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GREGARINES (mod. Lat. Gregarina, from...

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Originally appearing in Volume V12, Page 561 of the 1911 Encyclopedia Britannica.
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GREGARINES (mod. See also:Lat. Gregarina, from gregarius, See also:collecting in a See also:flock or See also:herd, grex) a large and abundant See also:order of See also:Sporozoa See also:Ectospora, in which a very high degree of morphological specialization and cytological differentiation of the See also:cell-See also:body is frequently found. On the other See also:hand, the See also:life-See also:cycle is, in See also:general, fairly See also:simple. Other See also:principal characters which distinguish Gregarines from allied Sporozoan parasites are as follows:—The fully-grown adult (trophozoite) is always " See also:free " in some See also:internal cavity, i.e. it is extracellular; in nearly all cases See also:prior to sporulation two Gregarines (associates) become attached to one another, forming a couple (See also:syzygy), and are surrounded by a See also:common cyst; inside the cyst the body of each See also:associate becomes segmented up into a number of sexual elements (gametes, See also:primary sporoblasts), which then conjugate in pairs; the resulting copula (zygote, definitive sporoblast) becomes usually a spore by the secretion of spore-membranes (sporocyst), its See also:protoplasm (sporoplasm) dividing up to See also:form the germs (sporozoites). F. Redi (1684) is said to have been the first to observe a Gregarine See also:parasite, but his claim to this See also:honour is by no means certain. Much later (1787) Cavolini described See also:Historical. and figured an indubitable Gregarine (probably the form now known as Aggregata conformis) from a Crustacean (Pachygrapsus), which, however, he regarded as a tapeworm. See also:Leon See also:Dufour, who in his researches on See also:insect See also:anatomy came across several See also:species of these parasites, also considered them as allied to the See also:worms and proposed the generic name of Gregarine. The unicellular nature of Gregarines was first realized by A. von See also:Kolliker, who from 1845–1848 added considerably to our know-ledge of the frequent occurrence and wide See also:distribution of these organisms. Further progress was due to F. See also:Stein who demonstrated about this See also:time the relation of the "pseudonavieellae" (spores) to the re-See also:production of the parasites.

Apart from the continually in-creasing number of known species, matters remained at about this See also:

stage for many years. It is, in fact, only since the closing years of the 19th See also:century that the See also:complete life-See also:history has been fully worked out; this has now been done in many cases, thanks to the researches of M. Siedlecki, L. Cuenot, L. Leger, 0. Duboscq, A. Laveran, M. From wasielewski's Sporozeenkunde, after See also:Pfeiffer. a U 11 e r y, F. C FIG. I.—a, Transverse See also:Section of See also:Intestine of Mealworm, infected with Gregarina (Clepsydeina) M e s n i l and polymorpha; b, See also:Part of a highly magnified. others, to whom also we owe most of our knowledge regarding the relations of the parasites to the cells of their See also:host during their See also:early development. Gregarines are essentially parasites of Invertebrates; they are not known to occur in any true Vertebrate although met with in occur- Ascidians. By far the greatest number of hosts is rence; furnished by the Arthropods.

Many members of the mode of various See also:

groups of worms (especially the Annelids) 3°fO°' also See also:harbour the parasites, and certain very interesting forms are found in Echinoderms; in the other classes, they either occur only sporadically or else are absent. Infection is invariably of the accidental (casual) type, by way of the alimentary See also:canal, the spores being usually swallowed by the host when feeding; a novel variation of this method has been described by See also:Woodcock (31) in the See also:case of a Gregarine parasitic in Cucumaria, where the spores are sucked up through the See also:cloaca into the See also:respiratory trees, by the inhalant current. The favourite See also:habitat is either the intestine (fig. I) or its diverticula (e.g. the Malpighian tubules), or the body-cavity. From Wasielewski, after Leger. In the latter case, after infection has occurred, the liberated germs at once See also:traverse the intestinal epithelium. They may come to See also:rest in the connective See also:tissue of the sub-mucosa (remain- 1 Figures 1, 2, 6, 7, 10, II, 12 and 16 are redrawn from Wasielewski's Sporozoenkunde, by permission of the author and of the publisher, Gustav See also:Fischer, See also:Jena.See also:ing, however, extracellular), grow considerably in that situation, and ultimately fall into the body-cavity (e.g. Diplocystis); or they may pass straightway into the body-cavity and Habitat there come into relation with some See also:organ or tissue (e.g. and effect, Monocystis of the See also:earthworm, which is for a time See also:intra- on host_ cellular in the spermatoblasts (fig. 4, c). In the case of intestinal Gregarines, the behaviour of the See also:young trophozoite with respect to the See also:epithelial cells of its host varies greatly. The parasite may remain only attached to the host-cell, never becoming actually intracellular (e.g. Pterocephalus); more usually it penetrates partially into it, the extracellular portion of the Gregarine, however, giving rise subsequently to most of the adult (e.g.

