摘要
本文采用组织学方法和透射电镜技术对处于不同发育阶段、不同生理状态长吻魷脑垂体的超微结构以及卵巢组织学结构进行了较详细的研究。结果表明:
长吻魷脑垂体包括腺垂体和神经垂体两个部分,腺垂体组织中有六种分泌细胞即促肾上腺皮质激素(ACTH)分泌细胞、催乳激素(PRL)分泌细胞、生长激素(STH)分泌细胞、促性腺激素(GTH)分泌细胞、促甲状腺激素(TSH)分泌细胞、促黑色素细胞刺激激素(MSH)分泌细胞;神经垂体组织中存在A型(A_1、A_2)和B型神经分泌纤维,A型分泌纤维轴突中具有不同类型的分泌颗粒,B型分泌纤维轴突中含有许多透明小囊泡。
生殖季节腺垂体中与生殖有关的许多腺细胞分泌活动旺盛。ACTH细胞分泌物颗粒数量、内质网及线粒体均发生明显变化;GTH细胞胞质中充满电子密度较高的大、小两种分泌颗粒;STH细胞处于活跃的合成和分泌状态,分泌颗粒呈现出由融合到释放的状态;TSH细胞胞质中分泌颗粒排空,且电子密度降低,出现大空泡。繁殖后,GTH细胞中的分泌颗粒出现排空;TSH细胞有空泡;STH细胞中的粗面内质网呈现围核环状排列。秋季,在GTH细胞中发现GTH_1和GTH_2两种细胞,GTH_1细胞具有大量的分泌颗粒,并呈围核分布,GTH_2细胞胞质中具有数量较少的小圆形分泌颗粒和大圆形颗粒;STH细胞的结构无明显变化。冬季,GTH细胞核为椭圆形,有少量的分泌颗粒散布在细胞核的周围;STH细胞分泌颗粒数量较少,胞核变形,合成和分泌活动比较微弱;TSH细胞胞质中的分泌颗粒相当多。ACTH、PRL、MSH三种分泌细胞在生殖前后变化不明显。
各时期神经垂体主要变化表现为:产前神经分泌纤维具有较多球形的无被膜分泌颗粒;产后神经分泌纤维轴突中的分泌颗粒呈现排空;秋季神经垂体中部分神经分泌纤维轴突可见到分泌颗粒,但数量相对较少;冬季神经垂体几乎没有变化。
长吻魷卵母细胞发育可分为6个时相,卵巢依其形态结构特征,亦可分为6个时期。 Ⅰ期性腺只存在于1龄个体,最小性成熟个体为3龄。卵巢的成熟系数全年在0.72%~4.65%变动,繁殖季节为每年的5~7月份,产后卵巢很快退回到Ⅱ期,以Ⅲ期卵巢越冬,属一次性产卵类型。
With the method of histologkal section and electronic microscope, we studied the ultra-structure of pituitary and ovary histology of Long-snout catfish at different development stages and physiological conditions. The results were as follows:
The pituitary of Long-snout catfish included adenohypophysis and neurohypophysis. The adenohypophysis have six-cell types in common,i.e adrenocorticotro-phs(ACT- H),lactot-rophs(PRL),somatotrophs(STH),gonadotrophs(GTH), thyrotrophs(TSH), and Melanotroph (MSH) cells. In neurohypophysis, there were three types of neurosecretory fibres, types A1,A2 and B.There were different types'granules in type A neurosecretory fibres axone.In type B neurosecretory fibres axone, there were many small transparent vesicles.
