At the present day nobody seriously questions the statement that the action of light upon organisms is primarily one of a chemical character. While this chemical action is of the utmost importance for organisms, the nutrition of which depends upon the action of chlorophyll, it becomes of less importance for organisms devoid of chlorophyll. Nevertheless, we find animals in which the formation of organs by regeneration is not possible unless they are exposed to light. An observation made by the writer on the regeneration of polyps in a hydroid, Eudendrium racemosum, at Woods Hole, may be mentioned as an instance of this. If the stem of this hydroid, which is usually covered with polyps, is put into an aquarium the polyps soon fall off. If the stems are kept in an aquarium where light strikes them during the day, a regeneration of numerous polyps takes place in a few days. If, however, the stems of Eudendrium are kept permanently in the dark, no polyps are formed even after an interval of some weeks; but they are formed in a few days after the same stems have been transferred from the dark to the light. Diffused daylight suffices for this effect.
Goldfarb, who repeated these experiments, states that an exposure of comparatively short duration is sufficient for this effect, it is possible that the light favours the formation of substances which are a prerequisite for the origin of polyps and their growth.
Of much greater significance than this observation are the facts which show that a large number of animals assume, to some extent, the colour of the ground on which they are placed. Pouchet found through experiments upon crustaceans and fish that this influence of the ground on the colour of animals is produced through the medium of the eyes. If the eyes are removed or the animals made blind in another way these phenomena cease.
The second general fact found by Pouchet was that the variation in the colour of the animal is brought about through an action of the nerves on the pigment-cells of the skin; the nerve-action being induced through the agency of the eye.
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 ****** 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.