极危孑遗物种东方水韭雌配子体及胚胎的解剖学观察
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摘要
以人工培养的国家一级保护植物东方水韭(Isoetes orientalis)为材料,采用切片技术对雌配子体和胚胎的发育进程进行解剖学观察研究,探讨其有性生殖过程及濒危机制。结果表明:(1)东方水韭大孢子3~5d萌发,成熟雌配子体呈球形,无假根,三裂缝处发育出多个颈卵器,成熟颈卵器只有颈壁细胞与颈沟细胞,无腹沟细胞。(2)多数雌配子体只发育出一个胚胎,偶见多胚共存现象;胚胎发育时期,第一叶原基相比第二、三叶原基发育迅速。(3)颈卵器部分组织常出现分化紊乱,导致雌配子体败育。该研究结果支持"根叶理论",并讨论了腹沟细胞的退化以及双胚共存机制,认为东方水韭雌配子体常停留在游离核阶段、颈卵器形态或位置不规则、卵细胞排列紊乱等可能是其败育的原因。
In this paper,the base of nation level protected species Isoetes orientalis is used as materials.With the method of slice technology,we anatomically observed the development process of female gametophytes and embryos,and discussed the reproduction and endangered mechanism.The results showed that:(1)Megaspore of I.orientalis would be germinated within 3-5d,the mature gametophytes were emerging spherical and forming a number of archegonias with triplets but no rhizoids.There were neck cells and neck canal cells in the archegonium,but no ventral canal cells.(2)Most of the female gametophytes were only developed into one embryo,occasionally into more embryos.During the development of embryo,the first primordium growth rapidly than the second and third primordiums.(3)The part tissue of archegonium often appear differentiation disorder and lead to female gametophyte abortion.The results supported the Phyllorhize Theory,also discussed the ventral canal cells degenerated and two embryos mechanism.We speculated that the gametophyte in I.orientalis usually stopped during free nuclei stage,morphology and anomaly site of archegonia and eggs arranged in disorder might be the causes of abortion.
In this paper,the base of nation level protected species Isoetes orientalis is used as materials.With the method of slice technology,we anatomically observed the development process of female gametophytes and embryos,and discussed the reproduction and endangered mechanism.The results showed that:(1)Megaspore of I.orientalis would be germinated within 3-5d,the mature gametophytes were emerging spherical and forming a number of archegonias with triplets but no rhizoids.There were neck cells and neck canal cells in the archegonium,but no ventral canal cells.(2)Most of the female gametophytes were only developed into one embryo,occasionally into more embryos.During the development of embryo,the first primordium growth rapidly than the second and third primordiums.(3)The part tissue of archegonium often appear differentiation disorder and lead to female gametophyte abortion.The results supported the Phyllorhize Theory,also discussed the ventral canal cells degenerated and two embryos mechanism.We speculated that the gametophyte in I.orientalis usually stopped during free nuclei stage,morphology and anomaly site of archegonia and eggs arranged in disorder might be the causes of abortion.
引文
[1]张宪春.中国植物志(第6卷第3分册)[M].北京:科学出版社.2004,6:220-223.
[2]ZHANG L B,TAYLOR W C.Isoetaceae[M]//Flora of China Editorial Committee.Flora of China.St.Louis Missouri,USA:Missouri Botanical Garden Press,2013,2-3:35-36.
[3]印红.中国珍稀濒危植物图鉴[M].北京:中国林业出版社,2013:32-36.
[4]张宪春,卫然,刘红梅,等.中国现代石松类和蕨类的系统发育与分类系统[J].植物学报,2013,48(2):119-137.ZHANG X C,WEI R,LIU H M,et al.Phylogeny and classification of the extant lycophytes and ferns from China[J].Chinese Bulletin of Botany,2013,48(2):119-137.
[5]中华人民共和国国务院.国家重点保护野生植物名录[J].植物杂志,1999,(5):4-11.STATE COUNCIL OF THE PEOPLES REPUBLIC OF CHI-NA.List of wild plants under state protection[J].Journal of Plant,1999,(5):4-11.
