雄性峨眉髭蟾“胡子”的组织结构特征
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摘要
采用石蜡切片与苏木精-伊红染色及扫描电镜,对雄性峨眉髭蟾Leptobrachium boringii的角质刺及其周边皮肤进行了显微结构和亚显微结构的观察。显微结构观察发现,峨眉髭蟾的角质刺属于皮肤衍生物,突起呈倒"V"形。角质刺由表皮和真皮构成,表皮为复层扁平上皮,可分成4层;最外层细胞角质化,细胞轮廓不清,被染成深红色。真皮由疏松结缔组织构成,分辨不出致密层与疏松层,其内未见皮肤腺,但有少量色素细胞与毛细血管分布。表皮嵴伸入到真皮层,在以往的无尾两栖类研究中未见报道。角质刺基部可见皮肤褶翻起将其包裹在内,皮肤褶向上延伸形成角质刺。扫描电镜观察表明,角质刺顶端呈锥形的"小山丘"状,表面可分辨出表皮细胞轮廓,细胞为呈覆瓦状排列的角质化细胞。角质刺与皮肤交界处为多边形的角质化细胞。角质化上皮细胞的上表面与下表面均具有凹凸不平的花纹结构,细胞之间以镶嵌的方式连接。
The microstructure and sub-microstructure of the keratinized nuptial spines and their relative peripheral skin of male Leptobrachium boringii were studied by general histological and scanning electron microscope(SEM) methods. The results showed that the spine of L. boringii is the skin derivative which rises up as an inverted V-like conical protuberance from the periheral skin area. Spine is composed of the external part of keratinized epidermis and inner part of dermis. The spine epidermis is stratified squamous epithelium and can be divided into 4 layers, of which the outermost layer is keratinized without cell outline and stained deeply red in color. The fold of skin encircles around the spine base and extends upward to form the spine. The spine epidermal ridge extends into the dermis under the epithermis. To the best of the authors' knowledge, this structure has never been reported in other studied frogs and toads. Furthermore, the underlying loose connective tissue of the epidermis makes up the spine dermis, where no stratum compactum and stratum spongiosum can be distinguished. In spine dermis, no skin glands were found but only a small number of pigment cells and capillary vessels. Under SEM observation, the tip of the spines looks like a conical "hill", and the outline of superficial epidermal cell is distinguishable in the surface. The cells arrange in imbricate pattern. At the junction of spine and its peripheric skin, the superficial epithermal cells are polygonous and keritinized. Both the upper and lower surfaces of keratinized epithelial cells have an uneven pattern. The cells are connected to each other with anchoring junctions.
The microstructure and sub-microstructure of the keratinized nuptial spines and their relative peripheral skin of male Leptobrachium boringii were studied by general histological and scanning electron microscope(SEM) methods. The results showed that the spine of L. boringii is the skin derivative which rises up as an inverted V-like conical protuberance from the periheral skin area. Spine is composed of the external part of keratinized epidermis and inner part of dermis. The spine epidermis is stratified squamous epithelium and can be divided into 4 layers, of which the outermost layer is keratinized without cell outline and stained deeply red in color. The fold of skin encircles around the spine base and extends upward to form the spine. The spine epidermal ridge extends into the dermis under the epithermis. To the best of the authors' knowledge, this structure has never been reported in other studied frogs and toads. Furthermore, the underlying loose connective tissue of the epidermis makes up the spine dermis, where no stratum compactum and stratum spongiosum can be distinguished. In spine dermis, no skin glands were found but only a small number of pigment cells and capillary vessels. Under SEM observation, the tip of the spines looks like a conical "hill", and the outline of superficial epidermal cell is distinguishable in the surface. The cells arrange in imbricate pattern. At the junction of spine and its peripheric skin, the superficial epithermal cells are polygonous and keritinized. Both the upper and lower surfaces of keratinized epithelial cells have an uneven pattern. The cells are connected to each other with anchoring junctions.