During propagation seasons, secretory activity of gland cells related to reprodution in pituitary was vigorous. Numbers of secretory granules, oplasmic reticulums, mitochondrias in ACTH cells all changed distinctly. GTH cytoplast were full of big and small, high density electron secretory granules.STH cells were in activity synthesis and secretion, secretory granules were from fusion to release. TSH cells cytoplast evacuated secretory granules, electron density decreased and vacuofes appeared. After propagation, GTH cells evacuated large number of secretory granules, TSH cells appeared vacuoles, rough endoplasmic reticulum in STH cells were circumnuclear.In autumn,there were GTH1 cells and GTH2 cells in GTH cells. A large number of secretory granules in GTH1 cells were circumnuclear; In GTH; cells cytoplast,there were two types of secretory granules,small circle and large circle, in small number.Structure of STH cells had no obvious changes. In winter,nuclear of GTH cells were elliptical and a small number o
f secretory granules scattered around nucleolus. Number of secretory granules in STH cells were relatively small, nucleolus distorted, activity of synthesis and secretion decreased. Secretory granules in TSH cells cytoplast were relatively small. Before and after propagation,changes of ACTH PRL and MSH cells were inapparent.
During all stages, main changes of eurohypophysis were as follows: There were global and no envelopes secretory granules in ncurosccrclory fibres before propagation; Neurosecretory fibres axone evacuated secretory granules after propagation.; Secretory granules could be seen in some of neurosecretory fibres axone in autumn,but the number was relatively small. In winter,
neurohypophysis had no changes.
According to the morphotogical and histological features, the development of ovary could be devided into six stages and the changes of the oocytes of the female also could be devided into six phases. Stage of ovaries could be seen in one -year old fishes only, and the smallest maturity age was three years old The mature coefficent of ovaries was 0.72%~4.65%,the spawning period was from May to July. After spawning, the ovaries quickly degenerated to the stage II and then devetoped to the stage III. The results of this paper indicated that Long-snout catfish spawned only once in a year.
引文
[1] 陈松林.鱼类生长内分泌学和鱼类养殖[J] .水产学报,1992,16(1):91-110.
[2] 王德寿,林浩然.鲶类生殖内分泌学研究[J] .水产学报,1996,20(2):151-158.
[3] Bell, WR.Studies on the endocrines of the teloests. The morphology of the hypophysis of goldfisk[J] .Anat.Res, 1937,70(1): 122.
[4] Charipper, HA.The morphology of the hypophysis in lower vertebrates,particularly fish and amphibia,with some notes on the cytology of the pituitary of goldfish and Necturrus maculesus[J] .Cold.Spr.Harb.Symp. 1937,5:151-164.
[6] Sokol,HW.Selective staining of the pituitary gland of the teleost fishes dermonstrating at least six tinctorial celltypes in the adeno-hypophysis[J] .Anat.Res,1953,117.
[7] Van Oodert.Gonadotropic cells in the pituitary of teleost,In:Richer, CJJ, Goos HJTh.Editors, Proceeding of the intemational symposium on reproductive physiology of fish[M] . Netherlands:Pudoc, Wageningenn, 1982,44-48.
[8] 上海水产学院主编.组织胚胎学[M] .北京农业出版社,1989,163-164.
[9] Scruggs,WM,The epithelial components and their seasonal changes in the pituitary gland of the carp and gold fish[J] . Morph, 1951,88:441-470.
[10] Olivereau,M.Hypophyse gland thyroide chez les poisons.Etude histophysiologique de quelquescorrelationns endocriniennes en particuliar chez salmo salarl[J] .Anne Biol, 1954,30:63-80.
[11] Kaul, S and Vollrath, L, Cytology.Cell and Tissue Research, 1974 (154),211.
[12] Nagahama, Y and Yamamoto,K.Morphological studies the chum salmon, Oncorhynchus kkefa.I;Fine structure of the adenohypophysis[J] .Bull. Fac.Fish.Hokkaido University, 1970, 20: 293-302.
[13] Van Eys,GJJM.Cytological localization of MSH,ACTH and Endorphin in the pars intermedia the cichlid teleost,Sarotherodon mossambicus[J] .Cell and Tissue Research, 1981,218:625-633.
[14] Benjiamins, M.Morphologicai studies on the growth hormone cells in the pituitary gland of the nine spined stick-ieback, Pungititus pungitius L.Z[J] .mikrosk-anat forsh, 1982,96(3):461-470.
