Nuclear condensation during spermiogenesis in the ostrich follows the
basic pattern established in other
vertebrates. The fine granular nuclear substance of early spermatids is
gradually replaced by numbers of
coarse dense granules which appear to arise by aggregation of smaller
dispersed elements of the chromatin.
The granules increase in size and eventually coalesce to form the compact
homogenous mass of chromatin
typical of the mature sperm. In ostrich spermatids, however, the aggregation
of the nuclear material
produces large numbers of longitudinally oriented rod-shaped structures in
addition to some granular
material. Although fibrillar chromatin has been observed during spermiogenesis
in a number of vertebrate
species, the hollow nature of the rod-shaped chromatin granules in ostrich
spermatids is a unique
phenomenon. The spiralisation of the chromatin material observed in ostrich
spermatids and in some other
nonpasserine birds is possibly related to the reduction in nuclear length
demonstrated during spermiogenesis
in these species. In common with other nonpasserine birds, spermiogenesis in
the ostrich is characterised by
the appearance both of a circular and a longitudinal manchette. The circular
manchette consists of a single
row of microtubules reinforced by additional peripherally arranged microtubules.
Links between adjacent
microtubules, and between the nucleolemma and some of the microtubules, are
evident. The longitudinal
manchette consists of arrays of interconnected microtubules arranged in
approximately 4–6 staggered, ill
defined rows. This structure seems to originate as a result of the rearrangement
of the microtubules of the
circular manchette and is only formed once the process of chromatin
condensation is well advanced. Based
on the sequence of morphological events observed during spermiogenesis in
the ostrich, it is concluded that
the circular manchette is responsible for the initial transformation in shape
of the spermatid nucleus.
Thereafter, the chromatin condenses independently within the confines of
the nucleolemma with the circular
manchette merely acting to maintain the shape of the nucleus while this
process is underway, to compress
the nuclear membrane, and possibly to orientate the subunits of the condensing
chromatin. The longitudinal
manchette appears to assist in the translocation of material during
spermatid elongation. There are
indications that the developing acrosome is instrumental in effecting
nuclear shaping of the apical
(subacrosomal) head region of the ostrich spermatid.