山溪鲵精巢的显微与超微结构研究
|
摘要
精子发生是发育生物学的重要内容之一，性激素对精子发生的调节是一种重要的调节方式。间质细胞是精巢中雄激素最主要的分泌细胞，对生精细胞的分化成熟起着至关重要的作用。支持细胞在生精细胞发育中，处于信息和物质交换的中心地位。因此，研究三者之间的关系一直是生殖内分泌学的核心问题。山溪鲵(Batrachuperus Pinchonii Davi)属两栖纲、有尾目、小鲵科，可做药用、食用，具有一定的经济开发价值。为深入探讨其精子发育机制，丰富其生殖生物学基础理论，为两栖动物资源开发奠定基础。本研究以山溪鲵为材料，就其精子发生和精子发育过程中支持细胞和间质细胞的关系进行了显微和超微结构研究。研究结果如下：
1 山溪鲵的精子发生属于不连续型的精子发生，有典型的季节变化。精子发生起始于4月，此时次级精原细胞占优势。在5月份，精巢小叶开始出现初级精母细胞。在6月的精巢中，初级精母细胞开始进行成熟分裂，形成次级精母细胞。在7月末精子细胞开始变态形成成熟精子。8月末成熟精子开始出现。9月份，精巢中出现大量精子。10月份，精巢中成熟精子开始减少。
2 在山溪鲵，4月份精巢尾部有一个特殊的结构一管泡结构。在管泡壁周围有分泌物，管泡中有成熟精子的存在。推断其具有分泌营养物质功能，可以贮藏精于过冬。在两栖动物精巢的尾部发现贮藏精子的特殊结构，目前未见报道。
3 山溪鲵精子与蟾蜍等无尾类精子结构比较相似，精子尾部具有轴丝纤维、轴丝和波动膜等结构。但山溪鲵有尾类轴丝旁纤维与轴丝微管的第八组临近，而蟾蜍等无尾类轴丝旁纤维与轴丝微管的第三组临近，与无尾类差异显著。
4 支持细胞的形态变化有明显季节变化。在精子形成时期，支持细胞胞质内有线粒体、滑面内质网和脂滴等类固醇激素分泌的细胞特征，可能分泌类固醇激素。支持细胞胞质内有糖原颗粒，对生精细胞有营养机能，并且有初级溶酶体和髓样小体，有吞噬作用。
5 在9月，支持细胞退化以后，间质细胞具有典型的类固醇激素分泌特征：胞质内具有管状嵴的线粒体、滑面内质网和脂滴。间质细胞的形态特征会随着季节变化而变化。
6 从形态上观察，支持细胞与间质细胞之间在机能上既相互抑制，又相互依赖。
Spermatogenesis is one of the most important section of developmental biology. Sex hormone is one of factors that regulate Spermatogenesis. Interstitial cell, that regulate germ cell differentiation and Spermatogenesis is the main testosterone source in the testis. The sertoli cell is the central point of exchange information and material. The relationship among the germ cell, sertoli cell and interstitial cell is the key problem in the reproductive endocrinology. The little salamander (Batrachuperus pinchonii) of urodele Amphibians can be used as medicine and food, and have some economic value. In order to reveal the developmental mechanism deeply, add documents to the reproductive biology, and lay the foundation of amphibians exploring. The microstructural and ultrastructural examination of the litter salamander testis was carried out to show the Spermatogenesis and the relationship between the sertoli cell and interstitial cell during the Spermatogenesis. The results as follows.
1 The testicular structure and reproductive cycle of B. pinchonii was studied .The testes of B.pinchonii are lobular type. In April, the testes were at the stage of spermatogonic proliferation, when primary spermatogonia divided and developed into secondary spermatogonia, which and sertoli cell form spermatogenic cysts. In May, the spermatocyte was observed in testis. In June, the spermatocyte was in the first meiotic division. In July, the testis entered spermiogenesis. During this stage, germ cells in the cysts developed into sperms finally. In August, the sperms appeared in cysts. In September, the testis was filled with the mature sperms. In October, sperm in the testis began to decrease. Spermatogenesis in this species follows an annual cycle.
2 In April, some special structure, which consists of columnar cells, was observed in the caudal testis. In this structure, there were some mature sperms and some material that nourish the sperms and make them go through the winter. The special structure, which can store sperm, was not reported till now,
3 The sperm of the little salamander is similar to that of toad in shape, both of them had the axonema, axial fiber and undulating membrane in their tail. But there is still some significant difference between them: the juxta-axonemal fiber is close to the eighth axial microtubular in the sperm of the little salamander, while the one is close
to the third one in the toad.