[6]LIU H,WANG Q F,TAYLOR W C.Isoetes orientalis(Isoetaceae),a new hexaploid quilwort from China[J].Novon,2005,15:164-167.
[7]SHANE W S,HICKEY R J.Comparative morphology of the glossopodia of three North American Isoetesligules[J].American Fern Journal,2005,95(3):94-114.
[8]ZHU W J,CHEN X,TANG J J,et al.Analyses on soil nutritional status and water pH value in natural habitat of endangered plant Isoetes orientalis[J].Journal of Plant Resources and Environment,2010,19(2):75-78.
[9]KAROL K G,ARUMUGANATHAN K,BOORE J L,et al.Complete plastome sequences of Equisetum arvense and Isoetes flaccida:implications for phylogeny and plastid genome evolution of early land plant lineages[J].BMC Evolutionary Biology,2010,10(1):321.
[10]KARRFALT E.Some observations on the reproductive anatomy of Isoetes andicola[J].American Fern Journal,1999,89(3):198-203.
[11]CHARLES L M.Morphology of the megagametophyte and the embryo sporophyte of Isoetes lithophila[J].Botanical Society of America,1933,20(4):217-233.
[12]COCHRAN G A,RENZAGLIA K S,TAYLOR W C.Ultrastructure of the spermatozoid of Isoetes[C].Proc.XVI Inter.Bot.Congress,1999:562.
[13]MACLUF C C,Morbelli M A.Giudice G E.Morphology and ultrastructure of megaspores and microspores of Isoetes savatieri Franchet(Lycophyta)[J].Review of Palaeobotany and Palynology,2003,126:197-209.
[14]HUANG SH F,CHIANG S H T.The development of the female gametophyte in Isoetes taiwanensis DeVol[J].Taiwania,1986,31:15-32.
[15]李婧,连永权,李范,等.中华水韭孢子囊发育研究[J].西北植物学报,2015,35(5):929-933.LI J,LIAN Y Q,LI F,et al.Studies on the sporangia development in Isoetes sinensis[J].Acta Bot.Boreali.-Occident.Sin.,2015,35(5):929-933.
[16]刘子玥,邢建娇,路靖,等.极危孑遗蕨类中华水韭有性生殖的形态学研究[J].西北植物学报,2015,35(5):922-928.LIU Z Y,XING J J,LU J,et al.Morphological study on the sexual reproductive of the critically endangered fern Isoetes sinensis[J].Acta Bot.Boreali.-Occident.Sin.,2015,35(5):922-928.
[17]沈玮,谭桂娟,刘子玥,等.云贵水韭雌配子体发育及其濒危机制的研究[J].热带亚热带植物学报,2015,23(5):511-517.SHEN W,TAN G J,LIU Z Y,et al.Famale gametophyte development and endangered mechanism research in Isoetes yunguiensis[J].Journal of Tropical and Subtropical Botany,2015,23(5):511-517.
[18]姜紫薇,刘保东,成晓,等.濒危孑遗蕨类植物云贵水蕨的胚胎发育[J].植物分类与资源学报,2015,37(5):531-536.JIANG Z W,LIU B D,CHENG X,et al.The embryonic development study on Isoetes yunguiensis,an endangered and relic pteridophyta[J].Plant diversity and resourse.,2015,37(5):531-536.
[19]刘婧宏,李范,邢建娇,等.中国4种水韭植物叶片分类学特征的比较观察[J].植物研究,2013,33(5):528-531.LIU J H,LI F,XING J J,et al.Comparative observation on the characteristics of leaves of four species in Chinese Isoetaceae[J].Bulletin of Botanical Research,2013,33(5):528-531.
[20]陈进明,王青锋.珍稀濒危蕨类植物东方水韭的遗传多样性[J].武汉植物学研究,2006,24(6):569-573.CHEN J M,WANG Q F.Genetic diversity in the rare and endangered fern Isoetes orientalis[J].Journal of Wuhan Botanical Research,2006,24(6):569-573.
