中国产桫椤类配子体在不同基质胁迫下的形态畸变多样性研究
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摘要
为了明确土壤污染对中国产桫椤科植物的生殖影响,以探讨中国桫椤类乃至其他濒危蕨类濒危的内在原因。该研究以中国产9种桫椤植物为材料,采用各植物原产地地表土壤培养孢子至配子体成熟,全程比较观察配子体畸变类型,统计畸变率;并分别取3个属的代表种笔筒树、大叶黑桫椤和中华桫椤的孢子,以MS培养基为对照,分别用5 mg·kg~(-1)Pb~(2+)和1 mg·kg~(-1)Cd~(2+)单因素胁迫培养配子体,比较观察各组的配子体形态畸变类型,统计畸变率。结果表明:(1)中国产桫椤类9种植物配子体培养过程中共出现17种形态畸变类型,其中4种为分化畸变,7种为假根畸变,4种为细胞畸变,2种为性器畸变。(2)不同培养基——对照组(MS培养基)、原产地土壤、Cd~(2+)胁迫、Pb~(2+)胁迫对中国产桫椤科9种植物配子体的平均畸变比例分别是2.28%、12.61%、31.58%和33.58%。(3) 9种桫椤的配子体畸变类型及其畸变率均与培养基的胁迫程度呈正相关关系,其配子体发育都受到了产地土壤污染的严重伤害。(4) 3种桫椤属植物配子体对土壤环境胁迫的耐受力强弱为黑桫椤属>白桫椤属>桫椤属。(5)桫椤科配子体的形态畸变多样性与胁迫条件之间具有稳定的对应关系。
In order to determine the reproductive effects of soil pollution on Chinese Cyatheaceae and explore the underlying causes of the endangered species of Chinese Cyatheaceae and other endangered ferns, were cultured the spores of nine species of Chinese Cyatheaceae to be mature gametophytes in the surface soil from their source area. The aberration types of gametophytes were observed and the aberration rate was calculated. And the spores of three representative species of genera, Sphaeropteris lepifera, Gymnosphaera gigantea and Alsophila costularis, were taken respectively. MS medium was taken as the control group, the gametophytes were cultured under univariate stress of 5 mg·kg~(-1) Pb~(2+)and 1 mg·kg~(-1) Cd~(2+), respectively, and then the types of morphological aberration of gametophytes and their aberration rates were compared and observed. The results showed that:(1) it was observed that 17 morphological aberrations occurred in the gametophyte culture of 9 kinds of Chinese Cyatheaceae, including four differentiation aberration, seven rhizoid aberration, four cell aberration and two reproductive organs aberration.(2) The aberration gametophytic ratio on different media——control group, soils from source area, Cd~(2+) stress, and Pb~(2+) stress was 2.28%, 12.61%, 31.58% and 33.58% in turn.(3) The types and rates of gametophyte aberration in all 9 types of Cyatheaceae were positively correlated with the stress degree of the medium, the development of their gametophytes has been seriously damaged by soil pollution.(4) Resistance of gametophytes of three species of Cyatheaceae to soil environmental stress from strong to weak, in turn, was Gymnosphaera, Sphaeropteris, and Alsophila.(5) There was a stable relationship between the diversity of morphological aberration and stress conditions of the gametophytes.
In order to determine the reproductive effects of soil pollution on Chinese Cyatheaceae and explore the underlying causes of the endangered species of Chinese Cyatheaceae and other endangered ferns, were cultured the spores of nine species of Chinese Cyatheaceae to be mature gametophytes in the surface soil from their source area. The aberration types of gametophytes were observed and the aberration rate was calculated. And the spores of three representative species of genera, Sphaeropteris lepifera, Gymnosphaera gigantea and Alsophila costularis, were taken respectively. MS medium was taken as the control group, the gametophytes were cultured under univariate stress of 5 mg·kg~(-1) Pb~(2+)and 1 mg·kg~(-1) Cd~(2+), respectively, and then the types of morphological aberration of gametophytes and their aberration rates were compared and observed. The results showed that:(1) it was observed that 17 morphological aberrations occurred in the gametophyte culture of 9 kinds of Chinese Cyatheaceae, including four differentiation aberration, seven rhizoid aberration, four cell aberration and two reproductive organs aberration.(2) The aberration gametophytic ratio on different media——control group, soils from source area, Cd~(2+) stress, and Pb~(2+) stress was 2.28%, 12.61%, 31.58% and 33.58% in turn.(3) The types and rates of gametophyte aberration in all 9 types of Cyatheaceae were positively correlated with the stress degree of the medium, the development of their gametophytes has been seriously damaged by soil pollution.(4) Resistance of gametophytes of three species of Cyatheaceae to soil environmental stress from strong to weak, in turn, was Gymnosphaera, Sphaeropteris, and Alsophila.(5) There was a stable relationship between the diversity of morphological aberration and stress conditions of the gametophytes.
