judgment remains in full vigour. The next faculties which usually suffer from the effects of age are the external senses, and the failure of sight and of hearing still farther contrihutes to the decline of the intellectual powers, hy withdrawing the occasions for their exercise. The actual demolition of the fabric commences whenever there is a considerable failure in the functions of assimilation: but the more immediate cause of the rapid extinction of life is usually the impediment which the loss of the sensorial power, necessary for maintaining the movements of the chest, creates to resj)iration. The heart, whose pulsations gave the first indications of life in the embryo, generally retains its vitality longer than any other organ; but its powers being dependent on the constant oxidation of the blood in the lungs, cannot survive the interruption of this function; and on the heart ceasing to throb, death may then be considered as complete in every part of the system. It is an important consideration, with reference to final causes, that generally long before the commencement of this *'Last scene of all, That ends this sti-ange eventful history," the power of feeling has wholly ceased, and the physical struggle is carried on by the vital powers alone, in the ab- sence of all consciousness of the sentient being, whose death may be said to precede, for some time, that of the body. In this, as well as in the gradual decline of the sensorial facul- ties, and the consequent diminution both of mental and of physical sensibility in advanced age, we cannot fail to re- cognise the wise ordinances of a superintending and bene- ficent providence, kindly smoothing the path along which we descend the vale of life, spreading a narcotic mantle over the bed of death, and giving to the last moments of de- parting sensation the tranquillity of approaching sleep. ( 437 ) CHAPTER IV. UNITY OF DESIGN. The inquiries on Animal and Vegetable Physiology in which we have been engaged, lead to the general conclusion that unity of design and identity of operation pervade the whole of nature; and they clearly point to one Great and only Cause of all things, arrayed in the attributes of infinite power, wisdom, and benevolence, whose mighty works ex- tend throughout the boundless regions of space, and whose comprehensive plans embrace eternity. In examining the manifold structures and diversified phe- nomena of living beings, we cannot but perceive that they are extensively, and perhaps universally connected by cer- tain laws oiJinalogy; a principle, the recognition of which has given us enlarged views of a multitude of important facts, which would otherwise have remained isolated and unintelligible. Hence naturalists, in arranging the objects of their study, according to their similarities and analogies, into classes, orders and genera, have but followed the foot- steps of Nature herself, who in all her operations combines the apparently opposite principles of general resemblance, and of specific variety; so that the races which she has united in the same group, though possessed of features in- dividually dilTerent, may easily be recognised bv their fa- mily likeness, as the offspring of a common parent. *• Fades non omnibus una; Nee (liversa tamcn; c[ualem decet esse sororuin." We have seen that in each of the two great divisions, or 438 UNITY OF DESIGN. klno-donis of organic nature, the same general objects are aimed at, and the same general plans are devised for their accomplishment; and, also, that in the execution of those plans similar means and agencies are employed. In each division there prevails a remarkable uniformity in the com- position and properties of their elementary textures, in the nature of their vital powers, in the arrangement of their or- gans, and in the laws of their production and development. The same principle of analogy may be traced, amidst endless modifications of detail, in all the subordinate groups into which each kingdom admits of being subdivided, both in re- spect to the organization and functions of the objects com- prehended in each assemblage, whether we examine the wonders of their mechanical fabric, or study the series of processes by which nutrition, sensation, voluntary motion, and reproduction are effected. To specify all the examples which misiht be adduced in confirmation of this obvious truth is here unnecessary; for it would be only to repeat the numerous facts already noticed in every cha])ter of this trea- tise, relative to eacli natural group of living beings: and it was, indeed, chiefly by the aid of such analogies, that we were enabled to connect and generalize those facts. We have seen that, in constructing each of the divisions so esta- blished, Nature appears to have kept in view a certain defi- nite type, or ideal standard, to which, amidst innumerable modifications, rendered necessary by the varying circum- stances and different destinations of each species, she always shows a decided tendency to conform. It would almost seem as if, in laying the foundations of each organized fab- ric, she had commenced by taking the exact copy of this primitive model; and, in building the superstructure, had allowed herself to depart from the original plan only for the purpose of accommodation to certain specific and ulterior objects, conformably with the destination of that particular race of created beings. Such, indeed, is the hypothetical principle, which, under the title of iLiiity of composition, has been adopted, and zealously pursued in all its conse- UNITY OF DESIGN. 