Gregarina); or lastly, in a few forms, the early development is entirely intracellular (e.g. Lankesteria, See also:

Steno pkara) The effects on the host are confined to the parasitized cells. These generally undergo at first marked See also:hypertrophy and alteration in See also:character; this See also:condition is succeeded by one of See also:atrophy, when the substance of the cell becomes in one way or another practically absorbed by the growing parasite (cf. also See also:COCCIDIA). Since, however, the Gregarines never over-run their hosts in the way that many other Sporozoa do (because of their lack, in general, of the See also:power of endogenous multiplication), the number of cells of any tissue attacked, even in the case of a strong infection, is only a From Lankester. very small percentage of the whole. In See also:short the hosts ds noi, as a See also:rule, suffer any appreciable inconvenience from the presence of the parasites. The body of a Gregarine is always of a definite shape, usually See also:oval C. From Lanke ter, after various authors. a-c,Trophozoites of Monocystis agilis. a and b, Young individuals showing changes of body-form. c, Older individual, still enveloped in a coat of spermatozoa. d, e, Trophozoites of M. magna attached to seminal See also:funnel of Lumbricus. See also:Goblet-shaped epithelial cells, in which the extremity of the parasite is inserted.

a or elongated; in one or two instances (e.g. Diplodina) it is spherical, Morpho- and, on the other hand, in Porospora (fig. 3) it is greatly See also:

drawn out and vermiform. In many adult Gregarines, See also:foy' the body is divided into two distinct but unequal regions or halves, the anterior part being known as the protomerite, the hinder, generally the larger, as the deutomerite. This feature is closely associated with another important morphological character, one which is observable, however, only during the earlier stages of growth and development, namely, the presence of a definite organ, the epimerite, which serves for the See also:attachment of the parasite to the host-cell (fig. 6). In those Gregarines (most intestinal forms) which become attached to an epithelial cell, the attachment occurs by means of a See also:minute See also:projection or See also:beak (rostrum) at the anterior end of the sporozoite, which pushes its way into the cell, followed by the first part of the growing germ. This portion of the body in-creases in See also:size much quicker at first than the rest (the extracellular part), more or less fills up the host-cell, and forms the well-See also:developed epimerite or secondary attaching organella. The extracellular part of the Gregarine next grows rapidly, and a transverse septum is formed at a short distance away from- (outside) the point where the body penetrates into the cell (fig. 6) ; this marks off the large deutomerite posteriorly (distally). Leger thinks that this See also:partition most likely owes its origin to trophic considerations, i.e. to the slightly different manner in which the two halves of the young parasite (the proximal, largely intracellular-part, and the distal, extracellular one) may be supposed to obtain their nutriment. In the case of the one See also:half, the host-cell supplies the nutriment, in that of the other, the intestinal liquid ; and the septum is, as it were, the expression of the conflicting limit between these two methods.

Nevertheless, the See also:

present writer does not think that See also:mechanical considerations should be altogether See also:left out of See also:account. The septum may also be, to some extent, an adaption for strengthening the body of the fixed parasite against lateral thrusts or strains, due to the impact of See also:foreign bodies (See also:food, &c.) in the intestine. At the point where the body becomes actually intracellular, it is constricted, and this constriction marks off the epimerite (internally) from the See also:middle portion (between this point and the septum), which is the protomerite. Further growth is restricted, practically, to the extracellular regions, and the epimerite often comes to appear ultimately as a small appendage at the anterior end of the protomerite. A Gregarine at this stage is known as a cephalont. Later on, the parasite breaks loose from the host-cell and becomes free in the lumen, the separation taking See also:place at the constriction between the protomerite and the epimerite; the latter is left behind in the remains of the host-cell, the former becomes the anterior part of the free trophozoite. In other Gregarines, however, those, namely, which pass inwards, ultimately becoming ` coelomic," as well as those which become entirely intracellular, no epimerite is ever developed, and, further, the body remains single or unseptate. These forms, which include, , for instance, Monocystis (fig. 4), Lankesteria, Diplocystis, are distinguished, as Acephalina or Aseptata (Haplocyta, Monocystida), ac-'cording to which character is referred.to, from the others, termed Cephalina or Septata (Polycystida). The two sets of terms are not, however, completely identical or interchangeable, for there are a few forms which possess an epimerite, but which lack the See also:division into protomerite and deutomerite, and are hence known as Pseudomonocystida; this condition may be See also:primitive (Dopiocystis) or (possibly) secondary, the partition having In course of time disappeared. Again, Stenophora is a septate formwhich has become, secondarily, completely intracellular during the young stages, and, doubtless correlated with this, shows no sign of an epimerite. With regard to the epimerites themselves, they are of all variety of form and shape and need not be described in detail (fig.