引文
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[2] 张宪春.中国植物志(第六卷第三分册)[M].北京:科学出版社,2004:249-274.
[3] 印红.中国珍稀濒危植物图鉴[M].北京:中国林业出版社,2013:13-26.
[4] 贵州省环境保护局.赤水桫椤自然保护区科学考察集[M].贵阳:贵州民族出版社,1990:1.
[5] 国家林业局,农业部.国家重点保护野生植物名录(第一批)[J].中华人民共和国国务院公报,2000,(13):39-47.State Forestry Administration&Ministry of Agriculture of the People’s Republic of China.Lists of wild plants that should be protected with special care at national level (First batch)[J].Gazette of the State Council of the People’s Republic of China,2000,(13):39-47.
[6] 鞠文彬,高信芬,包维楷.画稿溪国家级自然保护区珍稀植物桫椤种群结构与更新[J].植物科学学报,2014,32(2):113-121.JU W B,GAO X F,BAO W K.Population structure and regeneration of the endangered tree fern Alsophila spinulosa in Huagaoxi National nature reserve,Southwestern China[J].Plant Science Journal,2014,32(2):113-121.
[7] 周志琼,苏智先,廖永梅,等.桫椤的生物学研究进展.贵州师范大学学报[J].2004,22(3):100-103.ZHOU Z Q,SU Z X,LIAO Y M,et al.Advance of biological study on Alsophila spinulosa[J].Journal of Guizhou Normal University,2004,22(3):100-103.
[8] 董仕勇.广州市蕨类植物物种多样性研究[J].热带亚热带植物学报,2008,16(1):39-45.DONG S Y.A biodiversity study on the pteridophytes from Ghuangzhou City,Guangdong[J].Journal of Tropical and Subtropical Botany,2008,16(1):39-45.
[9] 徐德静,郭能彬,王鹏鹏,等.习水自然保护区桫椤种群结构与分布格局研究[J].西南大学学报,2014,36(11):93-98.XU D J,GUO N B,WANG P P,et al.Study on population structure and distribution patterns of Alsophila spinulosa in Xishui Nature Reserve[J].Journal of Southwest University,2014,36(11):93-98.
[10] 顾垒,闻丞,罗玫,等.中国最受关注濒危物种保护现状快速评价的新方法探讨[J].生物多样性,2015,23(5):583-590.GU L,WEN C,LUO M,et al.A rapid approach for assessing the conservation effectiveness of the most concerned endangered species in China[J].Biodiversity Science,2015,23(5):583-590.
[11] FU C H,HSIEH H M,CHEN C Y,et al.Ophiodiaporthe Cyatheae gen.et sp.nov.a diaporthalean pathogen causing a devastating wilt disease of Cyathea lepifera in Taiwan[J].Mycologia,2013,105(4):12.
[12] 王金娟,张宪春,刘保东,等.桫椤科三种植物配子体发育的研究[J].热带亚热带植物学报,2007,15(2):115-120.WANG J J,ZHANG X C,LIU B D,et al.Gametophyte development of three species in Cyatheaceae[J].Journal of Tropical and Subtropical Botany,2007,15(2):115-120.
[13] CHEN G J,CHENG X,LIU B D,et al.Gametophyte morphological variations of Sphaeropteris brunoniana (Cyatheaceae) under four culture conditions[J].Acta Botanica Yunnanica,2008,30(4):430-432.
