CO_2浓度升高对宁夏枸杞苗木光合特性及生物量分配影响
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摘要
为明确大气CO_2浓度升高对宁夏枸杞光合特性及生物量分配的影响,以宁夏枸杞苗木为试材,采用开顶气室(OTC)模拟自然环境大气CO_2浓度变化,设置3个CO_2浓度水平[CK(380±20μmol/mol)、TR1(570±20μmol/mol)、TR2(760±20μmol/mol)],分别于不同CO_2浓度处理90d、120d后,测定其净光合速率(P_n)、光响应曲线、CO_2响应曲线等相关指标及植株不同器官生物量。结果表明:(1)TR1及TR2下Pn于第一年处理至90d时均较CK显著升高(P<0.05),且TR1处理在120d时较CK显著降低;第二年处理90d时,TR1处理较CK下降了4.77%,处理120d时TR1、TR2均高于CK,但差异不显著。(2)随着CO_2浓度升高,两年中TR1、TR2处理的胞间CO_2浓度(C_i)较CK均显著升高,处理后120d时,气孔导度(G_s)较CK均显著下降;水分利用率(WUE)在第一年处理中均无显著变化,但在第二年处理120d时,TR1、TR2均较CK显著上升。(3)处理至90d和120d时,TR1、TR2组的宁夏枸杞苗木光饱和点、CO_2饱和点均高于CK,但TR2组初始羧化效率低于CK。(4)随着CO_2浓度升高,宁夏枸杞苗木地上部分生物量分配显著增加,地下部分生物量分配显著降低。研究发现,一定时间内适宜CO_2浓度升高可促进宁夏枸杞苗木光合作用,使得其地上部分生物量分配显著提高,地下部分生物量分配显著降低;但随着处理浓度升高及处理时间延长,其光合作用有下调趋势,表现为净光合速率、气孔导度、初始羧化效率等下降。
To explore the effects of elevated CO_2 on photosynthesis characteristics and biomass allocation in Lycium barbarum.With L.barbarum seedlings as research materials,we applied open top chamber(OTC),a facility to control changed CO_2 concentration to simulate natural environment in experiment.We conducted three gradients of CO_2 concentration,the control group is(380±20)μmol/mol(CK),which is ambient CO_2,and elevated was(570±20)μmol/mol(TR1),(760±20)μmol/mol(TR2).The effects of elevated CO_2 on photosynthesis and biomass allocation of L.barbarumseedlings were assessed by net photosynthetic rate(P_n),light response curves,CO_2 response curves and dry weight during seedlings exposed to all three treatments(CK,TR1,TR2)after 90 and 120 days in 2017 and 2018.Results showed:(1)when treated for 90 days in 2017,the TR1 and TR2 in Pnof L.barbarumseedlings were increased significantly than that of control(P<0.05).After 120 days,the Pnof TR1 was decreased significantly compared with CK(P<0.05).While P_n of TR1 was 4.77%lower than CK when we repeated the test after 90 days in2018(P>0.05).The treatment of TR1 and TR2 was higher in Pnthan CK after 120 days,but the difference between them was not significant(P>0.05).(2)With the rising of CO_2 concentration,it is similar that the intercellular CO_2 concentrations(C_i)of TR1 and TR2 treatment were soared respectively than ambient CO_2 in 2017 and 2018(P<0.05).With the time prolonging until 120 days,the stomatal conductance(G_s)of TR1 and TR2 dropped conspicuously than that of CK(P<0.05).No difference in water use efficiency(WUE)in 2017,but it raised obviously than control after 120 days in 2018.(3)At 90 days,the light saturation point and CO_2 saturation point of treatment TR1 and TR2 were all rising than control,which was maintained in 120 days,but the CO_2 assimilation rate of TR2 was declined.(4)The biomass allocation of L.barbarumseedlings increased signally in its aboveground with CO_2 rising,thus less allocated in underground organs(P<0.05).We concluded that slightly elevated CO_2 could promote the photosynthesis of L.barbarumseedlings,as a result of photosynthesis,the biomass accumulation in L.barbarum seedlings was tend to its aboveground organs,while underground was reduced remarkably.With the concentration and incubation time extending,the photosynthesis in L.barbarumseedlings showed a downregulation trend,which revealed by its Pn,Gsand photosynthetic CO_2 assimilation rate decreased.