7). In one or two cases, however, another variety of attaching organella is met with. Thus in Pterocephalus, only the rostrum of the sporozoite t 1 s 8 s 2 a From Wasielewski, after Leger. z, Gregarina longa. 6, Contetoides crinitus. 2, Sycia inopinata. 7, Geneiorhynchus monnieri. 3, Pileocephalus heerii. 8, Echinomera hispida. 4, Stylorhynchus longicollis. 9, Pterocephalus nobilis. 5, Beloides firmus.

penetrates into the host-cell, and no epimerite is formed. Instead, a number of See also:

fine See also:root-like processes are developed from near the anterior end, which pass in between the host-cells (fig. 5) and thus See also:anchor the parasite firmly. Similarly, in the curious Schizogregarinae, the anterior end of the (unseptate) body forms a number of stiff, irregular processes, which perform the same See also:function (fig. 8). It is to be noted that these processes are non-motile, and not in any way comparable to pseudopodia, to which they were formerly likened. A very interesting and remarkable morphological peculiarity has been recently described by Leger (18) in the case of a new Gregarine, Taeniocystis. In this form the body is elongated and metamerically segmented, recalling that of a segmented See also:worm, the adult trophozoites possessing numerous partitions or segments (each corresponding to the septum between the proto- and deuto-merite in an See also:ordinary Polycystid), which See also:divide up the cytoplasm into roughly equal compartments. Leger thinks only the deutomerite becomes thus segmented, the protomerite remaining small and undivided. The See also:nucleus remains single, so that there is no question as to the unicellular or individual nature of the entire See also:animal. The general cytoplasm usually consists of distinct ectoplasm and endoplasm, and is limited by a membrane or cuticle (epicyte), secreted by the former. The cuticle varies considerably Minute in thickness, being well developed in active, intestinal forms, but very thin and delicate in non-motile coelomic structure• forms (e.g.

Diplodina). In the former case it may show See also:

longitudinal striations. The cuticle also forms the hooks or spines of many epimerites. The ectoplasm usually shows (fig. 9A) a differentiation into two layers, an See also:outer, firmer layer, clear and hyaline, the sarcocyte, and an inner layer, the myocyte, which is formed of a network of muscle-fibrillae (mainly longitudinal and transverse, fig. 9B). The sarcocyte alone constitutes the septum, traversing the endoplasm, in septate Gregarines. The See also:myonemes are undoubtedly the agents responsible for the active gregarinoid " movements (of flexion and contraction) to be observed After Leger and fiagenmiiller, from Lankester's in many forms. The See also:Treatise on See also:Zoology. See also:peculiar gliding movements FIG. 8.-Three Individuals (G) of were formerly thought to Ophryocystis schneideri, attached to be produced by the extru- See also:wall of Malpighian tubule of Blaps sp. See also:sion of a gelatinous See also:thread p, Syncytial protoplasm of the tubule; posteriorly, but Crawley (8) c, See also:Cilia lining the lumen. has recently ascribed them to a complicated See also:succession of See also:wave-like contractions of the myocyte layer.

This view is supported by the fact that certain coelomic forms, like Diplodina and others, which either lack muscle-fibrils or else show no ectoplasmic differentiation at all, are non-motile. The endoplasm, or nutritive plasm, consists of a semi-fluid See also:

matrix in which are embedded vast See also:numbers of grains and spherules of various kinds and of all sizes, representing an See also:accumulation of food-material which is being stored up prior to See also:reproduction. The largest and most abundant grains are of a sub-stance termed See also:para-glycogen, a See also:carbohydrate; in addition, flattened tom`'( After Siedlecki, from Lankester's Treatise on Zoology. From Wasielewski, after Leger. lenticular platelets, of an albuminoid character, and highly-refringent granules often occur. The nucleus is always lodged in the endoplasm, and, in the septate forms, in the deutomeritic half of the body. It is normally spherical and always limited by a distinct nuclear membrane, which itself often contains chromatin. The most See also:char- acteristic feature of the nucleus is the deeply-staining, more or less vacuolated spherical karyosome (consisting of chromatin intimately See also:bound up with a plastinoid basis) which is invariably present. In one or two instances (e.g. Diplocystis schneideri) the nucleus has more than one karyosome. All the chro- matin of the nucleus is not, how- ever, confined to the karyosome, .some being in the form of grains in the nuclear See also:sap; and in some cases at any See also:rate (e.g. Diplodina, Lankesteria) there is a well-marked After Schewiakoff, from Lankester's Treatise on Zoology.