4 In the spermiogenesis, there were smooth endoplasmic reticulum, lipid droplet and mitochondria in sertoli cell. These characteristics revealed that the sertoli cell might have the function of secreting steroid. In the cytoplasm, there were some glycogen that nourished the spermatozoa, and some primary lysomes, residual bodies and myelin body that showed the phagocytosis function.
5 In September, when the sperms were released, the sertoli cells seemed to degenerate and the interstitial cells transformed into the Leydig cell, there cells shows the ultrastructural characteristics of steroid-secreting cells: some developed endoplasmic reticulum, mitochondria with tubular cristae and lipid droplets. With the change of season, their morphology would change.
6 The sertoli cell and interstitial cells not only inhibited each other, but depended on each other as well.
1 山溪鲵的精子发生属于不连续型的精子发生，有典型的季节变化。精子发生起始于4月，此时次级精原细胞占优势。在5月份，精巢小叶开始出现初级精母细胞。在6月的精巢中，初级精母细胞开始进行成熟分裂，形成次级精母细胞。在7月末精子细胞开始变态形成成熟精子。8月末成熟精子开始出现。9月份，精巢中出现大量精子。10月份，精巢中成熟精子开始减少。
2 在山溪鲵，4月份精巢尾部有一个特殊的结构一管泡结构。在管泡壁周围有分泌物，管泡中有成熟精子的存在。推断其具有分泌营养物质功能，可以贮藏精于过冬。在两栖动物精巢的尾部发现贮藏精子的特殊结构，目前未见报道。
3 山溪鲵精子与蟾蜍等无尾类精子结构比较相似，精子尾部具有轴丝纤维、轴丝和波动膜等结构。但山溪鲵有尾类轴丝旁纤维与轴丝微管的第八组临近，而蟾蜍等无尾类轴丝旁纤维与轴丝微管的第三组临近，与无尾类差异显著。
4 支持细胞的形态变化有明显季节变化。在精子形成时期，支持细胞胞质内有线粒体、滑面内质网和脂滴等类固醇激素分泌的细胞特征，可能分泌类固醇激素。支持细胞胞质内有糖原颗粒，对生精细胞有营养机能，并且有初级溶酶体和髓样小体，有吞噬作用。
5 在9月，支持细胞退化以后，间质细胞具有典型的类固醇激素分泌特征：胞质内具有管状嵴的线粒体、滑面内质网和脂滴。间质细胞的形态特征会随着季节变化而变化。
6 从形态上观察，支持细胞与间质细胞之间在机能上既相互抑制，又相互依赖。
Spermatogenesis is one of the most important section of developmental biology. Sex hormone is one of factors that regulate Spermatogenesis. Interstitial cell, that regulate germ cell differentiation and Spermatogenesis is the main testosterone source in the testis. The sertoli cell is the central point of exchange information and material. The relationship among the germ cell, sertoli cell and interstitial cell is the key problem in the reproductive endocrinology. The little salamander (Batrachuperus pinchonii) of urodele Amphibians can be used as medicine and food, and have some economic value. In order to reveal the developmental mechanism deeply, add documents to the reproductive biology, and lay the foundation of amphibians exploring. The microstructural and ultrastructural examination of the litter salamander testis was carried out to show the Spermatogenesis and the relationship between the sertoli cell and interstitial cell during the Spermatogenesis. The results as follows.
1 The testicular structure and reproductive cycle of B. pinchonii was studied .The testes of B.pinchonii are lobular type. In April, the testes were at the stage of spermatogonic proliferation, when primary spermatogonia divided and developed into secondary spermatogonia, which and sertoli cell form spermatogenic cysts. In May, the spermatocyte was observed in testis. In June, the spermatocyte was in the first meiotic division. In July, the testis entered spermiogenesis. During this stage, germ cells in the cysts developed into sperms finally. In August, the sperms appeared in cysts. In September, the testis was filled with the mature sperms. In October, sperm in the testis began to decrease. Spermatogenesis in this species follows an annual cycle.
2 In April, some special structure, which consists of columnar cells, was observed in the caudal testis. In this structure, there were some mature sperms and some material that nourish the sperms and make them go through the winter. The special structure, which can store sperm, was not reported till now,
3 The sperm of the little salamander is similar to that of toad in shape, both of them had the axonema, axial fiber and undulating membrane in their tail. But there is still some significant difference between them: the juxta-axonemal fiber is close to the eighth axial microtubular in the sperm of the little salamander, while the one is close
to the third one in the toad.
4 In the spermiogenesis, there were smooth endoplasmic reticulum, lipid droplet and mitochondria in sertoli cell. These characteristics revealed that the sertoli cell might have the function of secreting steroid. In the cytoplasm, there were some glycogen that nourished the spermatozoa, and some primary lysomes, residual bodies and myelin body that showed the phagocytosis function.
