The Physiology of the Crayfish
THE MECHANISM BY WHICH THE PARTS OF THE LIVING ENGINE ARE SUPPLIED WITH THE MATERIALS NECESSARY FOR THEIR MAINTENANCE AND GROWTH.
AN analysis of such a sketch of the "Natural History of the Crayfish" as is given in the preceding chapter, shows that it provides brief and general answers to three questions. First, what is the form and structure of the animal, not only when adult, but at different stages of its growth? Secondly, what are the various actions of which it is capable? Thirdly, where is it found? If we carry our investigations further, in such a manner as to give the fullest attainable answers to these questions, the knowledge thus acquired, in the case of the first question, is termed the Morphology of the crayfish; in the case of the second question, it constitutes the Physiology of the animal; while the answer to the third question would represent what we know of its Distribution or Chorology. There remains a fourth problem, which can hardly be regarded as seriously under discussion, so long as knowledge has advanced no further than the Natural History stage; the question, namely,
how all these facts comprised under Morphology, Physiology, and Chorology have come to be what they are; and the attempt to solve this problem leads us to the crown of Biological effort, Ætiology. When it supplies answers to all the questions which fall under these four heads, the Zoology of Crayfish will have said its last word.
As it matters little in what order we take the first three questions, in expanding Natural History into Zoology, we may as well follow that which accords with the history of science. After men acquired a rough and general knowledge of the animals about them, the next thing which engaged their interest was the discovery in these animals of arrangements by which results, of a kind similar to those which their own ingenuity effects through mechanical contrivances, are brought about. They observed that animals perform various actions; and, when they looked into the disposition and the powers of the parts by which these actions are performed, they found that these parts presented the characters of an apparatus, or piece of mechanism, the action of which could be deduced from the properties and connections of its constituents, just as the striking of a clock can be deduced from the properties and connections of its weights and wheels.
Under one aspect, the result of the search after the rationale of animal structure thus set afoot is Teleology; or the doctrine of adaptation to purpose. Under another
aspect, it is Physiology; so far as Physiology consists in the elucidation of complex vital phenomena by deduction from the established truths of Physics and Chemistry, or from the elementary properties of living matter.
We have seen that the crayfish is a voracious and indiscriminate feeder; and we shall be safe in assuming that, if duly supplied with nourishment, a full-grown crayfish will consume several times its own weight of food in the course of the year. Nevertheless, the increase of the animal's weight at the end of that time is, at most, a small fraction of its total weight; whence it is quite clear, that a very large proportion of the food taken into the body must, in some shape or other, leave it again. In the course of the same period, the crayfish absorbs a very considerable quantity of oxygen, supplied by the atmosphere to the water which it inhabits; while it gives out, into that water, a large amount of carbonic acid, and a larger or smaller quantity of nitrogenous and other excrementitious matters. From this point of view, the crayfish may be regarded as a kind of chemical manufactory, supplied with certain alimentary raw materials, which it works up, transforms, and gives out in other shapes. And the first physiological problem which offers itself to us is the mode of operation of the apparatus contained in this factory, and the extent to which the products of its activity are to be accounted for by reasoning from known physical and chemical principles.
We have learned that the food of the crayfish is made up of very diverse substances, both animal and vegetable; but, so far as they are competent to nourish the animal permanently, these matters all agree in containing a peculiar nitrogenous body, termed protein, under one of its many forms, such as albumen, fibrin, and the like. With this may be associated fatty matters, starchy and saccharine bodies, and various earthy salts. And these, which are the essential constituents of the food, may be, and usually are, largely mixed up with other substances, such as wood, in the case of vegetable food, or skeletal and fibrous parts, in the case of animal prey, which are of little or no utility to the crayfish.
The first step in the process of feeding, therefore, is to reduce the food to such a state, that the separation of its nutritive parts, or those which can be turned to account, from its innutritious, or useless, constituents, may be facilitated. And this preliminary operation is the subdivision of the food into morsels of a convenient size for introduction into that part of the machinery in which the extraction of the useful products is performed.
The food may be seized by the pincers, or by the anterior chelate ambulatory limbs; and, in the former case, it is usually, if not always, transferred to the first, or second, or both of the anterior pairs of ambulatory limbs. These grasp the food, and, tearing it into pieces of the proper dimensions, thrust them between the external maxillipedes, which are, at the same time,
worked rapidly to and fro sideways, so as to bring their toothed edges to bear upon the morsel. The other five pairs of jaws are no less active, and they thus crush and divide the food brought to them, as it is passed between their toothed edges to the opening of the mouth.
