Читать книгу Animal Behaviour - C. Lloyd Morgan - Страница 7

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From what has already been said it may be inferred that our use of the term “behaviour” neither implies nor excludes the presence of consciousness. Few are prepared to contend that the iron filings in a magnetic field consciously group themselves in definite and symmetrical patterns, or that sand grains on a vibrating plate assemble along certain nodal lines because they are conscious of the effects of the bow by which the plate is set in sounding vibration. But where organic response falls under our observation, no matter how simple and direct that response may be, there is a natural tendency to suppose that the behaviour is conscious; and where the response is less simple and more indirect, this tendency is so strengthened as to give rise to a state of mind bordering on, or actually reaching, conviction. Nor is this surprising: for, in the first place, organic responses, even the simplest, are less obviously and directly related to the interplay of surrounding circumstances; and, in the second place, they are more obviously in relation to some purpose in the sense that they directly or indirectly contribute to the maintenance of life or the furtherance of well-being. Now where behaviour is complex and subserves an end which we can note and name, there arises the supposition that it may well be of the same nature as our own complex and conscious behaviour.

Take for example the behaviour of the Slipper-animalcule, Paramecium, one of the minute creatures known to zoologists as Protozoa. The whole animal is constituted by a single cell, somewhat less than one-hundredth of an inch in length, the form and behaviour of which may be readily studied under the microscope. Thousands may be obtained from water in which some hay has been allowed to rot. The surface of the Paramecium is covered with waving hair-like cilia, by which it is propelled through the water, while stiffer hairs may be shot out from the surface at any point where there is a local source of irritation, as indicated at the top of the accompanying figure. Two little sacs expand and contract, and serve to drain off water and waste products from the substance of the cell. Food is taken in at the end of the funnel, shown in the lower part of the figure. The cilia here work in such manner as to drive the particles into and down the tube, and on reaching its inner end these particles burst through into the semi-fluid substance, and circulate therein. Just above the funnel there are two bean-like bodies, the larger of which is known as the macronucleus, the smaller as the micronucleus.

Fig. 1.—Paramecium.

The process of multiplication is by “fission,” or the division of each Paramecium into two similar animalcules. Not infrequently, however, two Paramecia may be seen to approach each other and come together, funnel to funnel; and in each the nuclei undergo curious changes. The macronucleus breaks up, and is scattered. The micronucleus in each divides into four portions, of which three break up and disappear; while the fourth again divides into two parts, one to be retained and the other to be exchanged for the similar micronuclear product of the other Paramecium. The retained portion and that received in exchange then unite to form a new micronucleus. M. Maupas concludes from his careful observations that, in the absence of such “conjugation” in the mid-period of life, Paramecia pass into a state of senility which ends in decrepitude and death. If this be so, conjugation is in them necessary for the continuance of a healthy race.

Here we have what a zoologist would describe as a specialized mode of behaviour of the nuclei; and we have also the behaviour of the minute creatures (which contain the nuclei) as they approach each other and come together in conjugation. Can one wonder that the latter, at any rate, has been regarded as an example of conscious procedure? In truth we do not know in what manner and by what subtle influences the Paramecia are drawn together in conjugation. But it is scarcely logical to base on such ignorance any positive assertion as to conscious attraction. It is better to confess that here is a piece of organic behaviour, the exact conditions of which are at present unexplained.

We may take from the writings[1] of Dr. H. S. Jennings, of Harvard, some account of other modes of behaviour among Paramecia. They largely feed upon clotted masses of bacteria. If a number are placed upon a glass slip, together with a small bacterial clot, they will be seen to congregate around the clot and to feed upon it. All apparently press in so as to reach it, or get as near it as possible. And if a number be placed on another slide without any clot, they soon collect in groups in one or more regions, as in Fig. 2, III. It appears as if they were actuated by some social impulse leading them to crowd together and shun isolated positions. Nay, more; it seems as if, after thus collecting and crowding in to some centre of interest, the attractive influence gradually waned; the group spreads, and the Paramecia are less densely packed; the assembly scatters more and more, but still seems to be retained by an invisible boundary beyond which the little creatures do not pass.