5 In September, when the sperms were released, the sertoli cells seemed to degenerate and the interstitial cells transformed into the Leydig cell, there cells shows the ultrastructural characteristics of steroid-secreting cells: some developed endoplasmic reticulum, mitochondria with tubular cristae and lipid droplets. With the change of season, their morphology would change.
6 The sertoli cell and interstitial cells not only inhibited each other, but depended on each other as well.
引文
[1]丁允闽，叶世隽，陈咏梅．大鼠睾丸支持细胞在硫酸糖蛋白-2MRA水平的生后调节．解剖学报，1995，26(4)：379-382．
[2]尤永隆，林丹军．尼罗罗非鱼精于形成中核内囊泡的释放．动物学报，1998，44(3)：257-263．
[3]尤永隆，林丹军，陈莲云．大黄鱼的精子发生．动物学研究，2001，22(6)：461-466．
[4]方永强，翁幼竹，胡晓霞．性类固醇激素及其受体在文昌鱼性腺和神经系统中的分布．动物学报，2001，47(4)：398-403．
[5]方永强．文昌鱼 sertoli 细胞超微结构的进一步研究．动物学报，1991，37(2)：123-125．
[6]方永强，赵维信，魏华．文昌鱼类固醇激素水平与性腺发育相关性研究．科学通报，1993，38(8)：744-746．
[7]本特利，方永强译．脊椎动物比较内分泌学．北京人民卫生出版社，1984，43-47．
[8]石磊，汗夏华，高洪娟，刘以训．甾体激素合成快速调节蛋白 mRNA 在卵泡发育和闭锁过程的表达．动物学报，2000，46(3)：324-329．
[9]阳爱生，卞伟，刘运清．大鲵性腺发育的组织学观察．动物学报，1981，27(3)：240-246．
[10]何德奎，陈毅峰，蔡斌．纳木错裸鲤性腺发育的组织学研究．水生生物学报，2001，25(1)：1-9．
[11]吴美锡，傅福英．中国水蛇 Enhydfis chinensis (Cray)生殖周期与生殖型式的研究．动物学报，1989，35(1)：82-87
[12]吴美锡，傅福英．中国水蛇睾丸组织化学的研究．两栖爬行动物学报，1988，75-80．
[13]张旭晨，王所安．细鳞鱼精巢超微结构和精子发生．动物学报，1992，38(4)：355-357．
[14]张耀光，罗泉笙，钟明超．长吻鮠精巢发育的分期及精子的发生和形成．动物学研究，1992，13(3)：281-287．
[15]李远友，林浩然，沈放．蛙类下丘脑-脑垂体-性腺轴的内分泌调节．动物学研究，1998，19(1)：83-89．
[16]杨勇正，滕松山．大鼠睾丸间质细胞的小管结构及其功能的初步研究．动物学报，1983，29(2)：112-115．
[17]林丹军，尤永隆，陈莲云．卵胎生硬骨鱼褐菖蚰精巢的周期发育．动物学研究，2000，21(5)：337-342．
[18]林丹军，尤永隆．褐菖鲇精细胞晚期的变化及精子结构研究．动物学研究，1998，19(5)：359-366．
[19]林丹军，尤永隆．中国雨蛙精子形成的研究．动物学报，2000，46(4)：376-384．
[20]林丹军，尤永隆，钟秀容．中国雨蛙精子结构及其在系统发育上的意义．动物学研究，1999，5，20(3)：161-167．
[21]施瑔芳．鱼类性腺发育研究新进展．水生生物学报，1988，12(3)：248-255．
[22]祝辉，于宁妮，林敏，周作民，沙家豪．hCG对成年小鼠睾丸 Lcydig 细胞中睾酮生成酶的调节作用．解剖学报，2001，32(2)：146-150．
[23]钟慈声 主编．细胞和组织的超微结构．人民卫生出版社(北京)，1984，7-26．
[24]徐剑．西藏山溪鲵生殖腺的季节变化．动物学杂志，1995，30(3)：195-217．
[25]特纳．贝格纳尔著，刘一训译．普通内分泌学．科学出版社(北京)，1983．
[26]莫慧英．黑斑蛙和中华大蟾蜍精子的超显微结构研究．动物学研究，1985，6(4)：381-388．
[27]高建民，郑翠芳，张彦定，张秋金，林玮．渔游蛇生精上皮超微结构的季节性变化．解剖学报，2001，32(1)：76-79．
[28]黄海燕，朱立基，宗书东．精浆聚集素在大鼠体内分布的免疫组织化学研究．解剖学报1997，28(3)：290-293．
[29]管汀鹭，黄丹青，黄国屏．金鱼精巢的细胞结构与精子的发生和形成．水生生物学报，1990，14(3)：233-237．
[30]Aoki,A.,Fawett,D.W It ther a local feedback from the seminiferous tubules affecting activity of the Leydig cell? Biol. Reprod. 1972,19:144-158.
