遮阴和干旱对荩草克隆生长和有性繁殖及权衡关系的影响
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摘要
以可建植边坡草坪的野生荩草为材料,设置遮阴(5个水平)和干旱(4个水平)交互形成的20个组合处理,测定克隆生长性状和有性繁殖数量及生物量结构,分析不同遮阴和干旱组合处理下克隆效率指数(CEI)、繁殖分配(RA)、繁殖效率指数(REI)、繁殖指数(RI)、繁殖比率(RR)的差异,研究遮阴和干旱对荩草克隆生长和有性繁殖及权衡关系的影响,结果表明:1)遮阴和干旱对克隆子株数及克隆效率指数有显著影响,中、重度干旱和≥10%遮阴显著降低了子株数(P<0.05),干旱对CEI无影响、遮阴降低了CEI,交互作用极显著降低了基株的子株数和CEI(P<0.01)。2)遮阴和干旱对母株、子株和基株的花穗数和小花数,对繁殖器官生物量(ROB)、非繁殖器官生物量(NROB)和总生物量(TB)有显著影响(P<0.05),轻、中度干旱或10%～30%遮阴或轻度遮阴和轻度干旱组合可显著增加母株、子株、基株的花穗数和小花数,轻度干旱或10%遮阴或轻度干旱和轻度遮阴组合可显著增加TB和ROB,交互作用是影响基株花穗数和小花数、TB、ROB和NROB的主要因子,干旱对TB和NROB影响次之,遮阴对ROB影响较大。3)遮阴和干旱对RA、REI、RI和RR有显著影响(P<0.05),轻中度干旱或10%遮阴可提高基株的繁殖参数,重度干旱或≥30%遮阴度降低了有性繁殖(P<0.05),轻度遮阴和轻度干旱组合比单一处理更利于有性繁殖,轻度干旱可缓解中重度遮阴对有性繁殖的影响,繁殖参数主要受交互作用影响,遮阴次之,干旱较小。4)荩草可依据遮阴和干旱胁迫程度,进行克隆生长和有性繁殖权衡,遮阴≥30%可延长克隆生长10～34d,中、重度干旱缩短克隆生长而促进有性繁殖,≥10%遮阴可缓解重度干旱对有性繁殖的影响(P<0.05)。5)20个遮阴和干旱组合处理下荩草均能同时维持两种繁殖方式,水分充足和全光照利于克隆生长,轻中度干旱和轻度遮阴单一或组合处理利于有性繁殖,遮阴和干旱交互影响增加了荩草的两种繁殖能力。
Arthraxon hispidus,is a wild grass with potential for development as a turf plant for sloping ground.In this study,20 treatments were set up by a factorial combination of five shade levels(0,10%,30%,50%,70%)and four drought levels(adequate moisture,mild drought,moderate drought and severe drought).The purpose was to study the combined effects of shade and drought on growth of A.hispidus,and identify tradeoffs between clonal growth and sexual reproduction.Various data,including clonal growth traits,number of seed-heads,and biomass structure were gathered,and differences in cloning efficiency index(CEI),reproductive allocation(RA),reproductive efficiency index(REI),reproduction index(RI)and ratio of breeding(RR)under the 20 treatments were analyzed.It was found that:1)Shade and drought stress had significant effects on clonal daughter ramet number and CEI.Moderate and severe drought and over 10%shading significantly reduced daughter ramet number.Drought had no effect on CEI and shade decreased CEI,and a shade x CEI interaction(P<0.01)was observed,expressed as a reduced daughter ramet and genet number,and reduced CEI.2)Shading and drought had significant effects on flower spike numbers and floret numbers of mother ramets,daughter ramets and genets,reproductive organ biomass(ROB),non-reproductive organ biomass(NROB)and total biomass(TB)(P<0.05).Light and moderate drought or 10%-30% shade or the combination of mild shade and mild drought significantly increased flower spike number and floret number of mother ramets,and of daughter ramets and genets.Mild drought or 10%shading or the combination stress of mild drought and light shade all significantly increased TB and ROB.Moreover,a synergistic interaction between shade and drought strongly increased flower spike number and floret number of genets,and TB,ROB and NROB.The effect of drought on TB and NROB was followed,and shading had greater influence on ROB.3)Shading and drought significantly(P<0.05)affected RA,REI,RI and RR:mild and moderate drought or 10%shading significantly increased the genet reproductive parameters,but severe drought or more than 30%shading significantly reduced reproductive parameters(P<0.05).Compared with either single stress,the combined stresses of light shading and mild drought promoted sexual reproduction.Mild drought alleviated reduction in sexual reproduction observed under moderate and severe shading.Reproductive parameters typically displayed a synergistic effect,with shading the larger effect,and the drought effect smaller.4)A.hispidus displayed an altered balance between clonal growth and sexual reproduction,depending on the degree of shading and drought.Greater than 30%shade prolonged the period of clonal growth by 10-34 days.5)Under all 20 treatment combinations of shading and drought,A.hispidus maintained both clonal and sexual reproduction concurrently.Adequate water and light promoted clonal growth.Light and moderate drought,and mild shade alone or in combination with drought were treatments that promoted sexual reproduction.
