水深梯度对钝脊眼子菜生长和繁殖的影响
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摘要
水深是影响湿地植物生长和分布的一个重要限制性因子,该研究以具有典型异型叶性的钝脊眼子菜(Potamogeton octandrus)为对象,通过分析浅水处理(10 cm和30 cm)和深水处理(50 cm和70 cm) 4个水深梯度下幼苗生长、生物量及繁殖策略等,探讨钝脊眼子菜在不同水深条件下的适应机制和表型可塑性。结果表明,钝脊眼子菜植株到达水面后出现异型叶,相对生长率显著降低,且与水深梯度呈正相关。钝脊眼子菜的株高随着水深的增加呈现爆发式的增长,10 cm水深的总茎长显著低于其他水深处理。水深对节间数也有显著性影响,其中,30 cm组处理节间数最多;而深水处理组的节间长和生物量均显著高于浅水处理组。分蘖数在4组处理之间均表现出显著性差异,随着水深的增加呈现显著性递减。生物量和地上生物量分配则随着水深增加而明显增加。水深处理对有性生殖指标有显著性影响,水深的增加抑制其有性繁殖。其中,10 cm条件下无花序形成,50 cm水深下的花粉量、P/O比和花序数显著高于其他处理组,且深水处理的结实数和结实率均显著高于30 cm组。这表明钝脊眼子菜可通过调整形态可塑性和生物量分配,并采取不同的繁殖策略,以达到对水深的最佳适应,其中最适水深生长范围在50 cm左右。
Water depth is an important limiting factor affecting the growth and distribution of wetland plants. In this study,Potamogeton octandrus with typical heterophylly was selected as the research object. To investigate the adaptive mechanism and phenotypic plasticity of P. octandrus under different water depth conditions,the seedling growth,biomass,and reproduction strategies under the four water depth gradients of shallow water treatments( 10 cm and 30 cm)and deep water treatments( 50 cm and 70 cm) were studied. The results showed that heterophyllous leaves appeared after the plants transferred from underwater to aerial condition,and the relative growth rate decreased significantly,which was also positively correlated with the water depth gradient. The shoot height showed an explosive growth with the increase of water depth,and the stem length in 10 cm water depth was significantly lower than those in other water depth treatments. The water depth also had a significant effect on the number of internodes,among which the number of internodes was the most in the 30 cm treatment,while the internode length and biomass in the deep water treatments were significantly higher than those in the shallow water treatments. The number of branch showed significant differences among the four treatments,and showed a significant decrease with the increase of water depth. While the biomass and aboveground biomass allocation increased significantly with the increase of water depth. Water depth treatment had a significant effect on sexual reproduction indexes,and the increase of water depth inhibited the sexual reproduction. No inflorescence was formed under 10 cm treatment. The pollen production,P/O ratio and the number of inflorescence in 50 cm water depth were significantly higher than those in other treatments,moreover,the seed number and seed set of the deep water treatments were significantly higher than those of the 30 cm group. Comprehensive studies indicate that P. octandrus can be adapted to the water depth by adjusting the morphological plasticity and biomass allocation to adopt different reproduction strategies,and the optimum water depth range is about 50 cm.
Water depth is an important limiting factor affecting the growth and distribution of wetland plants. In this study,Potamogeton octandrus with typical heterophylly was selected as the research object. To investigate the adaptive mechanism and phenotypic plasticity of P. octandrus under different water depth conditions,the seedling growth,biomass,and reproduction strategies under the four water depth gradients of shallow water treatments( 10 cm and 30 cm)and deep water treatments( 50 cm and 70 cm) were studied. The results showed that heterophyllous leaves appeared after the plants transferred from underwater to aerial condition,and the relative growth rate decreased significantly,which was also positively correlated with the water depth gradient. The shoot height showed an explosive growth with the increase of water depth,and the stem length in 10 cm water depth was significantly lower than those in other water depth treatments. The water depth also had a significant effect on the number of internodes,among which the number of internodes was the most in the 30 cm treatment,while the internode length and biomass in the deep water treatments were significantly higher than those in the shallow water treatments. The number of branch showed significant differences among the four treatments,and showed a significant decrease with the increase of water depth. While the biomass and aboveground biomass allocation increased significantly with the increase of water depth. Water depth treatment had a significant effect on sexual reproduction indexes,and the increase of water depth inhibited the sexual reproduction. No inflorescence was formed under 10 cm treatment. The pollen production,P/O ratio and the number of inflorescence in 50 cm water depth were significantly higher than those in other treatments,moreover,the seed number and seed set of the deep water treatments were significantly higher than those of the 30 cm group. Comprehensive studies indicate that P. octandrus can be adapted to the water depth by adjusting the morphological plasticity and biomass allocation to adopt different reproduction strategies,and the optimum water depth range is about 50 cm.