439 quences, by many naturalists, of the liighcst eminence, on the continent. As the facts on wliicli tliis hypothesis is supported, and the views wliich it unfolds, are highly de- serving of attention, I sliall here briefly state them; but in so doing I shall beg to premise the caution that these views should, for the present, be regarded as hyj)othctical, and as by no means possessing the certainty of philosophical ge- neralizations. The hypothesis in question is countenanced, in the first place, by the supposed constancy with which, in all the ani- mals belonging to the same natural group, we meet with the same constituent elements of structure, in each respective system of organs, notwithstanding the utmost diversity which may exist in the forms of the organs, and in the uses to which they are applied. This jjrinciple has been most strikingly exemplified in the osteology of vertcbrated ani- mals; but its truth is also inferred from tlie examination of the mechanical fabric of Insects, Crustacea, and Arachnida; and it appears to extend also to the structures subservient to other functions, and particularly those of the nervous sys- tem. Thus Nature has provided for the locomotion of the serpent, not by the creation of new structures, foreign to the type of the vertebiata, but by empIo3'ing the ribs in this new office; and in giving wings to a lizard, she has extended these same bones to serve as supports to the superadded parts. In arming the elephant .with tusks, she has merely caused two of the teeth in the upper jaw to be develojied into these formidable weapons; and in providing it with an instrument of prehension, has only resorted to a greater elongation of the snout. The law of Gradation^ in conformity to which all the living, together with the extinct races, of organic nature, ar- range themselves more or less, into certain regular series, is one of the consequences which have been deduced from the hypothesis we are considering. Every fresh copy taken of the original type is supposed to receive some additional extension of its Jii\culties and endowments by the graduated 440 UNITY OF DESIGN. development of elements, which existed in a latent form in the primeval germ, and which are evolved, in succession, as nature advances in her course. Thus, we find that each new form which arises, in following the ascending scale of creation, retains a strong affinity to that which had preceded it, and also tends to impress its own features on those which immediately succeed; and thus their specific differences re- sult merely from the different extent and direction given to these organic developments; those of inferior races proceed- ing to a certain point only, and there stopping: while in be- ings of a higher rank they advance farther, and lead to all the observed diversities of conformation and endowments. It is remarked, in farther corroboration of these views, that the animals which occupy the highest stations in each series possess, at the commencement of their existence, forms exhibiting a marked resemblance to those presented in the permanent condition of the lowest animals in the same se- ries; and that, during the progress of their development, they assume, in succession, the characters of each tribe, cor- responding to their consecutive order in the ascending chain: so that the peculiarities which distinguish the higher ani- mal, on its attaining its ultimate and permanent form, arc those which it had received in its last stage of embryonic evolution. Another consequence of this hypothesis is that we may expect occasionally to meet, in inferior animals, with rudimental organs, which fi'om their imperfect development may be of little or no use to the individual, but which be- come available to some superior species, in which they are sufficiently perfected. The following are among the most remarkable facts in illustration of these propositions. In the series of Articulated Animals, of which the An- nelida constitute the lowest, and winged Insects the highest terms, we find that the larvae of the latter are often scarcely distinguishable, either in outward form, or in internal or- ganization, from Vermes of the lower orders; both being equally destitute of, or but imperfectly provided with ex- ternal instruments of locomotion; both having a distinct vas- UNITY OF DESIGN. 441 ciilar circulation, and mulllplc organs of digestion; and the central filaments of the nervous system in both being stud- ded with numerous pairs of equidistant ganglia. In tlie worm all these features remain as permanent characters of the order: in the insect they arc subsequently modified and altered during its progressive metamorphoses. The em- bryo of a crab resembles in appearance the permanent forms of the Myriapoda, and of the lower animals of its own class, but acquires, in the progress of its growth, new parts; while those already evolved become more and more concentrated, passing, in their progress, through all the forms of transition which characterize the intermediate tribes of Crustacea; till the animal attains its last state, and then exhibits the most developed condition of that particular type.* However different the conformations of the Fish, the Rep- tile, the Bird, and the warm-blooded quadruped, may be at the period of their maturity, they arc scarcely distinguisha- ble from one another in their embryonic state; and their ear- ly development proceeds for some time in the same man- ner. They all possess at first the characters of aquatic ani- mals; and the Frog even retains this form for a considerable period after it has left the egg. The young tadpole is in truth a fish, whether we regard the form and actions of its instruments of progressive motion, the arrangement of its organs of circulation and of respiration, or the condition of the central organs of its nervous system. We have seen by what gradual and curious transitions all tliese aquatic cha- racters are changed for those of a terrestrial quadruped, fur- nished with limbs for moving on the ground, and with lungs for breathing atmospheric air; and how the plan of circula- tion is altered from branchial to pulmonary, in proportion * This curious analogy is particularly obsciTabIc in the successive forms as- sumed by the nervous system, which exhibits a gradual passage from that of the Talitrus, to its ultimate greatest concentration in the Main. (See Fi- gures 439 and 441, p. 382 and 