Chapter X: "On the Imperfection of the Geological Record," and Chapter (4)

The mechanism and the conditions for the change in colouration were made clear through the beautiful investigations of Keeble and Gamble, on the colour-change in crustaceans. According to these authors the pigment-cells can, as a rule, be considered as consisting of a central body from which a system of more or less complicated ramifications or processes spreads out in all directions. As a rule, the centre of the cell contains one or more different pigments which under the influence of nerves can spread out separately or together into the ramifications. These phenomena of spreading and retraction of the pigments into or from the ramifications of the pigment-cells form on the whole the basis for the colour changes under the influence of environment. Thus Keeble and Gamble observed that Macromysis flexuosa appears transparent and colourless or grey on sandy ground. On a dark ground their colour becomes darker. These animals have two pigments in their chromatophores, a brown pigment and a whitish or yellow pigment; the former is much more plentiful than the latter. When the animal appears transparent all the pigment is contained in the centre of the cells, while the ramifications are free from pigment. When the animal appears brown both pigments are spread out into the ramifications. In the condition of maximal spreading the animals appear black.

This is a comparatively simple case. Much more complicated conditions were found by Keeble and Gamble in other crustaceans, e.g. in Hippolyte cranchii, but the influence of the surroundings upon the colouration of this form was also satisfactorily analysed by these authors.

While many animals show transitory changes in colour under the influence of their surroundings, in a few cases permanent changes can be produced. The best examples of this are those which were observed by Poulton in the chrysalids of various butterflies, especially the small tortoise-shell. These experiments are so well known that a short reference to them will suffice. Poulton (Poulton, E.B., "Colours of Animals" (The International Scientific Series), London, 1890, page 121.) found that in gilt or white surroundings the pupae became light coloured and there was often an immense development of the golden spots, "so that in many cases the whole surface of the pupae glittered with an apparent metallic lustre. So remarkable was the appearance that a physicist to whom I showed the chrysalids, suggested that I had played a trick and had covered them with goldleaf." When black surroundings were used "the pupae were as a rule extremely dark, with only the smallest trace, and often no trace at all, of the golden spots which are so conspicuous in the lighter form." The susceptibility of the animal to this influence of its surroundings was found to be greatest during a definite period when the caterpillar undergoes the metamorphosis into the chrysalis stage. As far as the writer is aware, no physico-chemical explanation, except possibly Wiener's suggestion of colour-photography by mechanical colour adaptation, has ever been offered for the results of the type of those observed by Poulton.

V. EFFECTS OF GRAVITATION.

(a) EXPERIMENTS ON THE EGG OF THE FROG.

Gravitation can only indirectly affect life-phenomena; namely, when we have in a cell two different non-miscible liquids (or a liquid and a solid) of different specific gravity, so that a change in the position of the cell or the organ may give results which can be traced to a change in the position of the two substances. This is very nicely illustrated by the frog's egg, which has two layers of very viscous protoplasm one of which is black and one white. The dark one occupies normally the upper position in the egg and may therefore be assumed to possess a smaller specific gravity than the white substance. When the egg is turned with the white pole upwards a tendency of the white protoplasm to flow down again manifests itself. It is, however, possible to prevent or retard this rotation of the highly viscous protoplasm, by compressing the eggs between horizontal glass plates. Such compression experiments may lead to rather interesting results, as O. Schultze first pointed out. Pflueger had already shown that the first plane of division in a fertilised frog's egg is vertical and Roux established the fact that the first plane of division is identical with the plane of symmetry of the later embryo. Schultze found that if the frog's egg is turned upside down at the time of its first division and kept in this abnormal position, through compression between two glass plates for about 20 hours, a small number of eggs may give rise to twins. It is possible, in this case, that the tendency of the black part of the egg to rotate upwards along the surface of the egg leads to a separation of its first cells, such a separation leading to the formation of twins.

T.H. Morgan made an interesting additional observation. He destroyed one half of the egg after the first segmentation and found that the half which remained alive gave rise to only one half of an embryo, thus confirming an older observation of Roux. When, however, Morgan put the egg upside down after the destruction of one of the first two cells, and compressed the eggs between two glass plates, the surviving half of the egg gave rise to a perfect embryo of half size (and not to a half embryo of normal size as before.) Obviously in this case the tendency of the protoplasm to flow back to its normal position was partially successful and led to a partial or complete separation of the living from the dead half; whereby the former was enabled to form a whole embryo, which, of course, possessed only half the size of an embryo originating from a whole egg.