[15] Nagahama, Y.Histo-physiological studies on the pituitary glands of some teleost fishes, with special reference to the classification of hormone-producing cells in the adenohypophysis [J] . Mem.Fac. Fish Hokkaido University. 1973,21:1-63.
[16] Baker.The cell types in the adenohypophysis of Indian catfish,Heteropneustes fossilis Bloch[J] .Cell Tissue Res, 1974,148:347-358.
[17] Boddinguis,J.The cell types of the adenohypophysis in the rainbow trout.A histogical study[M] . Oxfoed:Holywell Press Ltd, 1975.
[18] 施泉芳,张水元.草、鲢鱼脑垂体变化的组织学研究[A] 水生生物学集刊,1964,5(1):63—72.
[19] 王良臣,阎家本,刘修业.黄鳝垂体嗜碱性细胞组织化学的研究[A] .鱼类学论文集第五辑[C] ,1986,29—34.
[20] 方永强.赤点石斑鱼脑垂体组织生理学的研究[J] .台湾海峡,1990,9(1):69—72.
[21] 鲁双庆,刘少军草鱼性成熟前后垂体组织学和超微结构[J] .湖南师范大学自然科学学报,1996,19(3):76—83.
[22] 翁幼竹,林君卓,洪万树等.鲻鱼脑垂体组织生理学的研究[J] .台湾海峡,2000,19(2):192—196.
[23] Abrahama, M.Ultrastructure of the cell types and of the neurosecretory innervation in the pituitary of Mugil Vcephalus L.from freshwater,the sea, and hypersaline lagoon, Ⅱ. The rostral pars djstaliso [J] .Gen CompEndocrinol, 1967,17:334—350.
[24] Abraha Ma,M.Uitrastructure of the cell types and of the neurosecretory innervatjon in the pituitary of Mugilcephalus L.from freshwater, the sea, and hypersaline lagoon Ⅰ.The proximalpars distalis[J] .Gen.Comp. Endocrinol,1974,24:121—132.
[25] Abrahama, M. Ultrastructure of the cell types and of the neurosecretory innervation in the pituitary of Mugilcephalus from freshwater, the sea and hypersaline lagoon, Ⅲ. The neurointermediatelobc [J] . Gen.Comp.Endocrinol, 1976,29:,511—521.
[26] Batten,T.F.C and JN.Ball.Ultrastructure of the neurohypophysis of the teleost Poecilia latipinna in relation to neural control of the adenohypophysial cells [J] . Cell Tissue Res, 1977,185:409—433.
[27] Batten,T F C and J.N. Ball.Ultrastructure of the adenohypophysis in the teleost,Poccilia latipinna [J] . Cell Tissue Res, 1975, 161:239—261.
[28] 中国科学院动物研究所内分泌室细胞组、湖北长江水产研究所生殖组.丘脑下部促黄体素释放激素(LRH-A)对草鱼的催产作用:垂体促性腺细胞亚显微结构研究[J] .中国科学,1977(6):600—603.
[29] 方展强,何艾文.尼罗罗非鱼神经垂体的超微结构研究[J] .动物学报,1990,36(4):352—357.
[30] 方展强,何艾文.罗非鱼脑垂体结构研究Ⅰ:腺垂体前叶的超微结构[J] .水生生物学报,1993,17(3):252—257.
[31] 王良臣,阎家本,刘修业.梭鱼脑垂体超微结构[J] .水生生物学报,1985,9(2):159—163.
[32] 林浩然,张梅丽,张素梅等.鳗鲡繁殖生物学研究Ⅳ:人工催产过程中下鳗鲡的GTH分泌活动、性腺发育状况和脑垂体GTH细胞的超微结构[J] 水生生物学报,1987,11(4):320—329.
[33] Swanson P.et al.Seasonal effects of primozide and desyGly10(D Ala6)LHR-H ethylamine secretion in goldfish.Gen.Comp.Endocrinol. 1991,44:29-38.
[34] Lin, YWP.Fundulus Heteroclitus gonadotropins.3.Cloning and sequening of gonadotropin hormone(GTH) Ⅰ and Ⅱ β-subunits using the polymerase chain reaction.Molecular and Cellar Endocrinology. 1992, 85:127-135.
