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 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
certain amount of unused sporoplasm left over in the centre of the
spore, constituting the sporal residuum. It is important to 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.
[Illustration: FIG. 15.--Ripe Cyst of _Gregarina blattarum_, partially
emptied. (From Lankester.) a, Channels leading to the sporoducts; b,
Mass of spores still left in the cyst; c, Endocyst; d, The everted
sporoducts; e, Gelatinous ectocyst.]
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 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 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.
Classification.
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