As the alimentary canal stretches from the mouth, at one end, to the vent at the other, and, at each of these limits, is continuous with the wall of the body, we may conceive the whole crayfish to be a hollow cylinder, the cavity of which is everywhere closed, though it is traversed by a tube, open at each end (fig. 6). The shut cavity between the tube and the walls of the cylinder may be termed the perivisceral cavity; and it is so much filled up by the various organs, which are interposed between the alimentary canal and the body wall, that all that is left of it is represented by a system of irregular channels, which are filled with blood, and are termed blood sinuses. The wall of the cylinder is the outer wall of the body itself, to which the general name of integument may be given; and the outermost layer of this, again, is the cuticle, which gives rise to the whole of the exoskeleton. This cuticle, as we have seen, is extensively impregnated with lime salts; and, moreover, in consequence of its containing chitin, it is often spoken of as the chitinous cuticuta.
Having arrived at this general conception of the disposition of the parts of the factory, we may next proceed to consider the machinery of alimentation which is
contained within it, and which is represented by the various divisions of the alimentary canal, with its appendages; by the apparatus for the distribution of nutriment; and by two apparatuses for getting rid of those products which are the ultimate result of the working of the whole organism.
And here we must trench somewhat upon the province of Morphology, as some of these pieces of apparatus are complicated; and their action cannot be comprehended without a certain knowledge of their anatomy.
The mouth of the crayfish is a longitudinally elongated, parallel-sided opening, in the integument of the ventral or sternal aspect of the head. Just outside its lateral boundaries, the strong mandibles project, one on each side (fig 3, B; 4); their broad crushing surfaces, which are turned towards one another, are therefore completely external to the oral cavity. In front, the mouth is overlapped by a wide shield-shaped plate termed the upper lip, or labrum (figs. 3 and 6, lb); while, immediately behind the mandibles, there is, on each side, an elongated fleshy lobe, joined with its fellow by the posterior boundary of the mouth. These together constitute the metastoma (fig. 3, B; mt), which is sometimes called the lower lip. A short wide gullet, termed the oesophagus (fig. 6, oe), leads directly upwards into a spacious bag, the stomach, which occupies almost the whole cavity of the head. It is divided by a constriction into a large anterior chamber (cs), into the under face of which the
gullet opens, and a small posterior chamber (ps), from which the intestine (hg) proceeds.
In a man's stomach, the opening by which the gullet communicates with the stomach is called the cardia, while that which places the stomach in communication with the intestine is named the pylorus; and these terms having been transferred from human anatomy to that of the lower animals, the larger moiety of the crayfish's stomach is called the cardiac division, while the smaller is termed the pyloric division of the organ. It must be recollected, however, that, in the crayfish, the so-called cardiac division is that which is actually furthest from the heart, not that which is nearest to it, as in man.
The gullet is lined by a firm coat which resembles thin parchment. At the margins of the mouth, this strong lining is easily seen to be continuous with the cuticular exoskeleton; while, at the cardiac orifice, it spreads out and forms the inner or cuticular wall of the whole gastric cavity, as far as the pylorus, where it ends in certain valvular projections. The chitinous cuticle which forms the outermost layer of the integument is thus, as it were, turned in, to constitute the innermost layer of the walls of the stomach; and it confers upon them so great an amount of stiffness that they do not collapse when the organ is removed from the body. Furthermore, just as the cuticle of the integument is calcified to form the hard parts of the exoskeleton, so is the cuticle of the stomach calcified, or otherwise hardened, to give rise, in the first
place, to the very remarkable and complicated apparatus which has already been spoken of, as a sort of gastric mill or food-crusher; and, secondly, to a filter or strainer, whereby the nutritive juices are separated from the innutritious hard parts of the food and passed on into the intestine.