[31]Ari V.T Reproductive physiology of vertebrates. Cornell press, 1983,104-147.
[32]Asahina.K,Suzuki.K, Aida.K,Hibiya.T,and Tamaoki.B. Relationship between the structures and steriodogenic functions of the testes of the urohaze-goby (Glossogobius olivaceus).Gen.Comp.Endocrinol.1985,57:281-292.
[33]Bao.S.N., Dalton.G.C, Oliverira.S.F. Spermiogenesis in odontophrynus cultripes(amphibia anuraseptodactdae):ultrastructual and cytochemical studies of proteins
using E-PTA J. Morphol., 1991,307-314．
[34] Callard,G.V., canick,J,A. Pudney.J.,Estrogen. Synthesis in leydig cells: structrural-functional correlations in necturus testis. Biol. Reprod. 1980, 23;461-479．
[35] Camatini.M, Franchi.E, and Curtis.I.D, Ultrastructure of leydig cells in the African Green Monkey. J. Ultrastructure Research .1981, 76:224-234．
[36] Chieffi.P,Minucci.S,Cobellis.G,Fasano.S,Pierantoni,R.,Changes in proto-oncogene activity in the testis of the frog,(Rana csculenta)during the annual reproductive cycle. Gen.Comp.Endocrinol. 1995,99(2) :127-136．
[37] Christensen,A.K. The fine structure of testicular interstitial cells in guinea pigs. J.Cell Biol. 1965,26:911
[38] Christensen.AK.,Gillim.S.W. In the gonads, New york,Appleton-century-Crofts. 1969,415．
[39] Cobellis.G,Pierantioni.R,Fasano.S c-fos-and c-jun-like mRNA expression in frog (Rana csculenta) testis during the annual reproductive cycle. Gen. Comp. Endocrinol. 1997, 106(1) :23-29．
[40] Drosdowsky M,Menon KM,Forchielli E,et al. Requirements of the cholesterol-side-chaincleaving enzymes of rat testis mitochondria. Biochem Biophys Acta, 1965, 104(1) :354-358．
[41] Ezeasor.D.E, Light and electron microscopical obervation on the Leydig cells of the scrotal and abdominal testes of naturally unilateral cryptorchid west African dwaf goats. J. Anat.1985, 141:27-40．
[42] Fraile,B.,R.Paniagua,M.C.Rodriguez.,saez.F.J. Annual changes in the number, testosterone content,and Ultrastructure of glandular tissue cells of the testis in the marbled newts Triturus marmoratus. J.Anat.1989, 167:82-92．
[43] Fraile.B, Panlagua.R,Rodriguez.M.C,and Saez.F.J, Effect of photoperiod on testicular function in Blinded and non-Blinded marbled newt, Triturus marmoratusc (caudata, salamandridae). J. Morphol. 1989, 200:151-162．
[44] Fraile.B,saez.F.J,and panlagua.R,The cycle of follicuiar and interstitial cells (leydig cells)in the testis of the marbled newt, Triturus marmoratus (caudata,salamandridae). J. Morphol.1990, 204:89-101．
[45] Garnier.D.H, Androger and estrogen levels in the plasma of pleurodeles waltl michah, during the annual cycle.l.male cycle. Gen. Comp. Endocrinol. 1985, 58:376-385．
[46] Gibert,S.F. Development Biology. 5 th edition.sinauer Associates, 1997, 855-859．
[47] Jones P.R, and Butler.R.D. Spermatozoon ultrastructure of Platichthys flesus, Cells of ultrastructure and moleculecular structure research. 1988, 98:71-82．
[48] Karl.J,capel.B. Senoli cells of the mouse testis originate from the coelomic epithelium. Devel. Biol. 1998, 203:323-333．
