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Cover This month's cover illustration shows Spermiogenesis of the
rat
Drawing (prepared by Clermont and Rambourg, 1978) of
spermiogenesis of the rat, as seen with the light microscope in
sections stained with toluidine blue and periodic acid-Schiff. The
numbers of the drawings correspond to the steps of spermiogenesis as
proposed by Leblond and Clermont (1952). During the first three steps
of spermiogenesis, the Golgi phase, the juxtanuclear Golgi apparatus
elaborates proacrosomic granules, which fuse into a single acrosomic
granule that attaches itself to the nucleus. During steps 4 to 7, or
cap phase, the Golgi apparatus contributes to the formation of the
acrosome that spreads on the surface of the nucleus. During this phase
the mitochondria are along the plasma membrane, and the miniscule
centrioles bind to the nucleus at the pole opposite the acrosome and
give rise to the axoneme of the growing tail. During steps 8 to 14, the
acrosome phase, the spermatid becomes polarized, the chromatin
condenses, and the nucleus elongates to take its characteristic and
elegant sickle shape. Simultaneously, the Golgi apparatus stops adding
glycoproteins to the acrosome and migrates in the cytoplasmic lobe
that extends along the forming tail. During steps 15 to 19, the
maturation phase, the tail completes its formation as the mitochondria
migrate towards the proximal segment of the developing tail to form the mitochondrial sheath. The nucleus completes its condensation and acquires a thin shell, the perinuclear theca or perforatorium, on which
the acrosome binds. Finally, the cytoplasmic lobe flows apically, and
the bulk of the unused cytoplasm separates from the spermatid that then
becomes a spermatozoon. All these perfectly timed and integrated events
have been analyzed with the electron microscope (reviewed by Clermont,
Oko, and Hermo, 1993). Such studies underscore the complexity of these
cytological events, which presuppose a multitude of regulatory
mechanisms, biochemical and physiochemical in nature, most of which
have yet to be clarified.
by Yves Clermont