De, Deutomerite. s, Septum. en, Endoplasm. sc, Sarcocyte. c, Cuticle. m,f,Myocyte fibrils (cut across). g, Gelatinous layer. nuclear reticulum which is impregnated with granules and dots of chromatin. A sexual multiplication (schizogony) is only known certainly to occur in a few cases, one being in a Monocystid form, a species of Life- Gonospora, which is for a See also:

long time intracellular (Caullery Life- and Mesnil [4]), the rest among the Schizogregarinae, so history. named for this See also:reason, in which schizogonous fission takes place regularly during the free, trophic condition. Usually, the body divides up, by a See also:process of multiple fission (fig. to), into a few (up to eight) daughter-individuals; but in a new genus (Eleutheroschizon), Brasil (3) finds that a See also:great number of little merozoites are formed, and a large amount of vacuolated cytoplasm is left over unused. In the vast See also:majority of Gregarines, however, the life-cycle is limited to gametogony and sporogony. A very general, if not indeed universal, prelude to gametogony is the characteristic and important feature of the order, known as association, the biological significance of which has only lately been fully brought out (see H.

M. Woodcock 131]). In normal association, two individuals which are to be regarded as of opposite See also:

sex, come into See also:close contact with each other and remain thus the characters of the adult. attached. The manner in which the parasites join varies in different forms; the association may be end-to-end (terminal), either by like or by unlike poles, or it may be See also:side-to-side (lateral) (fig. 12). The couple (syzygy) thus formed may proceed forthwith to encystment and sporoblast-formation (Lankesteria, Monocystis), or may continue in the trophic phase for some time longer (Gregarina). In one or two instances (Zygocystis), association occurs as soon as the trophozoites become adult. This leads on to the interesting phenomenon of precocious association (neogamy), found in non-motile, coelomic Gregarines (e.g. Cystobia, Diplodina and Diplocystis), in which the See also:parasitism is most advanced. Woodcock (loc. cit.) has de-scribed and compared the different methods adopted to ensure a permanent See also:union, and the degree of neogamy attained, in these forms. Here it must suffice to say that, in the extreme condition (seen, for instance, in Diplodina minchinii) the union takes place very early in the life-history, between individuals which are little more than sporozoites, and is of a most intimate character, the actual cytoplasm of the two associates join- ing.

In such cases, there is absolutely nothing to indicate the " See also:

double " nature of the growing trophozoite, but the presence of the two nuclei which remain quite distinct. There can be little doubt that, in the great majority, if not in all Gregarines, association is necessary for subsequent sporulation to take place; i.e. that the cytotactic attraction imparts a develop-See also:mental stimulus to both partners, which is requisite for the formation of primary sporoblasts (gametes). This association is usually permanent; but in one or two cases (perhaps Gonospora sp.) temporary association may suffice. While association has fundamentally a reproductive (sexual) significance, in some cases, this function may be delayed or, as it were, temporarily suspended, the cytotactic attraction serving meanwhile a subsidiary purpose in trophic life. Thus, probably, are to be explained the curious multiple associations and long chains of Gregarines (fig. II) sometimes met with (e.g. Eirmocystis, Clepsydrina). Encystment is nearly always double, i.e. of an associated couple. Solitary encystment has been described, but whether successful See also:independent sporulation results, is uncertain; if it does, the encystment in such cases is, in all See also:probability, only after prior (temporary) association. In the case of free parasites, a well-developed cyst is secreted by the syzygy, which rotates and gradually becomes spherical. A thick, at first gelatinous, outer cyst-membrane (ectocyst) is laid down, and then a thin, but See also:firm internal one (endocyst). The cyst once formed, further development is quite independent of the host, and, in fact, often proceeds outside it.

