are useful, since they procure for the colony an advantage in the struggle for existence, and they are therefore preserved by natural selection. Even the sterility itself in this case is not disadvantageous, since the fertility of the true females has at the same time considerably increased. We may therefore regard the sterile forms of ants, which have gradually been adapted in several directions to varying functions, AS A CERTAIN PROOF that selection really takes place in the germ-cells of the fathers and mothers of the workers, and that SPECIAL COMPLEXES OF PRIMORDIA (IDS) are present in the workers and in the males and females, and these complexes contain the primordia of the individual parts (DETERMINANTS). But since all living entities vary, the determinants must also vary, now in a favourable, now in an unfavourable direction. If a female produces eggs, which contain favourably varying determinants in the worker-ids, then these eggs will give rise to workers modified in the favourable direction, and if this happens with many females, the colony concerned will contain a better kind of worker than other colonies.
I digress here in order to give an account of the intimate processes, which, according to my view, take place within the germ-plasm, and which I have called "GERMINAL SELECTION." These processes are of importance since they form the roots of variation, which in its turn is the root of natural selection. I cannot here do more than give a brief outline of the theory in order to show how the Darwin-Wallace theory of selection has gained support from it.
With others, I regard the minimal amount of substance which is contained within the nucleus of the germ-cells, in the form of rods, bands, or granules, as the GERM-SUBSTANCE or GERM-PLASM, and I call the individual granules IDS. There is always a multiplicity of such ids present in the nucleus, either occurring individually, or united in the form of rods or bands (chromosomes). Each id contains the primary constituents of a WHOLE individual, so that several ids are concerned in the development of a new individual.
In every being of complex structure thousands of primary constituents must go to make up a single id; these I call DETERMINANTS, and I mean by this name very small individual particles, far below the limits of microscopic visibility, vital units which feed, grow, and multiply by division. These determinants control the parts of the developing embryo—in what manner need not here concern us. The determinants differ among themselves, those of a muscle are differently constituted from those of a nerve-cell or a glandular cell, etc., and every determinant is in its turn made up of minute vital units, which I call BIOPHORS, or the bearers of life. According to my view, these determinants not only assimilate, like every other living unit, but they VARY in the course of their growth, as every living unit does; they may vary qualitatively if the elements of which they are composed vary, they may grow and divide more or less rapidly, and their variations give rise to CORRESPONDING variations of the organ, cell, or cell-group which they determine. That they are undergoing ceaseless fluctuations in regard to size and quality seems to me the inevitable consequence of their unequal nutrition; for although the germ-cell as a whole usually receives sufficient nutriment, minute fluctuations in the amount carried to different parts within the germ-plasm cannot fail to occur.
Now, if a determinant, for instance of a sensory cell, receives for a considerable time more abundant nutriment than before, it will grow more rapidly—become bigger, and divide more quickly, and, later, when the id concerned develops into an embryo, this sensory cell will become stronger than in the parents, possibly even twice as strong. This is an instance of a HEREDITARY INDIVIDUAL VARIATION, arising from the germ.
The nutritive stream which, according to our hypothesis, favours the determinant N by chance, that is, for reasons unknown to us, may remain strong for a considerable time, or may decrease again; but even in the latter case it is conceivable that the ascending movement of the determinant may continue, because the strengthened determinant now ACTIVELY nourishes itself more abundantly—that is to say, it attracts the nutriment to itself, and to a certain extent withdraws it from its fellow-determinants. In this way, it may—as it seems to me—get into PERMANENT UPWARD MOVEMENT, AND ATTAIN A DEGREE OF STRENGTH FROM WHICH THERE IS NO FALLING BACK. Then positive or negative selection sets in, favouring the variations which are advantageous, setting aside those which are disadvantageous.
In a similar manner a DOWNWARD variation of the determinants may take place, if its progress be started by a diminished flow of nutriment. The determinants which are weakened by this diminished flow will have less affinity for attracting nutriment because of their diminished strength, and they will assimilate more feebly and grow more slowly, unless chance streams of nutriment help them to recover themselves. But, as will presently be shown, a change of direction cannot take place at EVERY stage of the degenerative process. If a certain critical stage of downward progress be passed, even favourable conditions of food-supply will no longer suffice permanently to change the direction of the variation. Only two cases are conceivable; if the determinant corresponds to a USEFUL organ, only its removal can bring back the germ-plasm to its former level; therefore personal selection removes the id in question, with its determinants, from the germ-plasm, by causing the elimination of the individual in the struggle for existence. But there is another conceivable case; the determinants concerned may be those of an organ which has become USELESS, and they will then continue unobstructed, but with exceeding slowness, along the downward path, until the organ becomes vestigial, and finally disappears altogether.
The fluctuations of the determinants hither and thither may thus be transformed into a lasting ascending or descending movement; and THIS IS THE CRUCIAL POINT OF THESE GERMINAL PROCESSES.
This is not a fantastic assumption; we can read it in the fact of the degeneration of disused parts. USELESS ORGANS ARE THE ONLY ONES WHICH ARE NOT HELPED TO ASCEND AGAIN BY PERSONAL SELECTION, AND THEREFORE IN THEIR CASE ALONE CAN WE FORM ANY IDEA OF HOW THE PRIMARY CONSTITUENTS BEHAVE, WHEN THEY ARE SUBJECT SOLELY TO INTRA-GERMINAL FORCES.
The whole determinant system of an id, as I conceive it, is in a state of continual fluctuation upwards and downwards. In most cases the fluctuations will counteract one another, because the passive streams of nutriment soon change, but in many cases the limit from which a return is possible will be passed, and then the determinants concerned will continue to vary in the same direction, till they attain positive or negative selection-value. At this stage personal selection intervenes and sets aside the variation if it is disadvantageous, or favours—that is to say, preserves—it if it is advantageous. Only THE DETERMINANT OF A USELESS ORGAN IS UNINFLUENCED BY PERSONAL SELECTION, and, as experience shows, it sinks downwards; that is, the organ that corresponds to it degenerates very slowly but uninterruptedly till, after what must obviously be an immense stretch of time, it disappears from the germ-plasm altogether.
Thus we find in the fact of the degeneration of disused parts the proof that not all the fluctuations of a determinant return to equilibrium again, but that, when the movement has attained to a certain strength, it continues IN THE SAME DIRECTION. We have entire certainty in regard to this as far as the downward progress is concerned, and we must assume it also in regard to ascending variations, as the phenomena of artificial selection certainly justify us in doing. If the Japanese breeders were able to lengthen the tail feathers of the cock to six feet, it can only have been because the determinants of the tail-feathers in the germ-plasm had already struck out a path of ascending variation, and this movement was taken advantage of by the breeder, who continually selected for reproduction the individuals in which the ascending variation was most marked. For all breeding depends upon the unconscious selection of germinal variations.
Of course these germinal processes cannot be proved mathematically, since we cannot actually see the play of forces of the passive fluctuations and their causes. We cannot say how great these fluctuations are, and how quickly or slowly, how regularly or irregularly they change. Nor do we know how far a determinant must be strengthened by the passive flow of the nutritive stream if it is to be beyond the danger of unfavourable variations, or how far it must be weakened passively before it loses the power of recovering itself by its own strength. It is no more possible to bring forward actual proofs in this case than it was in regard to the selection-value of the initial stages of an adaptation. But if we consider that all heritable variations must have their roots in the germ-plasm, and further, that when personal selection does not intervene, that is to say, in the case of parts which have become useless, a degeneration of the part, and therefore also of its determinant must inevitably take