The Physiology of the Crayfish 
--THE MECHANISM BY WHICH THE LIVING ORGANISM ADJUSTS ITSELF TO SURROUNDING CONDITIONS AND REPRODUCES ITSELF.
IF the hand is brought near a vigorous crayfish, free to move in a large vessel of water, it will generally give a vigorous flap with its tail, and dart backwards out of reach; but if a piece of meat is gently lowered into the vessel, the crayfish will sooner or later approach and devour it.
If we ask why the crayfish behaves in this fashion, every one has an answer ready. In the first case, it is said that the animal is aware of danger, and therefore hastens away; in the second, that it knows that meat is good to eat, and therefore walks towards it and makes a meal. And nothing can seem to be simpler or more satisfactory than these replies, until we attempt to conceive clearly what they mean; and, then, the explanation, however simple it may be admitted to be, hardly retains its satisfactory character.
For example, when we say that the crayfish is "aware of danger," or "knows that meat is good to eat," what
do we mean by "being aware" and "knowing"? Certainly it cannot be meant that the crayfish says to himself, as we do, "This is dangerous," "That is nice;" for the crayfish, being devoid of language, has nothing to say either to himself or any one else. And if the Crayfish has not language enough to construct a proposition, it is obviously out of the question that his actions should be guided by a logical reasoning process, such as that by which a man would justify similar actions. The crayfish assuredly does not first frame the syllogism, "Dangerous things are to be avoided; that hand is dangerous; therefore it is to be avoided;" and then act upon the conclusion thus logically drawn.
But it may be said that children, before they acquire the use of language, and we ourselves, long after we are familiar with conscious reasoning, perform a great variety of perfectly rational acts unconsciously. A child grasps at a sweetmeat, or cowers before a threatening gesture, before it can speak; and any one of us would start back from a chasm opening at our feet, or stoop to pick up a jewel from the ground, "without thinking about it." And, no doubt, if the crayfish has any mind at all, his mental operations must more or less resemble those which the human mind performs without giving them a spoken or unspoken verbal embodiment.
If we analyse these, we shall find that, in many cases, distinctly felt sensations are followed by a distinct desire to perform some act, which act is accordingly performed;
while, in other cases, the act follows the sensation without one being aware of any other mental process; and, in yet others, there is no consciousness even of the sensation. As I wrote these last words, for example, I had not the slightest consciousness of any sensation of holding or guiding the pen, although my fingers were causing that instrument to perform exceedingly complicated movements. Moreover, experiments upon animals have proved that consciousness is wholly unnecessary to the carrying out of many of those combined movements by which the body is adjusted to varying external conditions.
Under these circumstances, it is really quite an open question whether a crayfish has a mind or not; moreover, the problem is an absolutely insoluble one, inasmuch as nothing short of being a crayfish would give us positive assurance that such an animal possesses consciousness;. and, finally, supposing the crayfish has a mind, that fact does not explain its acts, but only shows that, in the course of their accomplishment, they are accompanied by phenomena similar to those of which we are aware in ourselves, under like circumstances.
So we may as well leave this question of the crayfish's mind on one side for the present, and turn to a more profitable investigation, namely, that of the order and connexion of the physical phenomena which intervene between something which happens in the neighbourhood of the animal and that other something which responds to it, as an act of the crayfish.
Whatever else it may be, this animal, so far as it is acted upon by bodies around it and reacts on them, is a piece of mechanism, the internal works of which give rise to certain movements when it is affected by particular external conditions; and they do this in virtue of their physical properties and connexions.
Every movement of the body, or of any organ of the body, is an effect of one and the same cause, namely, muscular contraction. Whether the crayfish swims or walks, or moves its antennæ, or seizes its prey, the immediate cause of the movements of the parts which bring about, or constitute, these bodily motions is to be sought in a change which takes place in the flesh, or muscle, which is attached to them. The change of place which constitutes any movement is an effect of a previous change in the disposition of the molecules of one or more muscles; while the direction of that movement depends on the connexions of the parts of the skeleton with one another, and of the muscles with them.
The muscle of the crayfish is a dense, white substance; and if a small portion of it is subjected to examination it will be found to be very easily broken up into more or less parallel bundles of fine fibres. Each of these fibres is generally found to be ensheathed in a fine transparent membrane, which is called the sarcolemma, within which is contained the proper substance of the muscle. When quite fresh and living, this substance is soft and
semi-fluid, but it hardens and becomes solid immediately after death.
Examined, with high magnifying powers, in this
condition, the muscle-substance appears marked by very regular transverse bands, which are alternately opaque and transparent; and it is characteristic of the group of animals to which the crayfish belongs that their muscle-substance has this striped character in all parts of the body.
[Figure 19: Astacus fluviatilis--Muscular fibres]
A greater or less number of these fibres, united into one or more bundles, constitutes a muscle; and, except when these muscles surround a cavity, they are fixed at each end to the hard parts of the skeleton. The attachment is frequently effected by the intermediation of a dense, fibrous, often chitinous, substance, which constitutes the tendon (fig. 19, A; t) of the muscle.
The property of the living muscle, which enables it to be the cause of motion, is this: Every muscular fibre is capable of suddenly changing its dimensions, in such a manner that it shortens and becomes proportionately thicker. Hence the absolute bulk of the fibre remains practically unchanged. From this circumstance, muscular contraction, as the change of form of a muscle is called, is radically different from the process which commonly goes by the same name in other things, and which involves a diminution of bulk.
The contraction of muscle takes place with great force, and, of course, if the parts to which its ends are fixed are both free to move, they are brought nearer at the moment of contraction: if one only is free to move that is approximated to the fixed part; and if the muscular fibre
surrounds a cavity, the cavity is lessened when the muscle contracts. This is the whole source of motor power in the crayfish machine. The results produced by the exertion of that power depend upon the manner in which the parts to which the muscles are attached are connected with one another.