In certain coelomic Gregarines, on the other hand, which remain in very close relation with the host's tissues, little or nothing of an encystment-process on the part of the parasites is recognizable, the cyst-wall being formed by an enclosing layer of the host (Diplodina). The nuclear changes and multiplication which precede sporoblast-formation vary greatly in different Gregarines and can only be outlined here. In the formation of both sets of sexual elements (gametes) there is always a comprehensive nuclear See also:

purification or maturation. This elimination of a part of the nuclear material (to he distinguished as trophic or somatic, from the functional or germinal portion, which forms From Wasielewski, after the sexual nuclei) may occur at widely- Leger. different periods. In some cases (Lankes- FIG. 12.—Associations teria, Monocystis), a large part of the of Gonospora sparse. See also:original (sporont-) nucleus of each associate is at once got rid of, and the resulting (segmentation-) nucleus, which is highly-specialized, represents the sexual part. In other cases, again, the entire sporont-nucleus proceeds to division, and the distinction between somatic and germinal portions does not become See also:manifest until after nuclear multiplication has continued for some little time, when certain of the daughter-nuclei become altered in character, and ultimately degenerate, the See also:remainder giving rise to the sporoblast-nuclei (Diplodina, Stylorhynchus). Even after the actual sporoblasts (sex-cells) themselves are constituted, their nuclei may yet undergo a final maturation (e.g. Clepsydrina ovata); and in Monocystis, indeed, Brasil (2) finds that what is apparently a similar process is delayed until after conjugation and formation of the zygote (definitive sporoblast). Nuclear multiplication is usually indirect, the mitosis being, as a From Wasielewski, after A. See also:Schneider.

B. From Wasielewski, after Leger. rule, more elaborate in the earlier than in the later divisions. The attraction-See also:

spheres are generally large and conspicuous, sometimes consisting of a well-developed centrosphere, with or without centrosomic granules, at other times of very large centrosomes with a few astral rays. In those cases where the karyosome is retained, and the sporont-nucleus divides up as a whole, however, the earliest nuclear divisions are See also:direct; the daughter-nuclei being formed either by a process of simple constriction (e.g. Diplodina), or by a See also:kind of multiple fission or fragmentation (Gregarina and Selenidium spp.). Nevertheless, the later divisions, at any rate in Diplodina, are in-direct. By the time nuclear multiplication is well sivanced or completed, the bodies of the two See also:parent-Gregarines (associates) have usually become very irregular in shape, and produced into numerous lobes and processes. While in some forms (e.g. Monocystis, Urospora, Stylorhynchus) the two individuals remain fairly See also:separate and independent of each other, in others (Lankesteria) they become inter-twined and interlocked, often to a remarkable extent (Diplodina). The sexual nuclei next pass to the See also:surface of the processes and segments, where they take up a position of See also:uniform distribution. Around each, a small See also:area of cytoplasm becomes segregated, the whole often projecting as a little bud or hillock from the general surface.

These uninuclear protuberances are at length cut off as the sporoblasts or gametes. Frequently a large amount of the general protoplasm of each parent-individual is left over unused, constituting two cystal residua, which may subsequently fuse; in Diplodina, however, practically the whole cytoplasm is used up in the formation of the gametes. The sporoblasts themselves show all gradations from a condition of marked differentiation into male and See also:

female (anisogamy), to one of complete equality (isogamy). Anisogamy is most highly developed in Pterocephalus. Here, the male elements (microgametes) are minute, elongated and spindle-like in shape, with a minute rostrum anteriorly and a long flagellum posteriorly, and very active; the female elements (megagametes) are much larger, oblong to ovoid, and quite passive. In Stylorhynchus the difference between the conjugating gametes is not quite so pronounced (fig. 13), the male elements being of about the same bulk as the See also:females, but pyriform f, g, Stages in conjugation and attached to body of parent- nuclear union of the two individual. elements. b-d, Stages in development of h, Zygote (copula). motile male gamete. i, Spore, still with single e, Mature female gamete. nucleus and undivided sporoplasm. instead of See also:round, and possessing a distinct flagellum; a most interesting point_ about this parasite is that certain highly motile and spermatozoon-like male gametes are formed (fig. 13), which are, however, quite sterile and have acquired a subsidiary function. In other cases, again, the two kinds of See also:element exhibit either very slight See also:differences (Monocystis) or none (Urospora, Gonospora), in size and See also:appearance, the See also:chief distinction being in the nuclei, those of the male elements being smaller and chromatically denser than those of the females.