383.) Milne Edwards has lately traced a si- milar progression of development in the orgtuis of locomotion of the Crvista- cea. (Annales des Sciences Naturcllcs, xxx. 354.) Vol. II. 56 442 UNITY OF DESIGN. as the gills wither and the lungs are developed. If, while this change is going on, and while both sets of organs are too-other executing the function of aeration, all farther de- velopment were prevented, we should have an amphibious animal, fitted for maintaining life both in air and in water. It is curious that this precise condition is the permanent state of the Siren and the Pro/ew.v, animals which thus exemplify one of the forms of transition in the metamorphoses of the Frog. In the rudimental form of the feet of serpents, which are so imperfectly developed as to be concealed underneath the skin, and to be useless as organs of progressive motion, we have an example of the first stage of that process, which, when carried farther in the higher animals, gives rise to the limbs of quadrupeds, and which it would almost seem as if nature had instituted with a prospective view to these more im- proved constructions. Another, and a still more remarkable instance of the same kind, occurs in the rudimental teeth of the young of the Whale, which are concealed within the lower jaw, and which are afterwards removed, to give place to the curious filtering apparatus, which occupies the roof of the mouth, and which nature has substituted for that of teeth, as if new objects, superseding those at first pursued, had arisen in the progress of development. Birds, though destined to a very different sphere of ac- tion from either fishes or reptiles, are yet observed to pass, in the embryonic stage of their existence, through forms of transition, which successively resemble these inferior classes. The brain presents, in its earliest formation, a series of tu- bercles, placed longitudinally, like those of fishes, and only assuming its proper character at a later period. The res- piratory organs are at first branchiae, placed, like those of fishes, in the neck, where there are also found branchial apertures similar to those of the lamprey and the shark; and the heart and great vessels are constructed like those of the tadpole, with reference to a branchial circulation. In their conversion to the purposes of aerial respiration, they under- UNITY OF DESIGN. 443 go a series of changes precisely analogous to those of the tadpole. Mammalia, during the early periods of their development, are subjected to all the transformations which iiave been now described, commencing with an organization corrcsj)onding to that of the aquatic tribes, exhibiting not only branchiae, supported on branchial arches, but also branchial aj)crtures in the neck, and thence passing quickly to Ihe conditions of structure adapted to a terrestrial existence. The development of various parts of the system, more especially of the brain, the ear, the mouth, and the extremities, is carried still far- ther than in birds. Nor is the human cmbrj-o exempt from the same metamorphoses, possessing, at one period, branchiai and branchial apertures similar to those of the cartilaginous fishes,* a heart with a single set of cavities, and a brain con- sisting of a longitudinal series of tubercles; next losing its branchiae, and acquiring lungs, while the circulation is yet single, and thus imitating the condition of the reptile; then acquiring a double circulation, but an incomplete diaphragm, like birds; afterwards, appearing like a quadruped, with a caudal prolongation of the sacrum, and an intermaxillary bone; and, lastly, changing its structure to one adapted to the erect position, accompanied by a great expansion of the cerebral hemispheres, which extend backwards so as com- pletely to cover the cerebellum. Thus does the whole fab- ric arrive, by a gradual process of mutation, at an extent of elaboration and refinement, unattained by an}' other race of terrestrial beings, and which has been justly regarded as constituting the climax of organic development.t * These facts are given on the authorities of JIathkc, Baer, Huschke, Breschet, &c. Ann. des Sc. Naturellcs, xv. 266. See, also, the paper of Dr. A. Thomson, ah-eady quoted. t A popular opinion has long prevailed, even among the well informed, that misshapen or monstrous productions, or lusus naturx, as they were termed, exhibit but the freaks of nature, who w;is believed, on these occa- sions, capriciously to abandon her usual course, and to amuse hei-self in the production of grotesque beings, without any special object. But it is now found that all defective formations of this kind arc occasioned by the impcr- 444 UNITY OF DESIGN. It must, I think, be admittetl that the analogies, on which the hypothesis in question is founded, are numerous and striking; but great care should be taken not to carry it far- ther than the just interpretation of the facts themselves may warrant. It should be borne in mind that these facts are few, compared with the entire history of animal development; and that the resemblances which have been so ingeniously traced, arc partial only, and fall very short of that universa- lity, which alone constitutes the solid basis of a strictly phi- losophical theory. Wliatever may be the apparent simila- rity between one animal and another, during different peri- ods of their respective developments, there still exists spe- cific differences, establishing between ihem an impassable barrier of separation, and elYactually preventing any conver- sion of one species into another, however nearly the two may be mutually allied. The essential characters of each species, amidst occasional varieties, remain ever constant and immutable. Although gradations, to a greater or less ex- tent, may be traced among the races both of plants and ani- mals, yet in no case is the series strictly continuous; each step, however short, being in reality an abrupt transition from one type of conformation to another. In many in- stances the interval is considerable; as, for example, in the passage from the invertebrate to the vertebrated classes; and, indeed, in every instance where great changes in the nature and arrangement of the functions take place.