蕨类植物的克隆性及其生态功能
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摘要
通过对蕨类植物生活史特性进行探讨可知,克隆整合影响着蕨类植物的表型可塑性,同时在异质环境中特别是胁迫环境中起着重要的作用。克隆整合作用是植物适应环境的重要机制之一。通过克隆整合,可在克隆蕨类植物生长系统内共享局部可利用资源、缓解局部压力,从而促进克隆蕨类的生长和生存,在蕨类植物的生态适应中发挥着重要的意义。
Through relevant researches on the life history characteristics of fern plants,its was showed that clonal integration will affect the phenotypic plasticity of ferns,and meanwhile,it plays an important role in heterogeneous environments,especially stressful environments.Clonal integration is one of the important mechanisms for adapting to the environment.Through clonal integration,local available resources could be shared in the growth system of clonal fern,and local stress could be alleviated,thereby improving the growth and survival of cloned ferns,and it play an significant role in the ecological adaptation of ferns.
Through relevant researches on the life history characteristics of fern plants,its was showed that clonal integration will affect the phenotypic plasticity of ferns,and meanwhile,it plays an important role in heterogeneous environments,especially stressful environments.Clonal integration is one of the important mechanisms for adapting to the environment.Through clonal integration,local available resources could be shared in the growth system of clonal fern,and local stress could be alleviated,thereby improving the growth and survival of cloned ferns,and it play an significant role in the ecological adaptation of ferns.
引文
[1]周喜乐,张宏春,孙久琼,等.中国石松类和蕨类植物的多样性与地理分布[J].生物多样性,2016,24(1):102-107.
[2]严岳鸿,张宪春,马克平.中国蕨类植物多样性与地理分布[M].北京:科学出版社,2013.
[3]Schimel J P,Bennett J.Nitrogen mineralization:challenges of a changing paradigm[J].Ecology,2004,85(3):591-602.
[4]Schuettpelz E,Pryer K M.Evidence for a Cenozoic radiation of ferns in an angiosperm-dominated canopy[J].Proc Natl Acad Sci U S A,2009,106(27):11200-11205.
[5]Jr J E W,Rundel P W,Cardelús C L.The influence of life form on carbon and nitrogen relationships in tropical rainforest ferns[J].Oecologia,2007,153(2):225-232.
[6]陆树刚.蕨类植物学[M].北京:高等教育出版社,2007.
[7]Barrington D S.Ecological and historical factors in fern biogeography[J].Journal of Biogeography,1993,20(3):275-279.
[8]Given D R.Changing aspects of endemism and endangerment in pteridophyta[J].Journal of Biogeography,1993,20(3):293-302.
[9]Aarssen L W.Death without sex-the‘problem of the small’and selection for reproductive economy in flowering plants[J].Evolutionary Ecology,2008,22(3):279-298.
[10]Tiffney B H,Niklas K J.Clonal growth in land plants:a paleobotanical perspective[M].New York:Yale University Press,1985.
[11]James E W,Amber C C,Michele H N.A site for sori:Ecophysiology of fertile-sterile leaf dimorphy in ferns[J].American Journal of Botany,2016,103(5):845.
[12]Erschbamer B.Ecology and evolutionary biology of clonal plants[M].Springer Science+Business Media B.V.,2002.
[13]Vance R R.Clonal organisms:population biology and evolution of clonal organisms[J].Science,1987,235(4793):1264.
[14]Alpert P.Clonal integration increases performance of ramets of the fern Diplopterygium glaucum in an evergreen forest in southeastern China[J].Flora,2009,205(6):399-403.
[15]卢华正.云南热带和亚热带森林附生蕨类植物的克隆性及其生态适应策略[D].北京:中国科学院大学,2017.
[16]Carrie R A.The effects of rhizome severing and nutrient addition on growth and biomass allocation in Diphasiastrumdigitatum[J].American Fern Journal,2000,90(2):77.
[17]Laube S,Zotz G.Neither host-specific nor random:vascular epiphytes on three tree species in a panamanian lowland forest[J].Annals of Botany,2006,97(6):1103-1114.
[18]Bell D L,Galloway L F.Population differentiation for plasticity to light in an annual herb:Adaptation and cost[J].American Journal of Botany,2008,95(1):59-65.
[19]Sultan S E.Evolutionary implications of phenotypic plasticity in plants[M].New York:Plenum Press,1987.
