Mechanisms of mammalian sperm migration through the female reproductive tract and ovum vestments are described. The perspective is biophysical as well as biochemical and morphological, and the focus is upon the role of sperm motility in these processes. Sperm forward progression is characterized as an interactive process between the the cell and its environment, and the mediation of flagellar bend propagation by the physical properties of its surroundings is described. These properties, together with flagellar beat kinematics, sperm morphology, and surface properties, determine the magnitude of the forces generated by sperm and their consequent rate of progression. Sperm interactions with the cervical mucus, the cumulus oophorus, and the zona pellucida are described. The poorly understood affinity of the sperm surface for the macromolecules of the mucus, cumulus, and zona is stressed, as is the viscoelastic structural mechanical resistance of these biopolymers to sperm motion. The kinematics and consequences of hyperactivated sperm motion are presented, with emphasis on objective characterization of such motion (as a biomarker), along with analysis of the mechanical advantage that such motion may confer on spermatozoa during egg-vestment interaction.
Key words: sperm, transport, mucus, cumulus, zona pellucida, fertilization
INTRODUCTIONFlagellar movement is a fundamental expression of the vitality of a sperm cell, and is essential for its reproductive function. The active mechanism of flagellar motion produces a sequence of principal and reverse bends [Gibbons and Gibbons, 1974; Woolley, 19771. These can have varying degrees of curvature and can be initiated and propagated along the flagellum at different rates. Flagellar bending results in a distribution of local forces and torques against the surrounding fluid. In accordance with Newton's Third Law, these are resisted by equal and opposite local forces and torques. In order to satisfy Newton's Second Law, the summation of the local local external forces and torques over the sperm head and tail must be zero. As a consequence, there Received November 8, 1988; accepted November 17, 1988. Address reprint requests to David F. Katz In a general biophysical sense, mammalian sperm swim optimally; the principles of hydrodynamics and optimal control theory demonstrate that the smooth flagellar traveling waves, commonly seen in low-viscosity media, tend to maximize the sperm's propulsive velocity with respect to the hydrodynamic power output (Pironneau and Katz, 1974). Mammalian sperm must use their swimming ability to reach the oocyte-cumulus complex, and the forces due to flagellar motion assist them in penetrating the egg vestments to reach the oolemma.Our studies of these processes are hampered by their complexity and by apparent variation among species. Some of this biological variation is real; for example, the site of semen deposition may be the vagina, cervix, or uterus. These environments differ not only with respect to their distance from the site of ferti...