of the environment the necessity for the sexual process may be diminished and the event may be deferred for a long time, if not indefinitely. Hence it is quite possible that in many Protozoa the process of syngamy may be in abeyance, just as there are plants which can be propagated indefinitely by suckers or cuttings without ever setting seed; and it is possible that the inoculative or artificial transmission of parasitic Protozoa from one host to another, as in the case of pathogenic trypanosomes, without any apparent diminution in their vital powers, is an instance of this kind.
As a general rule, in order that syngamy may be attended by beneficial results to the organism, it is necessary that the two conjugating individuals should be from different strains, that is to say, they should not be nearly related by descent and parentage. Thus F. Schaudinn found that in order to observe the sexual union of the gametes of Foraminifera it was necessary to bring together gametes of distinct parentage. On the other hand it has been observed that in many Protozoa, especially in parasitic forms, syngamy takes place between individuals of common parentage. Thus in Amoeba coli, according to F. Schaudinn, a single individual becomes encysted and its nucleus divides into two; after each nucleus has undergone certain maturative changes they give rise to pronuclei which conjugate and initiate a new developmental cycle. Syngamy between sister individuals, or autogamy, as it has been termed, is not, however, confined to parasitic Protozoa; it has been observed in Actinosphaerium by R. Hertwig. The benefit to the organism, if any, arising from autogamy can only be supposed to result from the rearrangement and reconstitution of the nuclear apparatus. The frequent occurrence of autogamy suggests that in many Protozoa the nature of the environment diminishes the importance of the sexual process, at least so far as the mixture of nuclear material from distinct sources is concerned; and, since autogamy is most common in parasitic forms, this result may, in the light of G. N. Calkins's experiments, be ascribed in great part to the frequent changes of environment and nutrition to which parasitic forms, above all, are subject.
True syngamy consists, as has been said, of nuclear fusion or karyogamy. It rarely, if ever, happens, however, that such fusion takes place without the conjugating nuclei having undergone some process of reduction by elimination of a portion of the nuclear substance, in a manner analogous to the maturation of the germ-cells in the Metazoa. The chromatin thus eliminated may be cast out from the body of the organism as one or more so-called polar bodies; or may be absorbed in the cytoplasm; or may remain in the cytoplasm and be left over in the residual protoplasm in cases where syngamy is followed by a process of rapid multiplication by sporulation; but in all cases the chromatin removed from the nucleus is rejected in some way or other and plays no part in the subsequent development of the organism. The nuclei of the gametes which have completed this process of épuration nucléaire are then ripe for syngamic fusion and are termed pronuclei; the union of two pronuclei produces a single nucleus termed a synkaryon.
It is certain that in many, if not in all, cases the nuclear substance that is rejected as a preliminary to syngamy consists of somatic or vegetative chromatin; that is to say, of chromatin that has been functional in regulating the ordinary vital functions, metabolism, growth, reproduction, &c., during previous generations, and has become effete; while on the other hand the chromatin that persists to form the pronuclei is generative chromatin which has remained in reserve for the sexual act and has retained its peculiar powers and properties unimpaired. The truth of this explanation is extremely obvious in such forms as the Infusoria, where somatic and generative chromatin are concentrated into two distinct and entirely separate nuclei. In some Rhizopoda also the body contains one or more principal nuclei and a mass of chromidia, and it has been observed that as a preparation for syngamy the principal nuclei are eliminated and the pronuclei are formed from the chromidia; in such cases, therefore, it is reasonable to regard the principal nuclei as representing somatic chromatin, the chromidia as generative chromatin. In other cases, however, for example Actinosphaerium, the chromidia must be interpreted, from their behaviour, as somatic chromatin, and the principal nuclei as generative chromatin; hence R. Goldschmidt has proposed the special term sporetia for those chromidia which represent reserve generative chromatin. In the majority of Protozoa, however, the nuclear substance is not differentiated in such a way that it can be distinguished by any visible peculiarities into somatic and generative chromatin.
The process of reduction is not limited, apparently, to the elimination of somatic chromatin, but a portion of the generative chromatin is also cast off. Thus in the Infusoria not only the somatic macronucleus, but also a considerable portion of the generative micronucleus, is absorbed at each act of conjugation. The elimination of generative chromatin is perhaps of importance as a factor in heredity and the production of variations, or possibly for sex determination, as will be discussed below; it is difficult to suggest any other explanations for it, unless it be supposed that during the exercise of ordinary vital functions a portion of the generative chromatin be rendered effete as well as the somatic chromatin.
From the considerations set forth in the foregoing paragraphs it must be supposed that the synkaryon, the fusion-product of the two pronuclei in syngamy, consists at first purely of generative chromatin, which must speedily become differentiated into the regulative somatic chromatin of the ensuing generations and the generative chromatin held in reserve for the next act of syngamy. Such a differentiation can be actually observed in the Infusoria, where immediately after conjugation the synkaryon divides into one or more pairs of nuclei, each pair becoming the two unequally sized nuclei of an ordinary individual, sometimes with, even at this stage, an apparently wanton elimination of nuclear substance. Thus the somatic and generative chromatin of the Protozoa offer a certain analogy with the soma and germ-plasm of Metazoa; but in making such comparisons the distinction between a physiological analogy and a morphological homology should be borne clearly in mind.
It has been stated above that the two gametes of a given species of Protozoa may be perfectly similar and indistinguishable, or may be very different one from the other. The condition with similar gametes is termed isogamy, that with differentiated gametes anisogamy. Every transition can be found from complete isogamy and pronounced anisogamy in the Protozoa; in tracing, however, the evolution of specialized gametes it must be remembered that we are dealing only with visible morphological differences mainly of an adaptive nature, without prejudice to the question of the possible existence of a fundamental sexual antithesis in all gametes, present even when not perceptible. The sex philosopher O. Weininger has urged that sex is a fundamental attribute of living things, and that the living substance, protoplasm, consists of arrhenoplasm and thelyplasm united in varying proportions. Certain observations of F. Schaudinn tend to support this view; in Trypanosoma noctuae, for example, Schaudinn found that the process of reduction in one gamete took an opposite course to that which it took in the other gamete. In one gamete certain portions of the nucleus were retained and certain other portions rejected; in the maturation of the other gamete the portions rejected and the portions retained were the reverse. Hence Schaudinn was led to regard the indifferent individuals as essentially hermaphrodite in nature, and therefore capable of giving rise to gametes of either order by elimination of one or the other set of sexual elements; a theory which throws further light on the elimination of generative chromatin mentioned above. It is possible, therefore, that the gametes of Protozoa may possess sexual characters intrinsically different even when perfectly similar so far as can be perceived. It is very probable, for instance, that the isogamy in Gregarines is a state of things derived secondarily from a primitive condition of anisogamy (see Gregarines).
The simplest possible condition of the gametes is seen in the free-swimming Ciliata, forms which in other respects are the