[49] Kuwahara.S, Mizukami.T, Omura.M, Hagihara.M, linuma.Y, Shimizu.Y., Tamada.H., Tsukamoto Y, Nishida. T., Sasaki.F. Seasonal changes in the hypothalamo-pituitary-testes axis of the Japanese wood mouse (Apodemus speciosus). Anat. Rec. 2000, 260:366-372．
[50] Lin M,Harman.A,and Rodger.J.C. Spermiogenesis and spermiation in a marsupial,the tammar wallaby (Macropus eugenii). J.Anat. 1997, 190:377-395．
[51] Lou.Y.H and Takahashi.H, Spermiogenesis in the nile tilapia oreochromis niloticus with notes on a unique pattern of nuclear chromatin condensation. J. Morphol. 1989, 200:321-330．
[52] Mechan.T,Scnlatt.S,.O Bryan.M.K ,Kretser.D.M.de, and Loveland.K.L, Regulation of germ cell and sertoli cell development by activin ,follistatin,and FSH. Devel. biol. 2000, 220:225-237．
[53] Minucci.S.,Rienzo.G.D,,Sena.R.D,CoBellis.G,Meccarielo.R,Dierantoni.R.and Fasano.S.. Effects of multiple injections of ethane 1,2-dimethane sulphonate (EDS) on the frog Rana esculenta, testicular activity. J. Exp. Zool., 2000, 287:384-393．
[54] Moriya,M. Ch.Kataginiri Immunoelectron microscopic localization of sperm-specific nuclearbasic proteins during spermatogenesis in anuran amphibians. Develop Groth. Differ. 1991, 33(1) : 19-27．
[55] Norris.D.O,Norman.M.F, Pancak.M.K, Puvall.D, Seasonal variations in spematogenesis ,testicular weights, vasa deferentia,and androgen levels in neotenic male tiger salamandars, Ambystoma tigrinum. Gen. Comp. Endocrinol. 1985, 60:51-57．
[56] Pudney.J,Canick.J.A, Mak.P. Callard G.V., The differentiation of leydig cells,Steroidogenesis, and the spermatogenetic wave in the testis of Necturus maculosus. Gen.Comp.Endocrinol., 1983, 50:43-66．
[57] Pudney.J.,Callard.G. Organization of interstial tissue in the testis of the Salamander Necturus maculosus (Caudata:Proteidae).J. Morphol.1984, 181:87-95．
[58] Rastogi R.K. Serminal vesicles and sperm duct of the Indian catfish Mystus tengara
with particular reference to their seasonal cycle. Act Anat. 1969, 72(4) :624-639．
[59] Rastogi.R.R, Daunara.J, Ela.L., Krasavich.m.A, Reproduction in the Mexican leaf frog, Pachymedusa dacnicolor Iv. Spermatogenesis: a light and ultrasonic study. J. Morphol., 1988, 197:207-302．
[60] Rosenblum P.M.,Pudey.J.Callard I.P. Gonadal morphology enzyme histochemistry and Plasma sterid levels during the annual reproductive cycle of male and female brown bullhead catfish, Ictalurus nebulosus Lesueur. J. fish biol.1987, 31(3) :325-342．
[61] Saze.F.J,Madrid.J.F.,Aparicio.R. Carbohydrate moieties of the interstitial and glandular tissues of the amphibian Pleurodeles waltl testis shows by lectin histochemistry. J.Anat, 2001, 198:47-56
[62] Schafer.M.,Nayernia.K,Engel.W,and Schafer.U, Translational control in spermatogenesis. Develop, biol. 1995, 172:344-352．
[63] Shikita,M, and P.F.Hall. The action of human chorionic gonadotropin in vivo upon microsomal enzyme on immature rat testis. Biochem.Biophys. Actal. 1967, 36: 484．
[64] Soley.J.T. Nuclear morphogenesis and the role of manchette during spermiogenesis in the osterich (struthio camelus). J. Anat 1997, 190:563-576．
