The paradoxical character of this general observation lies in the fact that a low temperature retards development, and hence should be expected to have the opposite effect from that mentioned by Chun. Recent investigations have led to the result that life-phenomena are affected by temperature in the same sense as the velocity of chemical reactions. In the case of the latter van't Hoff had shown that a decrease in temperature by 10 degrees reduces their velocity to one half or less, and the same has been found for the influence of temperature on the velocity of physiological processes. Thus Snyder and T.B. Robertson found that the rate of heartbeat in the tortoise and in Daphnia is reduced to about one-half if the temperature is lowered 10 deg C., and Maxwell, Keith Lucas, and Snyder found the same influence of temperature for the rate with which an impulse travels in the nerve. Peter observed that the rate of development in a sea-urchin's egg is reduced to less than one-half if the temperature (within certain limits) is reduced by 10 degrees. The same effect of temperature upon the rate of development holds for the egg of the frog, as Cohen and Peter calculated from the experiments of O. Hertwig. The writer found the same temperature-coefficient for the rate of maturation of the egg of a mollusc (Lottia).
All these facts prove that the velocity of development of animal life in Arctic regions, where the temperature is near the freezing point of water, must be from two to three times smaller than in regions where the temperature of the ocean is about 10 deg C. and from four to nine times smaller than in seas the temperature of which is about 20 deg C. It is, therefore, exactly the reverse of what we should expect when authors state that the density of organisms at or near the surface of the ocean in polar regions is greater than in more temperate regions.
The writer believes that this paradox finds its explanation in experiments which he has recently made on the influence of temperature on the duration of life of cold-blooded marine animals. The experiments were made on the fertilised and unfertilised eggs of the sea-urchin, and yielded the result that for the lowering of temperature by 1 deg C. the duration of life was about doubled. Lowering the temperature by 10 degrees therefore prolongs the life of the organism 2 to the power 10, i.e. over a thousand times, and a lowering by 20 degrees prolongs it about one million times. Since this prolongation of life is far in excess of the retardation of development through a lowering of temperature, it is obvious that, in spite of the retardation of development in Arctic seas, animal life must be denser there than in temperate or tropical seas. The excessive increase of the duration of life at the poles will necessitate the simultaneous existence of more successive generations of the same species in these regions than in the temperate or tropical regions.
The writer is inclined to believe that these results have some bearing upon a problem which plays an important role in theories of evolution, namely, the cause of natural death. It has been stated that the processes of differentiation and development lead also to the natural death of the individual. If we express this in chemical terms it means that the chemical processes which underlie development also determine natural death. Physical chemistry has taught us to identify two chemical processes even if only certain of their features are known. One of these means of identification is the temperature coefficient. When two chemical processes are identical, their velocity must be reduced by the same amount if the temperature is lowered to the same extent. The temperature coefficient for the duration of life of cold-blooded organisms seems, however, to differ enormously from the temperature coefficient for their rate of development. For a difference in temperature of 10 deg C. the duration of life is altered five hundred times as much as the rate of development; and, for a change of 20 deg C., it is altered more than a hundred thousand times as much. From this we may conclude that, at least for the sea-urchin eggs and embryo, the chemical processes which determine natural death are certainly not identical with the processes which underlie their development. T.B. Robertson has also arrived at the conclusion, for quite different reasons, that the process of senile decay is essentially different from that of growth and development.
(b) CHANGES IN THE COLOUR OF BUTTERFLIES PRODUCED THROUGH THE INFLUENCE OF TEMPERATURE.
The experiments of Dorfmeister, Weismann, Merrifield, Standfuss, and Fischer, on seasonal dimorphism and the aberration of colour in butterflies have so often been discussed in biological literature that a short reference to them will suffice. By seasonal dimorphism is meant the fact that species may appear at different seasons of the year in a somewhat different form or colour. Vanessa prorsa is the summer form, Vanessa levana the winter form of the same species. By keeping the pupae of Vanessa prorsa several weeks at a temperature of from 0 deg to 1 deg Weismann succeeded in obtaining from the summer chrysalids specimens which resembled the winter variety, Vanessa levana.
If we wish to get a clear understanding of the causes of variation in the colour and pattern of butterflies, we must direct our attention to the experiments of Fischer, who worked with more extreme temperatures than his predecessors, and found that almost identical aberrations of colour could be produced by both extremely high and extremely low temperatures. This can be clearly seen from the following tabulated results of his observations. At the head of each column the reader finds the temperature to which Fischer submitted the pupae, and in the vertical column below are found the varieties that were produced. In the vertical column A are given the normal forms:
(Temperatures in deg C.)
0 to -20 0 to +10 A. +35 to +37 +36 to +41 +42 to +46
(Normal forms)
ichnusoides polaris urticae ichnusa polaris ichnusoides
(nigrita) (nigrita)
antigone fischeri io - fischeri antigone
(iokaste) (iokaste)
testudo dixeyi polychloros erythromelas dixeyi testudo
hygiaea artemis antiopa epione artemis hygiaea
elymi wiskotti cardui - wiskotti elymi
klymene merrifieldi atalanta - merrifieldi klymene
weismanni porima prorsa - porima weismanni
The reader will notice that the aberrations produced at a very low temperature (from 0 to -20 deg C.) are absolutely identical with the aberrations produced by exposing the pupae to extremely high temperatures (42 to 46 deg C.). Moreover the aberrations produced by a moderately low temperature (from 0 to 10 deg C.) are identical with the aberrations produced by a moderately high temperature (36 to 41 deg C.)