The foam production system of the male Japanese quail: characterization of structure and function

The research described here characterizes a unique neuromuscular system involved in reproductive behavior--the foam production system of the male Japanese quail (Coturnix japonica). Male quail produce a large amount of foam that is transferred to the female during copulation, enhancing male fertilization success. The source is the foam gland complex, a large sexually dimorphic, androgen sensitive, external protuberance of the dorsal cloaca, consisting of glandular units interdigitated with striated muscle fibers of the sphincter cloacae muscle (mSC). Electromyographic (EMG) analysis of mSC activity in freely moving males interacting with females revealed different characteristics of the EMG signal during copulation, voiding of excreta, and other mSC movement. The amount of mSC activity and also the amount of foam produced were greatly increased by the presence of a female behind a screen. Denervation of mSC eliminated normal mSC movement and also abolished foam production, confirming that mSC activity is the mechanism for foam production. The spinal cord locations of the motoneurons innervating the major cloacal muscles, including mSC, were determined by injecting cholera-toxin conjugated horseradish peroxidase into each muscle. Labelled somata with multiple primary dendrites were located in Area IX of the lateral motor column of synsacral segments 7, 8, or 9 or 8, 9, and 10. The motoneurons serving mSC were intermingled with those projecting to the other cloacal muscles, but there were differences in the rostralcaudal placement of these neural populations. Thus mSC activity is an integral part of the male's reproductive behavior, mSC activity can be socially stimulated, and mSC activity occurring in anticipation of copulation is likely to be functionally significant. Continued investigation of this highly accessible system has the potential to shed light on the mechanisms by which complex motor acts are produced and hormonally regulated.