Male intromittent organs serve two primary reproductive functions: the physical entry into the female body during copulation and the effective delivery of gametes resulting in internal fertilization. Here we present a histological examination of the adult male American alligator phallus semen delivery apparatus, the sulcus spermaticus. While the highly collagenous basal crurae and more distal shaft of the alligator penis contain the rigid structures that facilitate cloacal intromission, the sulcus is more functionally intricate. Here we show the sulcus spermaticus (an open groove that runs along the ventral aspect of the phallic shaft) to be a spatially heterogeneous reproductive structure containing a complex architecture of multiple tissue types. Sulcus morphology markedly changes from its proximal origin between the crurae to its distal exit from the phallus tip. At the proximal origin of the sulcus, the ductus deferens vent semen into an expanded lumen lined by a convoluted secretory epithelium. Along the length of the phallic shaft, an arrangement of longitudinally and radially oriented smooth muscle bundles may act via rhythmic contractions to produce peristaltic sperm conveyance through the sulcus. An extensive vascular network of blood and lymph vessels putatively engorges the sulcus tissues during reproductive activity, increasing tension on an internal network of connective tissues and leading to localized inflation and increased tissue rigidity. We hypothesize that this engorgement works to seal the sulcus groove and allow the structure to convey semen through a functionally closed tube. Further, numerous epithelial secretory cells contribute seminal fluids to the ejaculate and may aid in as yet uncharacterized aspects of sulcus functioning. Together, these observations establish that the sulcus spermaticus is far more than a simple furrow in the phallus shaft for sperm conduction: it contains elements that form a complex functional gamete delivery system.
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