* It is in vain to allege that the original continuity of the series is indi- cated by a few species presenting, in some respects, inter- mediate characters, such as the Ornitfwrhyncus, between feet development of some parts of the embryo, while the natural process is carried on in the rest of the system; and thus it happens that a resemblance may often be traced, in these malformations, with the type or the permanent condition of some inferior animal. Hence, all these apparent anomahcs are, in reality, in perfect harmony with the established laws of ort^anic develop- ment, and afford, indeed, striking confirmations of the truth of the theory here cxj^lained. • See a paper on tiiis subject, by Cuvier, in the Arm. dcs Sciences Na- turcUes, xx. 241. UNITY OF DESIGN. /J45 birds and mammalia, and the Cetacea, between fislies and warm-blooded quadrupeds: for these are but detached links of a broken chain, tending, indeed, to prove the unity of the designs of Nature, but showing also the specific character of each of her creative efforts. The pursuit of remote and often fanciful analogies has, by many of the continental phy- siologists, been carried to an unwarrantable and extravagant length: for the scope which is given to the imagination in these seductive speculations, by leading us far away from the path of philosophical induction, tends rather to obstruct than to advance the progress of real knowledge. I3y confining our inquiries to more legitimate objects, we shall avoid the delusion into which one of the disciples of this transcenden- tal school appears to have fallen, when he announces, with exultation, that the simple laws he has discovered have now explained the universe;* nor shall we be disposed to lend a patient ear to the more presumptuous reveries of another system-builder, who, by assuming that there exists in or- ganized matter an inherent tendency to perfectibility, fan- cies that he can supersede the operations of Divine agency. t Very different was the humble spirit of the great New- ton, who, struck with the immensity of nature, compared our knowledge of her operations, into which he had himself penetrated so deeply, to that of a child gathering pebbles on • *' L'univers est expllque, et nous le voyons; c'est un petit nombre de prlncipes generaux et feconds qui nous en ont donne la clef." Serrcs, Ann. des So. Nat. xi. 50. f Allusion is here made to the celebrated theory of Lamarck, as exposed in his "Pliilosophie Zoolog-ique. " He conceives that there was orig-inally no distinction of species, but that each race has, in the coui-se of ages, been derived from some other, less perfect than itself, by a spontaneous effort at improvement; and he supposes that infusorial animalcules, spontaneously formed out of organic molecules, gave birth, by successive ti-ansformations, to all other animals now existing on the globe. He believes that tribes, ori- ginally aquatic, acquired by their own efforts, prompted by their desire to walk, both feet and legs, fitting them for progression on tlie ground; and that these members, by the long contiinicd operation of the wish to flv, were transformed into wings, adapted to gratify that desire. If this be philoso- phy, it is such as might have emanated from the college of Laputa. 446 UNITY OF DESIGN. the sea-shore. Compared, indeed, with the magnitude of the universe, how narrow is the field of our perceptions, and how far distant from any approximation to a knowledge of the essence of matter, of the source of its powers, or even of the ultimate configurations of its parts! How remote from all human cognizance are the intimate properties of those imponderable agents, Light, Heat, and Electricity, which pervade space, and exercise so potent a control over all the bodies in nature! Doubtless, there exist around us, on every side, influences of a still more subtle kind, which " eye hath not seen, nor ear heard," neither can it enter into the heart or imagination of man to conceive. How scanty is our knowledge of the mind; how incomprehensible is its connexion with the body; how mysterious arc its secret springs, and inmost workings! What ineffable wonders would burst upon us, were we admitted to the perception of the spiritual world, now encompassed by clouds impervious to mortal vision! The Great Author of our being, who. while he has been pleased to confer on us the gift of reason, has prescribed cer- tain limits to its powers, permits us to acquire, by its exer- cise, a knowledge of some of the wondrous works of his creation, to interpret the characters of wisdom and of good- ness with which they are impressed, and to join our voice to the general chorus which proclaims " His Might, Ma- jesty, and Dominion." From the same gracious hand we also derive that unquenchable thirst for knowledge, which this fleeting life must ever leave unsatisfied; those endow- ments of the moral sense, with which the present constitu- tion of the world so ill accords; and that innate desire of per- fection which our present frail condition is so inadequate to fulfil. But it is not given to man to penetrate into the coun- sels, or fathom the designs of Omnipotence; for in directing his views into futurity, the feeble light of his reason is scat- tered and lost in the vast abyss. Although we plainly dis- cern intention in every part of the creation, the grand ob- ject of the whole is placed far above the scope of our com- UNITY OF DESIGN. 