[Figure 20: Astacus fluviatilis--The chela of the forceps]
One example of this has already been given in the curious mechanism of the gastric mill. Another may be found in the chela which terminates the forceps. If the
articulation of the last joint (fig. 20, dp) with the one which precedes it (prp) is examined, it will be found that the base of the terminal segment (dp) turns on two hinges (x), formed by the hard exoskeleton and situated at opposite points of the diameter of the base, on the penultimate segment; and these hinges are so disposed that the last joint can be moved only in one plane, to or from the produced angle of the penultimate segment (prp), which forms the fixed claw of the chela. Between the hinges, on both the inner and the outer sides of the articulation, the exoskeleton is soft and flexible, and allows the terminal segment to play easily through a certain arc. It is by this arrangement that the direction and the extent of the motion of the free claw of the chela are determined. The source of the motion lies in the muscles which occupy the interior of the enlarged penultimate segment of the limb. Two muscles, one of very great size (m), the other smaller (m'), are fastened by one end to the exoskeleton of this segment. The fibres of the larger muscle converge to be fixed into the two sides of a long flat process of the chitinous cuticula, on the inner side of the base of the terminal segment, which serves as a tendon (t); while those of the smaller muscle are similarly attached to a like process which proceeds from the outer side of the base of the terminal segment (t'). It is obvious that, when the latter muscle shortens it must move the apex of the terminal segment (dp) away from the end of the fixed claw; while, when the former contracts, the end of the terminal segment is brought towards that of the fixed claw.
A living crayfish is able to perform very varied movements with its pincers. When it swims backwards, these limbs are stretched straight out, parallel with one another, in front of the head; when it walks, they are usually carried like arms bent at the elbow, the "forearm" partly resting on the ground; on being irritated, the crayfish sweeps the pincers round in any direction to grasp the offending body; when prey is seized, it is at once conveyed, with a circular motion, towards the region of the mouth. Nevertheless, these very varied actions are all brought about by a combination of simple flexions and extensions, each of which is effected in the exact order, and to the exact extent, needful to bring the chela into the position required.
The skeleton of the stem of the limb which bears the chela is, in fact, divided into four moveable segments; and each of these is articulated with the segments on each side of it by a hinge of just the same character as that which connects the moveable claw of the chela with the penultimate segment, while the basal segment is similarly articulated with the thorax.
If the axes of all these articulations [note 1] were parallel, it is obvious that, though the limb might be moved as a whole through a considerable arc, and might be bent in various
degrees, yet all its movements would be limited to one plane. But, in fact, the axes of the successive articulations are nearly at right angles to one another; so that, if the segments are successively either extended or flexed, the chela describes a very complicated curve; and by varying the extent of flexion or extension of each segment, this curve is susceptible of endless variation. It would probably puzzle a good mathematician to say exactly what position should be given to each segment, in order to bring the chela from any given position into any other; but if a lively crayfish is incautiously seized, the experimenter will find, to his cost, that the animal solves the problem both rapidly and accurately.
[Figure 21: Astacus fluviatilis--Two of the abdominal somites]
The mechanism by which the retrograde swimming of the crayfish is effected, is no less easily analysed. The apparatus of motion is, as we have seen, the abdomen, with its terminal five-pointed flapper. The rings of the abdomen are articulated together by joints (fig. 21, x) situated a little below the middle of the height of the rings, at opposite ends of transverse lines, at right angles to the long axis of the abdomen.
Each ring consists of a dorsal, arched portion, called the tergum (fig. 21; fig. 36, p.142, t. XIX), and a nearly flat ventral portion, which is the sternum (fig. 36, st. XIX). Where these two join, a broad plate is sent down on each side, which overlaps the bases of the abdominal appendages, and is known as the pleuron (fig. 36, pl. XIX).
The sterna are all very narrow, and are connected together by wide spaces of flexible exoskeleton.
When the abdomen is made straight, it will be found that these intersternal membranes are stretched as far as they will yield. On the other hand, when the abdomen is bent up as far as it will go, the sterna come close together, and the intersternal membranes are folded.
The terga are very broad; so broad, in fact, that each overlaps its successor, when the abdomen is straightened or extended, for nearly half its length in the middle line; and the overlapped surface is smooth, convex, and
marked off by a transverse groove from the rest of the tergum as an articular facet. The front edge of the articular facet is continued into a sheet of flexible cuticula, which turns back, and passes as a loose fold to the hinder edge of the overlapping tergum (fig. 21). This tergal interarticular membrane allows the terga to move as far as they can go in flexion; whilst, in extreme extension, they are but slightly stretched. But, even if the intersternal membranes presented no obstacle to excessive extension of the abdomen, the posterior free edge of each tergum fits into the groove behind the facet in the next in such a manner, that the abdomen cannot be made more than very slightly concave upwards without breaking.
Thus the limits of motion of the abdomen, in the vertical direction, are from the position in which it is straight, or has even a very slight upward concavity, to that in which it is completely bent upon itself, the telson being brought under the bases of the hinder thoracic limbs. No lateral movement between the somites of the abdomen is possible in any of its positions. For, when it is straight, lateral movement is hindered not only by the extensive overlapping of the terga, but also by the manner in which the hinder edges of the pleura of each of the four middle somites overlap the front edges of their successors. The pleura of the second somite are much larger than any of the others, and their front edges overlap the small pleura of the first abdominal somite; and when the abdomen is much flexed, these pleura even
ride over the posterior edges of the branchiostegites. In the position of extension, the overlap of the terga is great, while that of the pleura of the middle somites is small. As the abdomen passes from extension to flexion, the overlap of the terga of course diminishes; but any decrease of resistance to lateral strains which may thus arise, is compensated by the increasing overlap of the pleura, which reaches its maximum when the abdomen is completely flexed.