引文
曹燕, 谢锋, 江建平. 2011. 齿突蟾属四个物种皮肤的组织学观察[J]. 四川动物, 30(2): 214-219.
樊均德, 许勤智, 沈正雄, 等. 2011. 峨眉髭蟾研究与保护现状[J]. 吉林农业, (7): 246.
费梁, 叶昌媛, 江建平, 等. 1995. 两栖爬行动物学研究(第4、5辑): 髭蟾属的分类及其系统发育的研究(两栖纲:锄足蟾科)[M].贵阳: 贵州科技出版社: 65-71.
费梁, 叶昌媛, 江建平. 2012. 中国两栖动物及其分布彩色图鉴[M]. 成都: 四川科学技术出版社: 457-472.
高凤娟, 李丕鹏, 王译唱, 等. 2016. 西藏蟾蜍皮肤的组织学观察[J]. 四川动物, 35(3): 444-451.
贾丽丽, 田川, 李淑兰. 2013. 繁殖期东北林蛙两性皮肤组织结构的差异性比较[J]. 中国农学通报, 29(2): 23-26.
雷飞宙, 江建平, 李成, 等. 2012. 角蟾亚科3物种皮肤的组织学观察[J]. 动物学杂志, 47(3): 20-27.
李炳蔚, 赵红平, 冯西桥. 2010. 牛角的结构与力学性能研究[C]// 北京力学会. 北京力学会第十六届学术年会论文集.
梁刚, 王琼霞. 2004. 隆肛蛙皮肤及其腺体的显微结构特征[J]. 动物学杂志, 39(4): 73-76.
林文达, 朱静, 郭爱伟, 等. 2008. 云南昆明地区三种蛙皮肤显微结构的比较[J]. 四川动物, 27(3): 420-421.
刘承钊, 胡淑琴, 赵尔宓. 1980. 髭蟾属Vibrissaphora和种的初步探讨及其与分类学有关问题的讨论[J]. 两栖爬行动物学报, 3(1): 1-9.
刘满樱, 肖向红, 徐佳佳, 等. 2007. 东北林蛙皮肤及其腺体组织形态学观察[J]. 野生动物, 28(4): 6-9.
马云海, 佟金, 周江, 等. 2008. 穿山甲鳞片表面的几何形态特征及其性能[J]. 电子显微学报, 27(4): 336-340.
邵华信, 吕学洗. 1991. 不同胎龄手掌侧皮肤皮嵴的分化特征[J]. 解剖学杂志, 14(1): 78-80.
佘平昌. 2016. 牦牛角和毛囊的形态发生及相关基因的表达分析[D]. 北京: 中国农业科学院.
唐伟文. 2015. 穿山甲鳞片宏-微观力学性能表征及其耐冲击性仿真[D]. 成都: 西南交通大学.
王大江, 金维哲, 吕学先, 等. 1991. 手指表皮嵴形成的基因作用机制[J]. 佳木斯医学院学报, 14(2): 94-95.
王溯, 陈晓蓉. 2014. 成年裸鼠和成人体表皮肤的比较组织形态学研究[J]. 安徽医科大学学报, 49(8): 1157-1160.
吴文英, 李丕鹏, 陆宇燕, 等. 2011. 黑龙江林蛙繁殖期两性皮肤组织结构的差异比较[J]. 野生动物, 32(3): 141-145.
徐平, 何勤, 龙娟, 等 . 2012. 泛发性连续性肢端皮炎一例及文献复习[J]. 贵州医药, 36(3): 267-268.
杨纯. 2004. 秦岭山区中国林蛙精巢的组织学和免疫细胞化学研究[D]. 西安: 陕西师范大学.
俞宝根. 2010. 棘胸蛙不同地区的两性异形及人工环境下繁殖行为研究[D]. 杭州: 浙江师范大学.
邹猛, 于用军, 张荣荣, 等. 2015. 仿牛角结构薄壁管吸能特性仿真分析[J]. 吉林大学学报(工学版), 45(6): 1863-1868.