[21]刘虹,王青锋.中国珍稀濒危植物东方水韭二歧根的发育[J].武汉大学学报,2006,52(6):757-762.LIU H,WANG Q F.Dichotomous root development in Isoetes orientalis,a rare and endangered quillwort in China[J].Journal of Wuhan University,2006,52(6):757-762.
[22]盛玉辉,关维元,徐永星,等.东方水韭对人工污水胁迫的生理响应[J].热带生物学报,2015,6(3):298-303.SHENG Y H,GUAN W Y,XU Y X,et al.Physiological response of Isoetes orientalis to artificial sewage water stress[J].Journal of Tropical Biology,2015,6(3):298-303.
[23]吴兆洪,秦仁昌.中国蕨类植物科属志[M].北京:科学出版社,1991:114-120.
[24]黄武杰,曹建国,王全喜.蕨配子体发育及卵发生的显微结构观察[J].植物研究,2011,31(2):159-163.HUANG W J,CAO J G,WANG Q X.Microstructural observations on the development of gametophytes and oogenesis in the Fern Pteridium aquilium var.latiusculum[J].Bulletin of Botanical Research,2011,31(2):159-163.
[25]国家环境保护局自然保护司.珍惜濒危植物保护与研究[M].北京:中国环境科学出版社,1991:127-135.
[26]石磊,刘宁,刘建武.蕨雌性生殖器官发育的初步研究[J].北京师范大学学报(自然科学版),2003,39(3):377-381.SHI L,LIU N,LIU J W.A preliminary study on femal reproductive organ development in Pteridium aquilium(L.)Kuhn var.latiusculun(Desv.)underw[J].Journal of Beijing Normal University(Nat.Sci.Edi.),2003,39(3):377-381.
[27]路靖,关维元,徐永星,等.中华水韭雌配子体及胚胎发育的初步研究[J].植物科学学报,2013,31(4):353-359.LU J,GUAN W Y,XU Y X,et al.Preliminary study on the megagametophyte and embryological development of Isoetes sinensis[J].Plant Science Journal,2013,31(4):353-359.
[28]BELL P R,MUHLETHALER K.The fine structure of the cells taking part in oogenesis in Pteridiun aquilinum(L.)Kuhn[J].Journal of Ultrastructure Research.,1962,7(5):452-466.
[29]邵文,张宪春,刘保东.海金沙胚胎发育的研究[J].植物研究,2007,27(2):141-144.SHAO W,ZHANG X C,LIU B D.Studies on the development of embryo in Lygodium japonicum(Thunb.)Sw[J].Bulletin of Botanical Research,2007,27(2):141-144.
[30]张笛,檀龙云,刘保东.荚果蕨胚胎发育的研究[J].植物研究,2011,31(1):29-33.ZHANG D,TAN L Y,LIU B D.Development of embryo in Matteuccia struthiopteris(L.)Todaro[J].Bulletin of Botanical Research,2011,31(1):29-33.
[31]NAYAR B K.Evolution of a cormophytic plant body in lower vascular plants[J].Palaeobotanist,1992,41:75-86.
[32]刘保东,成晓,张宪春.Isoetes asiatica生殖的培养观察[C]//中国植物学会七十周年年会论文摘要汇编.中国植物学会,2003.
[33]杨远媛.油松雌配子体游离核有丝分裂的细胞学研究[D].北京:北京林业大学,2008.
[2]ZHANG L B,TAYLOR W C.Isoetaceae[M]//Flora of China Editorial Committee.Flora of China.St.Louis Missouri,USA:Missouri Botanical Garden Press,2013,2-3:35-36.
[3]印红.中国珍稀濒危植物图鉴[M].北京:中国林业出版社,2013:32-36.
