养分浓度和光照强度对泥炭藓有性更新的影响
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摘要
作为优势植物,泥炭藓(Sphagnum)在泥炭沼泽中缺乏有性更新的原因尚不清楚。针对影响孢子萌发的光强和养分条件,以泥炭藓(S.palustre)为材料,通过室内孢子萌发实验,研究不同光强和养分浓度对孢子萌发率、萌发势及萌发指数的影响。4种培养基中,养分浓度高的营养液培养基中孢子萌发率最高,达到60%,其次为养分浓度与营养液相近的琼脂+营养液培养基,萌发率为48%,再次为养分水平很低的沼泽水培养基,萌发率约为30%,几乎无养分的蒸馏水培养基中萌发率最低,约为5%。萌发势和萌发指数亦呈现相同的规律。琼脂+营养液和营养液培养基较沼泽水和蒸馏水培养基孢子萌发时间提前约3天时间。增加光强使孢子萌发率仅提高10%。研究表明,低养分浓度和弱光照均不利于孢子萌发,相对而言,泥炭沼泽的贫营养特征应是限制泥炭藓有性更新的更重要因素。
The mechanism for rare sexual regeneration in Sphagnum palustre, the dominant plant in peatlands is not clear. Aiming at the two factors affecting spore germination, we chose S. palustre as study material and conducted an indoor spore germination experiment to explore the effect of light intensity and nutrient concentration on germination rate(GR), germination vigor and germination index. Among the four culture media, nutrient solution with suitable level of nutrients showed the greatest GR, 60%; agar + nutrient solution with highest level of nutrients showed higher GR, 48%; mire water with much less nutrients showed lower GR, 30%; and distilled water with the least nutrients showed the lowest GR, about 5%. Similar pattern was also observed in germination vigor and germination index. The initial germination time in nutrient solution and agar + nutrient solution was approximate 3 days earlier than that in mire water and distilled water. Enhancing light intensity only increased GR by 10% or so. This study indicates that less nutrient and light are detrimental to spore germination; the oligotrophy in peatlands is the main factor to limit sexual regeneration of Sphagnum.
The mechanism for rare sexual regeneration in Sphagnum palustre, the dominant plant in peatlands is not clear. Aiming at the two factors affecting spore germination, we chose S. palustre as study material and conducted an indoor spore germination experiment to explore the effect of light intensity and nutrient concentration on germination rate(GR), germination vigor and germination index. Among the four culture media, nutrient solution with suitable level of nutrients showed the greatest GR, 60%; agar + nutrient solution with highest level of nutrients showed higher GR, 48%; mire water with much less nutrients showed lower GR, 30%; and distilled water with the least nutrients showed the lowest GR, about 5%. Similar pattern was also observed in germination vigor and germination index. The initial germination time in nutrient solution and agar + nutrient solution was approximate 3 days earlier than that in mire water and distilled water. Enhancing light intensity only increased GR by 10% or so. This study indicates that less nutrient and light are detrimental to spore germination; the oligotrophy in peatlands is the main factor to limit sexual regeneration of Sphagnum.
引文
[1]吴鹏程.苔藓植物生物学[M].北京:科学出版社,1998:10–46.
[2]范庆书,赵建成,于树宏.苔藓植物孢子萌发与原丝体发育研究进展[J].植物学通报,2003,20(3):280–286.
[3]CRONBERG N.Reproductive biology of Sphagnum[J].Lindbergia,1993,17(4):69–82.
[4]SUNDBERG S,RYDIN H.Habitat requirements for establishment of Sphagnum from spores[J].Journal of Ecology,2002,90(2):268–278.
[5]BOLD H C.The prothallium of Sphagnum palustre L.[J].The Bryologist,1948,51(2):55–63.
[6]李秋萍,吴思苹,鲁蓓蓓,等.藓类植物孢子生活力的快速检测方法初探[J].西北植物学报,2013,33(10):2126–2130.
[7]NOGUCHI A.Germination of spores in two species of Sphagnum[J].Journal of Hattori Botanical Laboratory,1958,19(1):71–75.
[8]ANDERSON L E,CROSBY M R.The protonema of Sphagnum meridense(Hampe)C.Muell[J].The Bryologist,1965,68(1):47–54.
[9]SIMON E.Axenic cultures of Sphagnum on solid substrates[M]//Glime J M.Methods in Bryology.Nichinan:Hattori Botanical Laboratory,1988:35–40.
[10]RUDOLPH H,KIRCHHOFF M,GLIESMANN S.Sphagnum culture techniques[M]//Glime J M.Methods in Bryology.Nichinan:Hattori Botanical Laboratory,1988:25–34.
[11]李敏,李茜,胡晶晶,等.七种真藓科植物孢子萌发与原丝体发育研究[J].贵州师范大学学报,2010,28(4):17–20.
[12]李敏,赵建成,黄士良,等.垂蒴真藓原丝体发育特征的研究[J].西北植物学报,2005,25(2):363–367.