(b) EXPERIMENTS ON HYDROIDS.

A striking influence of gravitation can be observed in a hydroid, Antennularia antennina, from the bay of Naples. This hydroid consists of a long straight main stem which grows vertically upwards and which has at regular intervals very fine and short bristle-like lateral branches, on the upper side of which the polyps grow. The main stem is negatively geotropic, i.e. its apex continues to grow vertically upwards when we put it obliquely into the aquarium, while the roots grow vertically downwards. The writer observed that when the stem is put horizontally into the water the short lateral branches on the lower side give rise to an altogether different kind of organ, namely, to roots, and these roots grow indefinitely in length and attach themselves to solid bodies; while if the stem had remained in its normal position no further growth would have occurred in the lateral branches. From the upper side of the horizontal stem new stems grow out, mostly directly from the original stem, occasionally also from the short lateral branches. It is thus possible to force upon this hydroid an arrangement of organs which is altogether different from the hereditary arrangement. The writer had called the change in the hereditary arrangement of organs or the transformation of organs by external forces HETEROMORPHOSIS. We cannot now go any further into this subject, which should, however, prove of interest in relation to the problem of heredity.

If it is correct to apply inferences drawn from the observation on the frog's egg to the behaviour of Antennularia, one might conclude that the cells of Antennularia also contain non-miscible substances of different specific gravity, and that wherever the specifically lighter substance comes in contact with the sea-water (or gets near the surface of the cell) the growth of a stem is favoured; while contact with the sea-water of the specifically heavier of the substances, will favour the formation of roots.

VI. THE EXPERIMENTAL CONTROL OF ANIMAL INSTINCTS.

(a) EXPERIMENTS ON THE MECHANISM OF HELIOTROPIC REACTIONS IN ANIMALS.

Since the instinctive reactions of animals are as hereditary as their morphological character, a discussion of experiments on the physico-chemical character of the instinctive reactions of animals should not be entirely omitted from this sketch. It is obvious that such experiments must begin with the simplest type of instincts, if they are expected to lead to any results; and it is also obvious that only such animals must be selected for this purpose, the reactions of which are not complicated by associative memory, or, as it may preferably be termed, associative hysteresis.

The simplest type of instincts is represented by the purposeful motions of animals to or from a source of energy, e.g. light; and it is with some of these that we intend to deal here. When we expose winged aphides (after they have flown away from the plant), or young caterpillars of Porthesia chrysorrhoea (when they are aroused from their winter sleep) or marine or freshwater copepods and many other animals, to diffused daylight falling in from a window, we notice a tendency among these animals to move towards the source of light. If the animals are naturally sensitive, or if they are rendered sensitive through the agencies which we shall mention later, and if the light is strong enough, they move towards the source of light in as straight a line as the imperfections and peculiarities of their locomotor apparatus will permit. It is also obvious that we are here dealing with a forced reaction in which the animals have no more choice in the direction of their motion than have the iron filings in their arrangement in a magnetic field. This can be proved very nicely in the case of starving caterpillars of Porthesia. The writer put such caterpillars into a glass tube the axis of which was at right angles to the plane of the window: the caterpillars went to the window side of the tube and remained there, even if leaves of their food-plant were put into the tube directly behind them. Under such conditions the animals actually died from starvation, the light preventing them from turning to the food, which they eagerly ate when the light allowed them to do so. One cannot say that these animals, which we call positively helioptropic, are attracted by the light, since it can be shown that they go towards the source of the light even if in so doing they move from places of a higher to places of a lower degree of illumination.

The writer has advanced the following theory of these instinctive reactions. Animals of the type of those mentioned are automatically orientated by the light in such a way that symmetrical elements of their retina (or skin) are struck by the rays of light at the same angle. In this case the intensity of light is the same for both retinae or symmetrical parts of the skin.