[35] Van Der kraak,et al.Preperties of common carp gonadotropin Ⅰ and gonadotropin Ⅱ.Gen.Comp. Endocrinol. 1992,85:217-229.
[36] Koiode, Y, itoh, H and Kawauchih.Isolation and characterization of two distinct gonadotropins, GTH Ⅰ and GTH Ⅱ,from bonito(Katsuwonus plelamis)pituitary glands.Int.J.Peptide Protein Res. 1993,41:52-65.
[37] Ider, D R.L.S.Bazar and S.J.Hwang.Fish gonadotropins Ⅲ.Evidence for more than one gonadotropin in chum salmon pituitary glands.Endocrine Res.Common. 1975,2:237-249.
[38] Kawwauchi,H.et al.The duality of teleost gonadotropins[J] .Fish and Biochemistry. 1989,7:29-38.
[39] 林浩然.鱼类促性腺激素(GTH)的生理学和分子生物学[J] .鱼类学论文集(第六辑).科学出版社,1997.
[40] Goos,HJTh,R.De Leeuw, E.Burzawa-Gerad, M.Terlou and J.J.Richter. Purification of gonadotropin hormone from the pituitary of the Africation catfish,Clarias gariepinus
and the development of a homologous radioimmunoassay.Gen Comp Endocrinol. 1986,49; 404-413.
[41] Rebers.FEM.et al. Modulation of glycoprotein hormone a-and gonadotropin Ⅱ β-subunit mRNA levers in the pituitary of mature male African catfish,Clarias gariepinus[J] .Fish Physiol.Biochem. 1997,17:99-108.
[42] Bogerd J and H.J.Th.Goos.lsolation sequence analysis and expression of mRNAs encoding GTH-subunits of the African catfish.Program summary of Ⅻ international congress of comparative endocrinology.Toronto,Ontario, Canada [J] . Endocrinol. 1992,85:217-229.
[43] Kawwauchi,H.et al.The duality of teleost gonadotropins[J] .Fish and Biochemistry. 1989,7:29-38.
[44] Peute,J.Immunocytochemical evidence for peptidergic(GnRH) and dopaminergic innervation of the gonadotropic cells in the pituitary of the African catfish, Clarias gariepinus[J] . Gen.Comp Endocrinol, 1987,67:303-310.
[45] Mettews, SA.The seasonal cycle in the gonads of Fundulus[J] .Bial Bull, 1939, 75.
[46] 施泉芳.鲢鱼性腺周年变化的研究[J] .水生生物学集刊,1964,5(2):77-94.
[47] Yoakim E G. Corpora atretica in the Nile catfish.Schilbe Mystus, a histology and histochemical study.Bull.Fac[J] .Sci.Assiiut.Uni. 1975,4 (1):85—92.
[48] 刘筠.草鱼性腺发育的研究[J] .湖南师范学院学报(自然科学版),1962(1):1-23.
[49] 林光华,性成熟草鱼卵巢发育的年周期变化[J] .水生生物学报,1985,9(2):186-195.
[50] Davis T L O.Reproductive biology of the freshwater catfish, Tandanus tandanus Mitchell, in the Gwydir River,Australia, I structure of the gonads.Aust[J] .Mar.Freshwater Res. 1977,28:139-158.
[51] Ntiba M S. et al. Gonad maturation and spawning times of Siganus sutor off the Kenya coast: evidence for definite spawning seasons in a tropical fish[J] .Journal of Fish Biology. 1990,37:315-325.
[52] 张耀光,谢小军.南方鲶的繁殖生物学研究:性腺发育及周年变化[J] .水生生物学报,1996,20(1):8—16.
[53] 胡成钰,洪一江,林光华等.繁殖季节黄颡鱼性腺的研究[J] .江西科学,1993,11(3):159-163.
[54] 李建中,张轩杰,刘少军.异源四倍体鲫鲤的性腺发育研究[J] .水生生物学报,2002,26(2):116-122.