Fig. 2.—Behaviour of Paramecia (after Jennings).

Furthermore, if kept in a jar, the Paramecia crowd up towards the surface where the bacteria clots are floating; and if, beneath the cover glass of a slip on which they are under microscopic examination, a drop of liquid be introduced through a very fine tube, they will seem either to be attracted to it, as in Fig. 2, I., or repelled from it, as in Fig. 2, II., according to its nature. From alkaline liquids they are repelled; to slightly acid drops they are attracted, unless the acidity be too pungent. Heat and cold are alike repellent, and even a drop of pure distilled water forms an area into which the Paramecia do not enter.

With such facts before him, the incautious observer may be led to the conclusion that Paramecia are not only conscious, but endowed with intelligence and volition. Even M. Binet,[2] who occupies a position which should lead him to exercise more caution, tells us that there is not a single infusorian which cannot be frightened, and does not manifest its fear by rapid flight; he speaks of some of these unicellular animals as “endowed with memory and volition,” and possessed of “instinct of great precision;” and he describes the following stages:—

“(1) The perception of an external object;

“(2) The choice made between a number of objects;

“(3) The perception of their position in space;

“(4) Movements calculated either to approach the body and seize it, or to flee from it.”

But when we seem to have grasped his point of view, when we have catalogued the memory, fear, instinct, perception, choice and volition, the whole intelligent edifice crumbles; for we are told that “we are not in a position to determine whether these various acts are accompanied by consciousness, or whether they follow as simple physiological processes.” To most of us fear, memory, choice, volition, imply something more than simple physiological processes; they imply not only consciousness, but highly elaborated consciousness.

Dr. Jennings’s researches show that no such implication can be accepted unless we are prepared to cast aside the trammels of reasonable caution. In the first place, the whole matter of feeding appears to be referable to simple organic behaviour not necessarily involving consciousness. The cilia in the mouth-groove and funnel constantly wave in such a manner as to drive a current of water, together with any particles which float therein, towards the interior; and the particles are then engulphed, no matter what their composition may be. Digestible or indigestible, in they go. There is no selection of the one or rejection of the other. But, as we have seen, the Paramecia collect around a bacterial clot and feed upon it. Surely here there is selection of the nutritious! Apparently not. They collect in just the same way towards a piece of blotting-paper, cotton-wool, cloth, sponge, or other fibrous body, and remain assembled round such an innutritious centre just as long as round a bacterial clot. There seems to be no choice in the matter; contact with any substance gives rise, as an organic response, to the lessening or cessation of the regular movements in all the cilia except those of the mouth-groove and funnel. As the Paramecia swim hither and thither, first one, then another, then more, chance to come in contact with the bacterial clot, the blotting-paper, or other substance, and since the lashing of the cilia is then automatically lessened, there they stay; others find their way to the same spot in the course of their random movements, and they, too, stay; thus many soon collect.

But this does not account for the seemingly social assemblages of Paramecia where there is no such substance to arrest their progress. Dr. Jennings attributes this to the fact that a dilute solution of carbon-dioxide has, what we may call for the present, an attractive influence. If a bubble of air and a bubble of carbon dioxide be introduced into the water in which Paramecia are swimming beneath a cover-glass, the animalcules collect around the carbonic dioxide, but not around the air bubble. At first they press up close to the bubble of carbon dioxide, but gradually form a ring farther and farther from its limiting boundary. This is held to be due to the fact that it is only the dilute solution of carbonic acid that has the peculiar “attraction”—a stronger solution has a different effect. And, as the gas dissolves, the Paramecia collect in a ring just where the solution is sufficiently dilute.