[Figure 9: Astacus fluviatilis--Stomach, pyloric region]
The gastric mill begins in the hinder half of the cardiac division. Here, on the upper wall of the stomach, we see a broad transverse calcified bar (figs. 9-11, c) from the middle of the hinder part of which another bar (uc), united to the first by a flexible portion, is continued backwards in the middle line. The whole has, therefore, somewhat the shape of a cross-bow. Behind the first-mentioned piece, the dorsal wall of the stomach is folded in, in such a manner as to give rise to a kind of pouch; and the second piece, or what we may call the handle of the crossbow, lies in the front wall of this pouch. The end of this piece is dense and hard, and its free surface, which looks into the top of the cardiac chamber, is raised into two oval, flattened convex surfaces (t). connected by a transverse joint with the end of the handle of the crossbow, there is another solid bar, which ascends obliquely forwards in the back wall of the pouch (pp). The end which is articulated with the handle of the crossbow is produced into a strong reddish conical tooth (mt), curved forwards and bifurcated at the summit; consequently, when the cavity of the stomach is inspected from the fore part of the cardiac pouch (fig. 9, B), the two-pointed curved tooth (mt) is seen projecting behind the convex surfaces (t), in the middle line, into the interior of that cavity. The joint which connects the handle of the crossbow with the hinder middle piece is elastic; hence, if the two are straightened out, they return to their bent disposition as soon as they are released. The upper end of
the hinder middle piece (pp) is connected with a second flat transverse plate which lies in the dorsal wall of the pyloric chamber (p). The whole arrangement, thus far, may be therefore compared to a large cross-bow and a small one, with the ends of their handles fastened together by a spring joint, in such a manner that the handle of the one makes an acute angle with the handle of the other; while the middle of each bow is united with the middle of the other
[Figure 10: Astacus fluviatilis--Longitudinal section of the stomach]
There are two small pointed teeth, one under each of the lateral teeth, and each of these is supported by
a broad plate, hairy on its inner surface, which enters into the lateral wall of the cardiac chamber. There are various other simpler skeletal parts, but the most important are those which have been described; and these, from what has been said, will be seen to form a sort of hexagonal frame, with more or less flexible joints at the angles, and having the anterior and the posterior sides connected by a bent jointed middle bar. As all these parts are merely modifications of the hard skeleton, the apparatus is devoid of any power of moving itself. It is set in motion, however, by the same substance as that which gives rise to all the other bodily movements of the crayfish, namely, muscle. The chief muscles which move it are four very strong bundles of fibres. Two of these are attached to the front crosspiece, and proceed thence, upwards and forwards, to be fixed to the inner face of the carapace in the front part of the head (figs. 5, 6, and 12, ag). The two others, which are fixed into the hinder crosspiece and hinder lateral pieces, pass upwards and backwards, to be attached to the inner face of the carapace in the back part of the head (pg). When these muscles shorten, or contract, they pull the front and back crosspieces further away from one another; consequently, the angle between the handles becomes more open and the tooth which is borne on their ends travels downwards and forwards. But, at the same time, the angle between the side bars becomes more open and the lateral tooth of each side moves inwards till it crosses in front of the middle tooth, and strikes against this and the opposite lateral tooth, which has undergone a corresponding change of place. The muscles being now relaxed, the elasticity of the joints suffices to bring the whole apparatus back to its first position, when a new contraction brings about a new clashing of the teeth. Thus, by the alternate contraction and relaxation of these two pair of muscles, the three teeth are made to stir up and crush whatever is contained in the cardiac chamber. When the stomach is removed and in the front part of the cardiac chamber is cut away, the front cross-piece may be seized with one pair of forceps and the hind cross-piece with another. On slightly pulling the two, so as to imitate the acton of the muscles, the three teeth will be found to come together sharply, exactly in the manner described.
Works on mechanics are full of contrivances for the conversion of motion; but it would, perhaps, be difficult to discover among these a prettier solution of the problem; given a straight pull, how to convert it into three simultaneous convergent movements of as many points.
What I have called the filter is constructed mainly out of the chitinous lining of the pyloric chamber. The aperture of communication between this and the cardiac chamber, already narrow, on account of the constriction of the walls of the stomach at this point, is bounded at the sides by two folds; while, from below, a conical tongue-shaped process (figs. 6, 10, and 11, cpv), the surface of which is covered with hairs, further obstructs the opening. In the posterior half of the pyloric chamber, its side walls are, as it were, pushed in; and, above, they so nearly meet in the middle line, that a mere vertical chink is left between them; while even this is crossed by hairs set upon the two surfaces. In its lower half, however, each side wall curves outwards, and forms a cushion-shaped surface (fig. 10, cs) which looks downwards and inwards. If the
floor of the pyloric chamber were flat, a wide triangular passage would thus be left open in its lower half. But, in fact, the floor rises into a ridge in the middle, while, at the sides, it adapts itself to the shape of the two cushion-shaped surfaces; the result of which is that the whole cavity of the posterior part of the pyloric division of the stomach is reduced to a narrow three-rayed fissure. In transverse section, the vertical ray of this fissure is straight, while the two lateral ones are concave upwards (fig. 9, E). The cushions of the side walls are covered with short close-set hairs. The corresponding surfaces of the floor are raised into longitudinal parallel ridges, the edge of each of which is fringed with very fine hairs. As everything which passes from the cardiac sac to the intestine must traverse this singular apparatus, only the most finely divided solid matters can escape stoppage, so long as its walls are kept together.
Finally, at the opening of the pyloric sac into the intestine, the chitinous investment terminates in five symmetrically arranged processes, the disposition of which is such that they must play the part of valves in preventing any sudden return of the contents of the intestine to the stomach, while they readily allow of a passage the other way. One of these valvular processes is placed in the middle line above (figs. 10 and 11, v1). It is longer than the others and concave below. The lateral processes (v2,) of which there are two on each side, are triangular and flat.