It is obvious that longitudinal muscular fibres fixed into the exoskeleton, above the axes of the joints, must bring the centres of the terga of the somites closer together, when they contract; while muscular fibres attached below the axes of the joints must approximate the sterna. Hence, the former will give. rise to extension, and the latter to flexion, of the abdomen as a whole.
[Figure 22: Astacus fluviatilis--A longitudinal section of the body to show the principal muscles and their relations to the exoskeleton]
Now there are two pairs of very considerable muscles disposed in this manner. The dorsal pair, or the extensors of the abdomen (fig. 22, e.m), are attached in front to the side walls of the thorax, thence pass backwards into the abdomen, and divide into bundles, which are fixed to the inner surfaces of the terga of all the somites. The other pair, or the flexors of the abdomen (f.m) constitute a very much larger mass of muscle, the fibres of which are curiously twisted, like the strands of a rope. The front end of this double rope is fixed to a series of processes of the exoskeleton of the thorax, called apodemata, some of which roof over the sternal blood-sinuses
and the thoracic part of the nervous system; while, in the abdomen, its strands are attached to the sternal exoskeleton of all the somites and extend, on each side of the rectum, to the telson.
When the exoskeleton is cleaned by maceration, the abdomen has a slight curve, dependent upon the form and the degree of elasticity possessed by its different parts; and, in a living crayfish at rest, it will be observed that the curvature of the abdomen is still more marked. Hence it is ready either for extension or for flexion.
A sudden contraction of the flexor muscles instantly increases the ventral curvature of the abdomen, and
throws the tail fin, the two side lobes of which are spread out, forwards; while the body is propelled backwards by the reaction of the water against the stroke. Then the flexor muscles being relaxed, the extensor muscles come into play; the abdomen is straightened, but less violently and with a far weaker stroke on the water, in consequence of the less strength of the extensors and of the folding up of the lateral plates of the fin, until it comes into the position requisite to give full force to a new downward and forward stroke. The tendency of the extension of the abdomen is to drive the body forward; but from the comparative weakness and the obliquity of its stroke, its practical effect is little more than to check the backward motion conferred upon the body by the flexion of the abdomen.
[Figure 23: Astacus fluviatilis--Three nerve fibres, with the connective tissue in which they are imbedded]
Thus, every action of the crayfish, which involves motion, means the contraction of one or more muscles. But what sets muscle contracting? A muscle freshly removed from the body may be made to contract in various ways, as by mechanical or chemical irritation, or by an electrical shock; but, under natural conditions, there is only one cause of muscular contraction, and that is the activity of a nerve. Every muscle is supplied with one or more nerves. These are delicate threads which, on microscopic examination, prove to be bundles of fine tubular filaments, filled with an apparently structureless substance of gelatinous consistency, the nerve fibres
(fig. 23). The nerve bundle which passes to a muscle breaks up into smaller bundles and, finally, into separate fibres, each of which ultimately terminates by becoming continuous with the substance of a muscular fibre (fig. 19, F.) Now the peculiarity of a muscle nerve, or motor nerve, as it is called, is that irritation of the nerve fibre at any part of its length, however distant from the muscle, brings about muscular contraction, just as if the muscle itself were irritated. A change is produced in the molecular condition of the nerve at the point of irritation; and this change is propagated along the nerve, until it reaches the muscle, in which it gives rise to that change in the arrangement of its molecules, the most obvious effect of which is the sudden alteration of form which we call muscular contraction.
[Figure 24: Astacus fluviatilis--Abdominal ganglia and ganglionic corpuscles]
If we follow the course of the motor nerves in a
direction away from the muscles to which they are distributed, they will be found, sooner or later, to terminate in ganglia (fig. 24 A. gl.c; fig. 25, gn. 1-13.) A ganglion is a body which is in great measure composed of nerve fibres; but, interspersed among these, or disposed around them, there are peculiar structures, which are termed ganglionic corpuscles, or nerve cells (fig. 24, B.) These are nucleated cells, not unlike the epithelial cells which have been already mentioned, but which are larger and often give off one or more processes. These processes, under favourable circumstances, can be traced into continuity with nerve fibres.
[Figure 25: Astacus fluviatilis--The central nervous system seen from above]
The chief ganglia of the crayfish are disposed in a longitudinal series in the middle line of the ventral aspect of the body close to the integument (fig. 25). In the abdomen, for example, six ganglionic masses are readily observed, one lying over the sternum of each somite, connected by longitudinal bands of nerve fibres, and giving off branches to the muscles. On careful examination, the longitudinal connecting bands, or commissures (fig. 24, co), are seen to be double, and each mass appears slightly bilobed. In the thorax, there are six, larger, double ganglionic masses, likewise connected by double commissures; and the most anterior of these, which is the largest (fig. 25, gn. 2), is marked at the sides by notches, as if it were made up of several pairs of ganglia, run together into one continuous whole. The front of this, two commissures (c) pass forwards, separating widely, to give room for the gullet (oes), which passes between them; while in front of the gullet, just behind the eyes, they unite with a transversely elongated mass of ganglionic substance (gn. 1), termed the brain, or cerebral ganglion. [see End note 10]
All the motor nerves, as has been said, are traceable, directly or indirectly, to one or other of these thirteen sets of ganglia; but other nerves are given off from the ganglia, which cannot be followed into any muscle. In
fact, these nerves go either to the integument or to the organs of sense, and they are termed sensory nerves.