Blaustein AR, Belden LK. 2003. Amphibian defenses against ultraviolet-B radiation[J]. Evolution & Development, 5(1): 89-97.
Brodie ED, Nussbaum RA, Yang DT. 1990. Antipredator behavior of Chinese salamanders (Salamandridae)[J]. Zoological Research, 11(1): 7-16.
Chen WW, Zhang W, Zhou SC, et al. 2013. Insight into the validity of Leptobrachium guangxiense (Anura: Megophryidae): evidence from mitochondrial DNA sequences and morphological characters[J]. Zootaxa, 3641(1): 31-40.
Dockray GJ, Hopkins CR. 1975. Caerulein secretion by dermal glands in Xenopus laevis[J]. Journal of Cell Biology, 64(3): 724-733.
Dubois A, Ohler A. 1998. A new species of Leptobrachium (Vibrissaphora) from northern Vietnam, with a review of the taxonomy of the genus Leptobrachium (Pelobatidae, Megophyinae)[J]. Dumerilia, 4(1): 1-32.
Dubois A. 1980. Notes sur la systematique et la repartition des amphibiens anoures de Chine et des regions avoisinantes Ⅳ. Classification generique et subgenerique des Pelobatidae Megophryinae[J]. Bulletin Mensuel de la Société Linnéenne de Lyon, 49: 469-482.
Duellman WE, Trueb L. 1994. Biology of amphibians[M]. Baltimore and London: The Johns Hopkins University Press.
Fei L, Wu GF. 2004. Leptobrachium boringii[DB/OL]. [2018-08-20]. The IUCN red list of threatened species 2004: e.T57625A11665713.
Fei L, Ye CY. 2016. Amphibians of China (Ⅰ)[M]. Beijing: Science Press: 472-474.
Han MJ, An JY, Kim WS. 2010. Expression patterns of Fgf-8 during development and limb regeneration of the axolotl[J]. Developmental Dynamics, 220(1): 40-48.
Hieronymus TL,Witmer LM, Ridgely RC. 2006. Structure of white rhinoceros (Ceratotherium simum) horn investigated by X-ray computed tomography and histology with implications for growth and external form[J]. Journal of Morphology, 267: 1172-1176.
Hofer R, Mokri C. 2000. Photoprotection in tadpoles of the common frog, Rana temporaria[J]. Journal of Photochemistry & Photobiology B: Biology, 59(1-3): 48-53.
Hudson CM, Fu JZ. 2013. Male biased sexual size dimorphism, resource defense polygyny, and multiple paternity in the Emei moustache toad (Leptobrachium boringii)[J]. PLoS ONE, 8(6): e67502. DOI: 10.1371/journal.pone.0067502.
Hudson CM, He XJ, Fu JZ. 2010. Keratinized nuptial spines are used for male combat in the Emei moustache toad (Leptobrachium boringii)[J]. Asian Herpetological Research, 2(1): 142-148.
Julian F, Haddad CFB, Baêta D, et al. 2010. The phylogenetic relationships of the charismatic poster frogs, Phyllomedusinae (Anura, Hylidae)[J]. Cladistics, 26(3): 227-261.
Liu CC. 1945. New frogs from west China[J]. Journal of the West China Border Research Society, Series B, 15: 28-44.
Luna MC, Roy W, Mcdiarmid RW, et al. 2018. From erotic excrescences to pheromone shots: structure and diversity of nuptial pads in anurans[J]. Biological Journal of the Linnean Society, 124(3): 403-446.
Luna MC, Taboada C, Baeta D, et al. 2012. Structural diversity of nuptial pads in Phyllomedusinae (Amphibia: Anura: Hylidae)[J]. Journal of Morphology, 273(4): 712-724.
Matsui M, Hamidy A, Murphy RW, et al. 2010. Phylogenetic relationships of megophryid frogs of the genus Leptobrachium (Amphibia, Anura) as revealed by mtDNA gene sequences[J]. Molecular Phylogenetics and Evolution, 56: 259-272.