[14] 陈贵菊,成晓,刘保东,等.四种培养条件下白桫椤配子体的形态发育[J].云南植物研究,2008,30(4):430-432.CHEN G J,CHENG X,LIU B D,et al.Gametophyte morphological variations of Sphaeropteris brunoniana (Cyatheaceae) under four culture conditions[J].Acta Botanica Yunnanica,2008,30(4):430-432.
[15] 杜红红,李杨,李东.光照、温度和pH值对小黑桫椤孢子萌发及早期配子体发育的影响[J].生物多样性,2009,17(2):182-187.DU H H,LI Y,LI D.Effects of light,temperature and pH on spore germination and early gametophytic development of Alsophila metteniana[J].Biodiversity Science,2009,17(2):182-187.
[16] HUANG Y M,YAO M,YING,et al.Morphology of gametophytes and young sporophytes of Sphaeropteris lepifera[J].American Fern Journal,2000,90(4):127-137.
[17] 汪盛,刘恩德,夏漪,等.重金属铅对3种蕨类植物孢子萌发和配子体发育的影响[J].植物研究,2015,35(4):499-503.WANG S,LIU E D,XIA Y,et al.Effects of Pb2+ on spore germination and gametophyte development of three species of fern[J].Bulletin of Botanical Research,2015,35(4):499-503.
[18] 亢希然,范稚莲,乔西,等.无菌条件下砷对蜈蚣草配子体生长发育的影响[J].广西植物,2007,27(6):882-885.KANG X R,FAN Z L,QIAO X,et al.Effect of Arsenic on gametophytes development of Pteris vittata under aseptic condition[J].Guihaia,2007,27(6):882-885.
[19] MOMOSE S.Prothallia of the Japanese Ferns[M].Tokyo:University of Tokyo Press,1967:177-187.
[20] CHEN G J,CHENG X,LIU B D,et al.Comparative studies on gametophyte morphology and development of species of Cyatheaceae[J].American Fern Journal,2008,98(2):83-95.
[21] 方炎明,魏勇,张晓平,等.苔藓生物监测大气重金属污染研究进展[J].南京林业大学学报,2000,24(5):64-68.FANG Y M,WEI Y,ZHANG X P,et al.Advance in Bry-monotoring of atmosperic heavy metal pollution[J].Journal of Nanjing Forestry University,2000,24(5):64-68.
[22] 温瑀,穆立蔷.土壤铅、镉胁迫对4种绿化植物生长、生理及积累特性的影响[J].水土保持学报,2013,27(5):234-239.WEN Y,MU L Q.Effect of soil Pb,Cd stress on the growth,physiological and accumulating charateristics of four ornamental trees[J].Journal of Soil and Water Conservation,2013,27(5):234-239.
[23] 纪善博,向建英,刘保东,等.桫椤科笔筒树游动精子行为及形态观察[J].西北植物学报,2018,38(3):478-483.JI S B,XIANG J Y,LIU B D,et al.Behavior and morphology of Sphaeropteris lepifera (Cyatheaceae) swimming sperm[J].Acta Botanica Boreali-Occidentalia Sinica,2018,38(3):478-483.
[24] 陈勤,孙冲,方炎明.苔藓植物的生态环境指示作用[J].世界林业研究,2013,26(2):19-23.CHEN Q,SUN C,FANG Y M.Review on bio-monitoring of Bryophyte to environment changes[J].World Forestry Research,2013,26(2),19-23.
[25] 常福辰,施国新,吴国荣,等.汞、镉复合污染对金鱼藻的影响及其抗性机制的探讨[J].广西植物,2002,22(5):453-457.CHANG F C,SHI G X,WU G R,et al.Changes of cell membrance lipid peroxidation and activities of antioxidant enzyme systems of Ceratophyllum demersum L.in combinative pollution of Cd2+ and Hg2+[J].Journal of Nanjing Normal University,2002,22(5):453-457.
[26] 王慧忠,何翠屏,赵楠.铅对草坪植物生物量与叶绿素水平的影响[J].草业科学,2003,20(6):73-75.WANG H Z,HE C P,ZHAO N.Effect of lead on turfgrass biomass and chlorophyll level[J].Pratacultural Science,2003,20(6):73-75.
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