When a muscle is connected by its motor nerve with a ganglion, irritation of that ganglion will bring about the contraction of the muscle, as well as if the motor nerve itself were irritated. Not only so; but if a sensory nerve, which is in connexion with the ganglion, is irritated, the same effect is produced; moreover, the sensory nerve itself need not be excited, but the same result will take place, if the organ to which it is distributed is stimulated. Thus the nervous system is fundamentally an apparatus by which two separate, and it may be distant, parts of the body, are brought into relation with one another; and this relation is of such a nature, that a change of state arising in the one part is followed by the propagation of changes along the sensory nerve to the ganglion, and from the ganglion to the other part; where, if that part happens to be muscle, it produces contraction. If one end of a rod of wood, twenty feet long, is applied to a sounding-board, the sound of a tuning-fork held against the opposite extremity will be very plainly heard. Nothing can be seen to happen in the wood, and yet its molecules are certainly set vibrating, at the same rate as the tuning-fork vibrates; and when, after travelling rapidly along the wood, these vibrations affect the sounding-board, they give rise to vibrations of the molecules of the air, which reaching the ear, are converted into an audible note. So in the nerve tract:
no apparent change is effected in it by the irritation at one end; but the rate at which the molecular change produced travels can be measured; and, when it reaches the muscle, its effect becomes visible in the change of form of the muscle. The molecular change would take place just as much if there were no muscle connected with the nerve, but it would be no more apparent to ordinary observation than the sound of the tuning-fork is audible in the absence of the sounding-board.
If the nervous system were a mere bundle of nerve fibres extending between sensory organs and muscles, every muscular contraction would require the stimulation of that special point of the surface on which the appropriate sensory nerve ended. The contraction of several muscles at the same time, that is, the combination of movements towards one end, would be possible only if the appropriate nerves were severally stimulated in the proper order, and every movement would be the direct result of external changes. The organism would be like a piano, which may be made to give out the most complicated harmonies, but is dependent for their production on the depression of a separate key for every note that is sounded. But it is obvious that the crayfish needs no such separate impulses for the performance of highly complicated actions. The simple impression made on the organs of sensation in the two examples with which we started, gives rise to a train of complicated and accurately co-ordinated muscular contractions. To carry the analogy
of the musical instrument further, striking a single key gives rise, not to a single note, but to a more or less elaborate tune; as if the hammer struck not a single string, but pressed down the stop of a musical box.
It is in the ganglia that we must look for the analogue of the musical box. A single impulse conveyed by a sensory nerve to a ganglion, may give rise to a single muscular contraction, but more commonly it originates a series of such, combined to a definite end.
The effect which results from the propagation of an impulse along a nerve fibre to a ganglionic centre, whence it is, as it were, reflected along another nerve fibre to a muscle, is what is termed a reflex action. As it is by no means necessary that sensation should be a concomitant of the first impulse, it is better to term the nerve fibre which carries it afferent rather than sensory; and, as other phenomena besides those of molar motion may be the ultimate result of the reflex action, it is better to term the nerve fibre which transmits the reflected impulse efferent rather than motor.
If the nervous commissures between the last thoracic and the first abdominal ganglia are cut, or if the thoracic ganglia are destroyed, the crayfish is no longer able to control the movements of the abdomen. If the forepart of the body is irritated, for example, the animal makes no effort to escape by swimming backwards. Nevertheless, the abdomen is not paralysed, for, if it be irritated, it will flap vigorously. This is a case of pure
reflex action. The stimulus is conveyed to the abdominal ganglia through afferent nerves, and is reflected from them, by efferent nerves, to the abdominal muscles.
But this is not all. Under these circumstances it will be seen that the abdominal limbs all swing backwards and forwards, simultaneously, with an even stroke; while the vent opens and shuts with a regular rhythm. Of course, these movements imply correspondingly regular alternate contractions and relaxations of certain sets of muscles; and these, again, imply regularly recurring efferent impulses from the abdominal ganglia. The fact that these impulses proceed from the abdominal ganglia, may be shown in two ways: first, by destroying these ganglia in one somite after another, when the movements in each somite at once permanently cease; and, secondly, by irritating the surface of the abdomen, when the movements are temporarily inhibited by the stimulation of the afferent nerves. Whether these movements are properly reflex, that is, arise from incessant new afferent impulses of unknown origin, or whether they depend on the periodical accumulation and discharge of nervous energy in the ganglia themselves, or upon periodical exhaustion and restoration of the irritability of the muscles, is unknown. It is sufficient for the present purpose to use the facts as evidence of the peculiar co-ordinative function of ganglia.
The crayfish, as we have seen, avoids light; and the slightest touch of one of its antennæ gives rise to active motions of the whole body. In fact, the animal's
position and movements are largely determined by the influences received through the feelers and the eyes. These receive their nerves from the cerebral ganglia; and, as might be expected, when these ganglia are extirpated, the crayfish exhibits no tendency to get away from the light, and the feelers may not only be touched, but sharply pinched, without effect. Clearly, therefore, the cerebral ganglia serve as a ganglionic centre, by which the afferent impulses derived from the feelers and the eyes are transmuted into efferent impulses. Another very curious result follows upon the extirpation of the cerebral ganglia. If an uninjured crayfish is placed upon its back, it makes unceasing and well-directed efforts to turn over; and if everything else fails, it will give a powerful flap with the abdomen, and trust to the chapter of accidents to turn over as it darts back. But the brainless crayfish behaves in a very different way. Its limbs are in incessant motion, but they are "all abroad;" and if it turns over on one side, it does not seem able to steady itself, but rolls on to its back again.
