CHANGES IN THE SPERMATOZOON DURING FERTILIZATION IN HYDROIDES HEXAGONUS (ANNELIDA) : I. Passage of the Acrosomal Region through the Vitelline Membrane

In the previous paper the structure of the acrosomal region of the spermatozoon was described. The present paper describes the changes which this region undergoes during passage through the vitelline membrane. The material used consisted of moderately polyspermic eggs of Hydroides hexagonus, osmium-fixed usually 9 seconds after insemination. There are essentially four major changes in the acrosome during passage of the sperm head through the vitelline membrane. First, the acrosome breaks open apically by a kind of dehiscence which results in the formation of a well defined orifice. Around the lips of the orifice the edges of the plasma and acrosomal membranes are then found to be fused to form a continuous membranous sheet. Second, the walls of the acrosomal vesicle are completely everted, and this appears to be the means by which the apex of the sperm head is moved through the vitelline membrane. The lip of the orifice comes to lie deeper and deeper within the vitelline membrane. At the same time the lip itself is made up of constantly changing material as first the material of the outer zone and then that of the intermediate zone everts. One is reminded of the lip of an amphibian blastopore, which during gastrulation maintains its morphological identity as a lip but is nevertheless made up of constantly changing cells, with constantly changing outline and even constantly changing position. Third, the large acrosomal granule rapidly disappears. This disappearance is closely correlated with a corresponding disappearance of a part of the principal material of the vitelline membrane from before it, and the suggestion is made that the acrosomal granule is the source of the lysin which dissolves this part of the vitelline membrane. Fourth, in the inner zone the fifteen or so short tubular invaginations of the acrosomal membrane, present in the normal unreacted spermatozoon, lengthen considerably to become a tuft of acrosomal tubules. These tubules are the first structures of the advancing sperm head to touch the plasma membrane of the egg. It is notable that the surface of the acrosomal tubules which once faced into the closed acrosomal cavity becomes the first part of the sperm plasma membrane to meet the plasma membrane of the egg. The acrosomal tubules of Hydroides, which arise simply by lengthening of already existing shorter tubules, are considered to represent the acrosome filaments of other species.