典型高寒植物有性繁殖对长期增温的响应
|
摘要
近年来,全球气候变化影响植物群落已经成为研究热点,植物有性繁殖对环境条件微小变化的响应十分敏感。为探究高寒植物采取何种生长繁殖策略以适应全球气候变暖,从而预测高寒植被群落变化趋势,本研究利用开顶式生长室(Open-Top Chambers,OTCs)装置模拟增温,探究增温对高寒植物中常见的矮生嵩草(Kobresia humilis)、青藏苔草(Carex moorcroftii),和垂穗披碱草(Elymus nutans)有性繁殖特征的影响。结果表明:1)增温下矮嵩草繁殖枝数量、有性繁殖效力极显著降低(P<0.01),青藏苔草小穗长度显著增加(P<0.05),垂穗披碱草小穗长度在增温下极显著增加(P<0.01);2)增温导致矮嵩草和垂穗披碱草种子生产能力显著降低(P<0.05),青藏苔草种子产量变化不明显(P<0.05);3)3种植物发芽势在增温下呈现显著降低的趋势(P<0.05),但发芽率无显著变化(P<0.05);4)经热图(heatmap)分析,高寒植物有性繁殖对增温响应不一致,且增温严重抑制矮嵩草有性繁殖。综上,在气候变暖下,不同植物可能表现出不同的适应策略,矮嵩草有性繁殖受到增温的抑制,且最为敏感。
The grassland ecosystem accounts for 50.9% of the Qinghai-Tibet Plateau,which is a sensitive area of climate change and an ecologically fragile zone.It is an ideal place to study the response mechanism of grassland ecosystems to climate change.The sexual reproduction stage of plants is very sensitive to the small variation in environmental conditions.The response mechanism of alpine plant growth and reproduction strategies to global warming is still unrevealed.In order to explore reproduction strategies of alpine plants to adapt to global warming and guide the prediction of the trend of alpine vegetation communities,in this study,a simulated warming experiment was set up using the Open-Top Chambers(OTCs)-type device to explore the effect of warming on the sexual reproduction characteristics of apline representative plants:Kobresia humilis,Carex moorcroftii,and Elymus nutans.The results showed that:1)The numbers of reproductive shoots and the reproductive efficiency of Kobresia humilis significantly decreased(P<0.01);The effective spikelets length of Carex moorcroftii increased significantly(P<0.05),and the length of Elymus nutans spikelets increased significantly under the warming(P<0.01).2)Simulated warming caused a significant decrease in seed production capacity of Kobresia humilis(P<0.01)and Elymus nutans(P<0.05),and there was no significant difference in seed yield of Carex moorcroftii(P>0.05).3)The germination potential of three plants significantly decreased under the increase of temperature(P<0.05),but the germination rate did not change significantly(P>0.05).4)According to the heatmap analysis,under the influence of temperature increase,each index of Kobresiahumilishad highly negative responses to the warming effect.It is expected that the grassland with Kobresia humilis as the dominant vegetation would have the highest inhibitory degree of sexual reproduction under the effect of warming.In summary,under the climate warming,different plants showed different adaptation strategies.The sexual reproduction of Kobresia humilis was inhibited,and the response to warming was the most sensitive.
The grassland ecosystem accounts for 50.9% of the Qinghai-Tibet Plateau,which is a sensitive area of climate change and an ecologically fragile zone.It is an ideal place to study the response mechanism of grassland ecosystems to climate change.The sexual reproduction stage of plants is very sensitive to the small variation in environmental conditions.The response mechanism of alpine plant growth and reproduction strategies to global warming is still unrevealed.In order to explore reproduction strategies of alpine plants to adapt to global warming and guide the prediction of the trend of alpine vegetation communities,in this study,a simulated warming experiment was set up using the Open-Top Chambers(OTCs)-type device to explore the effect of warming on the sexual reproduction characteristics of apline representative plants:Kobresia humilis,Carex moorcroftii,and Elymus nutans.The results showed that:1)The numbers of reproductive shoots and the reproductive efficiency of Kobresia humilis significantly decreased(P<0.01);The effective spikelets length of Carex moorcroftii increased significantly(P<0.05),and the length of Elymus nutans spikelets increased significantly under the warming(P<0.01).2)Simulated warming caused a significant decrease in seed production capacity of Kobresia humilis(P<0.01)and Elymus nutans(P<0.05),and there was no significant difference in seed yield of Carex moorcroftii(P>0.05).3)The germination potential of three plants significantly decreased under the increase of temperature(P<0.05),but the germination rate did not change significantly(P>0.05).4)According to the heatmap analysis,under the influence of temperature increase,each index of Kobresiahumilishad highly negative responses to the warming effect.It is expected that the grassland with Kobresia humilis as the dominant vegetation would have the highest inhibitory degree of sexual reproduction under the effect of warming.In summary,under the climate warming,different plants showed different adaptation strategies.The sexual reproduction of Kobresia humilis was inhibited,and the response to warming was the most sensitive.