[65] Specker.J.L.,Moore.F.l. Annual cycle of plasma androgens and testicular composition in the rough-skinned newts, Taricha granulose. Gen.Comp.Endocrinol.1980, 42:297-303．
[66] Sprando.R.L and Russell. L.D. Spermiogenesis in the Bluegill (lepomis macrochirus): a study of cytoplasmic events including cell volume changes and cytoplasmic elimination. J. morphol. 1988, 198:165-177．
[67] Sprardo.R.l., Russell..L.D, Spermiogenis in the bullfrog (Rana catesbeiana): a study of cytoplasmic events including cell volume changes and cytoplasmic elimination, J. Morphol., 1998, 198:303-319．
[68] Uribe.M.C.A, Rios.G.G, and Brandon.R.A, Spermatogenesis in the urodele Ambystoma dumerilli., J. Morphol., 1994, 222:287-299．
[69] Wake.M.H, comparative Morphology of caecilian sperm (amphibia:Gymnophioha). J. morphol. 1994, 221:261-276．
[70] Xie.Q, Mackay.S, Ullmann.S.L,Gilmore.D.P Payne.A.P, Testis development in the opossum Monodelphis domestica . J. Anat. 1996, 189:393-406．
[71] Yizhong Yin, Stahl.B.C, Dewolf.W.C, Morgentaler.A, P53-mediated germ cell quality control in spermatogenesis, Develop, biol. 1998, 204:165-171．
[72] Yoakim E.G. Studies on the seminal Vesicle on the Nile catfish, Schilbe mystus. Bulletin Faculty of science assiut university. 1976, 5(1) : 107-118．
[2]尤永隆，林丹军．尼罗罗非鱼精于形成中核内囊泡的释放．动物学报，1998，44(3)：257-263．
[3]尤永隆，林丹军，陈莲云．大黄鱼的精子发生．动物学研究，2001，22(6)：461-466．
[4]方永强，翁幼竹，胡晓霞．性类固醇激素及其受体在文昌鱼性腺和神经系统中的分布．动物学报，2001，47(4)：398-403．
[5]方永强．文昌鱼 sertoli 细胞超微结构的进一步研究．动物学报，1991，37(2)：123-125．
[6]方永强，赵维信，魏华．文昌鱼类固醇激素水平与性腺发育相关性研究．科学通报，1993，38(8)：744-746．
[7]本特利，方永强译．脊椎动物比较内分泌学．北京人民卫生出版社，1984，43-47．
[8]石磊，汗夏华，高洪娟，刘以训．甾体激素合成快速调节蛋白 mRNA 在卵泡发育和闭锁过程的表达．动物学报，2000，46(3)：324-329．
[9]阳爱生，卞伟，刘运清．大鲵性腺发育的组织学观察．动物学报，1981，27(3)：240-246．
[10]何德奎，陈毅峰，蔡斌．纳木错裸鲤性腺发育的组织学研究．水生生物学报，2001，25(1)：1-9．
[11]吴美锡，傅福英．中国水蛇 Enhydfis chinensis (Cray)生殖周期与生殖型式的研究．动物学报，1989，35(1)：82-87
[12]吴美锡，傅福英．中国水蛇睾丸组织化学的研究．两栖爬行动物学报，1988，75-80．
[13]张旭晨，王所安．细鳞鱼精巢超微结构和精子发生．动物学报，1992，38(4)：355-357．
[14]张耀光，罗泉笙，钟明超．长吻鮠精巢发育的分期及精子的发生和形成．动物学研究，1992，13(3)：281-287．
[15]李远友，林浩然，沈放．蛙类下丘脑-脑垂体-性腺轴的内分泌调节．动物学研究，1998，19(1)：83-89．
[16]杨勇正，滕松山．大鼠睾丸间质细胞的小管结构及其功能的初步研究．动物学报，1983，29(2)：112-115．
[17]林丹军，尤永隆，陈莲云．卵胎生硬骨鱼褐菖蚰精巢的周期发育．动物学研究，2000，21(5)：337-342．
[18]林丹军，尤永隆．褐菖鲇精细胞晚期的变化及精子结构研究．动物学研究，1998，19(5)：359-366．
[19]林丹军，尤永隆．中国雨蛙精子形成的研究．动物学报，2000，46(4)：376-384．
[20]林丹军，尤永隆，钟秀容．中国雨蛙精子结构及其在系统发育上的意义．动物学研究，1999，5，20(3)：161-167．
[21]施瑔芳．鱼类性腺发育研究新进展．水生生物学报，1988，12(3)：248-255．
[22]祝辉，于宁妮，林敏，周作民，沙家豪．hCG对成年小鼠睾丸 Lcydig 细胞中睾酮生成酶的调节作用．解剖学报，2001，32(2)：146-150．
[23]钟慈声 主编．细胞和组织的超微结构．人民卫生出版社(北京)，1984，7-26．
[24]徐剑．西藏山溪鲵生殖腺的季节变化．动物学杂志，1995，30(3)：195-217．
[25]特纳．贝格纳尔著，刘一训译．普通内分泌学．科学出版社(北京)，1983．
[26]莫慧英．黑斑蛙和中华大蟾蜍精子的超显微结构研究．动物学研究，1985，6(4)：381-388．
[27]高建民，郑翠芳，张彦定，张秋金，林玮．渔游蛇生精上皮超微结构的季节性变化．解剖学报，2001，32(1)：76-79．
[28]黄海燕，朱立基，宗书东．精浆聚集素在大鼠体内分布的免疫组织化学研究．解剖学报1997，28(3)：290-293．
[29]管汀鹭，黄丹青，黄国屏．金鱼精巢的细胞结构与精子的发生和形成．水生生物学报，1990，14(3)：233-237．
[30]Aoki,A.,Fawett,D.W It ther a local feedback from the seminiferous tubules affecting activity of the Leydig cell? Biol. Reprod. 1972,19:144-158.
