or elongated; in one or two instances (e.g. Diplodina) it is spherical,
and, on the other hand, in Porospora (fig. 3) it is greatly
drawn out and vermiform. In many adult Gregarines,
the body is divided into two distinct but unequal regions
Morphology.
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 attachment of the
parasite to the host-cell
(fig. 6).
|
| After Siedlecki, from Lankester’s Treatise on Zoology. |
| Fig. 5.—Part of a section through the apparatus of fixation of a Pterocephalus, showing root-like processes extending from the Gregarine between the epithelial cells. g, Head of Gregarine; r, Root-like processes; ep, Epithelial cells. |
In those Gregarines (most intestinal forms) which become attached to an epithelial cell, the attachment occurs by means of a minute projection or 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 increases in size much quicker at first than the rest (the extracellular part), more or less fills up the host-cell, and forms the well-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). Léger thinks that this 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 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 present writer does not think that mechanical considerations should be altogether left out of 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 foreign bodies (food, &c.) in the intestine.
|
| From Wasielewski, after Léger. |
| Fig. 6.—Corycella armata, Léger. a, Cephalont; b, Epimerite in host-cell; c, Sporont. |
At the point where the body becomes actually intracellular, it is constricted, and this constriction marks off the epimerite (internally) from the 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 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), according 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 division into protomerite and deutomerite, and are hence known as Pseudomonocystida; this condition may be primitive (Doliocystis) or (possibly) secondary, the partition having in course of time disappeared. Again, Stenophora is a septate form which has become, secondarily, completely intracellular during the young stages, and, doubtless correlated with this, shows no sign of an epimerite.
| ||
| From Wasielewski, after Léger. | ||
| Fig. 7.—Forms of Epimerites. | ||
| 1, Gregarina longa. | 6, Cometoides crinitus. | |
| 2, Sycia inopinata. | 7, Geneiorhynchus monnieri. | |
| 3, Pileocephalus heerii. | 8, Echinomera hispida. | |
| 4, Stylorhynchus longicollis. | 9, Pterocephalus nobilis. | |
| 5, Beloides firmus. | ||
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 penetrates into the host-cell, and no epimerite is formed. Instead, a number of fine root-like processes are developed from near the anterior end, which pass in between the host-cells (fig. 5) and thus 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 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.
|
| After Léger and Hagenmüller, from Lankester’s
Treatise on Zoology. |
| Fig. 8.—Three Individuals (G) of Ophryocystis schneideri, attached to wall of Malpighian tubule of Blaps sp. p, Syncytial protoplasm of the tubule; c, Cilia lining the lumen. |
A very interesting and remarkable morphological peculiarity has been recently described by Léger (18) in the case of a new Gregarine, Taeniocystis. In this form the body is elongated and metamerically segmented, recalling that of a segmented worm, the adult trophozoites possessing numerous partitions or segments (each corresponding to the septum between the proto- and deuto-merite in an ordinary Polycystid), which divide up the cytoplasm into roughly equal compartments. Léger thinks only the deutomerite becomes thus segmented, the protomerite remaining small and undivided. The nucleus remains single, so that there is no question as to the unicellular or individual nature of the entire 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 in thickness, being well developed in active, intestinal forms, but very thin and delicate in non-motile coelomic Minute structure. forms (e.g. Diplodina). In the former case it may show 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 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 myonemes are undoubtedly the agents responsible for the active “gregarinoid” movements (of flexion and contraction) to be observed in many forms. The peculiar gliding movements were formerly thought to be produced by the extrusion of a gelatinous thread posteriorly, but Crawley (8) has recently ascribed them to a complicated succession of 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 matrix in which are embedded vast numbers of grains and spherules of various kinds and of all sizes, representing an accumulation of food-material which is being stored up prior to reproduction. The largest and most abundant grains are of a substance termed para-glycogen, a carbohydrate; in addition, flattened