[55] 刘筠.中国养殖鱼类繁殖生理学[M] .北京农业出版社,1993,22-40.
[56] UPADNYAYA N.Ovarian cycle of a freshwater catfish Mystus vittatus Bloch.with a special mate on the steroidogenic sites Z.Mikrosk[J] .Anat Forsch Leipzig.- 1985,99(5):845-854.
[57] 张耀光,罗王泉笙,何学福.长吻鮠卵巢发育和周年变化及繁殖习性研究[J] .动物学研究,1994,15(2):42-48.
[58] 林浩然.鱼类生理学[M] .广东高等教育出版社,1999:146-202.
[59] Chang, JP, A.F.Cook and R.E.Peter.Influence of catcheelamines on gonadotropin secretion in goldfish,carasius auraius[J] .Gen.Comp.Endocrinol. 1983,49:22-31.
[60] Peter, RE.Changs, J.P, et al.Interaction of catecholamines and GnRH in regulation of gonadotropin secretion in teleost fish[J|.Recent Prog. Horm.Res. 1986,42:513-548.
[61] 王德寿,林浩然,HJ. Th. Goos.大鳍鳠促性腺激素分泌调控的研究[J] .动物学报 1998,44(3):322-328.
[62] De Leeuw, R.H.J.Th.Goos and P.G.WJ.VanOordt.The dopaminergic inhibition of the gonadotropin-releasing hormone from pituitaries of the African catfish,Clarias gariepinus[J] .Gen.Comp.Endocrinoi. 1986,63:171-177.
[63] 吴嘉敏,姜仁良,张继平.长吻鮠人工催产及脑垂体促性腺激素细胞的超微结构的变化[J] .水产学报,1999(3):248-253.
[64] 邓晓川.长吻鮠养殖技术[J] .内陆水产,2000,26(5):29-30.
[65] 林浩然.激素和人工诱导鱼类繁殖[J] .生物学通报,1999,34(8):1-3.
[66] Andrey L.Polenov etal.The hypothalamo-hypophysial system of the wild Carp,Cyprinus carpio Structure and ultrastructure of the posterior neurohypophysis. Cells Tissue Res. 1984,237:139-147.
[67] 方展强.尼罗罗非鱼在不同滲透压和温度环境中脑垂体及性腺发育的显微及超微结构观察.华南师范大学硕士研究生毕业论文,1988.
[68] 方昆阳.鲶人工繁殖机制的研究.华南师范大学硕士研究生毕业论文,1998.
[69] 郑文彪,陈民.冬季革胡子鲶中腺垂体细胞超微结构的研究[J] .动物学研究,1994,15(4):31-35.
[70] Belenky MA, Morphofunvtiomal evolution of the neurohemal regions of the hypo-
thalamo-hypophyseal system.In:Bargmann WmOksvhe A, Polenov A, Springer, Berlin rleidelberg New York, 1978,44-51.
[71] Cavaco,JEB.Androgen-induced changes in Leydig cells ultrastructure and stero-Ⅰ dogenesis in juvenile African catfish, Clarias gariepinus. University Utrecht.PhD.Dissertation. 1998.
[72] Olivier KRE,Peter PD and Harry C.Eeffects of Monosodium L-Glutamate on Pituitary in nervation Goldfish. Gen.Comp.Endocrinol, 1983 (51):338-346.
[73] Belenky MA.Functional cytomorphology of the neurohypophysis and foundations of its development in phylogenesiis and ontogenesis of vertebrates.Dr Sc Thesis, Leningrad. 1981.
[74] Evertt,JW.The mammalin hypothalamo-hypophysial In The Endocrine Hypothalamus(S.L.Jeffcoateand J.S.M.Hutchinson,eds) [J] .Aca-demic Press, New-York,1978,1-33.
[75] Joseph,HA and Knigge KM.The endocrine hypothalamus, recent anatomical studies.In The Hypothalamus, Raven, Neq York. 1978,15-47.
