Reproduction, Fertility and Development Reproduction, Fertility and Development Society
Vertebrate reproductive science and technology

Testicular descent, sperm maturation and capacitation. Lessons from our most distant relatives, the monotremes

Heath Ecroyd A , Brett Nixon B , Jean-Louis Dacheux C and Russell C. Jones B D

A School of Biological Sciences, University of Wollongong, NSW 2522, Australia.

B Discipline of Biological Sciences, School of Life and Environmental Sciences, University of Newcastle, NSW 2308, Australia.

C UMR INRA-CNRS 6175, F-37380 Nouzilly, France.

D Corresponding author. Email:

Reproduction, Fertility and Development 21(8) 992-1001
Submitted: 31 March 2009  Accepted: 17 May 2009   Published: 30 October 2009


The present review examines whether monotremes may help to resolve three questions relating to sperm production in mammals: why the testes descend into a scrotum in most mammals, why spermatozoa are infertile when they leave the testes and require a period of maturation in the specific milieu provided by the epididymides, and why ejaculated spermatozoa cannot immediately fertilise an ovum until they undergo capacitation within the female reproductive tract. Comparisons of monotremes with other mammals indicate that there is a need for considerable work on monotremes. It is hypothesised that testicular descent should be related to epididymal differentiation. Spermatozoa and ova from both groups share many of the proteins that are thought to be involved in gamete interaction, and although epididymal sperm maturation is significant it is probably less complex in monotremes than in other mammals. However, the monotreme epididymis is unique in forming spermatozoa into bundles of 100 with greatly enhanced motility compared with individual spermatozoa. Bundle formation involves a highly organised interaction with epididymal proteins, and the bundles persist during incubation in vitro, except in specialised medium, in which spermatozoa separate after 2–3 h incubation. It is suggested that this represents an early form of capacitation.

Additional keywords: epididymis, fertilisation.


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