黄土丘陵区6种侧蒴藓类植物营养繁殖特征
|
摘要
侧蒴藓类植物是苔藓植物中体型较大的类群,形态结构复杂,在水土保持、水源涵养等方面有巨大的应用潜力。该研究以黄土丘陵区6种常见侧蒴藓类植物为材料,采用人工气候室培养,研究了不同侧蒴藓类植物营养繁殖特征及种间差异。结果显示:(1)6种侧蒴藓类植物具有较强的繁殖能力,平均10d和17d均萌发新配子体和原丝体;营养繁殖中,6种藓新配子体萌发时间早于原丝体,其中青藓(Brachythecyum albicans)的新配子体萌发最早,镰叶灰藓(Hypnum bambergeri)萌发最晚。(2)6种藓新配子体的发枝长度、发枝数量在培养过程中均呈先增加后平稳的趋势,发枝长度在萌发后第21天均趋于平稳,发枝数量在第28天时均趋于平稳;藓盖度除柳叶藓(Amblystegium serpens)在培养结束前继续增加外其余藓种在生长42d后趋于平稳,其中青藓的发枝长度、发枝数量、盖度增幅最明显,分别较萌发前增加4.73mm、1.53条、46.22%。(3)6种藓盖度、新配子体发枝长度及发枝数量的增长速率差异显著,其中青藓3个指标的增长速率均显著高于其他藓种(P<0.05),分别为0.97%·d-1、0.24mm·d-1和0.101条·d-1。发枝长度增加速率最慢的是鳞叶藓(Taxiphyllum taxirameums),为0.17mm·d-1、发枝数量增加速率最慢的是镰叶灰藓,为0.041条·d-1、盖度增加速率最慢的是荫地绢藓(Entodon caliginosus),为0.46%·d-1。结果表明,黄土丘陵区6种常见侧蒴藓类植物通过人工培养均可进行营养繁殖,但营养繁殖存在种间差异,其中青藓的营养繁殖能力优势显著。本研究明确了侧蒴藓类植物营养繁殖特征及人工培养的可能性,可为该藓类植物在水土保持及生态恢复中的应用提供科学依据。
Pleurocarpous mosses,agroup of bryophytes with large size,have complex morphological structure and play important roles in soil and water conservation.In this study,vegetative propagation of six species of pleurocarpous mosses that collected from the typical Hilly Loess Plateau region were investigatedby artificial cultivation experiment,so as to determine their characteristics and inter-specific differences.The results showed that:(1)the six species of moss exhibited strong regenerative capacities.The germination times of the new gametophyte and protonema averaged out to 10 th and 17 th day of cultivation,respectively.Furthermore,the germination time of gametophyte of the six mosses was earlier than that of proteome.Among the six mosses,Brachyhecyum albicans was the first one that germinated gametophyte and Hypnum bambergeri was the last one.(2)During cultivation period,the branch length,branch number and coverage of the six mosses showed an increasing trend along with the increasing of cultivation time,and stabilized,thereafter.The branch length and number of the six mosses tended to be stable in the21 st and 28 th day after germination,respectively.Coverage of the six mosses,except Amblystegium serpens,whose coverage continued to increase,began to stabilize after 42-day cultivation.B.albicans showed the largest increase of branch length,branch number and coverage,which were 4.73 mm,1.53 and46.22%,respectively.(3)The average rate of coverage,branch length and branch number showed significant differences between moss species.The three indices of B.albicans are 0.97%·d-1,0.24 mm·d-1 and 0.101 shoot·d-1,respectively.They are significantly higher than those of other species(P<0.05).The slowest rate of branch length,branch number and coverage were observed in Taxiphyllum taxirameums(0.17 mm·d-1),Hypnum bambergeri(0.041 shoot·d-1)and Entodon caliginosus(0.46%·d-1).The results showed a strong reproductive and artificial cultivation capacity of the six species of common pleurocarpous moss from Hilly Loess Plateau region.Further,B.albicans showed an advantage over other species.This is as earlier report on the reproductive and cultivation capacity on the pleurocarpous mosses in the region,which may provide support for the application of such moss species in soil and water conservation and ecological restoration.
