In the vast 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 [31]). In
normal association, two individuals which are to be regarded as of
opposite sex, come into close contact with each other and remain thus
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 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 parasitism is most advanced. Woodcock (_loc. cit._) has
described and compared the different methods adopted to ensure a
permanent 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 joining. In such cases, there is absolutely
nothing to indicate the "double" nature of the growing trophozoite,
but the presence of the two nuclei which remain quite distinct.
[Illustration: From Wasielewski, after Leger.
FIG. 11.--_Eirmocystis spp._ a, b, Associations of two and three
Gregarines; c, Chain of five parasites; p, Primite; s, Satellites.]
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
developmental 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. 11) sometimes met with (e.g.
_Eirmocystis_, _Clepsydrina_).
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