引文
[1]IPCC,RWGII.Climate Change 2001:Impact,Adaptation and Vulner-ability[M].London:Cambridge University Press,2001:12-28
[2]高懋芳,邱建军.青藏高原主要自然灾害特点及分布规律研究[J].干旱区资源与环境,2011,25(08):101-106
[3]谢高地,鲁春霞,冷允法,等.青藏高原生态资产的价值评估[J].自然资源学报,2003,(02):189-196
[4]古琛,赵天启,王亚婷,等.短花针茅生长和繁殖策略对载畜率的响应[J].生态环境学报,2015,24(12):1962-1968
[5]Sherry R A,Zhou X,Gu S,et al.Divergence of reproductive phenology under climate warming[J].Proceedings of the National Academy of Sciences of the United States of America,2007,104(1):198-202
[6]李英年,薛晓娟,王建雷,等.典型高寒植物生长繁殖特征对模拟气候变化的短期响应[J].生态学杂志,2010,29(4):624-629
[7]Cleland E E,Chiariello N R,Loarie S R,et al.Diverse responses of phenology to global changes in a grassland ecosystem[J].Proceedings of the National Academy of Sciences of the United States of America,2006,103(37):13740-13744
[8]Benlloch-Gonzalez M,Sanchez-Lucas R,Bejaoui M A,et al.Global warming effects on yield and fruit maturation of olive trees growing under field conditions[J].Scientia Horticulturae,2019,249(3),162-167
[9]高福光.增温和施氮对短花针茅有性繁殖的影响[D].内蒙古:内蒙古农业大学,2010:6-7
[10]蒋小雪,金飚.气候变化对植物有性生殖影响的研究进展[J].西北植物学报,2012,32(10):2139-2150
[11]曹素珍.青藏高原高寒草甸常见植物出苗及生长繁殖对增温的响应研究[D].兰州:兰州大学,2018:4-5
[12]Klady R A,Henry G H R,Lemay V.Changes in high arctic tundra plant reproduction in response to long-term experimental warming[J].Global Change Biology,2011,17(4):1611-1624
[13]Ganjurjav H,Gao Q,Gornish E S,et al.Differential response of alpine steppe and alpine meadow to climate warming in the central Qinghai-Tibetan Plateau[J].Agricultural&Forest Meteorology,2016(223):233-240
[14]耿晓东,旭日,魏达.多梯度增温对青藏高原高寒草甸温室气体通量的影响[J].生态环境学报,2017,26(3):445-452
[15]张维,贺亚玲,吴泽昂,等.模拟增温对梭梭光合生理生态特征的影响[J].草地学报,2017,25(2):296-302
[16]吴楚,芦光新,德科加,等.人工增温条件下不同氮素形态比对重齿风毛菊气体交换及其群落生物量的影响[J].草地学报,2015,23(6):1129-1135
[17]周兴民,赵新全,周华坤,等.模拟增温效应对矮嵩草草甸影响的初步研究[J].植物生态学报,2000,24(5):547-553
[18]刘凤娇,孙玉军.林下植被生物量研究进展[J].世界林业研究,2011,24(2):53-58
[19]方精云,王襄平,沈泽昊,等.植物群落清查的主要内容、方法和技术规范[J].生物多样性,2009,17(06):533-548
[20]Panetta A M,Stanton M L,Harte J.Climate warming drives local extinction:Evidence from observation and experimentation[J].Science Advances,2018,4(2):18-19
[21]邓自发,谢晓玲,王启基,等.高寒草甸藏嵩草种群繁殖对策的研究[J].应用与环境生物学报,2001,7(04):332-334
[22]罗弦,潘远智,杨学军,等.低温层积处理对4种苔草种子休眠与萌发的影响[J].草业学报,2010,19(03):117-123
[23]Perry D A.Handbook of vigour test methods[M].3th ed.Zurich:International Seed Testing Association:1995,25(3),117-118
[24]周华坤,赵新全,周立,等.不同放牧强度对鹅绒委陵菜克隆生长特征的影响[J].西北植物学报,2006,26(5):1021-1029
[25]王志保,孙梅,刘振亚等.滇西北高原典型湿地湖滨带优势植物种子繁殖对增温的响应[J].应用生态学报,2018,29(03):696-704
[26]Shiping Wang,Changshun Wang,Jichuang Duan,et al.Timing and duration of phenological sequences of alpine plants along an elevation gradient on the Tibetan plateau[J].Agricultural and Forest Meteorology,2014,189(5):220-228