If anything is put between the chelæ of an uninjured crayfish, while on its back, it either rejects the object at once, or tries to make use of it for leverage to turn over. In the brainless crayfish a similar operation gives rise to a very curious spectacle. [note 2] If the object, whatever it
be--a bit of metal, or wood, or paper, or one of the animal's own antennulæ--is placed between the chelæ of the forceps, it is at once seized by them, and carried backwards; the chelate ambulatory limbs are at the same time advanced, the object seized is transferred to them, and they at once tuck it between the external maxillipedes, which, with the other jaws, begin vigorously to masticate it. Sometimes the morsel is swallowed; sometimes it passes out between the anterior jaws, as if deglutition were difficult. It is very singular to observe that, if the morsel which is being conveyed to the mouth by one of the forceps is pulled back, the forceps and the chelate ambulatory limbs of the other side are at once brought forward to secure it. The movements of the limbs are, in short, adjusted to meet the increased resistance. [see End note 11]
All these phenomena cease at once, if the thoracic ganglia are destroyed. It is in these, therefore, that the simple stimulus set up by the contact of a body with, for example, one of the forceps, is translated into all the surprisingly complex and accurately co-ordinated movements, which have been described. Thus the nervous system of the crayfish may be regarded as a system of co-ordinating mechanisms, each of which produces a certain action, or set of actions, on the receipt of an appropriate stimulus.
When the crayfish comes into the world, it possesses in its neuro-muscular apparatus certain innate
potentialities of action, and will exhibit the corresponding acts, under the influence of the appropriate stimuli. A large proportion of these stimuli come from without through the organs of the senses. The greater or less readiness of each sense organ to receive impulses, of the nerves to transmit them, and of the ganglia to give rise to combined impulses, is dependent at any moment upon the physical condition of these parts; and this, again, is largely modified by the amount and the condition of the blood supplied. On the other hand, a certain number of these stimuli are doubtless originated by changes within the various organs which compose the body, including the nerve centres themselves.
When an action arises from conditions developed in the interior of an animal's body, inasmuch as we cannot perceive the antecedent phenomena, we call such an action "spontaneous;" or, when in ourselves we are aware that it is accompanied by the idea of the action, and the desire to perform it, we term the act "voluntary." But, by the use of this language, no rational person intends to express the belief that such acts are uncaused or cause themselves. "Self-causation" is a contradiction in terms; and the notion that any phenomenon comes into existence without a cause, is equivalent to a belief in chance, which one may hope is, by this time, finally exploded.
In the crayfish, at any rate, there is not the slightest reason to doubt that every action has its definite physical
cause, and that what it does at any moment would be as clearly intelligible, if we only knew all the internal and external conditions of the case, as the striking of a clock is to any one who understands clockwork.
The adjustment of the body to varying external conditions, which is one of the chief results of the working of the nervous mechanism, would be far less important from a physiological point of view than it is, if only those external bodies which come into direct contact with the organism [note 3] could affect it; though very delicate influences of this kind take effect on the nervous apparatus through the integument.
It is probable that the setæ, or hairs, which are so generally scattered over the body and the appendages, are delicate tactile organs. They are hollow processes of the chitinous cuticle, and their cavities are continuous with narrow canals, which traverse the whole thickness of the cuticle, and are filled by a prolongation of the subjacent proper integument. As this is supplied with nerves, it is likely that fine nerve fibres reach the bases of the hairs, and are affected by anything which stirs these delicately poised levers.
There is much reason to believe that odorous bodies affect crayfish ; but it is very difficult to obtain experimental evidence of the fact. However, there is a good deal of analogical ground for the supposition that some peculiar structures, which are evidently of a sensory
nature, developed on the under side of the outer branch of the antennule, play the part of an olfactory apparatus.
[Figure 26: Astacus fluviatilis--Right antennule, olfactory appendages, auditory sac]
Both the outer (fig. 26 A. ex) and the inner (en) branches of the antennule are made up of a number of delicate ring-like segments, which bear fine setæ (b) of the ordinary character.
[Figure 27: Astacus fluviatilis--Auditory sac and hair]
The inner branch, which is the shorter of the two, possesses only these setæ; but the under surface of each of the joints of the outer branch, from about the seventh or eighth to the last but one, is provided with two bundles of very curious appendages (fig. 27, A, B, C, a), one in front and one behind. These are rather more than 1-200th of an inch long, very delicate, and shaped like a spatula, with a rounded handle and a flattened somewhat curved blade, the end of which is sometimes truncated, sometimes has the form of a prominent papilla. There is a sort of joint between the handle and the blade, such as is found between the basal and the terminal parts of the ordinary setæ, with which, in fact, these processes entirely correspond in their essential structure. A soft granular tissue fills the interior of each of these problematical structures, to which Leydig, their discoverer, ascribes an olfactory function.
It is probable that the crayfish possesses something analogous to taste, and a very likely seat for the organ of this function is in the upper lip and the metastoma; but if the organ exists it possesses no structural peculiarities by which it can be identified.
There is no doubt, however, as to the special recipients of sonorous and luminous vibrations; and these are of particular importance, as they enable the nervous machinery to be affected by bodies indefinitely remote from it, and to change the place of the organism in relation to such bodies.
Sonorous vibrations are enabled to act as the stimulants of a special nerve (fig. 25, a'n) connected with the brain, by means of the very curious auditory sacs (fig. 26, A, au) which are lodged in the basal joints of the antennules.
Each of these joints is trihedral, the outer face being convex; the inner, applied to its fellow, flat; and the upper, on which the eyestalk rests, concave. On this upper face there is a narrow elongated oval aperture, the outer lip of which is beset with a flat brush of long close-set setæ, which lie horizontally over the aperture, and effectually close it. The aperture leads into a small sac (au) with delicate walls formed by a chitinous continuation of the general cuticula. The inferior and posterior wall of the sac is raised up along a curved line into a ridge which projects into its interior (fig. 27, A, r). Each side of this ridge is beset with a series of delicate setæ (as), the longest of which measures about 1/50th of an inch; they thus form a longitudinal band bent upon itself. These auditory setæ project into the fluid contents of the sac, and their apices are for the most part imbedded in a gelatinous mass, which contains irregular particles of sand
and sometimes of other foreign matter. A nerve (n n',) is distributed to the sac, and its fibres enter the bases of the hairs, and may be traced to their apices, where they end in peculiar elongated rod-like bodies (fig. 27, C). Here is an auditory organ of the simplest description.