Pleurocarpous mosses,agroup of bryophytes with large size,have complex morphological structure and play important roles in soil and water conservation.In this study,vegetative propagation of six species of pleurocarpous mosses that collected from the typical Hilly Loess Plateau region were investigatedby artificial cultivation experiment,so as to determine their characteristics and inter-specific differences.The results showed that:(1)the six species of moss exhibited strong regenerative capacities.The germination times of the new gametophyte and protonema averaged out to 10 th and 17 th day of cultivation,respectively.Furthermore,the germination time of gametophyte of the six mosses was earlier than that of proteome.Among the six mosses,Brachyhecyum albicans was the first one that germinated gametophyte and Hypnum bambergeri was the last one.(2)During cultivation period,the branch length,branch number and coverage of the six mosses showed an increasing trend along with the increasing of cultivation time,and stabilized,thereafter.The branch length and number of the six mosses tended to be stable in the21 st and 28 th day after germination,respectively.Coverage of the six mosses,except Amblystegium serpens,whose coverage continued to increase,began to stabilize after 42-day cultivation.B.albicans showed the largest increase of branch length,branch number and coverage,which were 4.73 mm,1.53 and46.22%,respectively.(3)The average rate of coverage,branch length and branch number showed significant differences between moss species.The three indices of B.albicans are 0.97%·d-1,0.24 mm·d-1 and 0.101 shoot·d-1,respectively.They are significantly higher than those of other species(P<0.05).The slowest rate of branch length,branch number and coverage were observed in Taxiphyllum taxirameums(0.17 mm·d-1),Hypnum bambergeri(0.041 shoot·d-1)and Entodon caliginosus(0.46%·d-1).The results showed a strong reproductive and artificial cultivation capacity of the six species of common pleurocarpous moss from Hilly Loess Plateau region.Further,B.albicans showed an advantage over other species.This is as earlier report on the reproductive and cultivation capacity on the pleurocarpous mosses in the region,which may provide support for the application of such moss species in soil and water conservation and ecological restoration.
引文
[1]李秀芹.河北省侧蒴藓类植物研究[D].石家庄:河北师范大学,2002.
[2]吴玉环,黄国宏,高谦,等.苔藓植物对环境变化的响应及适应性研究进展[J].应用生态学报,2001,12(6):943-946.WU Y H,HUANG G H,GAO Q,et al.Research advance in response and adaptation of bryophytes to environmental change[J].Chinese Journal of Applied Ecology,2001,12(6):943-946.
[3]白学良,赵连梅,孙维,等.贺兰山苔藓植物物种多样性、生物量及生态学作用的研究[J].内蒙古大学学报(自然科学版),1998,29(1):118-124.BAI X L,ZHAO L M,SUN W,et al.A preliminary study on the species diversity,phytomass and ecological effect of bryophytes in Helan Mountain,China[J].Journal of Inner Mongolia University(Natural Science Edition),1998,29(1):118-124.
[4]郭水良,谢小伟,曹同.长白山苔藓植物研究进展与展望[J].吉林林业科技,2002,31(5):1-10.GUO S L,XIE X W,CAO T.Advance and prospect of researches on bryophytes in Changbai Mountain of northeast China[J].Jilin Forestry Science and Technology,2002,31(5):1-10.
[5]陈奇伯,张洪江,解明曙.森林枯落物及其苔藓层阻延径流速度研究[J].北京林业大学学报,1996,18(1):1-5.CHEN Q B,ZHANG H J,XIE M S.Study on runoff velocity retardation by forest litter and moss[J].Journal of Beijing Forestry University,1996,18(1):1-5.
[6] ZHAO Y G,QIN N Q,WEBER B,et al.Response of biological soil crusts to raindrop erosivity and underlying influences in the hilly Loess Plateau region,China[J].Biodiversity and Conservation,2014,23:1 669-1 686.