引文
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CHEN JR,WANG QL,LI M,et al.,2011.Morphological responses of Carex argyi to water fluctuation in Swan Islet wetland[J].Plant Sci J,29(4):474-479.[陈静蕊,王秋林,黎明,等,2011.红穗苔草对天鹅洲湿地淹水时间变化的形态学响应[J].植物科学学报,29(4):474-479.]
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CHRITENSEN HF,JENSEN KS,1995.Comparative kinetics of photosynthesis in floating and submerged Potamogeton leaves[J].Aquat Bot,51(1):121-134.
CLEVERING OA,HUNDSCHEID M,1998.Plastic and nonplastic variation in growth of newly established clones of Scirpus maritimus L.(Bolboschoenus)grown at different water depths[J].Aquat Bot,62(1):1-17.
EEWARDS AL,LEE DW,RICHARDS JH,2003.Responses to a fluctuating environment:Effects of water depth on growth and biomass allocation in Eleocharis cellulosa Torr.(Cyperaceae)[J].Can J Bot,81(9):964-975.
GALLOWAY LF,1995.Response to natural environmental heterogeneity:Maternal effects and selection on life-history characters and plasticities in Mimulus guttatus[J].Evolution,49(6):1095-1107.
GRACE JB,1989.Effects of water depth on Typha latifolia and Typha domingensis[J].Am J Bot,76(5):762-768.
HE W,WANG GX,YANG WB,et al.,2009.Growth response of Potamogeton crispus to water depth gradient[J].Chin JEcol,28(7):1224-1228.[何伟,王国祥,杨文斌,等,2009.水深梯度对菹草生长的影响[J].生态学杂志,28(7):1224-1228.]
HILLS JM,MURPHY KJ,PULFORD ID,et al.,1994.Amethod for classifying European riverine wetland ecosystems using functional vegetation groups[J].Funct Ecol,8(2):242-252.
HUTCHINGS MJ,KROON HD,1994.Forging in plants-the role of morphological plasticity in resource acquisition[J].Advan Ecol Res,25:159-238.
KENDE H,VAN KE,CHO HT,1998.Deepwater Rice:Amodel plant to study stem elongation[J].Plant Physiol,118(4):1105-1110.
LI YF,2015.Effects of water levels on the growth and reproductive characteristics of dominant plants in the Dongting Lake wetlands[D].Changsha:Central South University of Forestry and Technology.[李亚芳,2015.洞庭湖湿地克隆植物繁殖分配对水位的响应[D].长沙:中南林业科技大学.]
LIU Y,FU WL,CAO Y,et al.,2017.Study on the functional traits of submerged macrophytes[J].Plant Sci J,35(3):444-451.[刘洋,付文龙,操瑜,等,2017.沉水植物功能性状研究的思考[J].植物科学学报,35(3):444-451.]
LUO WB,XIE YH,SONG FB,2007.Survival strategies of wetland plants in flooding environments[J].Chin J Ecol,26(9):1478-1485.[罗文泊,谢永宏,宋凤斌,2007.洪水条件下湿地植物的生存策略[J].生态学杂志,26(9):1478-1485.]
MINORSKY PV,2003.Heterophylly in aquatic plants[J].Plant Physiol,133:1671-1672.
MONY C,MERCIER E,BONIS A,et al.,2010.Reproductive strategies may explain plant tolerance to inundation:A mesocosm experiment using six marsh species[J].Aquat Bot,92(2):99-104.
PHILBRICK CT,LES DH,1996.Evolution of aquatic angiosperm reproductive systems[J].Bioscience,46(11):813-826.
SLATER J,1999.Growth and resource allocation in response to flooding in the emergent sedge Bolboschoenus medianus[J].Aquat Bot,63(2):145-160.
STRAND JA,SEB W,2001.Morphological plastic responses to water depth and wave exposure in an aquatic plant(Myriophyllum spicatum)[J].J Ecol,89(2):166-175.