[Figure 11: Astacus fluviatilis--View of the roof of the stomach and of the mid-gut]
The cuticular lining which gives rise to all the complicated apparatus which has just been described, must not be confounded with the proper wall of the stomach, which invests it, and to which it owes it origin, just as the cuticle of the integument is produced by the soft
true skin which lies beneath it. The wall of the stomach is a soft pale membrane containing variously disposed muscular fibres; and, beyond the pylorus, it is continued into the wall of the intestine.
It has already been mentioned that the intestine is a slender and thin-walled tube, which passes straight through the body almost without change, except that it becomes a little wider and thicker-walled near the vent. Immediately behind the pyloric valves, its surface is quite smooth and soft (figs. 9, 10, and 12, mg), and its floor presents a relatively large aperture, the termination of the bile duct (fig. 12, bd, fig. 10, hp.), on each side. The roof is, as it were, pushed out into a short median pouch or cæcum (cæ). Behind this, its character suddenly changes, and six squarish elevations, covered with a chitinous cuticle, encircle the cavity of the intestine (r). From each. of these, a longitudinal ridge, corresponding with a fold of the wall of the intestine, takes its rise, and passes, with a slight spiral twist, to its extremity (hg). Each of these ridges is beset with small papillæ, and the chitinous lining is continued over the whole to the vent, where it passes into the general cuticle of the integument, just as the lining of the stomach is continuous with the cuticle of the integument at the mouth. The alimentary canal may, therefore, be distinguished into a fore and a hind-gut (hg), which have a thick internal lining of cuticular membrane; and a very short mid-gut (mg), which has no thick cuticular layer. It will be of
importance to recollect this distinction by-and-by, when the development of the alimentary canal is considered.
[Figure 11: Astacus fluviatilis--A dissection of a male specimen from the right side]
If the treatment to which the food is subjected in the alimentary apparatus were of a purely mechanical nature, there would be nothing more to describe in this part of the crayfish's mechanism. But, in order that the nutritive matters may be turned to account, and undergo the chemical metamorphoses, which eventually change them into substances of a totally different character, they must pass out of the alimentary canal into the blood. And they can do this only by making their way through the walls of the alimentary canal; to which end they must either be in a state of extremely fine division, or they must be reduced to the fluid condition. In the case of the fatty matters, minute subdivision may suffice; but the amylaceous substances and the insoluble protein compounds, such as the fibrin of flesh, must be brought into a state of solution. Therefore some substances must be poured into the alimentary canal, which, when mixed with the crushed food, will play the part of a chemical agent, dissolving out the insoluble proteids, changing the amyloids into soluble sugar, and converting all the proteids into those diffusible forms of protein matter, which are known as peptones.
The details of the processes here indicated, which may be included under the general name of digestion, have only quite recently been carefully investigated in the crayfish; and we have probably still much to
learn about them; but what has been made out is very interesting, and proves that considerable differences exist between crayfishes and the higher animals in this respect.
The physiologist calls those organs, the function of which is to prepare and discharge substances of a special character, glands; and the matter which they elaborate is termed their secretion. On the one side, glands are in relation with the blood, whence they derive the materials which they convert into the substances characteristic of their secretion; on the other side, they have access, directly or indirectly, to a free surface, on to which they pour their secretion as it is formed.
Of such glands, the alimentary canal of the crayfish is provided with a pair, which are not only of very large size, but are further extremely conspicuous, on account of their yellow or brown colour. These two glands (figs. 12 and 13, lr) are situated beneath, and on each side of, the stomach and the anterior part of the intestine, and answer in position to the glands termed liver and pancreas in the higher animals, inasmuch as they pour their secretion into the mid-gut. These glands have hitherto always been regarded as the liver, and the name may be retained, though their secretion appears rather to correspond with the pancreatic fluid than with the bile of the higher animals. [see End note 7]
[Figure 13: Astacus fluviatilis--The alimentary canals and livers seen from above]
Each liver consists of an immense number of short tubes, or cæca, which are closed at one end, but open at the other into a general conduit, which is termed their duct. The mass of the liver is roughly divided into
three lobes, one anterior, one lateral, and one posterior; and each lobe has its main duct, into which all the tubes composing it open. The three ducts unite together into a wide common duct (bd), which opens, just behind the pyloric valves, into the floor of the mid-gut. Hence the apertures of the two hepatic ducts are seen, one on each side, in this part of the alimentary canal when it is laid open from above. Every cæcum of the liver has a thin outer wall, lined internally by a layer of cells, constituting what is termed an epithelium; and, at the openings of the hepatic ducts, this epithelium passes into a layer of somewhat similar structure, which lines the mid-gut, and is continued through the rest of the alimentary canal, beneath the cuticula. Hence the liver may be regarded as a much divided side pouch of the mid-gut.