It retains, in fact, throughout life, the condition of a simple sac or involution of the integument, such as is that of the vertebrate ear in its earliest stage.
The sonorous vibrations transmitted through the water in which the crayfish lives to the fluid and solid contents of the auditory sac are taken up by the delicate hairs of the ridge, and give rise to molecular changes which traverse the auditory nerves and reach the cerebral ganglia.
The vibrations of the luminiferous ether are brought to bear upon the free ends of two large bundles of nerve fibres, termed the optic nerves (fig. 25, on), which proceed directly from the brain, by means of a highly complex eye. This is an apparatus, which, in part, sorts out the rays of light into as many very small pencils as there are separate endings of the fibres of the optic nerve, and, in part, serves as the medium by which the luminous vibrations are converted into molecular nerve changes.
[Figure 28: Astacus fluviatilis--Eye-stalk]
The free extremity of the eyestalk presents a convex, soft, and transparent surface, limited by an oval contour. The cuticle in this region, which is termed the cornea, (fig. 28, a), is, in fact, somewhat thinner and less distinctly laminated than in the rest of the eyestalk, and it contains no calcareous matter. But it is directly continuous with the rest of the exoskeleton of the eyestalk, to which it stands in somewhat the same relation as the soft integument of an articulation does to the adjacent hard parts.
The cornea is divided into a great number of minute, usually square facets, by faint lines, which cross it from side
to side nearly at right angles with one another. A longitudinal section shows that both the horizontal and the vertical contours of the cornea are very nearly semicircular, and that the lines which mark off the facets merely arise from a slight modification of its substance between the facets. The outer contour of each facet forms part of the general curvature of the outer face of the cornea; the inner contour sometimes exhibits a slight deviation from the general curvature of the inner face, but usually nearly coincides with it.
When a longitudinal or a transverse section is taken through the whole eyestalk, the optic nerve (fig. 28, A, op) is seen to traverse its centre. At first narrow and cylindrical, it expands towards its extremity into a sort of bulb (B, g), the outer surface of which is curved in correspondence with the inner surface of the cornea. The terminal half of the bulb contains a great quantity of dark colouring matter or pigment, and, in section, appears as what may be termed the inner dark zone (f). Outside this, and in connection with it, follows a white line, the inner white zone (e), then comes a middle dark zone (d); outside this an outer pale band, which may be called the outer white zone (e), and between this and the cornea (a) is another broad band of dark pigment, the outer dark zone (b).
When viewed under a low power, by reflected light, this outer dark zone is seen to be traversed by nearly parallel straight lines, each of which starts from the boundary between two facets, and can be followed inwards through the outer white zone to the middle dark zone. Thus the whole substance of the eye between the outer surface of the bulb of the optic nerve and the inner surface of the cornea is marked out into as many segments as the cornea has facets; and each segment has the form of a wedge or slender pyramid, the base of which is four-sided, and is applied against the inner surface of
one of the facets of the cornea, while its summit lies in the middle dark zone. Each of these visual pyramids consists of an axial structure, the visual rod, invested by a sheath. The latter extends inwards from the margin of each facet of the cornea, and contains pigment in two regions of its length, the intermediate space being devoid of pigment. As the position of the pigmented regions in relation to the length of the pyramid is always the same, the pigmented regions necessarily take the form of two consecutive zones when the pyramids are in their natural position.
The visual rod consists of two parts, an external crystalline cone (fig. 28, B, cr), and an internal striated spindle (sp). The crystalline cone consists of a transparent glassy-looking substance, which may be made to split up longitudinally into four segments. Its inner end narrows into a filament which traverses the outer white zone, and, in the middle dark zone, thickens into a four-sided spindle-shaped transparent body, which appears transversely striated. The inner end of this striated spindle narrows again, and becomes continuous with nerve fibres which proceed from the surface of the optic bulb.
The exact mode of connection of the nerve-fibres with the visual rods is not certainly made out, but it is probable that there is direct continuity of substance, and that each rod is really the termination of a nerve fibre.
Eyes having essentially the same structure as that of
the crayfish are very widely met with among Crustacea and Insecta, and are commonly known as compound eyes. In many of these animals, in fact, when the cornea is removed, each facet is found to act as a separate lens; and when proper arrangements are made, as many distinct pictures of external objects are found behind it as there are facets. Hence the notion suggested itself that each visual pyramid is a separate eye, similar in principle of construction to the human eye, and forming a picture of so much of the external world as comes within the range of its lens, upon a retina supposed to be spread out on the surface of the crystalline cone, as the human retina is spread over the surface of the vitreous humour.
But, in the first place, there is no evidence, nor any probability, that there is anything corresponding to a retina on the outer face of the crystalline cone; and secondly, if there were, it is incredible that, with such an arrangement of the refractive media as exists in the cornea and crystalline cones, rays proceeding from points in the external world should be brought to a focus in correspondingly related points of the surface of the supposed retina. But without this no picture could be formed, and no distinct vision could take place. It is very probable, therefore, that the visual pyramids do not play the part of the simple eyes of the Vertebrata, and the only alternative appears to be the adoption of a modification of the theory of mosaic vision, propounded many years by Johannes Müller.
[Figure 29:--Diagram showing the course of rays of light through the visual rods]
Each visual pyramid, isolated from its fellows by its coat of pigment, may be supposed, in fact, to play the part of a very narrow straight tube, with blackened walls, one end of which is turned towards the external world, while the other incloses the extremity of one of the nerve fibres. The only light which can reach the latter, under these circumstances, is such as proceeds from points which lie in the direction of a straight line represented by the produced axis of the tubes.