[31]Ari V.T Reproductive physiology of vertebrates. Cornell press, 1983,104-147.
[32]Asahina.K,Suzuki.K, Aida.K,Hibiya.T,and Tamaoki.B. Relationship between the structures and steriodogenic functions of the testes of the urohaze-goby (Glossogobius olivaceus).Gen.Comp.Endocrinol.1985,57:281-292.
[33]Bao.S.N., Dalton.G.C, Oliverira.S.F. Spermiogenesis in odontophrynus cultripes(amphibia anuraseptodactdae):ultrastructual and cytochemical studies of proteins
using E-PTA J. Morphol., 1991,307-314．
[34] Callard,G.V., canick,J,A. Pudney.J.,Estrogen. Synthesis in leydig cells: structrural-functional correlations in necturus testis. Biol. Reprod. 1980, 23;461-479．
[35] Camatini.M, Franchi.E, and Curtis.I.D, Ultrastructure of leydig cells in the African Green Monkey. J. Ultrastructure Research .1981, 76:224-234．
[36] Chieffi.P,Minucci.S,Cobellis.G,Fasano.S,Pierantoni,R.,Changes in proto-oncogene activity in the testis of the frog,(Rana csculenta)during the annual reproductive cycle. Gen.Comp.Endocrinol. 1995,99(2) :127-136．
[37] Christensen,A.K. The fine structure of testicular interstitial cells in guinea pigs. J.Cell Biol. 1965,26:911
[38] Christensen.AK.,Gillim.S.W. In the gonads, New york,Appleton-century-Crofts. 1969,415．
[39] Cobellis.G,Pierantioni.R,Fasano.S c-fos-and c-jun-like mRNA expression in frog (Rana csculenta) testis during the annual reproductive cycle. Gen. Comp. Endocrinol. 1997, 106(1) :23-29．
[40] Drosdowsky M,Menon KM,Forchielli E,et al. Requirements of the cholesterol-side-chaincleaving enzymes of rat testis mitochondria. Biochem Biophys Acta, 1965, 104(1) :354-358．
[41] Ezeasor.D.E, Light and electron microscopical obervation on the Leydig cells of the scrotal and abdominal testes of naturally unilateral cryptorchid west African dwaf goats. J. Anat.1985, 141:27-40．
[42] Fraile,B.,R.Paniagua,M.C.Rodriguez.,saez.F.J. Annual changes in the number, testosterone content,and Ultrastructure of glandular tissue cells of the testis in the marbled newts Triturus marmoratus. J.Anat.1989, 167:82-92．
[43] Fraile.B, Panlagua.R,Rodriguez.M.C,and Saez.F.J, Effect of photoperiod on testicular function in Blinded and non-Blinded marbled newt, Triturus marmoratusc (caudata, salamandridae). J. Morphol. 1989, 200:151-162．
[44] Fraile.B,saez.F.J,and panlagua.R,The cycle of follicuiar and interstitial cells (leydig cells)in the testis of the marbled newt, Triturus marmoratus (caudata,salamandridae). J. Morphol.1990, 204:89-101．
[45] Garnier.D.H, Androger and estrogen levels in the plasma of pleurodeles waltl michah, during the annual cycle.l.male cycle. Gen. Comp. Endocrinol. 1985, 58:376-385．
[46] Gibert,S.F. Development Biology. 5 th edition.sinauer Associates, 1997, 855-859．
[47] Jones P.R, and Butler.R.D. Spermatozoon ultrastructure of Platichthys flesus, Cells of ultrastructure and moleculecular structure research. 1988, 98:71-82．
[48] Karl.J,capel.B. Senoli cells of the mouse testis originate from the coelomic epithelium. Devel. Biol. 1998, 203:323-333．
[49] Kuwahara.S, Mizukami.T, Omura.M, Hagihara.M, linuma.Y, Shimizu.Y., Tamada.H., Tsukamoto Y, Nishida. T., Sasaki.F. Seasonal changes in the hypothalamo-pituitary-testes axis of the Japanese wood mouse (Apodemus speciosus). Anat. Rec. 2000, 260:366-372．
[50] Lin M,Harman.A,and Rodger.J.C. Spermiogenesis and spermiation in a marsupial,the tammar wallaby (Macropus eugenii). J.Anat. 1997, 190:377-395．
[51] Lou.Y.H and Takahashi.H, Spermiogenesis in the nile tilapia oreochromis niloticus with notes on a unique pattern of nuclear chromatin condensation. J. Morphol. 1989, 200:321-330．