This automatic orientation is determined by two factors, first a peculiar photo-sensitiveness of the retina (or skin), and second a peculiar nervous connection between the retina and the muscular apparatus. In symmetrically built heliotropic animals in which the symmetrical muscles participate equally in locomotion, the symmetrical muscles work with equal energy as long as the photo-chemical processes in both eyes are identical. If, however, one eye is struck by stronger light than the other, the symmetrical muscles will work unequally and in positively heliotropic animals those muscles will work with greater energy which bring the plane of symmetry back into the direction of the rays of light and the head towards the source of light. As soon as both eyes are struck by the rays of light at the same angle, there is no more reason for the animal to deviate from this direction and it will move in a straight line. All this holds good on the supposition that the animals are exposed to only one source of light and are very sensitive to light.

Additional proof for the correctness of this theory was furnished through the experiments of G.H. Parker and S.J. Holmes. The former worked on a butterfly, Vanessa antiope, the latter on other arthropods. All the animals were in a marked degree positively heliotropic. These authors found that if one cornea is blackened in such an animal, it moves continually in a circle when it is exposed to a source of light, and in these motions the eye which is not covered with paint is directed towards the centre of the circle. The animal behaves, therefore, as if the darkened eye were in the shade.

(b) THE PRODUCTION OF POSITIVE HELIOTROPISM BY ACIDS AND OTHER MEANS AND THE PERIODIC DEPTH-MIGRATIONS OF PELAGIC ANIMALS.

When we observe a dense mass of copepods collected from a freshwater pond, we notice that some have a tendency to go to the light while others go in the opposite direction and many, if not the majority, are indifferent to light. It is an easy matter to make the negatively heliotropic or the indifferent copepods almost instantly positively heliotropic by adding a small but definite amount of carbon-dioxide in the form of carbonated water to the water in which the animals are contained. If the animals are contained in 50 cubic centimetres of water it suffices to add from three to six cubic centimetres of carbonated water to make all the copepods energetically positively heliotropic. This heliotropism lasts about half an hour (probably until all the carbon-dioxide has again diffused into the air.) Similar results may be obtained with any other acid.

The same experiments may be made with another freshwater crustacean, namely Daphnia, with this difference, however, that it is as a rule necessary to lower the temperature of the water also. If the water containing the Daphniae is cooled and at the same time carbon-dioxide added, the animals which were before indifferent to light now become most strikingly positively heliotropic. Marine copepods can be made positively heliotropic by the lowering of the temperature alone, or by a sudden increase in the concentration of the sea-water.

These data have a bearing upon the depth-migrations of pelagic animals, as was pointed out years ago by Theo. T. Groom and the writer. It is well known that many animals living near the surface of the ocean or freshwater lakes, have a tendency to migrate upwards towards evening and downwards in the morning and during the day. These periodic motions are determined to a large extent, if not exclusively, by the heliotropism of these animals. Since the consumption of carbon-dioxide by the green plants ceases towards evening, the tension of this gas in the water must rise and this must have the effect of inducing positive heliotropism or increasing its intensity. At the same time the temperature of the water near the surface is lowered and this also increases the positive heliotropism in the organisms.

The faint light from the sky is sufficient to cause animals which are in a high degree positively heliotropic to move vertically upwards towards the light, as experiments with such pelagic animals, e.g. copepods, have shown. When, in the morning, the absorption of carbon-dioxide by the green algae begins again and the temperature of the water rises, the animals lose their positive heliotropism, and slowly sink down or become negatively heliotropic and migrate actively downwards.

These experiments have also a bearing upon the problem of the inheritance of instincts. The character which is transmitted in this case is not the tendency to migrate periodically upwards and downwards, but the positive heliotropism. The tendency to migrate is the outcome of the fact that periodically varying external conditions induce a periodic change in the sense and intensity of the heliotropism of these animals. It is of course immaterial for the result, whether the carbon-dioxide or any other acid diffuse into the animal from the outside or whether they are produced inside in the tissue cells of the animals. Davenport and Cannon found that Daphniae, which at the beginning of the experiment, react sluggishly to light react much more quickly after they have been made to go to the light a few times. The writer is inclined to attribute this result to the effect of acids, e.g. carbon-dioxide, produced in the animals themselves in consequence of their motion. A similar effect of the acids was shown by A.D. Waller in the case of the response of nerve to stimuli.

The writer observed many years ago that winged male and female ants are positively helioptropic and that their heliotropic sensitiveness increases and reaches its maximum towards the period of nuptial flight. Since the workers show no heliotropism it looks as if an internal secretion from the sexual glands were the cause of their heliotropic sensitiveness. V. Kellogg has observed that bees also become intensely positively heliotropic at the period of their wedding flight, in fact so much so that by letting light fall into the observation hive from above, the bees are prevented from leaving the hive through the exit at the lower end.