Penrose LS, Ohara PT. 1973. The development of the epidermal ridges[J]. Journal of Medical Genetics, 10: 201-208.
Pienaar DJ, Hall-Martin AJ, Hitchins PM. 1991. Horn growth rates of free-ranging white and black rhinoceros[J]. KOEDOE: African Protected Area Conservation and Science, 34(2): 97-105.
Rachlow JL, Berger J. 1997. Conservation implications of patterns of horn regeneration in dehorned white rhinos[J]. Conservation Biology, 11: 84-91.
Rao DQ, Wilkinson JA. 2008. Phylogenetic relationships of the mustache toads inferred from mtDNA sequences[J]. Molecular Phylogenetics and Evolution, 46: 61-73.
Seki T, Kikuyama S, Yanaihara N. 1989. Development of Xenopus laevis skin glands producting 5-hydroxytryptamine and caerulein[J]. Cell Tissue Research, 258: 483-489.
Seki T, Kikuyama S, Yanaihara N. 1995. Morphology of the skin glands of the crab-eating frog (Rana cancrivora)[J]. Zoological Society of Japan, 12: 623-626.
Shine R. 1979. Sexual selection and sexual dimorphism in the Amphibia[J].Copeia, 2: 297-306.
Wells KD. 2007. The ecology and behaviour of amphibians[M]. Chicago: University of Chicago Press.
Young HE. 1977. Epidermal ridge gormation furing limb regeneration in the adult salamander, Ambystoma annulatum[J]. Journal of the Arkansas Academy of Science, 31: 107-109.
Zheng YC, Deng D, Li SQ, et al. 2010. Aspects of the breeding biology of the Omei mustache toad (Leptobrachium boringii): polygamy and paternal care[J]. Amphibia-Reptilia, 31: 183-194.
Zheng YC, Li SQ, Fu JZ. 2008. A phylogenetic analysis of the frog genera Vibrissaphora and Leptobrachium, and the correlated evolution of nuptial spine and reversed sexual size dimorphism[J]. Molecular Phylogenetics and Evolution, 46: 695-707.
樊均德, 许勤智, 沈正雄, 等. 2011. 峨眉髭蟾研究与保护现状[J]. 吉林农业, (7): 246.
费梁, 叶昌媛, 江建平, 等. 1995. 两栖爬行动物学研究(第4、5辑): 髭蟾属的分类及其系统发育的研究(两栖纲:锄足蟾科)[M].贵阳: 贵州科技出版社: 65-71.
费梁, 叶昌媛, 江建平. 2012. 中国两栖动物及其分布彩色图鉴[M]. 成都: 四川科学技术出版社: 457-472.
高凤娟, 李丕鹏, 王译唱, 等. 2016. 西藏蟾蜍皮肤的组织学观察[J]. 四川动物, 35(3): 444-451.
贾丽丽, 田川, 李淑兰. 2013. 繁殖期东北林蛙两性皮肤组织结构的差异性比较[J]. 中国农学通报, 29(2): 23-26.
雷飞宙, 江建平, 李成, 等. 2012. 角蟾亚科3物种皮肤的组织学观察[J]. 动物学杂志, 47(3): 20-27.
李炳蔚, 赵红平, 冯西桥. 2010. 牛角的结构与力学性能研究[C]// 北京力学会. 北京力学会第十六届学术年会论文集.
梁刚, 王琼霞. 2004. 隆肛蛙皮肤及其腺体的显微结构特征[J]. 动物学杂志, 39(4): 73-76.
林文达, 朱静, 郭爱伟, 等. 2008. 云南昆明地区三种蛙皮肤显微结构的比较[J]. 四川动物, 27(3): 420-421.
刘承钊, 胡淑琴, 赵尔宓. 1980. 髭蟾属Vibrissaphora和种的初步探讨及其与分类学有关问题的讨论[J]. 两栖爬行动物学报, 3(1): 1-9.
刘满樱, 肖向红, 徐佳佳, 等. 2007. 东北林蛙皮肤及其腺体组织形态学观察[J]. 野生动物, 28(4): 6-9.