[20]Dong M,During H J,Werger M J A.Clonal plasticity in response to nutrient availability in the pseudoannual herb,Trientalis europaea L.[J].Plant Ecology,1997,131(2):233-239.
[21]Dong M,Werger D M J A.Morphological responses to nutrient availability in four clonal herbs[J].Vegetatio,1996,123(2):183-192.
[22]Slade A J,Hutchings M J.Clonal integration and plasticity in foraging behaviour in Glechoma hederacea[J].Journal of Ecology,1987,75(4):1023-1036.
[23]Slade A J,Hutchings M J.The Effects of light intensity on foraging in the clonal herb Glechoma hederacea[J].Journal of Ecology,1987,75(1):95-112.
[24]Hutchings K M J.Morphological plasticity in clonal plants:the foraging concept reconsidered[J].Journal of Ecology,1995,83(1):143-152.
[25]Niva M,Svensson B M,Karlsson P S.Effects of light and water availability on shoot dynamics of the stoloniferous plant Linnaea borealis[J].coscience,2006,13(3):318-323.
[26]Watkins J E,Cardelús C.Habitat differentiation of ferns in a lowland tropical rain forest[J].American Fern Journal,2009,99(3):162-175.
[27]Watkins J E,Mack M C,Mulkey S S S.Ecological and evolutionary consequences of desiccation tolerance in tropical fern gametophytes[J].New Phytologist,2007,176(3):708-717.
[28]Winkler M,Hietz P.High gene flow in epiphytic ferns despite habitat loss and fragmentation[J].Conservation Genetics (Print),2011,12(6):1411-1420.
[29]Zotz G,Hietz P,Schmidt G.Small plants,large plants:the importance of plant size for the physiological ecology of vascular epiphytes[J].Journal of Experimental Botany,2001,52(363):2051.
[30]Song Y B,Yu F H,Keser L H,et al.United we stand,divided we fall:a meta-analysis of experiments on clonal integration and its relationship to invasiveness[J].Oecologia (Berlin),2013,171(2):317-327.
[31]Kawai H,Kanegae T,Christensen S,et al.Responses of ferns to red light are mediated by an unconventional photoreceptor[J].Nature,2003,421(6920):287-290.
[32]Schneider H.Morphology and anatomy of roots in the filmy fern tribe Trichomaneae H.Schneider (Hymenophyllaceae,Filicatae) and the evolution of rootless taxa[J].Botanical Journal of the Linnean Society,2000,132(1):29-46.
[33]Benzing D H.Vascular epiphytes[M].Forest Canopies,2004:175-211.
[34]Dubuisson J Y,Schneider H,Hennequin S.Epiphytism in ferns:diversity and history[J].Comptesrendus- Biologies,2009,332(2):120-128.
[35]Zhang Q,Chen J W,Li B G,et al.The effect of drought on photosynthesis in two epiphytic and two terrestrial tropical fern species[J].Photosynthetica (Prague),2009,47(1):128-132.
[36]Magyar G,Kun A,Oborny B,et al.Importance of plasticity and decision-making strategies for plant resource acquisition in spatio-temporally variable environments[J].New Phytologist,2007,174(1):182-193.
[37]Wiens,J A.Population responses to patchy environments[J].Annual Review of Ecology & Systematics,1976,7(7):81-120.
[38]董鸣.克隆植物生态学[M].北京:科学出版社,2011.
[39]Alpert P.Division of labour in clonal plants[J].Ecology & Evolution of Clonal Plants,1997:137-154.
[40]Cook R E.Clonal plant populations[J].Amer Sci,1983,71(3):244-253.
[41]Friedman D,Alpert P.Reciprocal transport between ramets increases growth of Fragaria chiloensis when light and nitrogen occur in separate patches but only if patches are rich[J].Oecologia,1991,86(1):76-80.
[42]Kroon H D,Groenendael J V.The ecology and evolution of clonal plants[M].Leiden:Backhuys Publishers,1997.
[43]Michael J,Hutchings D K.Patchy habitats,division of labour and growth dividends in clonal plants[J].Trends in Ecology & Evolution,1997,12(10):390-394.
[44]Evans J P.The effect of local resource availability and clonal integration on ramet functional morphology in Hydrocotyle bonariensis[J].Oecologia,1992,89(2):265-276.
[45]Kleunen M V,Stuefer J F.Quantifying the effects of reciprocal assimilate and water translocation in a clonal plant by the use of steam-girdling[J].Oikos,1999,85(1):135-145.