[7]吴玉环,程国栋,高谦.苔藓植物的生态功能及在植被恢复与重建中的作用[J].中国沙漠,2003,23(3):215-220.WU Y H,CHENG G D,GAO Q.Bryophyte's ecology functions and its significances in revegetation[J].Journal of Desert Research,2003,23(3):215-220.
[8] CHAPIN F S,OECHEL W C,CLEVE K V,et al.The role of mosses in the phosphorus cycling of an Alaskan black spruce forest[J].Oecologia,1987,74(2):310-315.
[9] LIETH H.Primary productivity of the major vegetation units of the world[J].Ecology Study,1975,14(2):203-214.
[10]叶吉,郝占庆,戴冠华.长白山暗针叶林苔藓植物生物量的研究[J].应用生态学报,2014,15(5):737-740.YE J,HAO Z Q,DAI G H.Bryophyte biomass in dark coniferous forest of Changbai Mountain[J].China Journal of Applied Ecology,2014,15(5):737-740.
[11]吴鹏程.苔藓植物生物学[M].北京:科学出版社,1998.
[12] BELLAND R J.Multivariate study of moss distributions in relation to environment in the Gulf of St.Lawrence region Canada[J].Canadian Journal of Botany,2005,83(3):243-263.
[13]刚永运.几种药用苔藓植物的培养及生物学特性研究[D].北京:首都师范大学,2001.
[14]周政.梵净山生物协同进化与适应世界遗产价值研究-以苔藓植物及树干附生苔藓的螨类为例[D].贵阳:贵州师范大学,2017.
[15]余韵.黄土丘陵区人工培育生物结皮对坡面水蚀的影响研究[D].南昌:江西农业大学,2014.
[16]陈彦芹,赵允格,冉茂勇.4种营养物质对藓结皮形成发育的影响[J].西北农林科技大学学报(自然科学版),2011,39(5):44-50.CHEN Y Q,ZHAO Y G,RAN M Y.Influence of 4nutrients on the development of moss crust[J].Journal of Northwest A&F University(Natural Science Edion.),2011,39(5):44-50.
[17]卜崇峰,蔡强国,张兴昌,等.黄土结皮的发育机理与侵蚀效应研究[J].土壤学报,2009,46(1):16-23.BU C F,CAI Q G,ZHANG X C,et al.Mechanism and erosion effect of development of soil crust of loess[J].Acta Pedologica Sinica.2009,46(1):16-23.
[18]白学良,王瑶,徐杰,等.沙坡头地区固定沙丘结皮层藓类植物的繁殖和生长特性研究[J].中国沙漠,2003,23(2):23-29.BAI X L,WANG Y,XU J,et al.Characteristics of reproduction and growth of mosses in the soil crust of fixed dunes in Shapotou Area[J].Journal of Desert Research,2003,23(2):23-29.
[19]杜宝明.大灰藓(Hypnum plumaeforme)的栽培和抗旱性研究[D].杭州:浙江农林大学,2011.
[20]巨斌.中国藓类植物部分类群地理分布格局及其分布区预测的研究[D].上海:上海师范大学,2014.
[21]吴鹏程,贾渝.中国苔藓植物的地理分区及分布类型[J].植物资源与环境学报,2006,15(1):1-8.WU P C,JIA Y.The regionalization and distribution types of the bryophytes in China[J].Journal of Plant Resources and Environment,2006,15(1):1-8.
[22]白学良.贺兰山苔藓植物彩图志[M].银川:宁夏人民出版社,2014:369-441.
[23]白学良.贺兰山苔藓植物[M].银川:宁夏人民出版社,2009:211-258.
[24]《中国高等植物彩色图鉴》编委会.中国高等植物彩色图鉴第一卷苔藓植物[M].北京:科学出版社,2015:174-219.
[25]张力.中国常见植物野外识别手册:苔藓册[M].北京:商务印书馆出版社,2016:196-324.