Lastly, in Lankesteria, Gregarina, Clepsydrina, Diplocystis and Diplodina complete isogamy is found, there being no apparentdifference whatever between the conjugating elements. Nevertheless, these forms are also to be regarded as instances of binary sexuality and not merely of See also:

exogamy; for it is practically certain that this condition of isogamy is derived from one of typical anisogamy, through a stage such as is seen in Gonospora, &c. And, similarly, just as in all instances where the formation of differentiated gametes has been observed, the origin of the two conjugates is from different associates (parent-sporonts), and all the elements arising from the same parent are of the same sex, so it is doubtless the case here. The actual union is brought about or facilitated by the well-known phenomenon termed the danse See also:des sporoblastes, which is due to various causes. In the case of highly-differentiated gametes (Pterocephalus), the actively motile microgametes See also:rush about here and there, and seek out the female elements. In Stylorhynchus, Leger has shown that the function of the sterile male gametes is to bring about, by their vigorous movements, the me"See also:lee sexuelle. In the forms where the gametes are isogamous or only slightly differentiated and (probably) not of themselves motile, other factors aid in producing the necessary commingling. Thus in Gregarina sp. from the mealworm, the unused somata or cystal residua become amoeboid and send out processes which drive the peripherally-situated gametes round in the cyst; in some cases where the residual See also:soma becomes liquefied (Urospora) the movements of the host are considered to be sufficient; and lastly, in Diplodina, owing to the extent to which the inter-See also:twining process is carried, if each gamete is not actually contiguous to a suitable See also:fellow-conjugant, a very slight See also:movement or mutual attraction will bring two such, when liberated, into contact. An unusual modification of the process of sporoblast-formation and conjugation, which occurs in Ophryocystis, must be mentioned. Here encystment of two associates takes place as usual; the sporontnucleus of each, however, only divides twice, and one of the daughter-nuclei resulting from each division degenerates. Hence only one sporoblast-nucleus, representing a See also:quarter of the original nuclear-material, persists in each half. Around this some of the cytoplasm condenses, the rest forming a residuum.

The sporoblast or gamete thus formed is completely isogamous and normally conjugates with the like one from the other associate, when a single zygote results which becomes a spore containing eight sporozoites, in the ordinary manner. Sometimes, however, the septum between the two halves of the cyst does not break down, in which case parthenogenesis occurs, each sporoblast developing by itself into a small spore. The two conjugating elements unite completely, cytoplasm with cytoplasm and nucleus with nucleus, to form the definitive sporoblast or zygote. The protoplasm assumes a definite outline, generally that of an ovoid or See also:

barrel, and secretes a delicate membrane, the ectospore. This subsequently becomes thickened, and often produced into rims, spines or processes, giving rise to the characteristic appearance of the Gregarine spore. Internal to the ectocyst, another, thinner membrane, the endocyst, is also laid down. These two membranes form the spore-wall (sporocyst). Meanwhile the contents of the spore have been undergoing division. By successive divisions, usually mitotic, the zygote-nucleus gives rise to eight daughter-nuclei, each of which becomes the nucleus of a sporozoite. Next, the sporoplasm becomes split longitudinally, around each nucleus, and thus eight sickle. shaped (falciform) sporozoites are formed. There is usually a After LEger, from Lankester's Treatise on Zoology. The chief families, with representative genera, are as follows: Porosporidae, with Porospora gigantea, at present thought to be gymnosporous; Gregarinidae (Clepsydrinidae), with Gregarina, Clepsydrina, Eirmocystis, Hyelospore, Cmenidospore, Stenophora; Didymophyidae, with Didymophyes; Dactylophoridae, with Dactylophorus, Pterocephalus, Echinomera, Rhopalonia; Actinocephalidae with Actinocephalus, Pyxinia, Coleorhynchus, Stephanophora, Legeria, Stictospora, Pileocephalus, Sciadophora; Acanthosporidae with Acanthospore, Corycella, Cometoides; Menosporidae with Menospore, Hoplorhynchus; Stylorhynchidee, with Stylorhynchus, Lophocephalus; Doliocystidae with Doliocystis; and Taeniocystidae, with Taeniocystis.

The curious gftus Selenidium is somewhat apart. Tribe B.—Acephaline (practically See also:

equivalent to Aseptata, Haplocyta). The body never possesses an epimerite and is non-septate. Chiefly coelomic parasites of " worms," Holothurians and See also:insects. The Aseptata have not been so completely arranged in families as the Septata. Leger has distinguished two well-marked ones, but the remaining genera still want classifying more in detail. Fam. Gonosporidae, with Gonospora, Diplodina; and Urosporidae, with Urosopore, Cystobia, Lithocystis, Ceratospora; the genera Monocystis, Diplocystis Lankesteria and Zyeocystis probably constitute another; Pterospore and, again, Syncystis are distinct; lastly, certain forms, e.g. Zygosoma, Anchora (Anchorina), are incompletely known. There remains for mention the remarkable parasite, recently described by J. Nusbaum (24) under the appropriate name of Schaudinnella henleae, which inhabits the intestine of Henlea leptodera. Briefly enumerated, the principal features in the life-cycle are as follows.