Arthraxon hispidus,is a wild grass with potential for development as a turf plant for sloping ground.In this study,20 treatments were set up by a factorial combination of five shade levels(0,10%,30%,50%,70%)and four drought levels(adequate moisture,mild drought,moderate drought and severe drought).The purpose was to study the combined effects of shade and drought on growth of A.hispidus,and identify tradeoffs between clonal growth and sexual reproduction.Various data,including clonal growth traits,number of seed-heads,and biomass structure were gathered,and differences in cloning efficiency index(CEI),reproductive allocation(RA),reproductive efficiency index(REI),reproduction index(RI)and ratio of breeding(RR)under the 20 treatments were analyzed.It was found that:1)Shade and drought stress had significant effects on clonal daughter ramet number and CEI.Moderate and severe drought and over 10%shading significantly reduced daughter ramet number.Drought had no effect on CEI and shade decreased CEI,and a shade x CEI interaction(P<0.01)was observed,expressed as a reduced daughter ramet and genet number,and reduced CEI.2)Shading and drought had significant effects on flower spike numbers and floret numbers of mother ramets,daughter ramets and genets,reproductive organ biomass(ROB),non-reproductive organ biomass(NROB)and total biomass(TB)(P<0.05).Light and moderate drought or 10%-30% shade or the combination of mild shade and mild drought significantly increased flower spike number and floret number of mother ramets,and of daughter ramets and genets.Mild drought or 10%shading or the combination stress of mild drought and light shade all significantly increased TB and ROB.Moreover,a synergistic interaction between shade and drought strongly increased flower spike number and floret number of genets,and TB,ROB and NROB.The effect of drought on TB and NROB was followed,and shading had greater influence on ROB.3)Shading and drought significantly(P<0.05)affected RA,REI,RI and RR:mild and moderate drought or 10%shading significantly increased the genet reproductive parameters,but severe drought or more than 30%shading significantly reduced reproductive parameters(P<0.05).Compared with either single stress,the combined stresses of light shading and mild drought promoted sexual reproduction.Mild drought alleviated reduction in sexual reproduction observed under moderate and severe shading.Reproductive parameters typically displayed a synergistic effect,with shading the larger effect,and the drought effect smaller.4)A.hispidus displayed an altered balance between clonal growth and sexual reproduction,depending on the degree of shading and drought.Greater than 30%shade prolonged the period of clonal growth by 10-34 days.5)Under all 20 treatment combinations of shading and drought,A.hispidus maintained both clonal and sexual reproduction concurrently.Adequate water and light promoted clonal growth.Light and moderate drought,and mild shade alone or in combination with drought were treatments that promoted sexual reproduction.
引文
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[18] Liu J P,You M H,Zhang L H,et al.External supports affect the photosynthetic characteristics and biomass allocation of the climbing plant Humulus scandens.Acta Ecologica Sinica,2015,35(18):6032-6040.刘金平,游明鸿,张丽慧,等.不同支持物对攀援植物—葎草雌雄株光合特性及生物量结构的影响.生态学报,2015,35(18):6032-6040.
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[2] Zhang J,Zhang M R,Zhao Y K.Advances in researches of phenotypic plasticity of the clonal plant.Journal of Inner Mongolia Agricultural University,2008,29(4):271-275.张瑾,张明如,赵银宽.克隆植物表型可塑性研究若干进展.内蒙古农业大学学报,2008,29(4):271-275.
[3] Wang C A,Li D Z,Zhu Z L,et al.Research methods for physiological integration of clonal plants and their application.Chinese Journal of Applied&Environmental Biology,2006,12(4):581-588.王长爱,李德志,朱志玲,等.克隆植物生理整合作用的研究方法及其应用.应用与环境生物学报,2006,12(4):581-588.
[4] Liu J P,Zhang X Q,You M H,et al.Diversification of expansion capacity and efficiency of cloned Hemarthria compressa.Chinese Journal of Eco-Agriculture,2007,15(6):12-17.刘金平,张新全,游明鸿,等.扁穗牛鞭草克隆单株地上空间拓展能力与拓展效益多样性研究.中国生态农业学报,2007,15(6):12-17.
[5] Fan X,Cai J,Liu J P,et al.Effects of partial shading on morphological plasticity and biomass allocation of Potentilla anserina L.Acta Prataculturae Sinica,2016,25(3):172-180.樊星,蔡捡,刘金平,等.局部遮光对鹅绒委陵菜基株形态塑性及生物量配置的影响.草业学报,2016,25(3):172-180.