[13]李敏,赵建成,黄士良,等.中华缩叶藓孢子萌发与原丝体发育特征研究[J].西北植物学报,2006,26(8):1521–1525.
[14]冯璐,汤袁,卜兆君,等.长白山哈泥泥炭地持久孢子库的实验证据[J].植物研究,2013,33(2):248–251.
[15]冯璐,卜兆君,李振新,等.长白山哈泥泥炭地丘间生境泥炭藓孢子的寿命[J].生态学报,2015,35(9):2993–2997.
[16]GLIME J M.Ecophysiology of development:Spore germination[J].Bryophyte Ecology,2015,1(5):10–40.
[17]DURING H J.Longevity of spores of Funaria hygrometrica in chalk grassland soil[J].Lindbergia,1987,12(2/3):132–134.
[18]Bu Zhao Jun,Chen Xu,Rydin H,et al.Performance of four mosses in a reciprocal transplant experiment:implications for peatland succession in NE China[J].Journal of Bryology,2013,35(3):220–227.
[19]SUNDBERG S.Spore rain in relation to regional sources and beyond[J].Ecography,2013,36(36):364–373.
[20]BOATMAN D J,LARK P M.Inorganic nutrition of the protonemata of Sphagnum papillosum Lindb.,S.magellanicum Brid.and S.cuspidatum Ehrh[J].New Phytologist,1971,70(6):1053–1059.
[21]HAYWARD P M,CLYMO R S.The growth of Sphagnum:experiments on,and simulation of,some effects of light flux and water-table depth[J].Journal of Ecology,1983,71(3):845–863.
[22]SKRE O,OECHEL W C,MILLER P M.Moss leaf water content and solar radiation at the moss surface in a mature black spruce forest in central Alaska[J].Canadian Journal of Forest Research,1983,13(5):860–868.
[23]VEPRASKAS M J,FAULKNER S P.Redox chemistry of hydric soils[M]//J L Richardson,M J Vepraskas.Wetland Soils:Genesis,Hydrology,Landscapes,and Classification.Boca Raton:Lewis Publishers,2001:85–106.
[24]WIKLUND K.Phosphorus concentration and p H in decaying wood affect establishment of the red-listed moss Buxbaumia viridis[J].Canadian Journal of Botany,2003,81(6):541–549.
[25]KRUPA J.Studies on the physiology of germination of spores of Funaria hygrometrica(Sibth.).II.Dependence of germination on the intensity of white light and the role of photosynthesis in this process[J].Acta Societatis Botanicorum Poloniae,2015,34(4):687–695.
[26]DUCKETT J G,BURCH J,FLETCHER P W,et al.Pressel S in vitro cultivation of bryophytes:a review of practicalities,problems,progress and promise[J].Journal of Bryology,2004,26(1):3–20.
[27]MOGENSEN G S.The biological significance of morphological characters in bryophytes:The spore[J].Bryologist,1981,84(2):187–207.
[28]GEMMRICH A R.Keimungsinduktion durch salzionen,licht und gibberellinsaure bei sporen von Marchantia polymorpha L.[J].Flora,1976,165(2):479–480.
[29]CLYMO R S.The growth of Sphagnum:some effects of environment[J].Journal of Ecology,1974,61(3):849–869.
[30]VICHEROVA E,HAJEK M,HAJEK T.Calcium intolerance of fen mosses:Physiological evidence,effects of nutrient availability and successional drivers[J].Perspectives in Plant Ecology Evolution&Systematics,2015,17(5):347–359.
[31]BHATLA S C.Moss protonema differentiation[M].Research Studies Press,Wiley,1994:98.
[32]VALANNE N.The germination phases of moss spores and their control by light[J].Annales Botanici Fennici,1966,3(1):1–60.
[33]MACIEL A S,DA S,PORTO K C,et al.Effect of light and water availability on spore germination and protonemal growth of the Neotropical moss Thamniopsis incurva(Pilotrichaceae)[J].Cryptogamie Bryologie,2009,30(2):243–257.
[2]范庆书,赵建成,于树宏.苔藓植物孢子萌发与原丝体发育研究进展[J].植物学通报,2003,20(3):280–286.
[3]CRONBERG N.Reproductive biology of Sphagnum[J].Lindbergia,1993,17(4):69–82.
[4]SUNDBERG S,RYDIN H.Habitat requirements for establishment of Sphagnum from spores[J].Journal of Ecology,2002,90(2):268–278.
[5]BOLD H C.The prothallium of Sphagnum palustre L.[J].The Bryologist,1948,51(2):55–63.
[6]李秋萍,吴思苹,鲁蓓蓓,等.藓类植物孢子生活力的快速检测方法初探[J].西北植物学报,2013,33(10):2126–2130.
[7]NOGUCHI A.Germination of spores in two species of Sphagnum[J].Journal of Hattori Botanical Laboratory,1958,19(1):71–75.