The young trophozoites (aseptate) are attached to the intestinal cells, but practically entirely extracellular. Association is very primitive in character and indiscriminate; it takes place indifferently between individuals which will give rise to gametes of the same or opposite sex. Often it is only temporary; at other times it is multiple, several adults becoming more or less enclosed in a gelatinous investment. Nevertheless, in no case does true encystment occur, the sex-cells being developed practically free. The female gametes are large and See also:

egg-like; the See also:males, minute and sickle-like, but with no flagellum and apparently non-motile. While many of the zygotes (" amphionts ") resulting from copulation pass out to the exterior, to infect a new host, others, possessing a more delicate investing-membrane, penetrate in between the intestinal cells, producing a further infection (auto-infection). Numerous sporozoites are formed in each zygote. It will be seen that Schaudinnella is a practically unique form. While, on the one hand, it recalls the Gregarines in many ways, on the other hand it differs widely from them in several characteristic features, being primitive in some respects, but highly specialized in others, so that it cannot be properly included in the order. Schaudinnella rather represents a primitive Ectosooran parasite, which has proceeded upon a See also:line of its own, intermediate between the Gregarines and Coccidia. 4. M.

Caullery and F. Mesnil, " Sur une Gregarine presentant une phase de multiplication asporulee," C.R. Ac. Sci. 126, p. 262 (1898) ; 5. M. Caullery and F. Mesnil, " Le Parasitisme intracellulaire des Gregarines," op. cit. 132, p. 220 (1901) ; 6. M.

Caullery and F. Mesnil, Sur une mode particuliere de division nucleaire chez See also:

les Gregarines," See also:Arch. anat. microsc. 3, p. 146, 1 pl. (1900) ; 7. M. Caullery and F. Mesnil, " Sur quelques parasites internes des Annelides," Misc. biol. (See also:Tray. Stat. Wimereux), 9, p. 8o, t pl.

(1899); 7a. J. Cecconi, " Sur l'Anchorina sagittata, &c.," Arch. Protistenk. 6, p. 230, 2 pls. (1905) ; 8. H. Crawley, " Progressive Movement of Gregarines," P. Ac. Philad. 54, p.

4, 2 pls. (1902), also op. cit. 57, p. 89 (1905) ; 9. H. Crawley, " See also:

List of the Polycystid Gregarines of the U.S.," op. cit. 55, pp. 41, 632, 4 pls. (1903); 10. L. Cuenot, Recherches sur revolution et la conjugaison des Gregarines," Arch. biol. 17,.p.

581, 4 pls. (1901); 11. A. Laveran and F. Mesnil, " Sur quelques particularites de revolution d'une Gregarine et la reaction de la cellule-hote," C.R. See also:

Soc. Biol. 52, p. 554, 9 See also:figs. (Iwo); 12. L. Leger, " Recherches sur Ies Gregarines," Tebl. zool.

3, p. i., 22 pls. (1892); 13. L. Leger, " Contribution a la connaissance des Sporozoaires, &c.," See also:

Bull. Sci. See also:France, 30, p; 240, 3 pls. (1897) ; 14. L. Leger, " Sur un nouveau Sporozoaire Schizocystis), &c.," C.R. Ac. Sci. 131, p.

722 (1900) ; 15. L. Leger, " La Reproduction • sexuee chez les Ophryocystis," E. c. p. 761 (1900) ; 16. L. Leger, " Sur une nouvelle Gregarine (Aggregate coelomica,), &c.," op. cit. 132, p. 1343 (1901); 17. L. Leger, " La Reproduction sexuee chez les Stylorhynchus," Arch. Protistenk. 3, p.

304, 2 pis. (1904) ; 18. L. Leger, " Etude sur Taeniocystis mire (Leger), &c.," op. See also:

cat. 7, p. 307, 2 pls. (1906) ; 19. L. Leger and O. Duboscq, " La Reproduction sexuee chez Pierocephalus," Arch. zool. exp. (N. et R.) (4) 1, p. 141, II figs.

(1903); 20. L. Leger and O. Duboscq, " Aggregata vagans, n. sp., &c.," E. c. p. 147, 6 figs. (1903) ; 21. L. See also:

Leer and O. Duboscq, " Les Gregarines et 1'e ithelium intestinal, &c., ' Arch. parasitol. 6, p. 377, 4 pls. (1902); 22.