SULTAN SE,2000.Phenotypic plasticity for plant development,function and life history[J].Trends Plant Sci,5(12):537-542.
VAN DRUNEN WE,DORKEN ME,2012.Trade-offs between clonal and sexual reproduction in Sagittaria latifolia(Alismataceae)scale up to affect the fitness of entire clones[J].New Phytol,196(2):606-616.
VOESENEK LACJ,BLOM CWPM,1989.Growth responses of Rumex species in relation to submergence and ethylene[J].Plant Cell Environ,12(4):433-439.
VRETARE V,WEISNER SEB,2000.Influence of pressurized ventilation on performance of an emergent macrophyte(Phragmites australis)[J].J Ecol,88(6):978-987.
WANG HY,CHEN JK,ZHOU J,1999.Influence of water level gradient on plant growth,reproduction and biomass allocation of wetland plant species[J].Acta Phytoecol Sin,23(3):78-83.[王海洋,陈家宽,周进,1999.水位梯度对湿地植物生长繁殖和生物量分配的影响[J].植物生态学报,23(3):78-83.]
WANG QL,CHEN JR,LIU F,et al.,2014.Morphological changes and resource allocation of Zizania latifolia(Griseb.)Stapf in response to different submergence depth and duration[J].Flora,209(5-6):279-284.
WELLS CL,PIGLIUCCI M,2000.Adaptive phenotypic plasticity:The case of heterophylly in aquatic plants[J].Perspect Plant Ecol Evol Syst,3(1):1-18.
WU HY,CAO Y,GUO ZC,et al.,2017.Effects of waterlogging stress on growth and physiological characteristics of Phalaris arundinacea seedlings[J].Guihaia,37(9):1161-1167.[吴海英,曹昀,国志昌,等,2017.淹水胁迫对虉草幼苗生长和生理的影响[J].广西植物,37(9):1161-1167.]
YAN DL,JIN SH,XIA GH,et al.,2013.Response of clonal growth and biomass allocation of 3 aquatic plant species to different inundation levels[J].J SW For Coll,33(2):10-14.[闫道良,金水虎,夏国华,等,2013.3种湿地植物克隆生长及其生物量对不同水淹的响应[J].西南林业大学学报,33(2):10-14.]
CHEN JR,WANG QL,LI M,et al.,2011.Morphological responses of Carex argyi to water fluctuation in Swan Islet wetland[J].Plant Sci J,29(4):474-479.[陈静蕊,王秋林,黎明,等,2011.红穗苔草对天鹅洲湿地淹水时间变化的形态学响应[J].植物科学学报,29(4):474-479.]
CHEN ZY,WANG GX,WU XD,et al.,2011.Ecological adaptability of Potamogeton crispus under different water depths[J].J Lake Sci,23(6):942-948.[陈正勇,王国祥,吴晓东,等,2011.不同水深条件下菹草(Potamogeton crispus)的适应对策[J].湖泊科学,23(6):942-948.]
CHRITENSEN HF,JENSEN KS,1995.Comparative kinetics of photosynthesis in floating and submerged Potamogeton leaves[J].Aquat Bot,51(1):121-134.
CLEVERING OA,HUNDSCHEID M,1998.Plastic and nonplastic variation in growth of newly established clones of Scirpus maritimus L.(Bolboschoenus)grown at different water depths[J].Aquat Bot,62(1):1-17.
EEWARDS AL,LEE DW,RICHARDS JH,2003.Responses to a fluctuating environment:Effects of water depth on growth and biomass allocation in Eleocharis cellulosa Torr.(Cyperaceae)[J].Can J Bot,81(9):964-975.
GALLOWAY LF,1995.Response to natural environmental heterogeneity:Maternal effects and selection on life-history characters and plasticities in Mimulus guttatus[J].Evolution,49(6):1095-1107.
GRACE JB,1989.Effects of water depth on Typha latifolia and Typha domingensis[J].Am J Bot,76(5):762-768.
HE W,WANG GX,YANG WB,et al.,2009.Growth response of Potamogeton crispus to water depth gradient[J].Chin JEcol,28(7):1224-1228.[何伟,王国祥,杨文斌,等,2009.水深梯度对菹草生长的影响[J].生态学杂志,28(7):1224-1228.]
HILLS JM,MURPHY KJ,PULFORD ID,et al.,1994.Amethod for classifying European riverine wetland ecosystems using functional vegetation groups[J].Funct Ecol,8(2):242-252.