[26] SAXENA DK,HARINDER S.Uses of bryophytes[J].Resonance,2004,9(6):56-65.
[27]朱瑞良,王幼芳,熊李虎.苔藓植物研究进展(Ⅰ)——我国苔藓植物研究现状与展望[J].西北植物学报,2002,22(2):444-451.ZHU R L,WANG Y F,XIONG L H.Progress in bryological researchⅠ.Status and prospects of bryological research in China[J].Acta Botanica Boreali-Occidentalia Sinica,2002,22(2):444-451.
[28] SHAW J.A new approach to the experimental propagation of bryophytes[J].Taxon,1986,35:671-675.
[29]田桂泉,白学良,徐杰,等.腾格里沙漠固定沙丘藓类植物结皮层的自然恢复及人工培养试验研究[J].植物生态学报,2005,29(1):164-169.TIAN G Q,BAI X L,XU J,et al.Experimental studies on natural regeneration and artificial cultures of moss crusts on fixed dunes in the Tengger Desert[J].Chinese Journal of Plant Ecology,2005,29(1):164-169.
[30]杨永胜.黄土高原苔藓结皮的快速培育及其对逆境的生理响应研究[D].西安:中国科学院教育部水土保持与生态环境研究中心,2015.
[31] SCHONBECK M W,BEWLEY J D.Responses of the moss Tortula ruralis to desiccation treatments.II.Variations in desiccation tolerance[J].Canadian Journal of Botany,1981,59(59):2 707-2 712.
[32]赵建成.绿色流苏藓的形态特征与地理分布[J].干旱区研究,1993,10(1):49-52.ZHAO J C.A preliminary study on the morphological characters and geographical distribution of Grossidium chlornotos(Brid).Limpr[J].Arid Zone Research,1993,10(1):49-52.
[33]田桂泉,白学良,徐杰,等.固定沙丘生物结皮层藓类植物形态结构及其适应性研究[J].河套大学学报,2009,25(2):249-255.
[34] RINCON E,GRIME J P,RINCON E.An analysis of seasonal patterns of bryophyte growth in a natural habitat[J].Journal of Ecology,1989,77(1):447-455.
[35] PROCTORM C F.Physiological ecology:Water relation,light and temperature responses,carbon balance[M]//In A.J E Smith(ed)Bryophyte Ecology.London:Chapman and Hall,1982,333-382.
[36] COTT G A W.Desert bryophytes[C]//In:Smith A J E(ed)Bryophyte Ecology.Chapman and Hall,London,1982,105-122.
[37] MCCALL K K,MARTIN C E.Chlorophyll concentrations and photosynthesis in three forest understory mosses in northeastern Kansas[J].Bryologist,1991,94(1):25-29.
[38]张元明,曹同,潘伯荣.干旱与半干旱地区苔藓植物生态学研究综述[J].生态学报,2002,22(7):1 129-1 134.ZHANG Y M,CAO T,PAN B R.A review on the studies of bryophyte ecology in arid and semi-arid areas[J].Acta Ecologica Sinica,2002,22(7):1 129-1 134.
[39]吴鹏程.苔藓植物生物学[M].北京:科学出版社,1998:23-148.
[2]吴玉环,黄国宏,高谦,等.苔藓植物对环境变化的响应及适应性研究进展[J].应用生态学报,2001,12(6):943-946.WU Y H,HUANG G H,GAO Q,et al.Research advance in response and adaptation of bryophytes to environmental change[J].Chinese Journal of Applied Ecology,2001,12(6):943-946.
[3]白学良,赵连梅,孙维,等.贺兰山苔藓植物物种多样性、生物量及生态学作用的研究[J].内蒙古大学学报(自然科学版),1998,29(1):118-124.BAI X L,ZHAO L M,SUN W,et al.A preliminary study on the species diversity,phytomass and ecological effect of bryophytes in Helan Mountain,China[J].Journal of Inner Mongolia University(Natural Science Edition),1998,29(1):118-124.