[27]Wolkovich E M,Cook B I,Allen J M,et al.Warming experiments underpredict plant phenological responses to climate change[J].Nature,2012,485(7399):494-497
[28]Jiang L L,Wang S P,Meng F D,et al.Relatively stable response of fruiting stage to warming and cooling relative to other phenological events[J].Ecology,2016,97(8),1961-1969
[29]李英年,赵亮,赵新全,等.5年模拟增温后矮嵩草草甸群落结构及生产量的变化[J].草地学报,2004(03):236-239
[30]Hedhly A,JoséI,Hormaza,et al.Global warming and sexual plant reproduction[J].Trends in Plant Science,2009,14(1):0-36
[31]Cao Y,Xiao Y,Huang H,et al.Simulated warming shifts the flowering phenology and sexual reproduction of Cardamine hirsuta under different Planting densities[J].Scientific Reports,2016,6(1):27835
[32]Leubnermetzger G.Seed dormancy and the control of germination[J].New Phytologist,2010,171(3):501-523
[33]赵玉红,魏学红,沈振西,等.模拟增温效应对西藏苔草繁殖生态的影响[J].生态环境学报,2010,19(08):1783-1788
[34]Yang Z L,Zhang Q,Su F L,et al.Daytime warming lowers community temporal stability by reducing the abundance of dominant,stable species[J].Global Change Biology,2017,23(1):154-163
[35]Alward R.D.Grassland Vegetation Changes and Nocturnal Global Warming[J].Science,1999,283(5399):229-231
[36]赵建中,彭敏,刘伟,等.矮嵩草草甸不同功能群主要植物种生长特征与地表温度的相关性分析[J].西北植物学报,2012,32(03):559-565
[37]庞晓瑜,袁秀锦,王奥,等.模拟增温和功能群去除对岷江冷杉幼苗存活和生长的影响[J].应用生态学报,2018,29(3):687-695
[2]高懋芳,邱建军.青藏高原主要自然灾害特点及分布规律研究[J].干旱区资源与环境,2011,25(08):101-106
[3]谢高地,鲁春霞,冷允法,等.青藏高原生态资产的价值评估[J].自然资源学报,2003,(02):189-196
[4]古琛,赵天启,王亚婷,等.短花针茅生长和繁殖策略对载畜率的响应[J].生态环境学报,2015,24(12):1962-1968
[5]Sherry R A,Zhou X,Gu S,et al.Divergence of reproductive phenology under climate warming[J].Proceedings of the National Academy of Sciences of the United States of America,2007,104(1):198-202
[6]李英年,薛晓娟,王建雷,等.典型高寒植物生长繁殖特征对模拟气候变化的短期响应[J].生态学杂志,2010,29(4):624-629
[7]Cleland E E,Chiariello N R,Loarie S R,et al.Diverse responses of phenology to global changes in a grassland ecosystem[J].Proceedings of the National Academy of Sciences of the United States of America,2006,103(37):13740-13744
[8]Benlloch-Gonzalez M,Sanchez-Lucas R,Bejaoui M A,et al.Global warming effects on yield and fruit maturation of olive trees growing under field conditions[J].Scientia Horticulturae,2019,249(3),162-167
[9]高福光.增温和施氮对短花针茅有性繁殖的影响[D].内蒙古:内蒙古农业大学,2010:6-7
[10]蒋小雪,金飚.气候变化对植物有性生殖影响的研究进展[J].西北植物学报,2012,32(10):2139-2150
[11]曹素珍.青藏高原高寒草甸常见植物出苗及生长繁殖对增温的响应研究[D].兰州:兰州大学,2018:4-5
[12]Klady R A,Henry G H R,Lemay V.Changes in high arctic tundra plant reproduction in response to long-term experimental warming[J].Global Change Biology,2011,17(4):1611-1624