[46]Stuefer J F.Two types of division of labour in clonal plants:benefits,costs and constraints[J].Perspect Plant EcolEvol Syst,1998,1(1):47-60.
[47]Dong M.Clonal growth in plants in relation to resource heterogeneity:Foraging behavior[J].Acta Botanica Sinica,1996,38(10):828-835.
[48]Guo W,Song Y B,Yu F H J P O.Heterogeneous light supply affects growth and biomass allocation of the understory fern Diplopterygium glaucum at high patch contrast[J].2011,6(11):e27998.
[49]Lu H Z,Song L,Liu W Y,et al.Survival and growth of epiphytic ferns depend on resource sharing[J].Frontiers in Plant Science,2016,7:416.
[50]Lu H Z,Liu W Y,Yu F H,et al.Higher clonal integration in the facultative epiphytic fern Selliguea griffithiana growing in the forest canopy compared with the forest understorey[J].Ann Bot,2015,116(1):113-122.
[51]Song Y B,Yu F H,Lidewij H,et al.United we stand,divided we fall:a meta-analysis of experiments on clonal integration and its relationship to invasiveness[J].Oecologia,2013,171(2):317-327.
[52]Alpert P.Nitrogen sharing among ramets increases clonal growth in Fragaria chiloensis[J].Ecology,1991,72(1):69-80.
[53]Roiloa S R,Retuerto R.Clonal integration in Fragaria vesca growing in metal-polluted soils:parents face penalties for establishing their offspring in unsuitable environments[J].Ecological Research,2012,27(1):95-106.
[54]Reeves R D,Troumbis A Y.Hypotheses,mechanisms and trade-offs of tolerance and adaptation to serpentine soils:from species to ecosystem level[J].Biological Reviews,2010,83(4):495-508.
[55]Maestri E,Marmiroli M,Visioli G,et al.Metal tolerance and hyperaccumulation:Costs and trade-offs between traits and environment[J].Environmental & Experimental Botany,2010,68(1):1-13.
[56]Baker A J M.Accumulators and excluders strategies in the response of plants to heavy metals[J].Journal of Plant Nutrition,1981,3(1-4):643-654.
[57]Brady K U,Kruckeberg A R,Jr H D B.Evolutionary ecology of plant adaptation to serpentine soils[J].Annual Review of Ecology Evolution & Systematics,2005,36(1):243-266.
[58]Koivunen S,Saikkonen K,Vuorisalo T,et al.Heavy metals modify costs of reproduction and clonal growth in the stoloniferous herb Potentilla anserina[J].Evolutionary Ecology,2004,18(5-6):541-561.
[59]Outridge P M,Hutchinson T C.Induction of cadmium tolerance by acclimation transferred between ramets of the clonal fern Salvinia minima Baker[J].New Phytologist,1991,117(4):597-605.
[60]Hartnett D C,Bazzaz F A.Physiological integration among intraclonal ramets in Solidago canadensis[J].Ecology,1983,64(4):779-788.
[61]Sutherland W J,Stillman R A.The foraging tactics of plants[J].Oikos,1988,52(3):239-244.
[2]严岳鸿,张宪春,马克平.中国蕨类植物多样性与地理分布[M].北京:科学出版社,2013.
[3]Schimel J P,Bennett J.Nitrogen mineralization:challenges of a changing paradigm[J].Ecology,2004,85(3):591-602.
[4]Schuettpelz E,Pryer K M.Evidence for a Cenozoic radiation of ferns in an angiosperm-dominated canopy[J].Proc Natl Acad Sci U S A,2009,106(27):11200-11205.
[5]Jr J E W,Rundel P W,Cardelús C L.The influence of life form on carbon and nitrogen relationships in tropical rainforest ferns[J].Oecologia,2007,153(2):225-232.
[6]陆树刚.蕨类植物学[M].北京:高等教育出版社,2007.
[7]Barrington D S.Ecological and historical factors in fern biogeography[J].Journal of Biogeography,1993,20(3):275-279.
[8]Given D R.Changing aspects of endemism and endangerment in pteridophyta[J].Journal of Biogeography,1993,20(3):293-302.
[9]Aarssen L W.Death without sex-the‘problem of the small’and selection for reproductive economy in flowering plants[J].Evolutionary Ecology,2008,22(3):279-298.
[10]Tiffney B H,Niklas K J.Clonal growth in land plants:a paleobotanical perspective[M].New York:Yale University Press,1985.
