Membrane-substrate contact under the spermatozoon of Caenorhabditis elegans, a crawling cell that lacks filamentous actin.

Caenorhabditis elegans spermatozoa use a single, persistent pseudopod to crawl at about 20 micrometers/min but, unlike other types of crawling cells, sperm lack both filamentous actin and myosin. Interference reflection microscopy has revealed that sperm form broad grey areas of contact, analogous to the close contacts that have been described underneath other crawling eukaryotic cells, between their pseudopods and their substrate. Individual sperm change the size, shape and pattern of their substrate attachments as they crawl but we found no correlation between the extent of underside of the cell in contact with the substrate and the velocity of locomotion. Two predominant attachment patterns were observed: (1) a single broad contact extending from the front of the pseudopod nearly to the rear of the cell; and (2) two separate contact sites, one under the front of the pseudopod and one under the cell body. Occasionally, under cells exhibiting the second type of attachment pattern, portions of the anterior contact separated and remained stationary relative to the substrate while the cell moved forward. This observation, as well as the continuous change in shape of the contact areas, suggests that sperm continually form new contacts near the tip of the pseudopod and release these contacts backwards. In extreme cases, sperm were able to crawl with only the front of the pseudopod in contact with the substrate. Therefore, we propose that sperm locomotion depends on the interaction of several key events (traction, propulsion, membrane insertion) occurring at the leading edge of the pseudopod.