447 prehension. It is impossible, however, to conceive that this enormous expenditure of power, this vast accumulation of contrivances and of machinery, and this profusion of ex- istence resulting from them, can thus, from age to age, be prodigally lavished, without some ulterior end. Is Man, the favoured creature of nature's bounty, " The paragon of animals,'' whose spirit holds communion with celestial powers, formed but to perish with the wreck of his bodily frame? Are generations after generations of his race doomed to follow in endless succession, rolling darkly down the stream of time, and leaving no track in its pathless ocean? Are the operations of Almighty power to end with the pre- sent scene? May we not discern, in the spiritual constitu- tion of man, the traces of higher powers, to which those he now possesses are but preparatory; some embryo faculties which raise us above this earthly habitation? Have we not in the imagination, a power but little in harmony with the fetters of our bodily organs; and bringing within our view purer conditions of being, exempt from the illusions of our senses and the infirmities of our nature, our elevation to which will eventually prove that all these unsated desires of knowledge, and all these ardent aspirations after moral good, were not implanted in us in vain? Happily there has been vouchsafed to us, from a higher source, a pure and heavenly light to guide our faltering steps, and animate our fainting spirit, in this dark and dreary search: revealing those truths which it imports us most of all to know, giving to morality higher sanctions, elevating our hopes and our affections to nobler objects than belong to earth, and inspiring more exalted themes of thanksgiving and of praise. f*. 1 IV D E X. Abdojien of insects, i. 230. Aberration, chromatic, ii. 385. Aberration of parallax, ii. 325, SM. Aberration, spherical, ii. 324, 333. Absorption, vegetable, ii. 21, 23. Absorption, animal, ii. 17, 250. Absorption, lacteal, ii. 104, Absorption of shell, i. 174. Acalepha, i. 142 ; ii. 210. Acarus, i. 212. Achatina zebra, i. 175. Achromatic power, ii. 335. Acephala, i. 159 ; ii. 88, 215. Acetabulum, i. 282. Acid secretions, ii. 39. Ac rid a, ii. 155. Acridium, i. 236. Acoustic principles, ii. 294. Actinia, i. 136, 146 ; ii. 75, 272, 337, 412, 415, Adipose substance, i. 97. Adductor muscle, i. 160. Aeration of sap, ii. 28. Aeration, animal, ii. 31, 428. iEschna, i. 247. Affinities, organic, ii. 13. Agastric medusER, ii. 70. Age of trees, i. 73. Age, effects of, ii. 434. Agouti, i. 344. Agrion, ii. 174. Air-bladder, i. 298. Air cells of plants, i. 67. Air cells of birds, ii. 234. Air, rarefaction of, in birds, i. 384. Air tubes in plants, i. 05. Albumen, i. 85. Alburnum, i. 73; ii. 36. Algse, ii. 21. Alimentary canal, ii. 82, Alimentary canal, Jbrmation of, ii. 427. Vol. II. Alitrunk, i. 243-. Alligator, i. 317, 319 ; ii. 290. Amble, i. 342. Ambulacra, i. 148. Amici, i. 68; ii, 42, Amphibia, i. 303,336. Amphisbaena, i. 310, 311. Amphitrite, i. 201. Anabas, ii. 219. Analogy, law of, i. 49; ii. 437, Anarrhichas, ii. 96. Anchylosis, i. 267. Ancillaria, i, 175, Anemone, sea, i. 146. Angler, i. 293; ii. 276. Anguis, i, 310, 315, Animal functions, i. 42. Animal organization, i. 79. Animalcules. See Infusoria. Annelida, i. 194; ii. ISO, 213, 272^ 338. Annular vessels, i. 66. Anodon, i. 169. Ant, ii. 274, 341, 343. Ant-eater, i. 361; ii, 10(). Antelope, ii, 109, 285, Antelope, horn of, i, 3.5.5. Antenna?, i, 206; ii. 273, Antennulffi, ii. 93. Anther, ii. 418, Anthias, ii. 219. Anthophora, i, 247. Ant i pathos, i. 125, Antler of deer, i. 351. Antrum ma.xillare, ii. 284. Aorta, ii. 83, 426, Aphrodite, ii, 77, 91, 213, Aplysia, ii. 95, 124, 3^-*. Apodcs, i. 294, Apterous insects, i. 212, Aquatic animals, i. 11^^ Aquatic plants, ii. 41. 5/ 450 INDEX. Aquatic larvce, i. 2*20. Aijualic insects!, i, 2'S7. Aquatic birJs, i. 407. Aquatic respiration, ii. 210. Aqueous humour, ii. ^^27. Arachnida, i. 202; ii. 91, 235, 270, 34:3, 412. Aranea. See Spider. Arbor vitaj, ii. 393. Arcnicola, i. 199; ii. 211. Ar^ouauta, i. 165. Aristotle, ii. 394. Aristotle, lantern of, ii. 90. Ann, human, i. 375. Armadillo, ii. 271. Arteries, i. 44; ii. 82. Articulata, i. 193. Ascaris, ii. 86, 380. Ascidia, i. 106; ii. 212. Ass, i. 356. Assimilation, i. 43; ii. 16. Astacus, ii. 308, 346. Asterias, i. 147; ii. 76, 151, 171, 212, 272, 387, 412. Ateles, i. 278, 368. Atlas of Lion, i, 365. Atmosphere, purification of, ii. 32. Atmospheric respiration, ii. 221. Atriplex, ii. 40. AuJouin, i. 207, 228, 230; ii. 177, 226, 381. Audubon, ii. 288. Auricle, ii. 82, 187. Auricula, i. 1*^1. Avicula, i. 171. Axilla) of plants, i. 76; ii. 413. AxelotI, ii. 231. Babiroussa, ii. 105. Bacculite, i. 192. Baer, ii. 338, 429, 443. Bdker, ii. 337. Balaina, See Whale. Balance of affinities, ii. 13. Balistes, i. 300. Banks, i. 314. Barbels offish, ii. 276. Bark, formation of, i. 73. Barnacle, i. 185 ; ii. 212. Bat, i. 380;ii. 101,399. Batrachia, i. 303; ii. 418. Batracho.-permum, ii. 41. Bauer, i. 59. Bear, ii. 108. Beard of oyster, ii. 215. Beaver, i. 361; ii. 110, 135, 141. Bee, i. 247; ii. 275. Belchier, i. 269. Bell (Sir C.) ii. 375. Bell (Thomas) i. 333 ; ii. 290. Bellini, ii. 279. Berberis, i. 100. Berkeley, ii. 366. Beroe, i. 144, 149. Berzelius, ii. 21. Bicuspid teeth, ii. 107. Bipes canal iculatus, i. 317. Birds, i. 382; ii. 97, 234, 287, ct passim. Blind-worm, i. 315, 317. Blood, ii. 237. Blood vessels, ii. 201. Blunienhach, ii. 302. Boa, i. 310, 311. JJoar, i. 53; ii. 105, 118. Bombyx, i. 21-5, 217, 222; ii. 343. Bone, i. 89, 256, 263. Bonnet, i. 51; ii. 20, 62, 70, 182, 337. Borelli, i. 405. Bosc, i. 115. Boslock, ii. 237. Bound of deer, i. 342. Bowerhank, ii. 174. Boyle, ii. 19. Bractea), i. 78. Bradypus, i. 333; ii. 204. Brain, i. 40; ii. 260, 390, 404. Brain, formation of, ii. 423. Branchiffi, ii. 192,210,214. Brassica, ii. 40, 44. Braula, ii. 341. Bresehet, ii. 303. Brewster, I 169; ii. 333, 349. Brocken, spectre of, ii. 374. Broussonnet, ii. 412. Bru^uiere, i. 115, 179. Bryophyllum, ii. 411. Buccinum, i. 158, 167, 175; ii- 95, 215. Buckland, ii. 149. Buds,i. 73; ii. 413. Buffon,'!. 137; ii. 372,415. Bulb of hair, i. 93. Bulboffeather, i. 397. Bulbus arteriosus, ii. 196. Bulbiilus glandulosus, ii. 135, Bulimus, I. 180. Bulla, ii. 123. Burrowing of the mole, i. 361. INDEX. 