[6] Peng H,Sun B X.Revision of the aristolochia herb in Guizhou.Guizhou Science,1991,(4):287-291.彭华,孙必兴.贵州荩草植物的修订.贵州科学,1991,(4):287-291.
[7] Liu J P.Effects of sectional slope on reproductive ramets quantitative characteristics and reproductive investment of wild Arthraxon hispidus.Pratacultural Scinence,2013,30(10):1602-1607.刘金平.坡度对野生荩草分株特征及生殖分配的影响.草业科学,2013,30(10):1602-1607.
[8] Liu J P,Zhang H Y.The study on morphologic diversity of wild Arthraxon hispidus from Nanchong.Anhui Agricultural Sciences,2008,36(12):4926-4929.刘金平,张海燕.南充地区野生荩草种质资源形态多样性研究.安徽农业科学,2008,36(12):4926-4929.
[9] Zhang X J,Cai J,Liu J P,et al.Analysis about component traits and biomass sllocation of Arthraxon hispidus in different shading degree.Southwest China Journal of Agricultural Science,2015,28(6):2720-2725.张小晶,蔡捡,刘金平,等.不同遮阴度对荩草构件性状和生物量分配影响的差异性分析.西南农业学报,2015,28(6):2720-2725.
[10] Zhao Y,Cai J,Li Y,et al.The analysis of the response of three cool season turf grasses to temperature and shade synergy.Grassland and Livestock,2015,(2):28-33.赵艳,蔡捡,李莹,等.3种冷季型草坪草对温度与遮荫协同作用响应的差异性分析.草业与畜牧,2015,(2):28-33.
[11] Sheng L J.Phenotypic plasticity in response to light and patterns of carbon physiological integration in Zoysia japonica Steud.,a clonal plant.Shanghai:East China Normal University,2007.盛丽娟.克隆植物结缕草对光照的表型可塑性响应与碳素整合格局研究.上海:华东师范大学,2007.
[12] Sheng H Y,Li J Q,Yang Y Z,et al.Response of phenotypic plasticity of Potentilla anserina L.to soil moisture.Agricultural Research in the Arid Areas,2004,22(3):119-122.盛海彦,李军乔,杨银柱,等.土壤水分对鹅绒委陵菜表型可塑性的影响.干旱地区农业研究,2004,22(3):119-122.
[13] Marshall C.Source-sink relation of interconnected ramets//van Groenendael J,de Kroon H.Clonal growth in plants regulation and function.The Hague:SPB Academic Publishing,1990:23-42.
[14] Luo D,Qian Y Q,Liu J X,et al.Phenotypic responses of a clonal plant(Buchloe dactyloides)to nutrient heterogeneity.Acta Prataculturae Sinica,2014,23(3):104-109.罗栋,钱永强,刘俊祥,等.克隆植物野牛草对异质营养的表型可塑性响应.草业学报,2014,23(3):104-109.
[15] Eriksson O.Ramet behavior and population growth in the clonal herb Potentilla anserina.Journal of Ecology,1988,76:522-536.
[16] Stuefer J F,Huber H.Differential effects of light quantity and spectral light quality on growth,morphology and development of two stoloniferous Potentillaspecies.Oecologia,1998,117:1-8.
[17] Zhang Y F,Zhang D Y.Asexual and sexual reproductive strategies in clonal plants.Journal of Plant Ecology,2006,30(1):174-183.张玉芬,张大勇.克隆植物的无性与有性繁殖对策.植物生态学报,2006,30(1):174-183.
[18] Liu J P,You M H,Zhang L H,et al.External supports affect the photosynthetic characteristics and biomass allocation of the climbing plant Humulus scandens.Acta Ecologica Sinica,2015,35(18):6032-6040.刘金平,游明鸿,张丽慧,等.不同支持物对攀援植物—葎草雌雄株光合特性及生物量结构的影响.生态学报,2015,35(18):6032-6040.
[19] Xie R J,Zhang X J,Liu J P,et al.Synergistic effects of drought and shade on component morphology and biomass allocation of Arthraxon hispidus.Pratacultural Scinence,2017,34(7):1496-1505.谢瑞娟,张小晶,刘金平,等.干旱和遮阴对荩草构件形态及生物量分配的影响.草业科学,2017,34(7):1496-1505.
[20] Xie R J,Zhang X J,Liu J P,et al.Synergy effects of shade and drought on photosynthetic characteristics of Arthraxon hispidus.Acta Prataculturae Sinica,2017,26(10):64-76.谢瑞娟,张小晶,刘金平,等.遮阴和干旱对荩草光合特性影响的协同作用.草业学报,2017,26(10):64-76.
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