[4]张宪春,卫然,刘红梅,等.中国现代石松类和蕨类的系统发育与分类系统[J].植物学报,2013,48(2):119-137.ZHANG X C,WEI R,LIU H M,et al.Phylogeny and classification of the extant lycophytes and ferns from China[J].Chinese Bulletin of Botany,2013,48(2):119-137.
[5]中华人民共和国国务院.国家重点保护野生植物名录[J].植物杂志,1999,(5):4-11.STATE COUNCIL OF THE PEOPLES REPUBLIC OF CHI-NA.List of wild plants under state protection[J].Journal of Plant,1999,(5):4-11.
[6]LIU H,WANG Q F,TAYLOR W C.Isoetes orientalis(Isoetaceae),a new hexaploid quilwort from China[J].Novon,2005,15:164-167.
[7]SHANE W S,HICKEY R J.Comparative morphology of the glossopodia of three North American Isoetesligules[J].American Fern Journal,2005,95(3):94-114.
[8]ZHU W J,CHEN X,TANG J J,et al.Analyses on soil nutritional status and water pH value in natural habitat of endangered plant Isoetes orientalis[J].Journal of Plant Resources and Environment,2010,19(2):75-78.
[9]KAROL K G,ARUMUGANATHAN K,BOORE J L,et al.Complete plastome sequences of Equisetum arvense and Isoetes flaccida:implications for phylogeny and plastid genome evolution of early land plant lineages[J].BMC Evolutionary Biology,2010,10(1):321.
[10]KARRFALT E.Some observations on the reproductive anatomy of Isoetes andicola[J].American Fern Journal,1999,89(3):198-203.
[11]CHARLES L M.Morphology of the megagametophyte and the embryo sporophyte of Isoetes lithophila[J].Botanical Society of America,1933,20(4):217-233.
[12]COCHRAN G A,RENZAGLIA K S,TAYLOR W C.Ultrastructure of the spermatozoid of Isoetes[C].Proc.XVI Inter.Bot.Congress,1999:562.
[13]MACLUF C C,Morbelli M A.Giudice G E.Morphology and ultrastructure of megaspores and microspores of Isoetes savatieri Franchet(Lycophyta)[J].Review of Palaeobotany and Palynology,2003,126:197-209.
[14]HUANG SH F,CHIANG S H T.The development of the female gametophyte in Isoetes taiwanensis DeVol[J].Taiwania,1986,31:15-32.
[15]李婧,连永权,李范,等.中华水韭孢子囊发育研究[J].西北植物学报,2015,35(5):929-933.LI J,LIAN Y Q,LI F,et al.Studies on the sporangia development in Isoetes sinensis[J].Acta Bot.Boreali.-Occident.Sin.,2015,35(5):929-933.
[16]刘子玥,邢建娇,路靖,等.极危孑遗蕨类中华水韭有性生殖的形态学研究[J].西北植物学报,2015,35(5):922-928.LIU Z Y,XING J J,LU J,et al.Morphological study on the sexual reproductive of the critically endangered fern Isoetes sinensis[J].Acta Bot.Boreali.-Occident.Sin.,2015,35(5):922-928.
[17]沈玮,谭桂娟,刘子玥,等.云贵水韭雌配子体发育及其濒危机制的研究[J].热带亚热带植物学报,2015,23(5):511-517.SHEN W,TAN G J,LIU Z Y,et al.Famale gametophyte development and endangered mechanism research in Isoetes yunguiensis[J].Journal of Tropical and Subtropical Botany,2015,23(5):511-517.
[18]姜紫薇,刘保东,成晓,等.濒危孑遗蕨类植物云贵水蕨的胚胎发育[J].植物分类与资源学报,2015,37(5):531-536.JIANG Z W,LIU B D,CHENG X,et al.The embryonic development study on Isoetes yunguiensis,an endangered and relic pteridophyta[J].Plant diversity and resourse.,2015,37(5):531-536.