马云海, 佟金, 周江, 等. 2008. 穿山甲鳞片表面的几何形态特征及其性能[J]. 电子显微学报, 27(4): 336-340.
邵华信, 吕学洗. 1991. 不同胎龄手掌侧皮肤皮嵴的分化特征[J]. 解剖学杂志, 14(1): 78-80.
佘平昌. 2016. 牦牛角和毛囊的形态发生及相关基因的表达分析[D]. 北京: 中国农业科学院.
唐伟文. 2015. 穿山甲鳞片宏-微观力学性能表征及其耐冲击性仿真[D]. 成都: 西南交通大学.
王大江, 金维哲, 吕学先, 等. 1991. 手指表皮嵴形成的基因作用机制[J]. 佳木斯医学院学报, 14(2): 94-95.
王溯, 陈晓蓉. 2014. 成年裸鼠和成人体表皮肤的比较组织形态学研究[J]. 安徽医科大学学报, 49(8): 1157-1160.
吴文英, 李丕鹏, 陆宇燕, 等. 2011. 黑龙江林蛙繁殖期两性皮肤组织结构的差异比较[J]. 野生动物, 32(3): 141-145.
徐平, 何勤, 龙娟, 等 . 2012. 泛发性连续性肢端皮炎一例及文献复习[J]. 贵州医药, 36(3): 267-268.
杨纯. 2004. 秦岭山区中国林蛙精巢的组织学和免疫细胞化学研究[D]. 西安: 陕西师范大学.
俞宝根. 2010. 棘胸蛙不同地区的两性异形及人工环境下繁殖行为研究[D]. 杭州: 浙江师范大学.
邹猛, 于用军, 张荣荣, 等. 2015. 仿牛角结构薄壁管吸能特性仿真分析[J]. 吉林大学学报(工学版), 45(6): 1863-1868.
Blaustein AR, Belden LK. 2003. Amphibian defenses against ultraviolet-B radiation[J]. Evolution & Development, 5(1): 89-97.
Brodie ED, Nussbaum RA, Yang DT. 1990. Antipredator behavior of Chinese salamanders (Salamandridae)[J]. Zoological Research, 11(1): 7-16.
Chen WW, Zhang W, Zhou SC, et al. 2013. Insight into the validity of Leptobrachium guangxiense (Anura: Megophryidae): evidence from mitochondrial DNA sequences and morphological characters[J]. Zootaxa, 3641(1): 31-40.
Dockray GJ, Hopkins CR. 1975. Caerulein secretion by dermal glands in Xenopus laevis[J]. Journal of Cell Biology, 64(3): 724-733.
Dubois A, Ohler A. 1998. A new species of Leptobrachium (Vibrissaphora) from northern Vietnam, with a review of the taxonomy of the genus Leptobrachium (Pelobatidae, Megophyinae)[J]. Dumerilia, 4(1): 1-32.
Dubois A. 1980. Notes sur la systematique et la repartition des amphibiens anoures de Chine et des regions avoisinantes Ⅳ. Classification generique et subgenerique des Pelobatidae Megophryinae[J]. Bulletin Mensuel de la Société Linnéenne de Lyon, 49: 469-482.
Duellman WE, Trueb L. 1994. Biology of amphibians[M]. Baltimore and London: The Johns Hopkins University Press.
Fei L, Wu GF. 2004. Leptobrachium boringii[DB/OL]. [2018-08-20]. The IUCN red list of threatened species 2004: e.T57625A11665713.
Fei L, Ye CY. 2016. Amphibians of China (Ⅰ)[M]. Beijing: Science Press: 472-474.
Han MJ, An JY, Kim WS. 2010. Expression patterns of Fgf-8 during development and limb regeneration of the axolotl[J]. Developmental Dynamics, 220(1): 40-48.
Hieronymus TL,Witmer LM, Ridgely RC. 2006. Structure of white rhinoceros (Ceratotherium simum) horn investigated by X-ray computed tomography and histology with implications for growth and external form[J]. Journal of Morphology, 267: 1172-1176.