451 Cabbage, ii. 40, 44. Cachalot, i. 334 ; ii. 105. Caeca, ii. 78, 150. Csecilia, ii. 351. Calamary, i. 182. Callionymiis, ii. 354. Calosoma, i. 227. Cambium, ii. 3.5. Camel, i. 87; ii. 129, 144. Cameleopard, i. 332, 344; ii. 101. Camera obscura, ii. 324. Camerated shells, i. 100. Campanularin, ii. 170. Camper, ii. 310, 314, 394. Canada rat, ii. 130. Cancelli, i. 262. Cannon bone, i. 348. Capibara, ii. 118. Capillaries, ii. 190. Capsular ligaments, i. 86. Caput Medusae, i. 1-55. Carapace, i 207, 321. Carbon, non-absorption of, ii. 20. Carbonic acid, ii. 28, 239. Cardia, ii. 133. Card i urn, i. 102, 162, 163, 164. Carduus, i. 100. Carinated sternum, i. 390. Carlisle, i. 296, 301 ; ii. 204, 399. Carnivora, i. 364; ii. 52, 108. Carp, i. 286, 298. Carpus, i. 282. Cartilag-e, i. 88. Caruncle, lacrymal, ii. 331. Cams, i.257; ii. 151, 158, 174, 182, 356. Cassowary, i. 404; ii. 161. Cat, ii. 278, 355. Caterpillar, i. 217, 224; ii. 342. Caudal vertebrae, i. 281. Cavolini, i. 121. Celandine, ii. 41. Cells of plants, i. 60, 62. Cellular texture, animal, i. 81. Centaurea, i. 100. Cephalic franrjlion, ii. 381. Cephalo-thorax, i. 202. Cephalopoda, i. 186; ii. 159, 388. Cerambyx, i. 233; ii. 322, 323. Cercaria, i. 138; ii. 338. Cerebellum, ii. 391. Cerebral ganglion, ii. 381. Cerebral hemispheres, ii. 391. Cerithium, i. 180. Ceroxylon, ii. 40. Cetacea, i. 332, 333 ; ii. 105, 128, 140, 313, :m), 431. Chnbrier, i. 88,241. Chain of bcincr, i. 51 ; ii. 439. Chalcides, i. 311. 317. Chameleon, i. 320 ; ii. 97, 277, 351. Chara, ii. 42, 183. Chelidonium, ii. 41. Chelonia, i. \\Z\ ; ii. 97, 198, 229, 311. Chemistry, organic, ii. 12, 2Ji6. Chcseldcn, ii. 3()6. C/irvrruil, i. 97. Children, I 12,226; ii. 347. Chitinp, i. 226. Cliladni, ii. 296. Chondrilla, ii. 43. Choroid coat, ii, 326. Choroid gland, ii. 349. Chromatic aberration, ii. 33.5. Chronuile, i. 63. Chrysalis, i. 219. Chyle, ii. 82, 148. Chyme, ii. 132. Cicada, i. 240. Cicindela, ii. 154. Cilia, i. 99, 117, 119, 129, 144. 185. Ciliary ligament, ii. 327. Cimbex, i. 235. Cimex, ii. 93, Cineritious, ii. 39.5. Circulation, ii. 16, 167. Cirrhi, ii. 276. Cirrhopoda, i. 185. Classification, i. 50; ii. 437. Clausilia, ii. 226. Clausium, i. 183. Clavicle, i. 281, 293, 390. Claviger, ii. 341. Claw in lion's tail, i. 36(». Clio, i. 1%; ii. 103. Cinque t, ii. 3.51. Clypcaster, i. 155. Cobitis, ii. 221. Cobra do capcllo, i. 378; ii. 121. Coccygeal bone, i. 281. Cochlea, ii. 303. Cockchaffer. See Melolontha. Cockle, i. 162. See rardium. Cod, lens of, i. 56; ii. 394. Coenurus, ii. 65. Coexistence of forms, i. 49. Coffin-bone, i. IW). Coleoptcra, i. 245; ii. 271. 452 rNDEX. Collar-bone, i. 281. Colours of insects, i. 226. Colours, perceptions of, ii- 37^ Coluber, i.:311,:U2; ii. 120. Columella, i. 170; ii. 311. Comatula, i. 155. Commissures of brain, ii. 395. Coinparctli, ii. 177, 309. Complementary colours, ii. 373. Compound eyes, ii. 341. Concha of the ear, ii. 298. Condor, li. 235. Conyer eel, ii. 392. Conjjflomerate eyes, ii. 341. Conjunctiva, ii. 329. Consumption of animal matter, ii. 48. Contractility, muscular, i. 98. Conus, i. 181. Convolutions of the brain, ii. 393. Convolvulus, ii. 41. Cooper, ii. 308. Coracoid bone, i. 281, 390- Coral, i. 125. Coral islands, i. 26. Corium, i. 89. Cornea, ii. 326. Corneule, ii. 344. Cornu Ammonis, i. 192. Coronet bone, i. 356. Corpora quadrigemina, ii. 391. Corpus callosum, ii. 396. Corpus papillare, ii. 268. Cortical substance, ii. 395. Cossus, i. 214, 222, 249. Cotunnius, ii. 303. Cowrie, i. 179. Crab, i. 207; ii. 186, 214, 226, 347. €ranium, i. 278, 307, 326. Cranium of insects, i. 228. Craw, ii. 124. Cray-fish, ii. 308, 346. Cribriform plate, ii. 284. Crinoidca, i. 1.56. Crocodile, i. 317, 318, 320; ii. 105, 119,199,290,312,392. Crop, ii. 130. Cross-bill, ii. 98. Crotalus, i. 312. Crust, 1.89. Crusta pctrosa, ii. 113. Crustacea, i. 204; ii. 194, 211, 214, 381, 412. Cryptogamia, i. 64; ii. 416. fTystalline lens, i. 56; ii. 327. Crystalline needles in biliary ducts, ii. 159. Curculio, i.233. Cushions of insects, i. 235. Cuticle, vetretable, i. 67. Cuticle, animal, i. 90; ii. 268. Cuttle-fish. l Kieser, i. 61, 06. Kirby, i. 229 ; jL 293, 342. Knight, ii. 417. Knots in wood, ii. 413. Koala, i. 303. Kolpoda, i. 139. Ijabium of insects, ii. 93. Labnini of insects, ii. 93. Labyrinth, ii. 30;i Lacerta, i. 317. Lacrymal organs, ii. 330. Lacteals, ii. 82, 104. Lamarck, i. 115; ii. 71, 445. Lamina spiralis, ii. 305. Lamouroiix, i. 115. Lamprey, i. 289; ii. 88, 218, 310. Lancets of diptera, ii. 87. Language of insects, ii. 274. Lark, i. 401. Larva, i. 217, 2ia Lassaigne, i. 226 ; ii. 134. Latham, ii. 134. Latreille, i. 207; ii. 22.5, 276, .347. Laws of nature, L 20. Law of mortality, i. 44. Law of coexistence of forms, i. 50. liaw of gradation, ii. 439. Law of analogy, i. 49 ; ii. 437. Leach, i. 161. Leaves, ii. 27, 38. Leech. (»S'ec Hirudo.) Lemur, i. 367, 380; ii. 204, 3-55. Lens, crystalline, i. 56; ii. 327, 350, Lenticclla?, i. 78. Lepas, i. 185 ; ii. 212. Lepidoptera, i. 217, 249; ii. 87, 157. Lepisma, i. 212, 250. Lernsea, L 216 ; ii. 421, 426. Leuchft, ii. 340. Leucophra, ii. 73. Leuret, ii. 134. Lewenhoeck, i. 251 ; ii, 190. Libellula, i. 221, 247; ii. 343. Liber, i. 74 ; ii. 36. Lichen, ii. 21. Life, i. 39, 44. Ligaments, i. 86. Ligamentum nuchse, i. 87, 346. Light on plants, i. 76 ; ii. 27. Lignine, i. 63; ii. 36. Lilium, i. 68. Li max, ii. 95, 226. Limpet. See Patella. Link, i. 66. Lion, i. 87, 342, 365 ; ii. 101, 278, 392. Lister, ii. 169, 215. Liver, ii. 159, 248. Lizard, ii. 97, 277, 351, 412. Lobster, i. 208 ; ii. 123, 186, 214, 308, 383. Lobularia, i. 122. Loche, ii. 221. liOcomotion, i. 110. Locusta, ii, 91. Loligo, i. 188,283; ii. 195. Longevity of trees, ii. 434. Lophius, i. 293 ; ii. 276, 310. Loxia, ii. 98. Lucanus, i. 252. Lumbricus marinus, i. 199, 210. Lumbricus terrestris, ii. 77, 87^ 184, 213. Lungs, ii. 192,428. Lycopodium, i. 68. Lycoris, ii. 338. Lymphatics, ii. 250. Lymphatic hearts, ii. 251. Lyonet, i. 214, 222, 249. Macaire, ii. 43, 45, 47, 238. Macartney, i. 406; ii. 234, 235, 396. Macavoy, ii. 266. Mackerel, i. 295. Macleay, i. 52. Madder, i. 269. Madrepore, i. 126. Magendie, ii. 356, 375; Magilus, i. 180. Mam, ii. 194, 383. Malleus, ii. 302. Malpighi, ii. 268. Mamrna}, ii. 419, 431. Mammalia, i. 330 ; ii. 230, 312, 419. Man, i. 369; ii. 394. Man of war, Portuguese, i. 145. Manntus, ii. 105. Mandible, i. 206. Mantis, ii. 153. Mantle, i. 90, 172. Many-plies stomach, ii. 143. Marcet, ii. 47, 164, 238, 328. INDEX. 