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to two sets of forces-1, their initial energy and the direction of their aim; 2, the deflecting power of resisting objects or forces-or the different balls might roll with great velocity down a precipitous mountainside. In the first case velocity and direction of course would be determined largely by initial impulse; in the second, by the attraction of the earth and by the inequalities of its surface.
In evolution, environment, roughly speaking, corresponds to these deflecting or attracting external objects or forces; inherent tendencies to initial impulse. If we lay great weight on initial tendencies, inherent in protoplasm from the very beginning, we shall probably lay less stress on natural selection as a guiding, directing process.
The great botanist, Nägeli, has propounded a most ingenious and elaborate theory of evolution, as dependent mainly on inherent initial tendency. We can notice only one or two of its salient points. All development is, according to his view, due to a tendency in the primitive living substance toward more complete division of labor and greater complexity. This tendency, which he calls progression, or the tendency toward perfection, is the result of the chemical and molecular structure of the formative controlling protoplasm (idioplasm) of the body, and is transmitted with other parental traits from generation to generation. And structural complexity thus increases like money at compound interest. Development is a process of unfolding or of realization of the possibilities of this tendency under the stimulus of surrounding influences. Environment plays an essential part in his system. But only such changes are transmissible to
future generations as have resulted from modifications arising in the idioplasm. Descendants of plants which have varied under changed conditions revert, as a rule, to the old type, when returned to the old surroundings. And in the animal world effects of use and disuse are, according to his view, not transmissible.
Natural selection plays a very subordinate part. It is purely destructive. Given an infinity of place and nourishment—do away, that is, with all struggle and selection—and the living world would have advanced, purely by the force of the progressive tendency, just as far as it now has; only there would have survived an indefinite number of intermediate forms. It would have differed from our present living world as the milky way does from the starry firmament.
He compares the plant kingdom to a great, luxurious tree, branching from its very base, whose twigs would represent the present stage of our different species. Left to itself it would put out a chaos of innumerable branches. Natural selection, like a gardener, prunes the tree into shape. Childen might imagine that the gardener caused the growth; but the tree would have been broader and have branched more luxuriantly if left to itself.*
Every species must vary perpetually. Now this proposition is apparently not in accord with fact; for some have remained unchanged during immense periods. And natural selection, by removing the less fit, certainly appears to contribute to progress by raising the average of the species. The theory seems extreme and one-sided. And yet it has done great service by
* See Nägeli, "Theorie der Abstammungslehre," p. 18; also pp. 12, 118, 285.
calling in question the all-sufficiency of natural selection and the modifying power of environment, and by emphasizing, probably overmuch, the importance of initial inherent tendency, whose value has been entirely neglected by many evolutionists.
Lack of space compels us to leave unnoticed most of the exceedingly valuable suggestions of Nägeli's brilliant work.
It is still less possible to do any justice in a few words to Weismann's theory. Into its various modifications, as it has grown from year to year, we have no time to enter. And we must confine ourselves to his views of variation and heredity.
In studying protozoa we noticed that they reproduced by fission, each adult individual dividing into two young ones. There is therefore no old parent left to die. Natural death does not occur here, only death by violence or unfavorable conditions. The protozoa are immortal, not in the sense of the endless persistence of the individual, but of the absence of death. Heredity is here easily comprehensible, for one-half, or less frequently a smaller fraction, of the substance of the parent goes to form the new individual. There is direct continuity of substance from generation to generation.
But in volvox a change has taken place. The fertilized egg-cell, formed by the union of egg and spermatozoon, is a single cell, like the individual resulting from the conjugation or fusion of two protozoa. But in the many-celled individual, which develops out of the fertilized egg, there are two kinds of cells. 1. There are other egg-cells, like the first, each one of which can, under favorable conditions, develop into a
multicellular individual like the parent. And the germcells (eggs and spermatozoa) of volvox are immortal like the protozoa. But, 2, there are nutritive, somatic cells, which nourish and transport the germ-cells, and after their discharge die. These somatic cells, being mortal, differ altogether from the germ-cells and the protozoa. The protoplasm must differ in chemical, or molecular, or other structure in the two cases, and we distinguish the germ-plasm of the germ-cells, resembling in certain respects Nägeli's idioplasm, from somatoplasm, which performs most of the functions of the cell. The somatoplasm arises from, and hence must be regarded as a modification of, the germplasm. The germ-plasm can increase indefinitely in the lapse of generations, increase of the somatoplasm is limited.
When a new individual develops, a certain portion of the germ-plasm of the egg is set aside and remains unchanged in structure. This, increasing in quantity, forms the reproductive elements for the next generation. The germ-plasm, which does not form the whole of each reproductive element, but only a part of the nucleus, is thus an exceedingly stable substance. And there is a just as real continuity of germ-plasm through successive generations of volvox, or of any higher plants or animals, as in successive generations of protozoa.
In certain plants there is an underground stem or rootstock, which grows perennially, and each year produces a plant from a bud at its end. This underground rootstock would represent the continuous germ-plasm of successive generations; the plants which yearly arise from it would represent the successive generations of
adult individuals, composed mainly of somatoplasm. Or we may imagine a long chain, with a pendant attached to each tenth or one-hundredth link. The links of the chain would represent the series of generations of germ-cells; the pendants, the adults of successive generations.
But any leaf of begonia can be made to develop into a new plant, giving rise to germ-cells. Here there must be scattered through the leaves of the plant small portions of germ-plasm, which generally remain dormant, and only under special conditions increase and give rise to germ-cells.
A large part of the germ-plasm of the fertilized egg is used to give rise to the somatoplasm composing the different systems of the embryo and adult. Weismann's explanation of this change of germ-plasm into somatoplasm is very ingenious, and depends upon his theory of the structure of the germ-plasm; and this latter theory forms the basis of his theory of evolution. It would take too long to state his theory of the structure of germplasm, but an illustration may present fairly clear all that is of special importance
The molecules of germ-plasm are grouped in units, and these in an ascending series of units of continually increasing complexity, until at last we find the highest unit represented in the nucleus of the germcell. This grouping of molecules in units of increasing complexity is like the grouping of the men of an army in companies, regiments, brigades, divisions, etc.
To form the somatoplasm of the different tissues of the body, this complicated organization breaks up, as egg divides, into an ever-increasing number of cells.