[11]James E W,Amber C C,Michele H N.A site for sori:Ecophysiology of fertile-sterile leaf dimorphy in ferns[J].American Journal of Botany,2016,103(5):845.
[12]Erschbamer B.Ecology and evolutionary biology of clonal plants[M].Springer Science+Business Media B.V.,2002.
[13]Vance R R.Clonal organisms:population biology and evolution of clonal organisms[J].Science,1987,235(4793):1264.
[14]Alpert P.Clonal integration increases performance of ramets of the fern Diplopterygium glaucum in an evergreen forest in southeastern China[J].Flora,2009,205(6):399-403.
[15]卢华正.云南热带和亚热带森林附生蕨类植物的克隆性及其生态适应策略[D].北京:中国科学院大学,2017.
[16]Carrie R A.The effects of rhizome severing and nutrient addition on growth and biomass allocation in Diphasiastrumdigitatum[J].American Fern Journal,2000,90(2):77.
[17]Laube S,Zotz G.Neither host-specific nor random:vascular epiphytes on three tree species in a panamanian lowland forest[J].Annals of Botany,2006,97(6):1103-1114.
[18]Bell D L,Galloway L F.Population differentiation for plasticity to light in an annual herb:Adaptation and cost[J].American Journal of Botany,2008,95(1):59-65.
[19]Sultan S E.Evolutionary implications of phenotypic plasticity in plants[M].New York:Plenum Press,1987.
[20]Dong M,During H J,Werger M J A.Clonal plasticity in response to nutrient availability in the pseudoannual herb,Trientalis europaea L.[J].Plant Ecology,1997,131(2):233-239.
[21]Dong M,Werger D M J A.Morphological responses to nutrient availability in four clonal herbs[J].Vegetatio,1996,123(2):183-192.
[22]Slade A J,Hutchings M J.Clonal integration and plasticity in foraging behaviour in Glechoma hederacea[J].Journal of Ecology,1987,75(4):1023-1036.
[23]Slade A J,Hutchings M J.The Effects of light intensity on foraging in the clonal herb Glechoma hederacea[J].Journal of Ecology,1987,75(1):95-112.
[24]Hutchings K M J.Morphological plasticity in clonal plants:the foraging concept reconsidered[J].Journal of Ecology,1995,83(1):143-152.
[25]Niva M,Svensson B M,Karlsson P S.Effects of light and water availability on shoot dynamics of the stoloniferous plant Linnaea borealis[J].coscience,2006,13(3):318-323.
[26]Watkins J E,Cardelús C.Habitat differentiation of ferns in a lowland tropical rain forest[J].American Fern Journal,2009,99(3):162-175.
[27]Watkins J E,Mack M C,Mulkey S S S.Ecological and evolutionary consequences of desiccation tolerance in tropical fern gametophytes[J].New Phytologist,2007,176(3):708-717.
[28]Winkler M,Hietz P.High gene flow in epiphytic ferns despite habitat loss and fragmentation[J].Conservation Genetics (Print),2011,12(6):1411-1420.
[29]Zotz G,Hietz P,Schmidt G.Small plants,large plants:the importance of plant size for the physiological ecology of vascular epiphytes[J].Journal of Experimental Botany,2001,52(363):2051.
[30]Song Y B,Yu F H,Keser L H,et al.United we stand,divided we fall:a meta-analysis of experiments on clonal integration and its relationship to invasiveness[J].Oecologia (Berlin),2013,171(2):317-327.
[31]Kawai H,Kanegae T,Christensen S,et al.Responses of ferns to red light are mediated by an unconventional photoreceptor[J].Nature,2003,421(6920):287-290.
[32]Schneider H.Morphology and anatomy of roots in the filmy fern tribe Trichomaneae H.Schneider (Hymenophyllaceae,Filicatae) and the evolution of rootless taxa[J].Botanical Journal of the Linnean Society,2000,132(1):29-46.
[33]Benzing D H.Vascular epiphytes[M].Forest Canopies,2004:175-211.
[34]Dubuisson J Y,Schneider H,Hennequin S.Epiphytism in ferns:diversity and history[J].Comptesrendus- Biologies,2009,332(2):120-128.
[35]Zhang Q,Chen J W,Li B G,et al.The effect of drought on photosynthesis in two epiphytic and two terrestrial tropical fern species[J].Photosynthetica (Prague),2009,47(1):128-132.
[36]Magyar G,Kun A,Oborny B,et al.Importance of plasticity and decision-making strategies for plant resource acquisition in spatio-temporally variable environments[J].New Phytologist,2007,174(1):182-193.