457 Marginella, i. 179. Marmot, ii. 110. Marsigli, i. 115. Marsnpialia, ii. 199, 419. Marsupium, ii. 3.52. Mastication, ii. 104. Mastoid cells, ii. 301. Matrix of feather, i. 397. Matter, ii. 363. Maunoir, ii. 370. MaxillfD, ii. 93. Mayer, i. 310. Mayo, ii. 432. Meatus auditorius, ii. 299. Mechanical functions, i. 41. Meckel, i. 333 ; ii. 339. Medulla oblongata, ii. .391. Medullary substance, ii. 260. Medullary rays, i. 73. Medusa, i. 80, 143; ii. 51, 56, 05, 210, 337. Meibomian glands, ii. 331. Melolontha, i. 214 ; ii. 92, 154, 171, 223, 343, 34.5. Melophagus, ii. 341. Membrana nictitans, ii. 3-52, 353, Membrane, i. 83. Menobranchus, ii. 231. Mercurialis, ii. 44. Mergys, ii. 98. Merry-thought of fowl, i. 390. Mesembryanthemum, ii. 40. Mesenteric glands, ii. 164. Mesentery, ii. 82. Mesothorax, i. 229. Metacarpus, i. 282. Metals in plants, ii. 37. Metamorphoses, i. 216, 303 ; ii. 442, 443. Metatarsus, i. 282. Metathorax, i. 229. Milk, ii. 431. Millepedes, ii. 342. Millepora, i. 125. Mimosa, i. 100. Mint, ii. 20, 29. Mirandola, ii. 411. Mirbel, i. 62, 65. Mite, i. 212. Mitra, i. 180. Modiolus, ii. 30.5. Molar teeth, ii. 107. Moldenhawer, i. 66. Mole, i. 361,362; ii, 278, 356. Mole cricket, i. 241. Vol. II. Mollusca, i. 157; ii. 176,276,387. Monads", i. 25, 137 ; ii. 78, 409. Monkey, i. 367; ii. 110, 279, 4(K). Munoculus, ii. 34H. Monulhaluinous shell, i. 191. Monotrcmata, ii. 199. Munru, i. 97, 103; ii.217. Mordella, ii. :i43. Morpho, i. 249. Morren, ii. 182, 184. Mortality, i. 44; ii. 408. Mother of pearl, i. 169. Motion, voluntary, i. 41 ; ii. 375. Motion, vegetable, i. 1(X). Motor nerves, ii. 375. Mucous membrane, i. 90. Mucous glands, ii. 135. Mulberry, ii. 48. Mailer, i. 136 ; ii. 70, 251, :U0. Mullet, ii. 147. Multilocular shells, i. IfX). Multivalves, i. 185. Murscna, ii. 351, 31J2. Murex, i. 178, 182; ii. 95, 21.5, 340. Mus, ii. 130, 356. Musca, i. 235. Muscle, (shell fish,) i. 162, 164. Muscle, i. 97, 100, 214. Muscles of eye, ii. 328. Muscular power in plants, ii. 254- Muscular power in birds, i. 408. Mushroom, ii. 21. Musk shrew, ii. 101. Musical tone, ii. 297. My a, i. 163. Myriapoda, i.212; ii. 180, Myrmecophaga, ii. 100. Mysis Fabricii, i. 206. Mytilus, i. 162. Myxine, i. 283, 289 ; ii.«88, 351. Nacreous structure, i. 168. Nais, ii. 78, 182, 338, 412. Narwhal, i..53; ii. 105. Nature, i. 20, 25. Nautilus, i. 17.5, 191 ; ii. 194. Necrophorus, ii. 293. Needles in biliary ducts, ii. 159. Nereis, i. 19.5, 197, 201 ; ii. 182. Nerve, i. 40; ii. 261. Nervous system, ii. 260, 37"^. 3)^. Nervous ]K)wer, ii. 2-52. Nettle, ii. 40. Neuro-skeleton, i- 257. Neuroptcra, i. 215. 5S 458 INDEX. Newport, i. 247; ii. 78, 155, 157, 177, :385. Newt, ii. 311, 412. Nightshade, ii. 48. Nitrogen, ii. 18, 240. Nordmann, ii. 420, 426. Notonecta, i. 85, 238. Nursling sap, ii. 25. Nutrition, ii. 9, 15, 17, 47. Nutrition in lower orders, ii. 5^. Nutrition in higher orders, ii. K). Nutritive functions, i. 42. Nycteribia, ii. 341. Ovary, ii. 416, 417. Oviduct, ii. 418. Oviparous animals, ii. 419. Ovo-viviparous animals, ii. 419. Ovula, i. 179. Ovum, ii. 416. Owen, i. 388. Owl, ii. 235, 312, 354. Ox, iiorn of, i. 355. O.xygen, ii. 28. Oyster, i. 102, 161, 162. Oyster-catcher, ii. 98. Octopus, i. 188; ii. 348. Ocular spectra, ii. 301. Odier, i. 226. (Esophagus, ii. 76, 82, 129. Oken, i. 246, 279. Olfactory nerve, ii. 281. Olfactory lobes, ii. 391. OlivcT, i. 175, 180. Oniscus, ii. 382. Onocrotalus, i. 384. Operculum of Mollusca, i. 182. Operculum of fishes, ii. 217. Ophicephalus, ii. 219. Ophidia, i. 310. Ophiosaurus, i. 315, 317. Ophiura, i. 155. Opossum, ii. 101, 419. Optic axis, ii. ii54. Optic ganglion, ii. 345. Optic lobes, ii. 391. Opuntia, i. 100. Orache, ii. 40. Orbicular bone, ii. 303. Orbicular muscle, i. 105. Orchidese, i. 62. Organic Mechanism, i. 58, 79. Organic development, ii. 420. Ornithorhyncus, i. 276; ii. 101, 130, 277, 313, 350. Orobanche, ii. 45. Orthoceratite, i. 192. Orthoptera, i. 220, 245. Os hyoides, ii. 99, 216. Osier, i. 152, 162, 163, 199, 201. Osseous fabric, i. 256. Ossicula, tympanic, ii. 302. Ossification, i. 263, 383. Ostracion, i. 300. Ostrich, i. 388, 404,407; ii. 135, 162, 234, 390. Otter, sea, ii. 110. Paces of quadrupeds, i. 339. Pachydermata, i. 357; ii. 271, 278. Package of organs, i. 83. Pain, ii. 262. Palcmon, ii. 383. Paley, i. 83, 394 ; ii. 205. Palinurus, ii. 383. P(dlas, i. 115 ; ii. 244. Palms, i. 72. Palm squirrel, ii. 130. Palmer, ii. 28. Palpi, i. 206; ii. 93. Pancreas, ii. 160. Pander, ii. 425. Panniculus carnosus, i. 363. Panorpa, i. 231. Paper nautilus, i. 191. Papilio, i. 251 ; ii. 343. Papillce, ii. 269, 279. Par vagum, ii. 386. Parakeet, ii. 98. Parallax, aberration of, ii. 333. Parrot, ii. 130, 277. Pastern, i. 356. Patella, i. 167; ii. 167, 388. Patella of knee, i. 282. Patellaria, ii. 39. Paunch, ii. 142. Pearl, i. 169. Peccari, ii. 140. Pediculus, i. 212. Pelican, i. 383 ; ii. \^. Pelvis, i. 281. Pencil of rays, ii. 320. Penguin, i. 407. Penitentiary, ii. 138. Pennatula, i. 131 ; ii. 63. Penniform muscle, i. 103. Pentacrinus, i. 156. Perca, i. 93, 301 ; ii. 219, 291, 349, 392. Perception, i. 40; ii. 264, 358. INDEX. 