Suppose A-I to be nine such tubes, a-i the corresponding nerve fibres, and x y z three points from which light proceeds. Then it will be obvious that the only light
from x which will excite sensation, will be the ray which traverses B and reaches the nerve-fibre b, while that from y will affect only e, and that from x only h. The result, translated into sensation, will be three points of light on a dark ground, each of which answers to one of the luminous points, and indicates its direction in reference to the eye and its angular distance from the other two. [note 4]
The only modification needed in the original form of the theory of mosaic vision, is the supposition that part, or the whole, of the visual rod, is not merely a passive transmitter of light to the nerve-fibre, but is, itself, in someway concerned in transmuting the mode of motion, light, into that other mode of motion which we call nervous energy. The visual rod is, in fact, to be regarded as the physiological end of the nerve, and the instrument by which the conversion of the one form of motion into the other takes place; just as the auditory hairs are instruments by which the sonorous waves are converted into molecular movements of the substance of the auditory nerves. [see End note 12]
It is wonderfully interesting to observe that, when the so-called compound eye is interpreted in this manner,
the apparent wide difference between it and the vertebrate eye gives place to a fundamental resemblance. The rods and cones of the retina of the vertebrate eye are extraordinarily similar in their form and their relations to the fibres of the optic nerve, to the visual rods of the arthropod eye. And the morphological discrepancy, which is at first so striking, and which arises from the fact that the free ends of the visual rods are turned towards the light, while those of the rods and cones of the vertebrate eye are turned from it, becomes a confirmation of the parallel between the two when the development of the vertebrate eye is taken into account. For it is demonstrable that the deep surface of the retina in which the rods and cones lie, is really a part of the outer surface of the body turned inwards, in the course of the singular developmental changes which give rise to the brain and the eye of vertebrate animals.
Thus the crayfish has, at any rate, two of the higher sense organs, the ear and the eye, which we possess ourselves; and it may seem a superfluous, not to say a frivolous, question, if any one should ask whether it can hear and see.
But, in truth, the inquiry, if properly limited, is a very pertinent one. That the crayfish is led by the use of its eyes and ears to approach some objects and avoid others, is beyond all doubt; and, in this sense, most indubitably it can both hear and see. But it the question
means, do luminous vibrations give it the sensations of light and darkness, of colour and form and distance, which they give to us? and do sonorous vibrations produce the feelings of noise and tone, of melody and of harmony, as in us?--it is by no means to be answered hastily, perhaps cannot be answered at all, except in a tentative, probable way.
The phenomena to which we give the names of sound and colour are not physical things, but are states of consciousness, dependent, there is every reason to believe, on the functional activity of certain parts of our brains. Melody and harmony are names for states of consciousness which arise when at least two sensations of sound have been produced. All these are manufactured articles, products of the human brain; and it would be exceedingly hazardous to affirm that organs capable of giving rise to the same products exist in the vastly simpler nervous system of the crustacean. It would be the height of absurdity to expect from a meat-jack the sort of work which is performed by a Jacquard loom; and it appears to me to be little less preposterous to look for the production of anything analogous to the more subtle phenomena of the human mind in something so minute and rude in comparison to the human brain, as the insignificant cerebral ganglia of the crayfish.
At the most, one may be justified in supposing the existence of something approaching dull feeling in ourselves; and, to return to the problem stated in the
beginning of this chapter, so far as such obscure consciousness accompanies the molecular changes of its nervous substance, it will be right to speak of the mind of a crayfish. But it will be obvious that it is merely putting the cart before the horse, to speak of such a mind as a factor in the work done by the organism, when it is merely a dim symbol of a part of such work in the doing.
Whether the crayfish possesses consciousness or not, however, does not affect the question of its being an engine, the actions of which at any moment depend, on the one hand, upon the series of molecular changes excited, either by internal or by external causes, in its neuromuscular machinery; and, on the other, upon the disposition and the properties of the parts of that machinery. And such a self-adjusting machine, containing the immediate conditions of its action within itself, is what is properly understood by an automaton.
Crayfishes, as we have seen, may attain a considerable age; and there is no means of knowing how long they might live, if protected from the innumerable destructive influences to which they are at all ages liable.
It is a widely received notion that the energies of living matter have a natural tendency to decline, and finally disappear; and that the death of the body, as a whole, is the necessary correlate of its life. That all living things sooner or later perish needs no demonstration, but it would be difficult to find satisfactory grounds
for the belief that they must needs do so. The analogy of a machine that, sooner or later, must be brought to a standstill by the wear and tear of its parts, does not hold, inasmuch as the animal mechanism is continually renewed and repaired; and, though it is true that individual components of the body are constantly dying, yet their places are taken by vigorous successors. A city remains, notwithstanding the constant death-rate of its inhabitants; and such an organism as a crayfish is only a corporate unity, made up of innumerable partially independent individualities.
Whatever might be the longevity of crayfishes under imaginable perfect conditions, the fact that, notwithstanding the great number of eggs they produce, their number remains pretty much the same in a given district, if we take the average of a period of years, shows that about as many die as are born; and that, without the process of reproduction, the species would soon come to an end.
There are many examples among members of the group of Crustacea to which the crayfish belongs, of animals which produce young from internally developed germs, as some plants throw off bulbs which are capable of reproducing the parent stock; such is the case, for example, with the common water flea (Daphnia). But nothing of this kind has been observed in the crayfish; in which, as in the higher animals, the reproduction of the species is dependent upon the combination of two kinds of living
matter, which are developed in different individuals, termed males and females.
These two kinds of living matter are ova and spermatozoa, and they are developed in special organs, the ovary and the testis. The ovary is lodged in the female; the testis, in the male.