[8]ANDERSON L E,CROSBY M R.The protonema of Sphagnum meridense(Hampe)C.Muell[J].The Bryologist,1965,68(1):47–54.
[9]SIMON E.Axenic cultures of Sphagnum on solid substrates[M]//Glime J M.Methods in Bryology.Nichinan:Hattori Botanical Laboratory,1988:35–40.
[10]RUDOLPH H,KIRCHHOFF M,GLIESMANN S.Sphagnum culture techniques[M]//Glime J M.Methods in Bryology.Nichinan:Hattori Botanical Laboratory,1988:25–34.
[11]李敏,李茜,胡晶晶,等.七种真藓科植物孢子萌发与原丝体发育研究[J].贵州师范大学学报,2010,28(4):17–20.
[12]李敏,赵建成,黄士良,等.垂蒴真藓原丝体发育特征的研究[J].西北植物学报,2005,25(2):363–367.
[13]李敏,赵建成,黄士良,等.中华缩叶藓孢子萌发与原丝体发育特征研究[J].西北植物学报,2006,26(8):1521–1525.
[14]冯璐,汤袁,卜兆君,等.长白山哈泥泥炭地持久孢子库的实验证据[J].植物研究,2013,33(2):248–251.
[15]冯璐,卜兆君,李振新,等.长白山哈泥泥炭地丘间生境泥炭藓孢子的寿命[J].生态学报,2015,35(9):2993–2997.
[16]GLIME J M.Ecophysiology of development:Spore germination[J].Bryophyte Ecology,2015,1(5):10–40.
[17]DURING H J.Longevity of spores of Funaria hygrometrica in chalk grassland soil[J].Lindbergia,1987,12(2/3):132–134.
[18]Bu Zhao Jun,Chen Xu,Rydin H,et al.Performance of four mosses in a reciprocal transplant experiment:implications for peatland succession in NE China[J].Journal of Bryology,2013,35(3):220–227.
[19]SUNDBERG S.Spore rain in relation to regional sources and beyond[J].Ecography,2013,36(36):364–373.
[20]BOATMAN D J,LARK P M.Inorganic nutrition of the protonemata of Sphagnum papillosum Lindb.,S.magellanicum Brid.and S.cuspidatum Ehrh[J].New Phytologist,1971,70(6):1053–1059.
[21]HAYWARD P M,CLYMO R S.The growth of Sphagnum:experiments on,and simulation of,some effects of light flux and water-table depth[J].Journal of Ecology,1983,71(3):845–863.
[22]SKRE O,OECHEL W C,MILLER P M.Moss leaf water content and solar radiation at the moss surface in a mature black spruce forest in central Alaska[J].Canadian Journal of Forest Research,1983,13(5):860–868.
[23]VEPRASKAS M J,FAULKNER S P.Redox chemistry of hydric soils[M]//J L Richardson,M J Vepraskas.Wetland Soils:Genesis,Hydrology,Landscapes,and Classification.Boca Raton:Lewis Publishers,2001:85–106.
[24]WIKLUND K.Phosphorus concentration and p H in decaying wood affect establishment of the red-listed moss Buxbaumia viridis[J].Canadian Journal of Botany,2003,81(6):541–549.
[25]KRUPA J.Studies on the physiology of germination of spores of Funaria hygrometrica(Sibth.).II.Dependence of germination on the intensity of white light and the role of photosynthesis in this process[J].Acta Societatis Botanicorum Poloniae,2015,34(4):687–695.
[26]DUCKETT J G,BURCH J,FLETCHER P W,et al.Pressel S in vitro cultivation of bryophytes:a review of practicalities,problems,progress and promise[J].Journal of Bryology,2004,26(1):3–20.
[27]MOGENSEN G S.The biological significance of morphological characters in bryophytes:The spore[J].Bryologist,1981,84(2):187–207.
[28]GEMMRICH A R.Keimungsinduktion durch salzionen,licht und gibberellinsaure bei sporen von Marchantia polymorpha L.[J].Flora,1976,165(2):479–480.
[29]CLYMO R S.The growth of Sphagnum:some effects of environment[J].Journal of Ecology,1974,61(3):849–869.
[30]VICHEROVA E,HAJEK M,HAJEK T.Calcium intolerance of fen mosses:Physiological evidence,effects of nutrient availability and successional drivers[J].Perspectives in Plant Ecology Evolution&Systematics,2015,17(5):347–359.
[31]BHATLA S C.Moss protonema differentiation[M].Research Studies Press,Wiley,1994:98.
[32]VALANNE N.The germination phases of moss spores and their control by light[J].Annales Botanici Fennici,1966,3(1):1–60.
[33]MACIEL A S,DA S,PORTO K C,et al.Effect of light and water availability on spore germination and protonemal growth of the Neotropical moss Thamniopsis incurva(Pilotrichaceae)[J].Cryptogamie Bryologie,2009,30(2):243–257.