The epithelium is made up of nucleated cells, which are particles of simple living matter, or protoplasm, in the midst of each of which is a rounded body, which is termed the nucleus. It is these cells which are the seat of the manufacturing process which results in the formation of the secretion; it is, as it were, their special business to form that secretion. To this end they are constantly being newly formed at the summits of the cæca. As they grow, they pass down towards the duct and, at the same time, separate into their interior certain special products, among which globules of yellow fatty matter are very conspicuous. When these products are fully formed, what remains of the substance of the cells dissolves away, and the
yellow fluid accumulating in the ducts passes into the mid-gut. The yellow colour is due to the globules of fat. In the young cells, at the summit of the cæca, these are either absent, or very small, whence the part appears colourless. But, lower down, small yellow granules appear in the cells, and these become bigger and more numerous in the middle and lower parts. In fact, few glands are better fitted for the study of the manner in which secretion is effected than the crayfish's liver.
We may now consider the alimentary machinery, the general structure of which has been explained, in action.
The food, already torn and crushed by the jaws, is passed through the gullet into the cardiac sac, and there reduced to a still more pulpy state by the gastric mill. By degrees, such parts as are sufficiently fluid are drained off into the intestine, through the pyloric strainer, while the coarser parts of the useless matters are probably rejected by the mouth, as a hawk or an owl rejects his casts. There is reason to believe, though it is not certainly known, that fluids from the intestine mix with the food while it is undergoing trituration, and effect the transformation of the starchy and the insoluble protein compounds into a soluble state. At any rate, as soon as the strained-off fluid passes into the mid-gut it must be mixed with the secretion of the liver, the action of which is probably
similar to that of the pancreatic juice of the higher animals.
The mixture thus produced, which answers to the chyle of the higher animals, passes along the intestine, and the greater part of it, transuding through the walls of the alimentary canal, enters the blood, while the rest accumulates as dark coloured fæces in the hind gut, and is eventually passed out of the body by the vent. The fæcal matters are small in amount, and the strainer is so efficient that they rarely contain solid particles of sensible size. Sometimes, however, there are a good many minute fragments of vegetable tissue.
[Figure 14: Astacus fluviatilis--The corpuscles of the blood]
The blood of which the nutritive elements of the food
have thus become integral parts, is a clear fluid, either colourless, or of a pale neutral tint or reddish hue, which, to the naked eye, appears like so much water. But if subjected to microscopic examination, it is found to contain innumerable pale, solid particles, or corpuscles, which, when examined fresh, undergo constant changes of form (fig. 14). In fact, they correspond very closely with the colourless corpuscles which exist in our own blood; and, in its general characters, the crayfish's blood is such as ours would be if it were somewhat diluted and deprived of its red corpuscles. In other words, it resembles our lymph more than it does our blood. Left to itself it soon coagulates, giving rise to a pretty firm clot.
The sinuses, or cavities in which the greater part of the blood is contained, are disposed very irregularly in the intervals between the internal organs. But there is one of especially large size on the ventral or sternal side of the thorax (fig. 15, sc), into which all the blood in the body sooner or later makes its way. From this sternal sinus passages (av) lead to the gills, and from these again six canals (bcv), pass up on the inner side of the inner wall of each branchial chamber to a cavity situated in the dorsal region of the thorax, termed the pericardium (p), into which they open.
The blood of the crayfish is kept in a state of constant circulating motion by a pumping and distributing machinery, composed of the heart and of the arteries, with
their larger and smaller branches, which proceed from it and ramify through the body, to terminate eventually in the blood sinuses, which represent the veins of the higher animals.
[Figure 15: Astacus fluviatilis--A diagrammatic transverse section of the thorax through the twelfth somite, showing the course of the circulation of the blood]
When the carapace is removed from the middle of the region which lies behind the cervical groove, that is, when the dorsal or tergal wall of the thorax is taken away, a spacious chamber is laid open which is full of blood. This is the cavity already mentioned as the pericardium (fig. 15, p), though, as it differs in some respects from that which is so named in the higher animals, it will be better to term it the pericardial sinus.
[Figure 16: Astacus fluviatilis--The heart]
The heart (fig. 15, h), lies in the midst of this sinus. It is a thick muscular body (fig. 16), with an irregularly hexagonal contour when viewed from above, one angle of the hexagon being anterior and another posterior. The lateral angles of the hexagon are connected by bands of fibrous tissue (ac) with the walls of the pericardial sinus. Otherwise, the heart is free, except in so far as it is kept in place by the arteries which leave it and traverse the walls of the pericardium. One of these arteries (figs. 5, 12, and 16, saa), starting from the hinder part of the heart, of which it is a sort of continuation, runs along the middle line of the abdomen above the intestine, to which it gives off many branches. A second large artery starts from a dilatation, which is common to it with the foregoing, but passing directly downwards (figs. 12 and 15, sa, and fig. 16, st. a), either on the right or on the left side of the intestine,
traverses the nervous cord (figs. 12 and 15), and divides into an anterior (fig. 12, sa) and a posterior (iaa) branch, both of which run beneath and parallel with that cord. A third artery runs, from the front part of the heart, forwards in the middle line, over the stomach, to the eyes and fore part of the head (figs. 5, 12, and 16, oa); and two others diverge one on each side of this, and sweep round the stomach to the antennæ (aa). Behind these, yet two other arteries are given off from the under side of the heart, and supply the liver (ha). All these arteries branch out and eventually terminate in fine, so-called capillary, ramifications.