[13]Ganjurjav H,Gao Q,Gornish E S,et al.Differential response of alpine steppe and alpine meadow to climate warming in the central Qinghai-Tibetan Plateau[J].Agricultural&Forest Meteorology,2016(223):233-240
[14]耿晓东,旭日,魏达.多梯度增温对青藏高原高寒草甸温室气体通量的影响[J].生态环境学报,2017,26(3):445-452
[15]张维,贺亚玲,吴泽昂,等.模拟增温对梭梭光合生理生态特征的影响[J].草地学报,2017,25(2):296-302
[16]吴楚,芦光新,德科加,等.人工增温条件下不同氮素形态比对重齿风毛菊气体交换及其群落生物量的影响[J].草地学报,2015,23(6):1129-1135
[17]周兴民,赵新全,周华坤,等.模拟增温效应对矮嵩草草甸影响的初步研究[J].植物生态学报,2000,24(5):547-553
[18]刘凤娇,孙玉军.林下植被生物量研究进展[J].世界林业研究,2011,24(2):53-58
[19]方精云,王襄平,沈泽昊,等.植物群落清查的主要内容、方法和技术规范[J].生物多样性,2009,17(06):533-548
[20]Panetta A M,Stanton M L,Harte J.Climate warming drives local extinction:Evidence from observation and experimentation[J].Science Advances,2018,4(2):18-19
[21]邓自发,谢晓玲,王启基,等.高寒草甸藏嵩草种群繁殖对策的研究[J].应用与环境生物学报,2001,7(04):332-334
[22]罗弦,潘远智,杨学军,等.低温层积处理对4种苔草种子休眠与萌发的影响[J].草业学报,2010,19(03):117-123
[23]Perry D A.Handbook of vigour test methods[M].3th ed.Zurich:International Seed Testing Association:1995,25(3),117-118
[24]周华坤,赵新全,周立,等.不同放牧强度对鹅绒委陵菜克隆生长特征的影响[J].西北植物学报,2006,26(5):1021-1029
[25]王志保,孙梅,刘振亚等.滇西北高原典型湿地湖滨带优势植物种子繁殖对增温的响应[J].应用生态学报,2018,29(03):696-704
[26]Shiping Wang,Changshun Wang,Jichuang Duan,et al.Timing and duration of phenological sequences of alpine plants along an elevation gradient on the Tibetan plateau[J].Agricultural and Forest Meteorology,2014,189(5):220-228
[27]Wolkovich E M,Cook B I,Allen J M,et al.Warming experiments underpredict plant phenological responses to climate change[J].Nature,2012,485(7399):494-497
[28]Jiang L L,Wang S P,Meng F D,et al.Relatively stable response of fruiting stage to warming and cooling relative to other phenological events[J].Ecology,2016,97(8),1961-1969
[29]李英年,赵亮,赵新全,等.5年模拟增温后矮嵩草草甸群落结构及生产量的变化[J].草地学报,2004(03):236-239
[30]Hedhly A,JoséI,Hormaza,et al.Global warming and sexual plant reproduction[J].Trends in Plant Science,2009,14(1):0-36
[31]Cao Y,Xiao Y,Huang H,et al.Simulated warming shifts the flowering phenology and sexual reproduction of Cardamine hirsuta under different Planting densities[J].Scientific Reports,2016,6(1):27835
[32]Leubnermetzger G.Seed dormancy and the control of germination[J].New Phytologist,2010,171(3):501-523
[33]赵玉红,魏学红,沈振西,等.模拟增温效应对西藏苔草繁殖生态的影响[J].生态环境学报,2010,19(08):1783-1788
[34]Yang Z L,Zhang Q,Su F L,et al.Daytime warming lowers community temporal stability by reducing the abundance of dominant,stable species[J].Global Change Biology,2017,23(1):154-163
[35]Alward R.D.Grassland Vegetation Changes and Nocturnal Global Warming[J].Science,1999,283(5399):229-231
[36]赵建中,彭敏,刘伟,等.矮嵩草草甸不同功能群主要植物种生长特征与地表温度的相关性分析[J].西北植物学报,2012,32(03):559-565
[37]庞晓瑜,袁秀锦,王奥,等.模拟增温和功能群去除对岷江冷杉幼苗存活和生长的影响[J].应用生态学报,2018,29(3):687-695