Now carbon dioxide is a product of the organic waste of living substance; it is given off by active Paramecia. Where therefore many are collected together they form a centre of the production of this substance; and when other Paramecia come, in the course of their random movements, into such a centre they remain there and help to swell the numbers in the cluster. If Paramecia be placed in water to which a distinctly reddish tinge is given by mixing it with a small quantity of rosol—a substance which is decolourized by carbon dioxide, and is not injurious to Paramecia—it will be seen that, where the groups are collected, the reddish tinge fades and disappears. As the groups expand, and are less densely packed, the colourless area expands too: and the limits within which the group is circumscribed are also the limits of decolourization. Dr. Jennings considers it beyond question that the assembling of Paramecia is due to the presence in such assemblages of carbonic acid produced by the animals themselves. The first beginning of the crowd may be some small fragment of bacterial clot or other substance.

It would seem, then, that Paramecia are attracted by faintly acid solutions; and here at least there is, it may be urged, an element of choice. But even here, according to Dr. Jennings, there is not only no real choice, but not even any real attraction. What takes place, according to his observations, is briefly as follows. Suppose a faintly acid drop be inserted beneath the cover-glass. Paramecia may almost graze its boundary without being in any way affected by its presence. But in their random movements some, and eventually many, perhaps most, of the little animals chance to enter the faintly acid region; but there is no sign of reaction or response; they swim on across the drop until they reach its further margin. Here a reaction does take place. Instead of proceeding onwards, slowly revolving on its long axis, a Paramecium thus situated jerks backwards by a reversal of all the cilia, at the same time revolving on its axis in a direction opposite to that in which it was before turning. But the cilia of the mouth-groove resume their normal mode of working sooner than the others, and this causes the Paramecium to turn aside. It then goes ahead until it again reaches the boundary at another point, when the same behaviour is seen. The course of such a Paramecium is shown in Fig. 2, IV.

If, instead of a faintly acid drop, a little alkaline liquid be introduced beneath the cover-glass, the Paramecium similarly jerks backward and turns aside on reaching its outer boundary. The turning may carry it away from the alkali, as shown in Fig. 2, V.; but it just as often brings it again towards the drop, especially a large one. It seems to be a matter of chance which result follows. But eventually the little creature sails off, since each time it comes within the influence of the alkaline fluid it jerks back and turns. It appears, then, that when it is swimming in a normal solution a faintly acid liquid does not much modify its behaviour, but an alkaline fluid evokes a reversal of the cilia; and that when it is a slightly acid solution, not only does stronger acid cause reversal, but normal fluid produces a similar result. A reaction of essentially the same kind is in fact called forth by such different stimuli as chemical substances, water heated above the normal temperature, or cooled considerably below it, and fluids which cause changes of internal pressure within the substance of the cell. Nor does it matter where the stimulus is applied. If it be applied at the hinder end the infusorian still jerks backward, though this may drive it into a destructive solution and thus cause death. There is, however, some evidence of different behaviour in some infusorians according as the stimulus is here or there. In other words, the behaviour is to some extent related to the position of the part stimulated.

Furthermore, it may be gathered from Dr. Jennings’s account that there is nothing to lead us to suppose that such free living cells show any indication of what may be regarded as the keynote of intelligent behaviour. They do not profit by experience. They exhibit organic reactions which may be accompanied by some dim form of consciousness, but which do not seem to be under the guidance of such consciousness, if it exist.

One of the first lessons which the study of animal behaviour, in its organic aspect, should impress upon our minds is, that living cells may react to stimuli in a manner which we perceive to be subservient to a biological end, and yet react without conscious purpose—that is to say, automatically. The living cell assimilates food and absorbs oxygen, it grows and subdivides, it elaborates secretions, produces a skeletal framework or covering, rids itself of waste products, responds to stimuli in a definite fashion, moves hither and thither at random, its functional activities being stimulated or checked by many influences; and yet this varied life may give no evidence of a guiding consciousness: if purpose there be, it lies deeper than its protoplasm, deeper than the dim sentience which may be present or may be absent—we cannot tell which.