[37]Wiens,J A.Population responses to patchy environments[J].Annual Review of Ecology & Systematics,1976,7(7):81-120.
[38]董鸣.克隆植物生态学[M].北京:科学出版社,2011.
[39]Alpert P.Division of labour in clonal plants[J].Ecology & Evolution of Clonal Plants,1997:137-154.
[40]Cook R E.Clonal plant populations[J].Amer Sci,1983,71(3):244-253.
[41]Friedman D,Alpert P.Reciprocal transport between ramets increases growth of Fragaria chiloensis when light and nitrogen occur in separate patches but only if patches are rich[J].Oecologia,1991,86(1):76-80.
[42]Kroon H D,Groenendael J V.The ecology and evolution of clonal plants[M].Leiden:Backhuys Publishers,1997.
[43]Michael J,Hutchings D K.Patchy habitats,division of labour and growth dividends in clonal plants[J].Trends in Ecology & Evolution,1997,12(10):390-394.
[44]Evans J P.The effect of local resource availability and clonal integration on ramet functional morphology in Hydrocotyle bonariensis[J].Oecologia,1992,89(2):265-276.
[45]Kleunen M V,Stuefer J F.Quantifying the effects of reciprocal assimilate and water translocation in a clonal plant by the use of steam-girdling[J].Oikos,1999,85(1):135-145.
[46]Stuefer J F.Two types of division of labour in clonal plants:benefits,costs and constraints[J].Perspect Plant EcolEvol Syst,1998,1(1):47-60.
[47]Dong M.Clonal growth in plants in relation to resource heterogeneity:Foraging behavior[J].Acta Botanica Sinica,1996,38(10):828-835.
[48]Guo W,Song Y B,Yu F H J P O.Heterogeneous light supply affects growth and biomass allocation of the understory fern Diplopterygium glaucum at high patch contrast[J].2011,6(11):e27998.
[49]Lu H Z,Song L,Liu W Y,et al.Survival and growth of epiphytic ferns depend on resource sharing[J].Frontiers in Plant Science,2016,7:416.
[50]Lu H Z,Liu W Y,Yu F H,et al.Higher clonal integration in the facultative epiphytic fern Selliguea griffithiana growing in the forest canopy compared with the forest understorey[J].Ann Bot,2015,116(1):113-122.
[51]Song Y B,Yu F H,Lidewij H,et al.United we stand,divided we fall:a meta-analysis of experiments on clonal integration and its relationship to invasiveness[J].Oecologia,2013,171(2):317-327.
[52]Alpert P.Nitrogen sharing among ramets increases clonal growth in Fragaria chiloensis[J].Ecology,1991,72(1):69-80.
[53]Roiloa S R,Retuerto R.Clonal integration in Fragaria vesca growing in metal-polluted soils:parents face penalties for establishing their offspring in unsuitable environments[J].Ecological Research,2012,27(1):95-106.
[54]Reeves R D,Troumbis A Y.Hypotheses,mechanisms and trade-offs of tolerance and adaptation to serpentine soils:from species to ecosystem level[J].Biological Reviews,2010,83(4):495-508.
[55]Maestri E,Marmiroli M,Visioli G,et al.Metal tolerance and hyperaccumulation:Costs and trade-offs between traits and environment[J].Environmental & Experimental Botany,2010,68(1):1-13.
[56]Baker A J M.Accumulators and excluders strategies in the response of plants to heavy metals[J].Journal of Plant Nutrition,1981,3(1-4):643-654.
[57]Brady K U,Kruckeberg A R,Jr H D B.Evolutionary ecology of plant adaptation to serpentine soils[J].Annual Review of Ecology Evolution & Systematics,2005,36(1):243-266.
[58]Koivunen S,Saikkonen K,Vuorisalo T,et al.Heavy metals modify costs of reproduction and clonal growth in the stoloniferous herb Potentilla anserina[J].Evolutionary Ecology,2004,18(5-6):541-561.
[59]Outridge P M,Hutchinson T C.Induction of cadmium tolerance by acclimation transferred between ramets of the clonal fern Salvinia minima Baker[J].New Phytologist,1991,117(4):597-605.
[60]Hartnett D C,Bazzaz F A.Physiological integration among intraclonal ramets in Solidago canadensis[J].Ecology,1983,64(4):779-788.
[61]Sutherland W J,Stillman R A.The foraging tactics of plants[J].Oikos,1988,52(3):239-244.
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