Spermiogenesis pursues three major objectives: (1) The safeguard of the mal
e genome within the confines of a compact nucleus. (2) The accumulation of
enzymes in the acrosome of be released at fertilization. (3) The developmen
t of a sperm propelling tail consisting of an axoneme surrounded by a scaff
old of keratin-containing outer dense fibers and a fibrous sheath. Recent e
xperimental data indicate that three keratins-Sak57, Odf1 and Odf2- and oth
er proteins (the 26S proteasome and the Odf1-binding protein Spag4) are tem
porarily stored in the manchette before being sorted to the developing sper
m tail. These findings support a general model for the manchette as an ephe
meral structure timely developed and strategically positioned to provide a
transient storage to both structural and signaling proteins. Some of the pr
oteins are later sorted to the developing tail; others may participate in t
he reciprocal nuclear-cytoplasmic signaling pathways as the gene activity o
f the male genome gradually becomes silent. (C) 2001 Wiley-Liss, Inc.