And when the cells are incorporated in the body of one of the higher animals, instead of each preserving a free and nomad existence; when they become the multitudinous constituents of an organic republic with unity of plan and unity of biological end, then the behaviour of each is limited in range but perfected within that range, in subservience to the requirements of the more complex unity. The muscle cell contracts, the gland-cell secretes, the rods and cones of the retina respond to the waves of light, and all the normal responses of the special cells go on with such orderly regularity that the term behaviour seems scarcely applicable to reactions so stereotyped. But the physiologist and the physician know well that such uniformity of response is dependent on uniformity of conditions. A little dose of some drug will profoundly modify and render abnormal the procedure which was before so mechanical in its exactitude; and we are thus led to see how dependent the orderly behaviour really is on the maintenance of certain surrounding conditions.

Moreover, the existence of every cell in the body corporate is the outcome of a process of division involving a special mode of behaviour in the nucleus, of which we are only beginning to guess the meaning and significance, and of which we seek in vain to find an explanation in mechanical terms. And when we trace these divisions back to their primary source in the fertilized ovum, we find changes and evolutions in the nuclear matter of which it can only be said that the more they are studied the more complex and varied do they appear.

The egg, or ovum, is a single cell produced by the female, and varying much in size, according to the amount of food-yolk with which it is supplied. Like other cells, it has a nucleus, and this undergoes changes which are definitely related to the fertilization of the ovum, which we describe as the biological end. Such preparatory changes for a future contingency are especially characteristic of organic behaviour. There is nothing like it in the mineral kingdom. The nucleus divides into two parts, one of which passes out of the ovum and is lost. The nucleus again divides, and again one part passes out and is lost. Thus only one quarter of the original amount of nuclear matter remains. Now, division of the nucleus occurs whenever an animal cell divides; but in this case (apart from details which would here be out of place) there is this difference. During the ordinary division of cells there are found in the nucleus a definite number of curved rods, and this number is constant for any given species; but in the nucleus which remains in the ovum after three parts of its substance are lost, the number of rods has been reduced to half that which is common to the species. The egg is now ready for fertilization. A minute active cell, which is produced by the male, and which also has only half the normal number of rods, enters the ovum. The two nuclei approach each other, and give rise to the single nucleus of the fertilized ovum, which thus has the full number of rods—half of them derived from one parent, half from the other parent. The sperm cell of the male adds little to the store of protoplasm in the ovum; but it introduces a minute body, which seems to initiate subsequent divisions of the cell. The nature of these divisions may be seen in the accompanying diagrammatic figure. In A the cell is just preparing to divide. Above the nucleus is the minute body (centrosome) just spoken of, which has already divided. In the nucleus the matter of which the rods will be constituted is net-like. In B this net-work has taken on the new form of a coiled thread, while the divided body above is associated with a spindle of delicate fibres. In C the membrane round the nucleus has disappeared, and the coiled thread has broken up into curved rods (chromosomes), four of which are shown. The two halves of the minute body form the centres of radiating stars. In D each curved rod has split along its length, and the two parts are being drawn asunder towards the centres of the two stars; the cell itself is beginning to divide. In E the process is carried a step further, while in F the cell has completely divided into two: the rods have disappeared as such, and are replaced by a net-work; a new nuclear membrane has been formed, and the minute body has again divided preparatory to the further division of the cell.

Fig. 3.—Cell-division.

Such, stripped as far as possible of technicalities, are some of the facts concerning the behaviour of cells and their nuclei during the process of cell-multiplication. No good purpose would be subserved by pretending that we fully understand them. The splitting of the rods does indeed seem an efficient means to the end of securing a fair division of the nuclear substance, which, according to many biologists, is the organic bearer of hereditary qualities in the cells. But that is nearly all that we can say. Is the process accompanied by some form of sentience? We do not know. That it is controlled and guided by any consciousness in the cell is most improbable. But if it be a purely organic and unconscious process it should at least impress on our minds the fact that such organic behaviour may reach a high degree of delicacy and complexity.