centrosome during the division of the nucleus, and sometimes true centrosomes are present. Flagella also commonly arise from basal granules of a centrosomic nature, blepharoplasts in the correct sense of the term; these blepharoplasts are always in connexion with the nucleus, or with the kinetic nucleus if there is one distinct from the trophic nucleus, as in the genus Trypanosoma and allied forms.
Reproduction of the Protozoa.—The mode of reproduction in these organisms is the same as that of the cell generally, and takes always the form of fission of some kind; that is to say, of division of the body into smaller portions, each of which represents a young individual. The division of the body is preceded by that of the nucleus, if single, or of each nucleus in the cases where there are two different nuclei; if, however, more than one nucleus of the same kind be present, the nuclei may be simply shared amongst the daughter-individuals, this mode of division being known as plasmotomy. Other organs of the body may either, like the nucleus, undergo fission, or may be formed afresh in the daughter-individuals.
The division of the nucleus in Protozoa may take place by the direct method or by means of mitosis. Direct division, without mitosis, is of very common occurrence; the division may be simple or multiple, that is to say, into only two parts, or into a number of fragments formed simultaneously. An extreme case of multiple fission is seen in the formation of the microgametes of Coccidium schubergi, where the nucleus breaks up into a great number of chromidia, which become concentrated in patches to form the several daughter-nuclei. In some cases, on the other hand, multiple daughter-nuclei are formed by rapidly repeated simple division of the parent nucleus. The mode of division may be different in different nuclei of the same individual; thus in the Infusoria the macronucleus divides by direct division, the micronucleus by mitosis.
The mitosis of the Protozoa is far from being of the uniform stereotyped pattern seen in the Metazoa, but, as might have been expected, often shows a much simpler and more primitive condition. Centrosomes are often absent, and their place may be taken, as stated above, by other bodies. The nuclear membrane may be retained throughout the mitosis. Definite chromosomes can, as a rule, be made out, but the chromosomes are often very numerous and minute, without definite form, and divide irregularly. Much remains to be done in studying the mitosis of the Protozoa, but it is probable that wider knowledge will show many conditions intermediate between direct division and perfect mitosis.
The simplest method of fission in Protozoa is that termed binary, where the body divides into two halves, which may be equal and similar, so that the result is two sister-individuals impossible to distinguish as parent and offspring. In many cases of binary fission, however, the resulting daughter-individuals may be markedly unequal in size, so that one may be distinguished as the parent, the other as the offspring. If the daughter-individual be relatively very small, and formed in a more or less imperfect condition at first, the process is termed gemmation or budding. The buds formed in this way may be either external, formed on the surface of the body, or internal, that is, formed in special internal cavities, from which the offspring are later set free, as in many Acinetaria. Gemmation may be correlated with multiple nuclear fission in such a way that buds are formed over the whole body surface of the organism, which thereby undergoes a process of simultaneous multiple fission into numerous daughter-individuals. Rapid multiple fission of this kind is termed sporulation, and is a form of reproduction which is of common occurrence, especially in parasitic forms. Usually the central portion of the parent body remains over as a residual body (Restkörper) , but sometimes the parent organism is entirely resolved into the daughter-individuals, which are termed spores in a general way, but can be given special names in special cases (see Gregarines, Coccidia, &c.).
Life-cycles of the Protozoa.—It is probable that in all Protozoa, as in the Metazoa, the life-history takes its course in a series of recurrent cycles of greater or less extent, a fixed point, as it were, in the cycle being marked by the act of syngamy, or conjugation, which represents, apparently, a process for recuperation of the waning vital powers of the organism. It is true that in many types of Protozoa syngamy is not known as yet to occur, but in all species which have been thoroughly investigated syngamy in some form has been observed, and there is nothing to lead to the belief that the sexual process is not of universal occurrence in the Protozoa.
The life-cycle of a given species may be very simple or it may be extremely complex, the organism occurring under many different forms at different phases or periods of its development. The polymorphism of the Protozoa is best considered under three categories, according to the three main causes to which it is due, namely, first, polymorphism due to adaptation to different conditions of existence; secondly, polymorphism due to differences of size and structure during growth; thirdly, polymorphism due to the differentiation of individuals in connexion with the process of syngamy or sexual conjugation.
1. Polymorphism in Relation to Life-conditions.—As a protection against unfavourable conditions, or for other reasons, most Protozoa have the power of passing into a resting condition, during which the vital functions may be wholly or in part suspended. In the resting phase the animal usually becomes enveloped in a resistant membrane or cyst secreted by it, and is then said to be encysted. The formation of a cyst may be a response to conditions of various kinds. Very commonly it is formed to protect the organism against a change of medium, as in the case of freshwater forms liable to desiccation, or of parasites about to pass out of the bodies of their hosts. In other cases the organism passes into the resting state in order to absorb ingested nutriment or in order to enter upon reproductive phases.
As a preparation for encystment, organs of locomotion, if present, are retracted or cast off; contractile vacuoles cease to be formed; and the food-vacuoles disappear, usually by digestion of their contents and rejection of the waste residue. The body becomes rounded off and more or less spherical in form, and the protoplasm becomes denser, that is, less fluid and more opaque, but at the same time of diminished specific gravity, by loss of water. The cyst is then secreted at the surface as a layer of varying thickness and toughness. In the encysted condition many Protozoa are capable of being transported by the wind, a fact which explains their appearance in infusions and liquids exposed to the air. In favourable conditions the cysts germinate, that is to say, the envelope is dissolved and the contained organism or organisms are set free to enter upon the strenuous life once more.
In the Mycetozoa, organisms adapted to a semi-terrestrial life in moist surroundings, the protoplasm is capable, when desiccated, of passing into a tough condition resembling sealing-wax, which, when moistened, assumes again its normal appearance and active condition.
Resting phases, analogous to encystment, are seen in the spores of various forms, especially those of parasitic habit, which are commonly enclosed in tough, resistant envelopes or sporocysts, and enveloped as a protection against change of medium or of host. Within the sporocyst multiplication of the sporoplasm may take place to form more or fewer sporozoites. The sporocysts usually show definite symmetry and structure, infinitely variable in different species. In a suitable medium the spores germinate by rupture of the sporocysts and escape of the contents.
2. Polymorphism in Relation to Growth and Development.—In many species of Protozoa there is hardly any difference to be observed between different individuals during their active phases except in size. Those individuals about to multiply by fission are slightly above the normal in dimensions: on the
- The kinetic nucleus of Trypanosoma is sometimes, but in the writer’s opinion wrongly, named centrosome or blepharoplast; the bodies to which cytologists give these names are achromatic bodies; the kinetic nucleus is a true chromatic nucleus. The question of the centrosome in Protozoa is discussed by R. Goldschmidt and M. Popoff.