Luconi M, Krausz C, Forti G, Baldi E
Dipartimento di Fisiopatologia Clinica, Universita' di Firenze, Florence, Italy.
Biol Reprod. 1996 Jul;55(1):207-16. doi: 10.1095/biolreprod55.1.207.
Capacitation of spermatozoa, a complex process occurring after sperm ejaculation, is required to obtain fertilization of the oocyte in vivo and in vitro. Although most of the biochemical/ biophysical events that occur during capacitation in vitro have been characterized, the molecular mechanisms underlying these complex events are still obscure. Increases of intracellular free Ca2+ concentrations ([Ca2+]i) and protein tyrosine phosphorylation have previously been demonstrated during in vitro capacitation of human spermatozoa. In the present study we investigated the relationship between extracellular/intracellular Ca2+, protein tyrosine phosphorylation, and tyrosine kinase and phosphatase activities during sperm capacitation. We report that the increase in tyrosine phosphorylation of several protein bands that occurs during sperm capacitation is independent of the presence of Ca2+ in the external medium and, at least partially, of the increase in [Ca2+]i occurring during the process. Indeed, the spontaneous increase in phosphorylation was still present in Ca(2+)-free/EGTA-containing-medium and in the presence of the intracellular Ca2+ chelator BAPTA/AM. Moreover, phosphorylation of proteins and protein tyrosine kinase (PTK) activity was enhanced if spermatozoa were incubated in Ca(2+)-free medium, suggesting the presence of Ca(2+)-inhibited tyrosine kinase(s) in human sperm. This hypothesis is further substantiated by the lower tyrosine phosphorylation observed after incubation with the ionophore A23187 and the endoplasmic Ca(2+)-ATPase inhibitor thapsigargin, which promote Ca2+ influx in human sperm. The ability of the cells to undergo acrosome reaction in response to progesterone, which can be considered a functional endpoint of capacitation, was highly compromised when spermatozoa were incubated in Ca(2+)-free medium or in the presence of EGTA, confirming that Ca2+ is required for sperm capacitation. Conversely, in the presence of erbstatin, a inhibitor of tyrosine kinase activity, which blunts tyrosine phosphorylation during capacitation, response to progesterone was maintained, suggesting that tyrosine phosphorylation must be kept at a low level (physiologically by the presence of Ca2+ in the external medium, or pharmacologically by the presence of erbstatin) in order to obtain response to progesterone. This mechanism may be important in vivo during sperm transit in the female genital tract to ensure appropriate timing of full capacitation in the proximity of the oocyte.
精子获能是精子射出后发生的一个复杂过程,是体内和体外使卵母细胞受精所必需的。尽管体外获能过程中发生的大多数生化/生物物理事件已得到表征,但这些复杂事件背后的分子机制仍不清楚。此前已证明,在人类精子体外获能过程中,细胞内游离钙离子浓度([Ca2+]i)升高以及蛋白质酪氨酸磷酸化增加。在本研究中,我们调查了精子获能过程中细胞外/细胞内钙离子、蛋白质酪氨酸磷酸化以及酪氨酸激酶和磷酸酶活性之间的关系。我们报告称,精子获能过程中出现的几条蛋白带酪氨酸磷酸化增加与细胞外培养基中钙离子的存在无关,并且至少部分与该过程中发生的[Ca2+]i升高无关。事实上,在无钙/含乙二醇双四乙酸(EGTA)的培养基中以及存在细胞内钙离子螯合剂1,2-双(2-氨基苯氧基)乙烷-N,N,N',N'-四乙酸/乙酰甲酯(BAPTA/AM)的情况下,磷酸化的自发增加仍然存在。此外,如果精子在无钙培养基中孵育,蛋白质磷酸化和蛋白质酪氨酸激酶(PTK)活性会增强,这表明人类精子中存在受钙离子抑制的酪氨酸激酶。在用离子载体A23187和内质网钙离子-ATP酶抑制剂毒胡萝卜素孵育后观察到较低的酪氨酸磷酸化,进一步证实了这一假设,离子载体A23187和毒胡萝卜素可促进钙离子流入人类精子。当精子在无钙培养基中或存在EGTA的情况下孵育时,细胞对孕酮发生顶体反应的能力(可被视为获能的一个功能终点)受到严重损害,这证实了钙离子是精子获能所必需的。相反,在存在酪氨酸激酶活性抑制剂埃伯他汀的情况下,对孕酮的反应得以维持,埃伯他汀会抑制获能过程中的酪氨酸磷酸化,这表明为了获得对孕酮的反应,酪氨酸磷酸化必须保持在低水平(在生理上通过细胞外培养基中钙离子的存在,或在药理上通过埃伯他汀的存在)。这种机制在体内精子在女性生殖道中运输过程中可能很重要,以确保在卵母细胞附近完全获能的适当时间。