In the dorsal wall of the heart two small oval apertures are visible, provided with valvular lips (fig. 16, sa), which open inwards, or towards the internal cavity of the heart. There is a similar aperture in each of the lateral faces of the heart (la), and two others in its inferior face (ia), making six in all. These apertures readily admit fluid into the heart, but oppose its exit. On the other hand, at the origins of the arteries, there are small valvular folds, directed in such a manner as to permit the exit of fluid from the heart, while they prevent its entrance.
The walls of the heart are muscular, and, during life, they contract at intervals with a regular rhythm, in such a manner as to diminish the capacity of the internal cavity of the organ. The result is, that the blood which it contains is driven into the arteries, and necessarily forces into their smaller ramifications an equivalent amount of the blood which they already contained; whence, in the long run, the same amount of blood passes out of the ultimate capillaries into the blood sinuses. From the disposition of the blood sinuses, the impulse thus given to the blood which they contain is finally conveyed to the blood in the branchiæ, and a proportional quantity of that
blood leaves the branchiæ and passes into the sinuses which connect them with the pericardial sinus (fig. 15, bcv), and thence into that cavity. At the end of the contraction, or systole, of the heart, its volume is of course diminished by the volume of the blood forced out, and the space between the walls of the heart and those of the pericardial sinus is increased to the same extent. This space, however, is at once occupied by the blood from the branchiæ, and perhaps by some blood which has not passed through the branchiæ, though this is doubtful. When the systole is over, the diastole follows; that it to say, the elasticity of the walls of the heart and that of the various parts which connect it with the walls of the pericardium, bring it back to its former size, and the blood in the pericardial sinus flows into its cavity by the six apertures. With a new systole the same process is repeated, and thus the blood is driven in a circular course through all parts of the body.
It will be observed that the branchiæ are placed in the course of the current of blood which is returning to the heart; which is the exact contrary of what happens in fishes, in which the blood is sent from the heart to the branchiæ, on its way to the body. It follows, from this arrangement, that the blood which goes to the branchiæ is blood in which the quantity of oxygen has undergone a diminution, and that of carbonic acid an increase, as compared with the blood in the heart itself. For the
activity of all the organs, and especially of the muscles, is inseparably connected with the absorption of oxygen and the evolution of carbonic acid; and the only source from which the one can be derived, and the only receptacle into which the other can be poured, is the blood which bathes and permeates the whole fabric to which it is distributed by the arteries.
The blood, therefore, which reaches the branchiæ has lost oxygen and gained carbonic acid; and these organs constitute the apparatus for the elimination of the injurious gas from the economy on the one hand, and, on the other, for the taking in of a new supply of the needful "vital air," as the old chemists called it. It is thus that the branchiæ subserve the respiratory function.
The crayfish has eighteen perfect and two rudimentary branchiæ in each branchial chamber, the boundaries of which have been already described.
[Figure 17: Astacus fluviatilis--Podobranchiæ]
Of the eighteen perfect branchiæ, six (podobranchiæ) are attached to the basal joints of the thoracic limbs, from the last but one to the second (second maxillipede) inclusively (fig. 4, p.26, pdb, and fig. 17, A, B); and eleven (arthrobranchiæ) are fixed to the flexible interarticular membranes, which connect these basal joints with the parts of the thorax to which they are articulated (fig. 4, arb, arb', fig. 17, C). Of these eleven branchiæ, two are attached to the interarticular membranes of all the ambulatory legs but the last, (=6) and to those of the pincers and of the external maxillipedes, (=4) and one to that of the
second maxillipede. The first maxillipede and the last ambulatory limb have none. Moreover, where there are two arthrobranchiæ, one is more or less in front of and external to the other.
These eleven arthrobranchiæ are all very similar in structure (fig. 17, C). Each consists of a stem which contains two canals, one external and one internal, separated by a longitudinal partition. The stem is beset with a great number of delicate branchial filaments, so that it looks like a plume tapering from its base to its summit. Each filament is traversed by large vascular channels, which break up into a net-work immediately beneath the surface. The blood driven into the external canals of the stem (fig. 15, av) is eventually poured into the inner canal (ev), which again communicates with the channels (bcv) which lead to the pericardial sinus (p). In its course, the blood traverses the branchial filaments, the outer investment of each of which is an excessively thin chitinous membrane, so that the blood contained in them is practically separated by a mere film from the aërated water in which the gills float. Hence, an exchange of gaseous constituents readily takes place, and as much oxygen is taken in as carbonic acid is given out.