HUTCHINGS MJ,KROON HD,1994.Forging in plants-the role of morphological plasticity in resource acquisition[J].Advan Ecol Res,25:159-238.
KENDE H,VAN KE,CHO HT,1998.Deepwater Rice:Amodel plant to study stem elongation[J].Plant Physiol,118(4):1105-1110.
LI YF,2015.Effects of water levels on the growth and reproductive characteristics of dominant plants in the Dongting Lake wetlands[D].Changsha:Central South University of Forestry and Technology.[李亚芳,2015.洞庭湖湿地克隆植物繁殖分配对水位的响应[D].长沙:中南林业科技大学.]
LIU Y,FU WL,CAO Y,et al.,2017.Study on the functional traits of submerged macrophytes[J].Plant Sci J,35(3):444-451.[刘洋,付文龙,操瑜,等,2017.沉水植物功能性状研究的思考[J].植物科学学报,35(3):444-451.]
LUO WB,XIE YH,SONG FB,2007.Survival strategies of wetland plants in flooding environments[J].Chin J Ecol,26(9):1478-1485.[罗文泊,谢永宏,宋凤斌,2007.洪水条件下湿地植物的生存策略[J].生态学杂志,26(9):1478-1485.]
MINORSKY PV,2003.Heterophylly in aquatic plants[J].Plant Physiol,133:1671-1672.
MONY C,MERCIER E,BONIS A,et al.,2010.Reproductive strategies may explain plant tolerance to inundation:A mesocosm experiment using six marsh species[J].Aquat Bot,92(2):99-104.
PHILBRICK CT,LES DH,1996.Evolution of aquatic angiosperm reproductive systems[J].Bioscience,46(11):813-826.
SLATER J,1999.Growth and resource allocation in response to flooding in the emergent sedge Bolboschoenus medianus[J].Aquat Bot,63(2):145-160.
STRAND JA,SEB W,2001.Morphological plastic responses to water depth and wave exposure in an aquatic plant(Myriophyllum spicatum)[J].J Ecol,89(2):166-175.
SULTAN SE,2000.Phenotypic plasticity for plant development,function and life history[J].Trends Plant Sci,5(12):537-542.
VAN DRUNEN WE,DORKEN ME,2012.Trade-offs between clonal and sexual reproduction in Sagittaria latifolia(Alismataceae)scale up to affect the fitness of entire clones[J].New Phytol,196(2):606-616.
VOESENEK LACJ,BLOM CWPM,1989.Growth responses of Rumex species in relation to submergence and ethylene[J].Plant Cell Environ,12(4):433-439.
VRETARE V,WEISNER SEB,2000.Influence of pressurized ventilation on performance of an emergent macrophyte(Phragmites australis)[J].J Ecol,88(6):978-987.
WANG HY,CHEN JK,ZHOU J,1999.Influence of water level gradient on plant growth,reproduction and biomass allocation of wetland plant species[J].Acta Phytoecol Sin,23(3):78-83.[王海洋,陈家宽,周进,1999.水位梯度对湿地植物生长繁殖和生物量分配的影响[J].植物生态学报,23(3):78-83.]
WANG QL,CHEN JR,LIU F,et al.,2014.Morphological changes and resource allocation of Zizania latifolia(Griseb.)Stapf in response to different submergence depth and duration[J].Flora,209(5-6):279-284.
WELLS CL,PIGLIUCCI M,2000.Adaptive phenotypic plasticity:The case of heterophylly in aquatic plants[J].Perspect Plant Ecol Evol Syst,3(1):1-18.
WU HY,CAO Y,GUO ZC,et al.,2017.Effects of waterlogging stress on growth and physiological characteristics of Phalaris arundinacea seedlings[J].Guihaia,37(9):1161-1167.[吴海英,曹昀,国志昌,等,2017.淹水胁迫对虉草幼苗生长和生理的影响[J].广西植物,37(9):1161-1167.]
YAN DL,JIN SH,XIA GH,et al.,2013.Response of clonal growth and biomass allocation of 3 aquatic plant species to different inundation levels[J].J SW For Coll,33(2):10-14.[闫道良,金水虎,夏国华,等,2013.3种湿地植物克隆生长及其生物量对不同水淹的响应[J].西南林业大学学报,33(2):10-14.]