Hofer R, Mokri C. 2000. Photoprotection in tadpoles of the common frog, Rana temporaria[J]. Journal of Photochemistry & Photobiology B: Biology, 59(1-3): 48-53.
Hudson CM, Fu JZ. 2013. Male biased sexual size dimorphism, resource defense polygyny, and multiple paternity in the Emei moustache toad (Leptobrachium boringii)[J]. PLoS ONE, 8(6): e67502. DOI: 10.1371/journal.pone.0067502.
Hudson CM, He XJ, Fu JZ. 2010. Keratinized nuptial spines are used for male combat in the Emei moustache toad (Leptobrachium boringii)[J]. Asian Herpetological Research, 2(1): 142-148.
Julian F, Haddad CFB, Baêta D, et al. 2010. The phylogenetic relationships of the charismatic poster frogs, Phyllomedusinae (Anura, Hylidae)[J]. Cladistics, 26(3): 227-261.
Liu CC. 1945. New frogs from west China[J]. Journal of the West China Border Research Society, Series B, 15: 28-44.
Luna MC, Roy W, Mcdiarmid RW, et al. 2018. From erotic excrescences to pheromone shots: structure and diversity of nuptial pads in anurans[J]. Biological Journal of the Linnean Society, 124(3): 403-446.
Luna MC, Taboada C, Baeta D, et al. 2012. Structural diversity of nuptial pads in Phyllomedusinae (Amphibia: Anura: Hylidae)[J]. Journal of Morphology, 273(4): 712-724.
Matsui M, Hamidy A, Murphy RW, et al. 2010. Phylogenetic relationships of megophryid frogs of the genus Leptobrachium (Amphibia, Anura) as revealed by mtDNA gene sequences[J]. Molecular Phylogenetics and Evolution, 56: 259-272.
Penrose LS, Ohara PT. 1973. The development of the epidermal ridges[J]. Journal of Medical Genetics, 10: 201-208.
Pienaar DJ, Hall-Martin AJ, Hitchins PM. 1991. Horn growth rates of free-ranging white and black rhinoceros[J]. KOEDOE: African Protected Area Conservation and Science, 34(2): 97-105.
Rachlow JL, Berger J. 1997. Conservation implications of patterns of horn regeneration in dehorned white rhinos[J]. Conservation Biology, 11: 84-91.
Rao DQ, Wilkinson JA. 2008. Phylogenetic relationships of the mustache toads inferred from mtDNA sequences[J]. Molecular Phylogenetics and Evolution, 46: 61-73.
Seki T, Kikuyama S, Yanaihara N. 1989. Development of Xenopus laevis skin glands producting 5-hydroxytryptamine and caerulein[J]. Cell Tissue Research, 258: 483-489.
Seki T, Kikuyama S, Yanaihara N. 1995. Morphology of the skin glands of the crab-eating frog (Rana cancrivora)[J]. Zoological Society of Japan, 12: 623-626.
Shine R. 1979. Sexual selection and sexual dimorphism in the Amphibia[J].Copeia, 2: 297-306.
Wells KD. 2007. The ecology and behaviour of amphibians[M]. Chicago: University of Chicago Press.
Young HE. 1977. Epidermal ridge gormation furing limb regeneration in the adult salamander, Ambystoma annulatum[J]. Journal of the Arkansas Academy of Science, 31: 107-109.
Zheng YC, Deng D, Li SQ, et al. 2010. Aspects of the breeding biology of the Omei mustache toad (Leptobrachium boringii): polygamy and paternal care[J]. Amphibia-Reptilia, 31: 183-194.
Zheng YC, Li SQ, Fu JZ. 2008. A phylogenetic analysis of the frog genera Vibrissaphora and Leptobrachium, and the correlated evolution of nuptial spine and reversed sexual size dimorphism[J]. Molecular Phylogenetics and Evolution, 46: 695-707.