[76] 杨惠彬,王之贤.成年大鼠神经垂体垂体细胞超微结构观察[J] .解剖学报,1989,20(3):319-322.
[77] Jackson K A, Goldberg D,Ofir M,et al.Cloning, sequencing of GTH-Ⅰ, GTH-Ⅱ and GH cDNAs of the blue gouraami and their expression during the differrent stage of oogenesis[A] .Proceedings of the 6th International Symposium on the Reproductive Physiology of Fish[C] .Norway:Bergen, 1999,489.
[78] Schulz R W, Bogerd J, Bosma P T, et al.Physiological, morphological and molecular aspects of gonadotropins in fish with special reference to the African catfish, Clarias gariepinus[A] .Proceedings of the 5th International Symposium on the Reproductive Physiology of Fish[C] .Texas:Austin, 1995.2-6.
[79] Olivereau M,Nagahama Y.Immunocytochemistry of gonadotropic celiss in the pituitary of some teleost species[J] .Gen Comp Endocrinol, 1983,50:252-260.
[80] Peute, de Leeuw R,Goos H J Th,et al.Ultrastructure and immunolabeling of gonadotrops and thyrotropss in the pituitary of the African catfish,Clarias lazera[J] .Cells Tissue Res, 1984,238:95-103.
[81] Van Oodert.The cellsular origin of pituitary in teleosts.In:W S Hoar et al editors, Fish physiolog[M] .New York:Academic Press, 1983,246-256.
[82] 方永强,汪敏.丘脑下部促黄体释放激素类似物(促黄体素释放激素类似物)的作用能够机制Ⅲ.对罗非鱼脑垂体促性腺激素分泌细胞结构的影响[J] .动物学报,1983,29(2):124-130.
[83] 王文,王韫明,杜开河.团头鲂脑垂体的显微和超微结构以及其GTH在人工催产过程中的变化[J] .电子显微镜学报,1995,(3):170-176.
[84] 楼允东.组织胚胎学[M] .北京农业出版社 1981,173.
[85] 杨若宾,李禾,彭绍君.长吻鮠卵巢发育的组织学观察[J] .四川动物,1990,9(2):19-20.
[86] 汪小东,林浩然.硬骨鱼类卵母细胞最后成熟的调控[J] .水产学报,1998,22(1):72-77.
[87] 林浩然.关于硬骨鱼类生殖内分泌学的研究[J] .水生生物学集刊,1981,7,(3).
[88] 赵维信.诱导鲢鱼排卵的性类固醇含量的变化[J] .水生生物学报,1988,12:312—318.
[89] 林浩然.鱼类生殖生理学[M] (讲义).
[90] Billard, R,Fostier A, Jalabert B, Weil.C.Endocrine control the teleost reproductive cycle and its relation to external factors:salmonid and cyprinid models. In: Comparative Endocrinology. Elsevier North-Holland, Amsterdam. 1978,37-48.
[91] Jalabast B and Berton B.Evolution de la gonadotropin plasmatique GTH après tovulation chez la Truite are-en-ciei et influence de la retentation des ovules.C.R. Hebd.Seances Acad,Sei, Ser, D. 1980,290,799-801.
[92] 赵维信,姜仁良,黄世蕉等.鲤鱼脑垂体中促性腺激素含量的周年变化[J] .水产学报.1983,7(1):69-75.
[93] 王德寿,林浩然,H.J.Th.Goos.大鳍鳠脑垂体和血清促性腺激素水平的季节变化[J] .鱼类学论文集(第6辑),1997,22-27.
[94] Sokolowska, M. et al.Seasonal effects of pimozide and desyGly10 LRH-A ethylamine secretion in goidfish.Gen.Comp.Endocrinol[J] . 1985,57,472-479.
[95] 王黎,林浩然,林信伟.多巴胺能药物对鲤鱼促性腺激素分泌活动的影响[J] .动物学报,1997,43(1):74-79.
[96] 周仰璟.大鳍鳠的生物学资料[J] .动物学杂志,1983,(2):39-42.