459 Perch. See Perca. Perennibranchia, ii. 230. Perilymph, ii. 303. Periostracum, i. 172. Peristaltic motion, ii. 148. P^ron, i. 80; ii. 56. Pfaff, ii. 240. Phalanges, i. 282. Phalena, ii. 177, 343. Phanerog-amous plants, ii. 417. Phantasmagoria, ii. 374. Phantasmascope, ii. 368. Phaseolus, ii. 43. Phenakisticope, ii. 369. Philip, ii. 138, 256. Phoca, i. 336. Pholas, i. 161, 185. Phosphorescence of the sea, i. 143; ii. 50. Phrenology, ii. 397. Phyllosoma, ii. 382. Physalia, i. 145. Physiology, i. 30. Physsophora, i. 145. Phytozoa, i. 113. Pierard, ii. 145. Pigeon, ii. 131, 431. Pigmentum of skin, i. 90. Pigmentum of the eye, ii. 327. Pike, i. 296. Pileopsis, i. 182. Pineal gland, ii. 394. Pinna, i. 171, 164. Pistil, ii. 418. Pith of plants, i. 73. Pith of qui 11, i. 399. Placuna, i. 169. Planaria, ii. 86, 171, 181, 211, 338, 402. Planorbis, i. 166, 175. Plantigrada, i. 367. Plastron, i. 321. Pleurobranchus, ii. 159. Pleuronectes, i. 299 ; ii. 354. Plexus, nervous, ii. 255. Pliny, ii. 394. Plumula, ii. 422. Plumularia, ii. 170. Pneumo-branchisB, ii. 225. Pneumo-gastric nerve, ii. 386. Podura, i. 212. Poisers, i. 249. Poison of nettle, ii. 40. PoU,\. 165, 171. Pollen, ii. 418. Polygastrica, ii. 73. Polypi, i. 122 ; ii. 58, 63, 210, 272. Poly stoma, ii. 86. Polythalamous shell, i. 190. Pontia brassica, i. 249. Pontobdella, i. 195. Poppy, ii. 41. Porcupine quills, i. 95. Porcupine, i. 363 ; ii. 110, 140. Porifera, i. 113. Porpita, i. 144. Porpus, ii. 10.5, 140. Porterfield, i. 262. Potato, ii. 413. Prehension of food, ii. 86, 89. Priestley, ii. 28, 2:38, 240. Pristis, i. 53; ii. 122. Pritchard, ii. 174. Privet Flawk moth, ii. 157. Proboscis of insects, ii. 87. Proboscis of molltisca, ii. 9.5. Proboscis of Elephant, i. 359. Progressive motion, i. 112. Prologs, i. 223 Promontory of ear, ii. 302. Proteus, 1. 139 ; ii. 231, 442. Prothorax, i. 229. Praia, ii. .33, 36. Provengal, ii. 220. Proximate principles, ii. 12. Pterocern, i. 178. Pteropoda, i. 18.5. Pteropus, ii. 101. Pubic bone, i. 282. Pulmonary organs, ii. 19'2. Puncta lacrymalia, ii. 331. Punctnm .=;iliori=, ii. 425. Pupa, i. 217, 220. Pupil, ii. 327. Pupiparn, ii. ;341. Pyloric appnndicos, ii. 160. Pylorus, ii. 82, 133. Pyramidalis muscle, ii.353. Python, i. 310. Quadratus muscle, ii. 353. (iuadrumana, i. 367; ii. 110. (iuadrupedis, i. 336. Quaggn, i. 3,56. Uuail, i. 401. Quills of porcupine, i. 95. Quills of feathers, i. 392. quny, i. «0. Rabbit, i. 343 ; ii. IKK 1M> 460 INDEX. Racoon, i. 90. Radiata, i. 124. Radicles, ii. 422. Radius, i. 282. Ranunculus, i. 69. Rapp, ii. 337. Rat, ii. 110, 140. Ruthke, ii. 443. Rattle-snake, i. 312. Ray, i. 24. Ray, i. 292, 293, 294 ; ii. 354, 400. Rays of fins, i. 294. Razor-shell-fish, i. 162. Reamitr, i. 147, 149, 165, 173,208; ii. b7, 125, 134. Receptacles of food, ii. 130. Receptaculum chyli, ii. 82, 165. Reed of ruminants, ii. 143. Refraction, law of, ii. 321. Regeneration of claw, i. 212. Rennet, ii. 143- Reparation, ii. 10, 14, 412. Repetition of organs, i. 54. Reproduction, i. 45; ii. 408. Reptiles, i. 302; ii. 197. Resinous secretions, ii. 40. Respiration, i. 44; ii- 16, 191, 208. Rete muscosum, i. 90. Reticulated cells, i. 62. Reticule of Ruminants, ii. 142. Retina, ii. 266, 317, 327. Returning sap, ii. 32. Revelation, ii. 447. Reviviscence, i. 58 ; ii. 184. Rhea, i. 404. Rhinoceros, i. 3.56; ii. 101, 112, 271, 278, 355. Rhipiptcra, i. 246. Rliizostoma, ii. 67. Rhyncops, ii. 98. Ribs, i. 280; ii. 233. Ricinus, i. 212. Rings of annclida, i. 19.5. Rodcntia, i. 361; ii. 109, 112, 119, 128, 139, 354. Rcpsel, ii. 337. Rogpt, ii. 368, 373, 408. Rolando, ii. 429. Roosting, i. 405. Roots, i. 78 ; ii. 22. Ross, i, 27. Rostrum, ii. 93. Rotifer, i. 5<^, 140; ii. 70, 338, 379, 415. JRoux, ii. 400. Rudimental organs, i. 52 ; ii. 442. Rudolphi, i. 66. Rumjord, i. 67. Ruminantia, i. 345; ii. 142, 354. Rusconi, ii. 429. Sabella, i. 198. Sacculusofear, ii. 305. Sacrum, ii. 281. St. Ayiirp, ii. 200. 2. Voluntary motion, i. 41 ; ii. 375. Volute, i. 170; ii. 95, :U0. Volvo.x, i. 13^, 139; ii. 414. Voracity of hydra, ii. 61. Vorticella, i. 58, 136; ii. 74, 410. Vulture, ii. 131, 288. Wading birds,*!. 40.3, 408. Walking, i. 340, .374. Waller, ii. 100. Walrus, i. 336; ii. 105. Warfare, animal, i. 47; ii. 53. Warm-blooded circulation, ii. 199. Water, not the food of plants, ii. 19. Water-beetle. See Dytiscus. Water-boatman, i. 35, 238. Wax, vegetable, ii. 40. Web-footed bird.s i. 408. Weber, ii. 305, 339. Whale, i. .53; ii. 130, 313, 35.5,394. Whalebone, ii. 102. Wheel animalcule, i. 140. Wheel spokes, spectre of, ii. 368. Whelk. See Buccinum. Whiskers, ii. 278. Whorls of plants, i. 76. Whorls of shells, i. 176. Willow, i. a9. Wings, i. 243, 391. Winged insects, i. 214. * Withers, i. 257. Wolf-fish, ii. 96. Wollaston, i. 77 ; ii. 45, 346, 4()1. Wombat, i. 363. Woodhoiise, ii. 28. Woodpecker, ii. 98. Woody fibres, i. 64, 66. Worms. See Annelida and Eii- tozoa. Yarrell, ii. 98. Young, ii. 335. Zebra, i. 356. Zemni, ii. 3.56. Zoanthus, i. 123, 136. Zoocarpia, i. 119. Zoophytes, i. 113; ii. .3.37. .37^. Zostira, ii. 146. Zygodactyli, i. 404. THE END. m C lie Colldt CAREY, LEA, & BLAN CHARD, •HAVE LATELY PUBLISHED THE SECOND EDITION, VERY MUCH ALTERED AND IMPROVED, OF nrHIAN PHYSIOIiOGY, ILLUSTRATED BY NUMEROUS ENGRAVINGS. BY ROBLEY DUNGLISON, M. D., &c., &c. This work, although intended chiefly for the professional reader, is adapted to tile comprehension of every one, the anatomical and other descriptions being elucidated by wood-cuts, and by copperplate engravings. It comprises a full investigation of every function e.xecuted by ihe various organs of the body in health, and is calculated to con- vey accurate impressions regarding all the deeply interesting and mysterious phenomena that are associated with the life of man — both as an individual, and a species— and a knowledge of which is now regarded indispensable to the formation of the well educated gentleman. " It is the most complete and satisfactory system of Physiology in tl)e English language. ^t will add to the already high reputation of the aulhoi :'—Jimerican Journal of the Me- dical Sciences. " We feel warranted in recommending the work to the student of physiology, as being one of the very best text books with which we are acquainted ; while we are persuaded its very superior merits will command for it a place in every medical library."— 7fti