We notice also the reverse phenomenon, namely, that chemical changes produced in the animal destroy its heliotropism. The caterpillars of Porthesia chrysorrhoea are very strongly positively heliotropic when they are first aroused from their winter sleep. This heliotropic sensitiveness lasts only as long as they are not fed. If they are kept permanently without food they remain permanently positively heliotropic until they die from starvation. It is to be inferred that as soon as these animals take up food, a substance or substances are formed in their bodies which diminish or annihilate their heliotropic sensitiveness.

The heliotropism of animals is identical with the heliotropism of plants. The writer has shown that the experiments on the effect of acids on the heliotropism of copepods can be repeated with the same result in Volvox. It is therefore erroneous to try to explain these heliotropic reactions of animals on the basis of peculiarities (e.g. vision) which are not found in plants.

We may briefly discuss the question of the transmission through the sex cells of such instincts as are based upon heliotropism. This problem reduces itself simply to that of the method whereby the gametes transmit heliotropism to the larvae or to the adult. The writer has expressed the idea that all that is necessary for this transmission is the presence in the eyes (or in the skin) of the animal of a photo-sensitive substance. For the transmission of this the gametes need not contain anything more than a catalyser or ferment for the synthesis of the photo-sensitive substance in the body of the animal. What has been said in regard to animal heliotropism might, if space permitted, be extended, mutatis mutandis, to geotropism and stereotropism.

(c) THE TROPIC REACTIONS OF CERTAIN TISSUE-CELLS AND THE MORPHOGENETIC EFFECTS OF THESE REACTIONS.

Since plant-cells show heliotropic reactions identical with those of animals, it is not surprising that certain tissue-cells also show reactions which belong to the class of tropisms. These reactions of tissue-cells are of special interest by reason of their bearing upon the inheritance of morphological characters. An example of this is found in the tiger-like marking of the yolk-sac of the embryo of Fundulus and in the marking of the young fish itself. The writer found that the former is entirely, and the latter at least in part, due to the creeping of the chromatophores upon the blood-vessels. The chromatophores are at first scattered irregularly over the yolk-sac and show their characteristic ramifications. There is at that time no definite relation between blood-vessels and chromatophores. As soon as a ramification of a chromatophore comes in contact with a blood-vessel the whole mass of the chromatophore creeps gradually on the blood-vessel and forms a complete sheath around the vessel, until finally all the chromatophores form a sheath around the vessels and no more pigment cells are found in the meshes between the vessels. Nobody who has not actually watched the process of the creeping of the chromatophores upon the blood-vessels would anticipate that the tiger-like colouration of the yolk-sac in the later stages of the development was brought about in this way. Similar facts can be observed in regard to the first marking of the embryo itself. The writer is inclined to believe that we are here dealing with a case of chemotropism, and that the oxygen of the blood may be the cause of the spreading of the chromatophores around the blood-vessels. Certain observations seem to indicate the possibility that in the adult the chromatophores have, in some forms at least, a more rigid structure and are prevented from acting in the way indicated. It seems to the writer that such observations as those made on Fundulus might simplify the problem of the hereditary transmission of certain markings.

Driesch has found that a tropism underlies the arrangement of the skeleton in the pluteus larvae of the sea-urchin. The position of this skeleton is predetermined by the arrangement of the mesenchyme cells, and Driesch has shown that these cells migrate actively to the place of their destination, possibly led there under the influence of certain chemical substances. When Driesch scattered these cells mechanically before their migration, they nevertheless reached their destination.

In the developing eggs of insects the nuclei, together with some cytoplasm, migrate to the periphery of the egg. Herbst pointed out that this might be a case of chemotropism, caused by the oxygen surrounding the egg. The writer has expressed the opinion that the formation of the blastula may be caused generally by a tropic reaction of the blastomeres, the latter being forced by an outside influence to creep to the surface of the egg.

These examples may suffice to indicate that the arrangement of definite groups of cells and the morphological effects resulting therefrom may be determined by forces lying outside the cells. Since these forces are ubiquitous and constant it appears as if we were dealing exclusively with the influence of a gamete; while in reality all that it is necessary for the gamete to transmit is a certain form of irritability.

