HHMI & California Institute of Technology, Division of Biology, Pasadena, CA 91125 U.S.A. As the C. elegans male attempts to penetrate the hermaphrodite vulva with his spicules, the protractor muscles that are attached to these copulatory structures contract and relax rapidly such that the spicules prod the vulva slit at a frequency of 7.2 1.3 Hz. Phasic protractor muscle contractions persist until the vulva slit is partially penetrated. Once the vulva is breached, the protractor muscles contract tonically and the spicules extend completely into the hermaphrodite. The spicule muscles stay contracted for 75 20 seconds, sufficient time for sperm to transfer into the hermaphrodite. The male protractor muscles are innervated by the SPC motor neurons (2), and removal of these cells results in male impotency (1). The SPC neurons are essential for tonic, but not phasic spicule muscle contractions; the spicules of SPC-ablated males can not fully extend into the hermaphrodite, but can prod the vulva at a frequency of 5.1 1.1 Hz. The ACh agonists levamisole, nicotine, arecoline, and oxotremorine can stimulate the protractor muscles to contract; and the SPC motor neurons support the expression of the
unc-17-encoded VAChT. Therefore ACh most likely regulates the contractile state of the spicule muscles. The ACh agonists differentially require
egl-19 and
unc-68-encoded calcium channels to mediate the behavioral output; thus suggesting that calcium mobilization from these channels might have non-overlapping roles during copulation. During mating the spicules of
egl-19 males can prod the vulva at a frequency of 5.0 1.5 Hz, but they can not breach the vulva lips. In contrast, the spicules of
unc-68 males can insert into the hermaphrodite, but prior to penetration, the spicules prod the vulva at a frequency of 0.48 1.4 Hz. Therefore, the UNC-68 channel is utilized in phasic muscle contractions, and the EGL-19 channel is required for tonic muscle contractions. 1. Liu, K.S., and Sternberg, P.W. (1995).Neuron 14, 79-89. 2. Sulston, J.E., Albertson, D.G., and Thomson, J.N. (1980). Dev. Biol. 78, 542-576.