[Figure 30: Astacus fluviatilis--The female reproductive organs]
The ovary (fig. 30, ov) is a body of a trefoil form, which is situated immediately beneath, or in front of, the heart, between the floor of the pericardial sinus and the alimentary canal. From the ventral face of this organ two short and wide canals, the oviducts (od), lead down to the bases of the second pair of walking limbs, and terminate in the apertures (od') already noticed there.
[Figure 31: Astacus fluviatilis--The male reproductive organs]
The testis (fig. 31, t) is somewhat similar in form to the ovary, but, the three divisions are much narrower
and more elongated: the hinder median division lies under the heart; the anterior divisions are situated between the heart behind, and the stomach and the liver in front (figs. 5 and 12, t). From the point at which the three divisions join, proceed two ducts, which are termed the vasa deferentia (fig. 31, vd). These are very narrow, long, and make many coils before they reach the apertures upon the bases of the hindermost pair of walking limbs, by which they open externally (fig. 31, vd', and fig. 35, vd). Both the ovary and the testis are very much larger during the breeding season than at other times; the large brownish-yellow eggs become conspicuous in the ovary, and the testis assumes a milk-white colour, at this period.
[Figure 32: Astacus fluviatilis--Egg, ovisac]
[Figure 33: Astacus fluviatilis--Testis, spermatic cells]
The walls of the ovary are lined internally by a layer of
nucleated cells, separated from the cavity of the organ by a delicate structureless membrane. The growth of these cells gives rise to papillary elevations which project into the cavity of the ovary, and eventually become globular bodies attached by short stalks, and invested by the structureless membrane as a membrana propria (fig. 32, m). These are the ovisacs. In the mass of cells which becomes the ovisac, one rapidly increases in size and occupies the centre of the ovisac, while the others surround it as a peripheral coat (ep.). This central cell is the ovum. Its nucleus enlarges, and becomes what is called the germinal vesicle (g.v.). At the same time numerous small corpuscles, flattened externally and convex internally, appear in it and are the germinal spots (g.s.). The protoplasm of the cell, as it enlarges, becomes granular and opaque, assumes a deep brownish-yellow colour, and is thus converted into the yelk or vitellus (v.). As the egg grows, a structureless vitelline membrane is formed between the vitellus and the cells which line the ovisac, and incloses the egg, as in a bag. Finally, the ovisac bursts, and the egg, falling into the cavity of the ovary, makes its way down the oviduct, and sooner or later passes out by its aperture. When they leave the oviduct, the ova are invested by a viscous, transparent substance, which attaches them to the swimmerets of the female, and then sets; thus each egg, inclosed in a tough case, is firmly suspended by a stalk, which, on the one side, is continued into the substance of the case, while, on the other, it is fixed to the swimmeret. The swimmerets are kept constantly in motion, so that the eggs are well supplied with ærated water.
[Figure 34: Astacus fluviatilis--Different stages in the development of a spermatozoon from a seminal cell]
The testis consists of an immense number of minute spheroidal vesicles (fig. 33, A, a), attached like grapes to the ends of short stalks (b), formed by the ultimate ramifications of the vasa deferentia. The vesicles may, in fact, be regarded as dilatations of the ends and sides
of the finest branches of the ducts of the testis. The cavity of each vesicle is filled by the large nucleated cells which line its walls (fig. 33, B), and, as the breeding season approaches, these cells multiply by division. Finally, they undergo some very singular changes of form and internal structure (fig. 34, A-D), each becoming converted into a flattened spheroidal body, about 1/1700th of an inch in diameter, provided with a number of slender curved rays, which stand out from its sides (fig. 34, E-G). These are the spermatozoa. [see End note 13]
[Figure 35: Astacus fluviatilis--The last thoracic sternum, seen from behind, with the proximal ends of the appendages]
The spermatozoa accumulate in the testicular vesicles, and give rise to a milky-looking substance, which traverses the smaller ducts, and eventually fills the vasa deferentia. This substance, however, consists, in addition to the spermatozoa, of a viscid material, secreted by the walls of the vasa deferentia, which envelopes the spermatozoa, and gives the secretion of the testis the form and the consistency of threads of vermicelli.
The ripening and detachment of both the ova and the spermatozoa take place immediately after the completion of ecdysis in the early autumn; and at this time, which is the breeding season, the males seek the females with great avidity, in order to deposit the fertilizing matter contained in the vasa deferentia on the sterna of their hinder thoracic and anterior abdominal somites. There it adheres as a whitish, chalky-looking mass; but the manner in which the contained spermatozoa reach and enter the ova is unknown. The analogy
of what occurs in other animals, however, leaves no doubt that an actual mixture of the male and female elements takes place and constitutes the essential part of the process of impregnation.
Ova to which spermatozoa have had no access, give rise to no progeny; but, in the impregnated ovum, the young crayfish takes its origin in a manner to be described below, when the question of development is dealt with.______
[Author's Notes to Chapter 3]
[Note 1]: By axis of the articulation is meant a line drawn through the pair of hinges which constitute it.
[Note 2]: My attention was first drawn to these phenomena by my friend Dr. M. Foster, F.R.S., to whom I had suggested the desirableness of an experimental study of the nerve physiology of the crayfish.
[Note 3]: It may be said that, strictly speaking, only those external bodies which are in direct contact with the organism do affect it--as the vibrating ether, in the case of luminous bodies; the vibrating air or water, in the case of sonorous bodies; odorous particles, in the case of odorous bodies: but I have preferred the ordinary phraseology to a pedantically accurate periphrasis.
[Note 4]: Since the visual rods are strongly refracting solids, and not empty tubes, the diagram given in fig. 29 does not represent the true course of the rays, indicated by dotted lines, which fall obliquely on any cornea of a crayfish's eye. Such rays will be more or less bent towards the axis of the visual rod of that cornea; but whether they reach its apex and so affect the nerve or not will depend on the curvature of the cornea; its refractive index and that of the crystalline cone; and the relation between the length and the thickness of the latter.
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