[19]刘婧宏,李范,邢建娇,等.中国4种水韭植物叶片分类学特征的比较观察[J].植物研究,2013,33(5):528-531.LIU J H,LI F,XING J J,et al.Comparative observation on the characteristics of leaves of four species in Chinese Isoetaceae[J].Bulletin of Botanical Research,2013,33(5):528-531.
[20]陈进明,王青锋.珍稀濒危蕨类植物东方水韭的遗传多样性[J].武汉植物学研究,2006,24(6):569-573.CHEN J M,WANG Q F.Genetic diversity in the rare and endangered fern Isoetes orientalis[J].Journal of Wuhan Botanical Research,2006,24(6):569-573.
[21]刘虹,王青锋.中国珍稀濒危植物东方水韭二歧根的发育[J].武汉大学学报,2006,52(6):757-762.LIU H,WANG Q F.Dichotomous root development in Isoetes orientalis,a rare and endangered quillwort in China[J].Journal of Wuhan University,2006,52(6):757-762.
[22]盛玉辉,关维元,徐永星,等.东方水韭对人工污水胁迫的生理响应[J].热带生物学报,2015,6(3):298-303.SHENG Y H,GUAN W Y,XU Y X,et al.Physiological response of Isoetes orientalis to artificial sewage water stress[J].Journal of Tropical Biology,2015,6(3):298-303.
[23]吴兆洪,秦仁昌.中国蕨类植物科属志[M].北京:科学出版社,1991:114-120.
[24]黄武杰,曹建国,王全喜.蕨配子体发育及卵发生的显微结构观察[J].植物研究,2011,31(2):159-163.HUANG W J,CAO J G,WANG Q X.Microstructural observations on the development of gametophytes and oogenesis in the Fern Pteridium aquilium var.latiusculum[J].Bulletin of Botanical Research,2011,31(2):159-163.
[25]国家环境保护局自然保护司.珍惜濒危植物保护与研究[M].北京:中国环境科学出版社,1991:127-135.
[26]石磊,刘宁,刘建武.蕨雌性生殖器官发育的初步研究[J].北京师范大学学报(自然科学版),2003,39(3):377-381.SHI L,LIU N,LIU J W.A preliminary study on femal reproductive organ development in Pteridium aquilium(L.)Kuhn var.latiusculun(Desv.)underw[J].Journal of Beijing Normal University(Nat.Sci.Edi.),2003,39(3):377-381.
[27]路靖,关维元,徐永星,等.中华水韭雌配子体及胚胎发育的初步研究[J].植物科学学报,2013,31(4):353-359.LU J,GUAN W Y,XU Y X,et al.Preliminary study on the megagametophyte and embryological development of Isoetes sinensis[J].Plant Science Journal,2013,31(4):353-359.
[28]BELL P R,MUHLETHALER K.The fine structure of the cells taking part in oogenesis in Pteridiun aquilinum(L.)Kuhn[J].Journal of Ultrastructure Research.,1962,7(5):452-466.
[29]邵文,张宪春,刘保东.海金沙胚胎发育的研究[J].植物研究,2007,27(2):141-144.SHAO W,ZHANG X C,LIU B D.Studies on the development of embryo in Lygodium japonicum(Thunb.)Sw[J].Bulletin of Botanical Research,2007,27(2):141-144.
[30]张笛,檀龙云,刘保东.荚果蕨胚胎发育的研究[J].植物研究,2011,31(1):29-33.ZHANG D,TAN L Y,LIU B D.Development of embryo in Matteuccia struthiopteris(L.)Todaro[J].Bulletin of Botanical Research,2011,31(1):29-33.
[31]NAYAR B K.Evolution of a cormophytic plant body in lower vascular plants[J].Palaeobotanist,1992,41:75-86.
[32]刘保东,成晓,张宪春.Isoetes asiatica生殖的培养观察[C]//中国植物学会七十周年年会论文摘要汇编.中国植物学会,2003.
[33]杨远媛.油松雌配子体游离核有丝分裂的细胞学研究[D].北京:北京林业大学,2008.