By JACQUES LOEB, M.D.
Professor of Physiology in the University of California.
I. INTRODUCTORY REMARKS.
What the biologist calls the natural environment of an animal is from a physical point of view a rather rigid combination of definite forces. It is obvious that by a purposeful and systematic variation of these and by the application of other forces in the laboratory, results must be obtainable which do not appear in the natural environment. This is the reasoning underlying the modern development of the study of the effects of environment upon animal life. It was perhaps not the least important of Darwin's services to science that the boldness of his conceptions gave to the experimental biologist courage to enter upon the attempt of controlling at will the life-phenomena of animals, and of bringing about effects which cannot be expected in Nature.
The systematic physico-chemical analysis of the effect of outside forces upon the form and reactions of animals is also our only means of unravelling the mechanism of heredity beyond the scope of the Mendelian law. The manner in which a germ-cell can force upon the adult certain characters will not be understood until we succeed in varying and controlling hereditary characteristics; and this can only be accomplished on the basis of a systematic study of the effects of chemical and physical forces upon living matter.
Owing to limitation of space this sketch is necessarily very incomplete, and it must not be inferred that studies which are not mentioned here were considered to be of minor importance. All the writer could hope to do was to bring together a few instances of the experimental analysis of the effect of environment, which indicate the nature and extent of our control over life-phenomena and which also have some relation to the work of Darwin. In the selection of these instances preference is given to those problems which are not too technical for the general reader.
The forces, the influence of which we shall discuss, are in succession chemical agencies, temperature, light, and gravitation. We shall also treat separately the effect of these forces upon form and instinctive reactions.
II. THE EFFECTS OF CHEMICAL AGENCIES.
(a) HETEROGENEOUS HYBRIDISATION.
It was held until recently that hybridisation is not possible except between closely related species and that even among these a successful hybridisation cannot always be counted upon. This view was well supported by experience. It is, for instance, well known that the majority of marine animals lay their unfertilised eggs in the ocean and that the males shed their sperm also into the sea-water. The numerical excess of the spermatozoa over the ova in the sea-water is the only guarantee that the eggs are fertilised, for the spermatozoa are carried to the eggs by chance and are not attracted by the latter. This statement is the result of numerous experiments by various authors, and is contrary to common belief.
As a rule all or the majority of individuals of a species in a given region spawn on the same day, and when this occurs the sea-water constitutes a veritable suspension of sperm. It has been shown by experiment that in fresh sea-water the sperm may live and retain its fertilising power for several days. It is thus unavoidable that at certain periods more than one kind of spermatozoon is suspended in the sea-water and it is a matter of surprise that the most heterogeneous hybridisations do not constantly occur. The reason for this becomes obvious if we bring together mature eggs and equally mature and active sperm of a different family. When this is done no egg is, as a rule, fertilised. The eggs of a sea-urchin can be fertilised by sperm of their own species, or, though in smaller numbers, by the sperm of other species of sea-urchins, but not by the sperm of other groups of echinoderms, e.g. starfish, brittle-stars, holothurians or crinoids, and still less by the sperm of more distant groups of animals.
The consensus of opinion seemed to be that the spermatozoon must enter the egg through a narrow opening or canal, the so-called micropyle, and that the micropyle allowed only the spermatozoa of the same or of a closely related species to enter the egg.
It seemed to the writer that the cause of this limitation of hybridisation might be of another kind and that by a change in the constitution of the sea-water it might be possible to bring about heterogenous hybridisations, which in normal sea-water are impossible. This assumption proved correct.
Sea-water has a faintly alkaline reaction (in terms of the physical chemist its concentration of hydroxyl ions is about (10 to the power minus six)N at Pacific Grove, California, and about (10 to the power minus 5)N at Woods Hole, Massachusetts). If we slightly raise the alkalinity of the sea-water by adding to it a small but definite quantity of sodium hydroxide or some other alkali, the eggs of the sea-urchin can be fertilised with the sperm of widely different groups of animals, possibly with the sperm of any marine animal which sheds it into the ocean. In 1903 it was shown that if we add from about 0.5 to 0.8 cubic centimetre N/10 sodium hydroxide to 50cubic centimetres of sea-water, the eggs of Strongylocentrotus purpuratus (a sea-urchin which is found on the coast of California) can be fertilised in large quantities by the sperm of various kinds of starfish, brittle-stars and holothurians; while in normal sea-water or with less sodium hydroxide not a single egg of the same female could be fertilised with the starfish sperm which proved effective in the hyper-alkaline sea-water. The sperm of the various forms of starfish was not equally effective for these hybridisations; the sperm of Asterias ochracea and A. capitata gave the best results, since it was possible to fertilise 50 per cent or more of the sea-urchin eggs, while the sperm of Pycnopodia and Asterina fertilised only 2 per cent of the same eggs.