Role of the ionic environment and internal pH on sperm activity.

In most species, once formed in the testis, spermatozoa are bathed in a fluid where they remained immobile and with a very low level of metabolism. This immotile status is understandable in view of the need to preserve the sperm energy reserve and to decrease the risk of alteration to membranes, internal structures and biochemical compounds by endogenous oxidizing agents produced by mitochondrial activity. This quiescent phase can be of different lengths and finishes when the semen is released into the external environment where the spermatozoa become motile and metabolically active. For invertebrates, and some fish, sexual activity is generally seasonal and fertilization is external. Spermatozoa, once differentiated in the gonad, remain there completely quiescent until they are released into the external medium, which is either fresh water or sea water. Dilution of the testicular fluid surrounding the spermatozoa allows the initiation of motility and metabolism. In fact, this seminal fluid has an inhibitory effect on sperm activity. For birds and mammals (including humans), the situation is much more complex. In these species, sperm production is almost continuous although for some of them, seasonal variations occur. When spermatozoa are released from the Sertoli cells, they are rapidly exported from the testis to the epididymis where the composition of the surrounding medium is profoundly modified. For most species, the spermatozoa remain immobile in the lower part of the epididymis, even though they have gained the capability to be fully motile as shown by dilution in an adequate medium. In vivo, motility is activated when the spermatozoa are mixed with secretions from the different accessory glands during ejaculation. This paper will review the role played by environmental factors, such as ions, in the activation of sperm motility and metabolism of different species of invertebrates and vertebrates. Special attention is given to changes in sperm internal pH, its regulation and role in the activation of sperm axonemal movement.