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The nuclei of spermatozoa leaving the testies are transcritionally inactivated and are structurally stabilised by extensive association of sperm DNA with protamines, highly basic proteins that replace somatic histones during spermiogenesis. During sperm matureation within teh epididymis, protamine–SH is oxidised to -SS-(Bedford & Calvin, 1974).
Ryuzo Yanagimachi (Tue,) studied this question.