Molluscs, Arthropods and Annelids; the first named group furnishes by far the most hosts, the parasites being frequently met with in domestic animals, both birds and mammals. Following from the casual method of infection, the epithelium of Habitat: effects on host. the gut or of its appendages (e.g. the liver [Plate I., fig. 1]) is a very common seat of the parasitic invasion. But in many cases Coccidia are found in other organs, to which they are doubtless carried by lymphatic or circulatory channels. In Molluscs, they often occur in the kidneys (fig. 2); in Insects, they are met with as “coelomic” parasites, the fat-bodies, pericardial cells, &c., being a favourite habitat; even the testis is not free from their attentions in one or two instances, though the ovary appears always immune.
The parasite invariably destroys its host-cell completely. The latter is at first stimulated to abnormal growth and activity and becomes greatly hypertrophied, the nucleus also undergoing karyolytic changes (fig. 4). The fatty materials elaborated by the host-cell are rapidly used up by the Coccidian, as nourishment; and at length the weakened and disorganized cell is no longer able to assimilate but dies and is gradually absorbed by the parasite, becoming reduced to a mere enclosing skin or envelope. In some cases (ex. Cyclospora caryolytica of the mole) the parasite is actually intranuclear, the nucleus becoming greatly swollen and transformed into a huge vacuole containing it.
The effects of a Coccidian infection upon the host as a whole depend largely upon the extent to which endogenous multiplication of the parasites takes place. On the one hand, schizogony may be so limited in extent as not to cause appreciable injury to the host. This seems to be often the case in forms infecting Molluscs and Arthropods. On the other hand, where schizogony is rapid and prolonged, the results are often serious. For, although any one individual only causes the death of a single host-cell, yet the number of the parasites may be so enormously increased by this means, that the entire affected epithelium may be overrun and destroyed. Thus are occasioned grave attacks of coccidiosis, characterized by severe enteritis and diarrhoea, which may end fatally. In the case of the Vertebrates, secondary causes, resulting from the stoppage of the bile ducts, also help to produce death. There is, however, one factor in the endangered animal’s favour. Schizogony cannot go on indefinitely; it has a limit, dependent upon the supply of host-cells, and consequently of nutriment, available. As this shows signs of becoming exhausted, by the rapid multiplication of the parasites, the latter begin to make preparations for the exogenous cycle, inaugurated by gametogony. When conjugation has taken place and sporogony is begun, the danger to the host is at an end. So that, if the acute stage of the disease is once successfully passed, the regenerative capacity of the epithelium may be able to restore something like equilibrium to the deranged metabolism in time to prevent collapse.
Coccidium schubergi, parasitic in the intestine of a centipede (Lithobius forficatus), may be taken as an example of a Coccidian life-history (see Schaudinn, 1900): some of the more important variations exhibited by other forms will be Morphology and life-history. noted afterwards. The trophozoite, or actively-growing parasite, is an oval or rounded body (fig. 3, I.). The general cytoplasm shows no differentiation into ectoplasm and endoplasm; it is uniformly alveolar in character. The nucleus is relatively large, and possesses a distinct membrane and a well-marked reticulum in which are embedded grains of chromatin. Its most conspicuous feature is the large deeply-staining karyosome, which consists of the greater part of the chromatin of the nucleus intimately bound up with a plastinoid basis. When fully grown, the trophozoite (now a schizont) undergoes schizogony. Its nucleus divides successively to form a number of nuclei, which travel to the periphery, and there become more or less regularly disposed (fig. 3, II. and III.). The protoplasm in the neighbourhood of each next grows out, as a projecting bud, carrying the nucleus with it. In this manner are formed a number of club-shaped bodies, the merozoites, which are at length set free from the parent-body (IV.), leaving a certain amount of residual cytoplasm behind. By the rupture of the disorganized host-cell, the fully-formed merozoites are liberated into the intestinal lumen, and seek out fresh epithelial cells. Each is more or less sickle-shaped, and capable of active movements. Once inside a new host-cell, the merozoite grows to a schizont again.
After this course has been repeated several times, gametogony sets in, the trophozoites growing more slowly and becoming the parent-cells of the sexual elements (gametocytes), either male individuals (microgametocytes) or female ones (megagametocytes). A microgametocyte (fig. 3, VI. ♂) is characterized by its dense but finely reticular or alveolar cytoplasm, very different from the loose structure of that of a schizont. The male elements (microgametes) are formed in a manner essentially comparable to that in which the formation of merozoites takes place. Although the details of the nuclear changes and divisions vary somewhat, the end-result is similar, a number of little nuclear agglomerations being evenly distributed at the surface (VII. ♂) Each of these elongates considerably, becoming comma-shaped and projecting from the gametocyte. Nearly all the body of the male gamete (VIII. ♂) consists of chromatin, the cytoplasm only forming a very delicate zone or envelope around the nucleus. From the cytoplasm two long fine flagella grow out, one of which originates at the anterior end, the other, apparently, at the hinder end, acting as a rudder; but it is probable that this also is developed at the anterior end and attached to the side of the body. By means of their flagella the numerous microgametes break loose from the body of the microgametocyte and swim away in search of a female element.
A megagametocyte (VI. ♀) is distinguished by its rather different shape, being more like a bean than a sphere until ripe for maturation, and by the fact that it stores up in its cytoplasm quantities of reserve nutriment in the form of rounded refringent plastinoid grains. Each female gametocyte gives rise to only a single female element (megagamete), after a process of nuclear purification. The karyosome is expelled from the nucleus into the cytoplasm, where it breaks up at once into fragments (VII. ♀). Meanwhile the gametocyte is becoming spherical, and its changes in shape aid in setting it free from the shrivelled host-cell. The fragments of the karyosome, which are, as it were, squeezed out to the exterior, exert a powerful attraction upon the microgametes, many of which swarm round the now mature megagamete. The female nucleus (pronucleus) approaches the surface of the cell (VIII. ♀), and at this spot a little clear cytoplasmic prominence arises (cone of reception). On coming into contact with this protuberance (probably attracted to it by the female pronucleus), a microgamete adheres. Partly by its own movements and partly by the withdrawal of the cone of attraction, the male penetrates into the female element and fertilization is accomplished. Only one microgamete can thus pass into the megagamete, for immediately its entry is effected a delicate membrane is secreted around the copula (zygote), which effectually excludes other less fortunate ones. This membrane rapidly increases in thickness and becomes the oocyst (IX.), and the copula is now ready to begin sporogony.
Sporogony goes on indifferently either inside the host or after the cyst has been passed out with the faeces to the exterior. The definitive nucleus of the zygote (resulting from the intimate fusion of the male and female pronuclei, by means of a somewhat elaborate “fertilization-spindle” [X.]) gives rise by successive direct divisions to four nuclei (XII.), around which the protoplasm becomes segregated; these segments form the four sporoblasts. Around each sporoblast two membranes are successively secreted (exospore and endospore), which constitute the sporocyst (XIII.); the sporocyst and its contents forming the spore. The nucleus of each spore next divides, again directly, and this is followed by the division of the cytoplasm. As a final result, each of the four spores contains two germs (sporozoites), and a certain amount of residual protoplasm (fig. 3, IV.); this latter encloses a viscid, vacuole-like body, which aids in the subsequent dehiscence of the sporocyst. On being eaten by a fresh host, the wall of the oocyst is dissolved at a particular region by the
- It is important to note that in schizogony there is never any cyst or cyst-membrane formed around the parasite.