[4]郭水良,谢小伟,曹同.长白山苔藓植物研究进展与展望[J].吉林林业科技,2002,31(5):1-10.GUO S L,XIE X W,CAO T.Advance and prospect of researches on bryophytes in Changbai Mountain of northeast China[J].Jilin Forestry Science and Technology,2002,31(5):1-10.
[5]陈奇伯,张洪江,解明曙.森林枯落物及其苔藓层阻延径流速度研究[J].北京林业大学学报,1996,18(1):1-5.CHEN Q B,ZHANG H J,XIE M S.Study on runoff velocity retardation by forest litter and moss[J].Journal of Beijing Forestry University,1996,18(1):1-5.
[6] ZHAO Y G,QIN N Q,WEBER B,et al.Response of biological soil crusts to raindrop erosivity and underlying influences in the hilly Loess Plateau region,China[J].Biodiversity and Conservation,2014,23:1 669-1 686.
[7]吴玉环,程国栋,高谦.苔藓植物的生态功能及在植被恢复与重建中的作用[J].中国沙漠,2003,23(3):215-220.WU Y H,CHENG G D,GAO Q.Bryophyte's ecology functions and its significances in revegetation[J].Journal of Desert Research,2003,23(3):215-220.
[8] CHAPIN F S,OECHEL W C,CLEVE K V,et al.The role of mosses in the phosphorus cycling of an Alaskan black spruce forest[J].Oecologia,1987,74(2):310-315.
[9] LIETH H.Primary productivity of the major vegetation units of the world[J].Ecology Study,1975,14(2):203-214.
[10]叶吉,郝占庆,戴冠华.长白山暗针叶林苔藓植物生物量的研究[J].应用生态学报,2014,15(5):737-740.YE J,HAO Z Q,DAI G H.Bryophyte biomass in dark coniferous forest of Changbai Mountain[J].China Journal of Applied Ecology,2014,15(5):737-740.
[11]吴鹏程.苔藓植物生物学[M].北京:科学出版社,1998.
[12] BELLAND R J.Multivariate study of moss distributions in relation to environment in the Gulf of St.Lawrence region Canada[J].Canadian Journal of Botany,2005,83(3):243-263.
[13]刚永运.几种药用苔藓植物的培养及生物学特性研究[D].北京:首都师范大学,2001.
[14]周政.梵净山生物协同进化与适应世界遗产价值研究-以苔藓植物及树干附生苔藓的螨类为例[D].贵阳:贵州师范大学,2017.
[15]余韵.黄土丘陵区人工培育生物结皮对坡面水蚀的影响研究[D].南昌:江西农业大学,2014.
[16]陈彦芹,赵允格,冉茂勇.4种营养物质对藓结皮形成发育的影响[J].西北农林科技大学学报(自然科学版),2011,39(5):44-50.CHEN Y Q,ZHAO Y G,RAN M Y.Influence of 4nutrients on the development of moss crust[J].Journal of Northwest A&F University(Natural Science Edion.),2011,39(5):44-50.
[17]卜崇峰,蔡强国,张兴昌,等.黄土结皮的发育机理与侵蚀效应研究[J].土壤学报,2009,46(1):16-23.BU C F,CAI Q G,ZHANG X C,et al.Mechanism and erosion effect of development of soil crust of loess[J].Acta Pedologica Sinica.2009,46(1):16-23.
[18]白学良,王瑶,徐杰,等.沙坡头地区固定沙丘结皮层藓类植物的繁殖和生长特性研究[J].中国沙漠,2003,23(2):23-29.BAI X L,WANG Y,XU J,et al.Characteristics of reproduction and growth of mosses in the soil crust of fixed dunes in Shapotou Area[J].Journal of Desert Research,2003,23(2):23-29.
[19]杜宝明.大灰藓(Hypnum plumaeforme)的栽培和抗旱性研究[D].杭州:浙江农林大学,2011.
[20]巨斌.中国藓类植物部分类群地理分布格局及其分布区预测的研究[D].上海:上海师范大学,2014.