(d) FACTORS WHICH DETERMINE PLACE AND TIME FOR THE DEPOSITION OF EGGS.

For the preservation of species the instinct of animals to lay their eggs in places in which the young larvae find their food and can develop is of paramount importance. A simple example of this instinct is the fact that the common fly lays its eggs on putrid material which serves as food for the young larvae. When a piece of meat and of fat of the same animal are placed side by side, the fly will deposit its eggs upon the meat on which the larvae can grow, and not upon the fat, on which they would starve. Here we are dealing with the effect of a volatile nitrogenous substance which reflexly causes the peristaltic motions for the laying of the egg in the female fly.

Kammerer has investigated the conditions for the laying of eggs in two forms of salamanders, e.g. Salamandra atra and S. maculosa. In both forms the eggs are fertilised in the body and begin to develop in the uterus. Since there is room only for a few larvae in the uterus, a large number of eggs perish and this number is the greater the longer the period of gestation. It thus happens that when the animals retain their eggs a long time, very few young ones are born; and these are in a rather advanced stage of development, owing to the long time which elapsed since they were fertilised. When the animal lays its eggs comparatively soon after copulation, many eggs (from 12 to 72) are produced and the larvae are of course in an early stage of development. In the early stage the larvae possess gills and can therefore live in water, while in later stages they have no gills and breathe through their lungs. Kammerer showed that both forms of Salamandra can be induced to lay their eggs early or late, according to the physical conditions surrounding them. If they are kept in water or in proximity to water and in a moist atmosphere they have a tendency to lay their eggs earlier and a comparatively high temperature enhances the tendency to shorten the period of gestation. If the salamanders are kept in comparative dryness they show a tendency to lay their eggs rather late and a low temperature enhances this tendency.

Since Salamandra atra is found in rather dry alpine regions with a relatively low temperature and Salamandra maculosa in lower regions with plenty of water and a higher temperature, the fact that S. atra bears young which are already developed and beyond the stage of aquatic life, while S. maculosa bears young ones in an earlier stage, has been termed adaptation. Kammerer's experiments, however, show that we are dealing with the direct effects of definite outside forces. While we may speak of adaptation when all or some of the variables which determine a reaction are unknown, it is obviously in the interest of further scientific progress to connect cause and effect directly whenever our knowledge allows us to do so.

VII. CONCLUDING REMARKS.

The discovery of De Vries, that new species may arise by mutation and the wide if not universal applicability of Mendel's Law to phenomena of heredity, as shown especially by Bateson and his pupils, must, for the time being, if not permanently, serve as a basis for theories of evolution. These discoveries place before the experimental biologist the definite task of producing mutations by physico-chemical means. It is true that certain authors claim to have succeeded in this, but the writer wishes to apologise to these authors for his inability to convince himself of the validity of their claims at the present moment. He thinks that only continued breeding of these apparent mutants through several generations can afford convincing evidence that we are here dealing with mutants rather than with merely pathological variations.

What was said in regard to the production of new species by physico-chemical means may be repeated with still more justification in regard to the second problem of transformation, namely the making of living from inanimate matter. The purely morphological imitations of bacteria or cells which physicists have now and then proclaimed as artificially produced living beings; or the plays on words by which, e.g. the regeneration of broken crystals and the regeneration of lost limbs by a crustacean were declared identical, will not appeal to the biologist. We know that growth and development in animals and plants are determined by definite although complicated series of catenary chemical reactions, which result in the synthesis of a DEFINITE compound or group of compounds, namely, NUCLEINS.

The nucleins have the peculiarity of acting as ferments or enzymes for their own synthesis. Thus a given type of nucleus will continue to synthesise other nuclein of its own kind. This determines the continuity of a species; since each species has, probably, its own specific nuclein or nuclear material. But it also shows us that whoever claims to have succeeded in making living matter from inanimate will have to prove that he has succeeded in producing nuclein material which acts as a ferment for its own synthesis and thus reproduces itself. Nobody has thus far succeeded in this, although nothing warrants us in taking it for granted that this task is beyond the power of science.

XV. THE VALUE OF COLOUR IN THE STRUGGLE FOR LIFE. By E.B. Poulton.

Hope Professor of Zoology in the University of Oxford.