Foetal genital development in Hyrax capensis, a species with primary testicondia: proposal for the evolution of Hunter's gubernaculum

Abstract

Background: Control of testicular descent is poorly understood. There are a number of mammalian species in which testis descent does not occur, and the phenomenon is called testicondia. Analysis of foetal development of such species could contribute to a better understanding of the key events in anatomical development underlying testis descent. Specific attention is to be given to the development of the so-called gubernaculum of Hunter: a structure of complex architecture and composition, which extends from the caudal pole of the mesonephric remnants into the inguinal abdominal wall and which is present in most mammals but not in submammalian vertebrates.

Methods: Serially sectioned male and female foetuses of Hyrax capensis, a species in which testes remain close to the caudal pole of the kidneys throughout life, were analysed for architecture of the developing genital apparatus and the immediately surrounding structures. The results of this analysis were compared with those from a similar analysis of reptiles, of monotremata without testis descent, and of mammals with testis descent.

Results: Reptiles and Monotremata showed no gubernacular structures. Mammals with testis descent showed gubernacular growth and differentiation which varied between the sexes and with the stage of foetal development. Hyrax capensis foetuses showed the development of only one component of the gubernaculum: the gubernacular cord as a part of the mesonephric mesentery and extending between the caudal pole of the mesonephros (or mesonephric remnants in older foetuses) and the lateral bladder ligament. No part developed in the inguinal abdominal wall components, which are the primordia of the cremaster sacs in species with testis descent.

Conclusion: Hyrax capensis shows only partial development of the gubernacular structures, and, specifically, the primordia of the cremaster sacs remain absent. Thus, this species lacks a key anatomical condition for testis descent. Gubernacular architecture in Hyrax capensis seems of a degree intermediate between egg laying monotremata mammals and mammals with testis descent. A model is proposed within which to understand the development of the gubernacular components within the mammalian class.