[21]吴鹏程,贾渝.中国苔藓植物的地理分区及分布类型[J].植物资源与环境学报,2006,15(1):1-8.WU P C,JIA Y.The regionalization and distribution types of the bryophytes in China[J].Journal of Plant Resources and Environment,2006,15(1):1-8.
[22]白学良.贺兰山苔藓植物彩图志[M].银川:宁夏人民出版社,2014:369-441.
[23]白学良.贺兰山苔藓植物[M].银川:宁夏人民出版社,2009:211-258.
[24]《中国高等植物彩色图鉴》编委会.中国高等植物彩色图鉴第一卷苔藓植物[M].北京:科学出版社,2015:174-219.
[25]张力.中国常见植物野外识别手册:苔藓册[M].北京:商务印书馆出版社,2016:196-324.
[26] SAXENA DK,HARINDER S.Uses of bryophytes[J].Resonance,2004,9(6):56-65.
[27]朱瑞良,王幼芳,熊李虎.苔藓植物研究进展(Ⅰ)——我国苔藓植物研究现状与展望[J].西北植物学报,2002,22(2):444-451.ZHU R L,WANG Y F,XIONG L H.Progress in bryological researchⅠ.Status and prospects of bryological research in China[J].Acta Botanica Boreali-Occidentalia Sinica,2002,22(2):444-451.
[28] SHAW J.A new approach to the experimental propagation of bryophytes[J].Taxon,1986,35:671-675.
[29]田桂泉,白学良,徐杰,等.腾格里沙漠固定沙丘藓类植物结皮层的自然恢复及人工培养试验研究[J].植物生态学报,2005,29(1):164-169.TIAN G Q,BAI X L,XU J,et al.Experimental studies on natural regeneration and artificial cultures of moss crusts on fixed dunes in the Tengger Desert[J].Chinese Journal of Plant Ecology,2005,29(1):164-169.
[30]杨永胜.黄土高原苔藓结皮的快速培育及其对逆境的生理响应研究[D].西安:中国科学院教育部水土保持与生态环境研究中心,2015.
[31] SCHONBECK M W,BEWLEY J D.Responses of the moss Tortula ruralis to desiccation treatments.II.Variations in desiccation tolerance[J].Canadian Journal of Botany,1981,59(59):2 707-2 712.
[32]赵建成.绿色流苏藓的形态特征与地理分布[J].干旱区研究,1993,10(1):49-52.ZHAO J C.A preliminary study on the morphological characters and geographical distribution of Grossidium chlornotos(Brid).Limpr[J].Arid Zone Research,1993,10(1):49-52.
[33]田桂泉,白学良,徐杰,等.固定沙丘生物结皮层藓类植物形态结构及其适应性研究[J].河套大学学报,2009,25(2):249-255.
[34] RINCON E,GRIME J P,RINCON E.An analysis of seasonal patterns of bryophyte growth in a natural habitat[J].Journal of Ecology,1989,77(1):447-455.
[35] PROCTORM C F.Physiological ecology:Water relation,light and temperature responses,carbon balance[M]//In A.J E Smith(ed)Bryophyte Ecology.London:Chapman and Hall,1982,333-382.
[36] COTT G A W.Desert bryophytes[C]//In:Smith A J E(ed)Bryophyte Ecology.Chapman and Hall,London,1982,105-122.
[37] MCCALL K K,MARTIN C E.Chlorophyll concentrations and photosynthesis in three forest understory mosses in northeastern Kansas[J].Bryologist,1991,94(1):25-29.
[38]张元明,曹同,潘伯荣.干旱与半干旱地区苔藓植物生态学研究综述[J].生态学报,2002,22(7):1 129-1 134.ZHANG Y M,CAO T,PAN B R.A review on the studies of bryophyte ecology in arid and semi-arid areas[J].Acta Ecologica Sinica,2002,22(7):1 129-1 134.
[39]吴鹏程.苔藓植物生物学[M].北京:科学出版社,1998:23-148.