The six podobranchiæ, or gills which are attached to the basal joints of the legs, play the same part, but differ a good deal in the details of their structure from those which are fixed to the interarticular membranes. Each consists of a broad base (fig. 17, A and B; b) beset with many
fine straight hairs, or setæ (F), whence a narrow stem (st) proceeds. At its upper end this stem divides into two parts, that in front, the plume (pl), resembling the free end of one of the gills just described, while that behind, the lamina (l), is a broad thin plate, bent upon itself longitudinally in such a manner that its folded edge lies forwards, and covered with minute hooked setæ (G). The gill which follows is received into the space included between the two lobes or halves of the folded lamina (fig. 4, p. 26). Each lobe is longitudinally plaited into about a dozen folds. The whole front and outer face of the stem is beset with branchial filaments; hence, we may compare one of these branchiæ to one of the preceding kind, in which the stem has become modified and has given off a large folded lamina from its inner and posterior face.
The branchiæ now described are arranged in sets of three for each of the thoracic limbs, from the third maxillipede to the last but one ambulatory limb, and two for the second maxillipede, thus making seventeen in all (3 x 5 + 2 = 17); and, between every two there is found a bundle of long twisted hairs (fig. 17, A, cx.s; D and E), which are attached to a small elevation (t) on the basal joint of each limb. These coxopoditic setæ no doubt, serve to prevent the intrusion of parasites and other foreign matters into the branchial chamber. From the mode of attachment of the six branchiæ it is obvious that they must share in the movements of the basal joints of the
legs; and that, when the crayfish walks, they must be more or less agitated in the branchial chamber.
The eighteenth branchia resembles one of the eleven arthrobranchiæ in structure; but it is larger, and it is attached neither to the basal joint of the hindermost ambulatory limb, nor to its interarticular membrane, but to the sides of the thorax, above the joint. From this mode of attachment it is distinguished from the others as a pleurobranchia (fig. 4, plb. 14).
Finally, in front of this, fixed also to the walls of the thorax, above each of the two preceding pairs of ambulatory limbs, there is a delicate filament, about a sixteenth of an inch long, which has the structure of a branchial filament, and is, in fact, a rudimentary pleurobranchia (fig. 4, plb. 12, plb. 13).
The quantity of water which occupies the space left in the branchial chamber by the gills is but small, and as the respiratory surface offered by the gills is relatively very large, the air contained in this water must be rapidly exhausted, even when the crayfish is quiescent; while, when any muscular exertion takes place, the quantity of carbonic acid formed, and the demand for fresh oxygen, is at once greatly increased. For the efficient performance of the function of respiration, therefore, the water in the branchial chamber must be rapidly renewed, and there must be some arrangement by which the supply of fresh water may be proportioned to the demand. In many animals, the respiratory surface is
covered with rapidly vibrating filaments, or cilia, by means of which a current of water is kept continually flowing over the gills, but there are none of these in the crayfish. The same object is attained, however, in another way. The anterior boundary of the branchial chamber corresponds with the cervical groove, which, as has been seen, curves downwards and then forwards, until it terminates at the sides of the space occupied by the jaws. If the branchiostegite is cut away along the groove, it will be found that it is attached to the sides of the head, which project a little beyond the anterior part of the thorax, so that there is a depression behind the sides of the head--just as there is a depression, behind a man's jaw, at the sides of the neck. Between this depression in front, the walls of the thorax internally, the branchiostegite externally, and the bases of the forceps and external foot-jaws below, a curved canal is included, by which the branchial cavity opens forwards as by a funnel. Attached to the base of the second maxilla there is a wide curved plate (fig. 4, 6) which fits against the projection of the head, as a shirt collar might do, to carry out our previous comparison; and this scoop-shaped plate (termed the scaphognathite), which is concave forwards and convex backwards, can be readily moved backwards and forwards.
If a living crayfish is taken out of the water, it will be found that, as the water drains away from the branchial cavity, bubbles of air are forced out of its anterior opening.
Again, if, when a crayfish is resting quietly in the water, a little coloured fluid is allowed to run down towards the posterior opening of the branchial chamber, it will very soon be driven out from the anterior aperture, with considerable force, in a long stream. In fact, as the scaphognathite vibrates not less than three or four times in a second, the water in the funnel-shaped front passage of the branchial cavity is incessantly baled out; and, as fresh water flows in from behind to make up the loss, a current is kept constantly flowing over the gills. The rapidity of this current naturally depends on the more or less quick repetition of the strokes of the scaphognathite; and hence, the activity of the respiratory function can be accurately adjusted to the wants of the economy. Slow working of the scaphognathite answers to ordinary breathing in ourselves, quick working to panting.