L. Leger and O. Duboscq, Nouvelles Recherches sur See also:

Classification. certain amount of unused sporoplasm left over in the centre of the spore, constituting the sporal residuum. It is important to See also:note that in all known Gregarines, with one exception, the number of sporozoites in the spore is eight; the exception is Selenidium, in many ways far from typical, where the number is half, viz. four. Hitherto a variation from the general mode of spore-formation has been considered to occur in certain Crustacean Gregarines, the Aggregatidae and the Porosporidae. The spores of these forms have been regarded as gymnospores (naked), lacking the enveloping membranes (sporocyst) of the ordinary spores, and the sporozoites, consequently, as developed freely in the cyst. In the case of the first-named parasites, however, what was taken for sporogony has been proved to be really schizogony, and on other grounds these forms are, in the present writer's See also:opinion, preferably associated with the Coccidia (q.v.). With regard to the Porosporidae, also, it is quite likely that the gymnosporous cysts considered to belong to the Gregarine Porospora (as known in the trophic condition) have really no connexion with it, but represent the schizogonous See also:generation of some other form, similar to Aggregata; in which case the true spores of Porospora have yet to be identified. In the intestine of a fresh host the cysts rupture and the spores are liberated. This is usually largely brought about by the swelling of the residual protoplasm. Sometimes (e.g.

Gregarina) long tubular outgrowths, known as sporoducts (fig. 15), are developed from the residual protoplasm, for the passage of the spores to the exterior. The Gregarines are extremely numerous, and include several families, characterized, for the most part, by the form of the spores (fig. 16). The specialized Schizogregarinae are usually separated off from the rest as a distinct sub-order. SUB-ORDER I. Schizogregarinae. Forms in which schizogonic reproduction is of general occurrence during the extra-cellular, trophic phase. Three genera, Ophryocystis, Schizocystis and Eleutheroschizon, different peculiarities of which have been referred to above. Mostly parasitic in the intestine From Wasielews'ai, after Leger. a, Eirmocystis, Sphaerocystis, &c. f, Stylorhynchidae (type of). b, Echinomera, Pterocephalus, &c. g, Menosporidae.

c, Gregarina, &c. h, Gonospora terebellae. d, Beloides. i, Ceratospora. e, Ancyrophora. k, Urospora synaptae. or Malpighian tubules of insects. (In this type of parasite, as exemplified by Ophryocystis, the body was formerly wrongly considered as amoeboid, and hence this genus was placed in a See also:

special order, the Amoebosporidia.) SUB-ORDER II.—Eugregarinae. Schizogony very exceptional, only occurring during the intracellular phase, if at all. Gregarines fall naturally into two tribes,' described as cephalont and septate, or as acephalont and aseptate (haplocytic), respectively. In strictness, however, as already mentioned, these two sets of terms do not agree absolutely, and whichever set is 'adopted, the other must be taken into account in estimating the proper position of certain parasites. Here the cephalont or acephalont condition is regarded as the more primary and fundamental. Tribe A.—Cephalina (practically equivalent to Septata). See also:Save exceptionally, the body possesses an epimerite, at any rate during the early stages of growth, and is typically septate. Mostly intestinal parasites of Arthropods.

le. Gregarines, &c.," Arch. Protistenk. 4, p. 335, 2 pls. (1904); 23. M. Lithe, " Bau and Entwickelung der Gregarinen," t. c. p. 88, several figs. (1904) ; 24. J. Nusbaum, " Ober See also:

die .

. . Fortpflanzung einer . . . Gregarine, Schaudinnetla henleae," Zeit. wiss. Zool. 75, p. 281, pl. 22 (1903) ; 25. F. Paehler, " Uber die Morphologie, Fortpflanzung , von Gregarina ovate," Arch. Protistenk. 4, p.

64, 2 pls. (1904) ; 26. S. Prowazek, " Zur Entwickelung der Gregarinen," op. Cit., 1, p. 297, pl. 9 (1902); 27. A. Schneider (Various See also:

memoirs on Gregarines), See also:Tabl. zool. I and 2 (1886—1892); 28. H. Schnitzler, " Uber die Fortpflanzung von Clepsydrina ovate," Arch.

Protistenk. 6, p. 309, 2 pls. (1905); 29. M. Siedlecki, " Ober die geschlechtliche Vermehrung der Monocystis ascidiae," Bull. Ac. Cracovie, p. 515, 2 pls. (1900); 30. M. Siedlecki, " Contribution a l'etude des changements cellulaires provoquees See also:

par les Gregarines," Arch. anat. microsc.

4, p. 87, 9 figs. (1901); 31. H. M. Woodcock, " The Life-Cycle of Cystobia irregularis, &c.," Q.J.M. Sci. 5o, p. 1. 6 pls. (1906). (H.

M.

End of Article: GREGARINES (mod. Lat. Gregarina, from gregarius, collecting in a flock or herd, grex)

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