[52] Mechan.T,Scnlatt.S,.O Bryan.M.K ,Kretser.D.M.de, and Loveland.K.L, Regulation of germ cell and sertoli cell development by activin ,follistatin,and FSH. Devel. biol. 2000, 220:225-237．
[53] Minucci.S.,Rienzo.G.D,,Sena.R.D,CoBellis.G,Meccarielo.R,Dierantoni.R.and Fasano.S.. Effects of multiple injections of ethane 1,2-dimethane sulphonate (EDS) on the frog Rana esculenta, testicular activity. J. Exp. Zool., 2000, 287:384-393．
[54] Moriya,M. Ch.Kataginiri Immunoelectron microscopic localization of sperm-specific nuclearbasic proteins during spermatogenesis in anuran amphibians. Develop Groth. Differ. 1991, 33(1) : 19-27．
[55] Norris.D.O,Norman.M.F, Pancak.M.K, Puvall.D, Seasonal variations in spematogenesis ,testicular weights, vasa deferentia,and androgen levels in neotenic male tiger salamandars, Ambystoma tigrinum. Gen. Comp. Endocrinol. 1985, 60:51-57．
[56] Pudney.J,Canick.J.A, Mak.P. Callard G.V., The differentiation of leydig cells,Steroidogenesis, and the spermatogenetic wave in the testis of Necturus maculosus. Gen.Comp.Endocrinol., 1983, 50:43-66．
[57] Pudney.J.,Callard.G. Organization of interstial tissue in the testis of the Salamander Necturus maculosus (Caudata:Proteidae).J. Morphol.1984, 181:87-95．
[58] Rastogi R.K. Serminal vesicles and sperm duct of the Indian catfish Mystus tengara
with particular reference to their seasonal cycle. Act Anat. 1969, 72(4) :624-639．
[59] Rastogi.R.R, Daunara.J, Ela.L., Krasavich.m.A, Reproduction in the Mexican leaf frog, Pachymedusa dacnicolor Iv. Spermatogenesis: a light and ultrasonic study. J. Morphol., 1988, 197:207-302．
[60] Rosenblum P.M.,Pudey.J.Callard I.P. Gonadal morphology enzyme histochemistry and Plasma sterid levels during the annual reproductive cycle of male and female brown bullhead catfish, Ictalurus nebulosus Lesueur. J. fish biol.1987, 31(3) :325-342．
[61] Saze.F.J,Madrid.J.F.,Aparicio.R. Carbohydrate moieties of the interstitial and glandular tissues of the amphibian Pleurodeles waltl testis shows by lectin histochemistry. J.Anat, 2001, 198:47-56
[62] Schafer.M.,Nayernia.K,Engel.W,and Schafer.U, Translational control in spermatogenesis. Develop, biol. 1995, 172:344-352．
[63] Shikita,M, and P.F.Hall. The action of human chorionic gonadotropin in vivo upon microsomal enzyme on immature rat testis. Biochem.Biophys. Actal. 1967, 36: 484．
[64] Soley.J.T. Nuclear morphogenesis and the role of manchette during spermiogenesis in the osterich (struthio camelus). J. Anat 1997, 190:563-576．
[65] Specker.J.L.,Moore.F.l. Annual cycle of plasma androgens and testicular composition in the rough-skinned newts, Taricha granulose. Gen.Comp.Endocrinol.1980, 42:297-303．
[66] Sprando.R.L and Russell. L.D. Spermiogenesis in the Bluegill (lepomis macrochirus): a study of cytoplasmic events including cell volume changes and cytoplasmic elimination. J. morphol. 1988, 198:165-177．
[67] Sprardo.R.l., Russell..L.D, Spermiogenis in the bullfrog (Rana catesbeiana): a study of cytoplasmic events including cell volume changes and cytoplasmic elimination, J. Morphol., 1998, 198:303-319．
[68] Uribe.M.C.A, Rios.G.G, and Brandon.R.A, Spermatogenesis in the urodele Ambystoma dumerilli., J. Morphol., 1994, 222:287-299．
[69] Wake.M.H, comparative Morphology of caecilian sperm (amphibia:Gymnophioha). J. morphol. 1994, 221:261-276．
[70] Xie.Q, Mackay.S, Ullmann.S.L,Gilmore.D.P Payne.A.P, Testis development in the opossum Monodelphis domestica . J. Anat. 1996, 189:393-406．
[71] Yizhong Yin, Stahl.B.C, Dewolf.W.C, Morgentaler.A, P53-mediated germ cell quality control in spermatogenesis, Develop, biol. 1998, 204:165-171．
[72] Yoakim E.G. Studies on the seminal Vesicle on the Nile catfish, Schilbe mystus. Bulletin Faculty of science assiut university. 1976, 5(1) : 107-118．