A further self-adjustment of the respiratory apparatus is gained by the attachment of the six gills to the basal joints of the legs. For, when the animal exerts its muscles in walking, these gills are agitated, and thus not only bring their own surfaces more largely in contact with the water, but produce the same effect upon the other gills. [see End note 8]
The constant oxidation which goes on in all parts of the body not only gives rise to carbonic acid; but, so far as it affects the proteinaceous constituents, it produces
compounds which contain nitrogen, and these, like other waste products, must be eliminated. In the higher animals, such waste products take the form of urea, uric acid, hippuric acid, and the like; and are got rid of by the kidneys. We may, therefore, expect to find some organ which plays the part of a kidney in the crayfish; but the position of the structure, which there is much reason for regarding as the representative of the kidney, is so singular that very different interpretations have been put upon it.
[Figure 18: Astacus fluviatilis--Anterior part of the body, stomach, eosophagus, green gland]
On the basal joint of each antenna it is easy to see a small conical eminence with an opening on the inner side of its summit (fig. 18). The aperture (x) leads by a short canal into a spacious sac, with extremely delicate walls (s), which is lodged in the front part of the head, in front of and below the cardiac division of the stomach (cs). Beneath this, in a sort of recess, which corresponds with the epistoma, and with the base of the antenna, there is a discoidal body of a dull green colour, in shape somewhat like one of the fruits of the mallow, which is known as the green gland (gg). The sac narrows below like a wide funnel, and the edges of its small end are continuous with the walls of the green gland; they surround an aperture which leads into the interior of the latter structure, and conveys its products into the sac, whence they are excreted by the opening in the antennary papilla. The green gland is said to contain a substance termed guanin (so named because it is found in the guano which is the accumulated
excrement of birds), a nitrogenous body analogous in some respects to uric acid, but less highly oxidated and if this be the case, there can be little doubt that the green gland represents the kidney, and its secretion the urinary fluid, while the sac is a sort of urinary bladder. [see End note 9]
Restricting our attention to the phenomena which have now been described, and to a short period in the life of the crayfish, the body of the animal may be regarded as a factory, provided with various pieces of machinery, by means of which certain nitrogenous and other matters are extracted from the animal and vegetable substances which serve for food, are oxidated, and are then delivered out of the factory in the shape of carbonic acid gas, guanin, and probably some other products, with which we are at present unacquainted. And there is no doubt, that if the total amount of products given out could be accurately weighed against the total amount of materials taken in, the weight of the two would be found to be identical. To put the matter in its most general shape, the body of the crayfish is a sort of focus to which certain material particles converge, in which they move for a time, and from which they are afterwards expelled in new combinations. The parallel between a whirlpool in a stream and a living being, which has often been drawn, is as just as it is striking. The whirlpool is permanent, but the particles of water which constitute it are incessantly changing. Those which enter it, on the one side, are whirled around and temporarily constitute a part of its individuality; and as they leave it on the other side, their places are made good by new comers.
Those who have seen the wonderful whirlpool, three miles below the Fails of Niagara, will not have forgotten the heaped-up wave which tumbles and tosses, a very embodiment of restless energy, where the swift stream hurrying from the Falls is compelled to make a sudden turn towards Lake Ontario. However changeful in the contour of its crest, this wave has been visible, approximately in the same place, and with the same general form, for centuries past. Seen from a mile off, it would appear to be a stationary hillock of water. Viewed closely, it is a typical expression of the conflicting impulses generated by a swift rush of material particles.
Now, with all our appliances, we cannot get within a good many miles, so to speak, of the crayfish. If we could, we should see that it was nothing but the constant form of a similar turmoil of material molecules which are constantly flowing into the animal on the one side, and streaming out on the other.
The chemical changes which take place in the body of the crayfish, are doubtless, like other chemical changes, accompanied by the evolution of heat. But the amount of heat thus generated is so small and, in consequence of the conditions under which the crayfish lives, it is so easily carried away, that it is practically insensible. The crayfish has approximately the temperature of the surrounding medium, and it is, therefore, reckoned among the cold-blooded animals.
If our investigation of the results of the process of
alimentation in a well-fed Crayfish were extended over a longer time, say a year or two, we should find that the products given out were no longer equal to the materials taken in, and the balance would be found in the increase of the animal's weight. If we inquired how the balance was distributed, we should find it partly in store, chiefly in the shape of fat; while, in part, it had been spent in increasing the plant and in enlarging the factory. That is to say, it would have supplied the material for the animal's growth. And this is one of the most remarkable respects in which the living factory differs from those which we construct. It not only enlarges itself, but, as we have seen, it is capable of executing its own repairs to a very considerable extent.
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