To explore the effects of elevated CO_2 on photosynthesis characteristics and biomass allocation in Lycium barbarum.With L.barbarum seedlings as research materials,we applied open top chamber(OTC),a facility to control changed CO_2 concentration to simulate natural environment in experiment.We conducted three gradients of CO_2 concentration,the control group is(380±20)μmol/mol(CK),which is ambient CO_2,and elevated was(570±20)μmol/mol(TR1),(760±20)μmol/mol(TR2).The effects of elevated CO_2 on photosynthesis and biomass allocation of L.barbarumseedlings were assessed by net photosynthetic rate(P_n),light response curves,CO_2 response curves and dry weight during seedlings exposed to all three treatments(CK,TR1,TR2)after 90 and 120 days in 2017 and 2018.Results showed:(1)when treated for 90 days in 2017,the TR1 and TR2 in Pnof L.barbarumseedlings were increased significantly than that of control(P<0.05).After 120 days,the Pnof TR1 was decreased significantly compared with CK(P<0.05).While P_n of TR1 was 4.77%lower than CK when we repeated the test after 90 days in2018(P>0.05).The treatment of TR1 and TR2 was higher in Pnthan CK after 120 days,but the difference between them was not significant(P>0.05).(2)With the rising of CO_2 concentration,it is similar that the intercellular CO_2 concentrations(C_i)of TR1 and TR2 treatment were soared respectively than ambient CO_2 in 2017 and 2018(P<0.05).With the time prolonging until 120 days,the stomatal conductance(G_s)of TR1 and TR2 dropped conspicuously than that of CK(P<0.05).No difference in water use efficiency(WUE)in 2017,but it raised obviously than control after 120 days in 2018.(3)At 90 days,the light saturation point and CO_2 saturation point of treatment TR1 and TR2 were all rising than control,which was maintained in 120 days,but the CO_2 assimilation rate of TR2 was declined.(4)The biomass allocation of L.barbarumseedlings increased signally in its aboveground with CO_2 rising,thus less allocated in underground organs(P<0.05).We concluded that slightly elevated CO_2 could promote the photosynthesis of L.barbarumseedlings,as a result of photosynthesis,the biomass accumulation in L.barbarum seedlings was tend to its aboveground organs,while underground was reduced remarkably.With the concentration and incubation time extending,the photosynthesis in L.barbarumseedlings showed a downregulation trend,which revealed by its Pn,Gsand photosynthetic CO_2 assimilation rate decreased.
引文
[1]IPCC.Climate change 2013:The physical science basis[M].Cambridge:Cambridge University Press,2013.
[2]IPCC.Climate Change 2013:The physical science basis.Contribution of working group I to the fifth assessment report of the intergovernmental panel on climate change.Cambridge:Cambridge University Press,2013.
[3]种培芳,姬江丽,李毅,等.红砂(Reaumuria soongorica)对大气CO2浓度升高及降水变化的光合生理响应[J].中国沙漠,2017,37(4):714-723.ZHONG P F,JI J L,LI Y,et al.Photosynthetic physiology responses to elevated CO2concentration and changing precipitation in desert plant Reaumuria soongorica[J].Journal of Desert Research,2017,37(4):714-723.
[4]徐明怡,倪红伟.CO2浓度升高对植物光合作用的影响[J].国土与自然资源研究,2016,(2):83-86.XU M Y,NI H W.Effects of elevated CO2concentration on plant photosynthesis[J].Territory&Natural Resources Study,2016,(2):83-86.
[5]JIANG Y L,XU Z Z,ZHOU G S.Elevated CO2can modify the response to a water status gradient in a steppe grass:from cell organelles to photosynthetic capacity to plant growth[J].BMC Plant Biology,2016,16:157.
[6]WANG L,FENG Z Z,et al.Effects of elevated atmospheric CO2on physiology and yield of wheat(Triticum aestivum L.):A meta-analytic test of current hypotheses[J].Agriculture,Ecosystems and Environment 2013,178:57-63.
[7]高霁,郝兴宇,居煇,等.自由大气CO2浓度升高对夏大豆光合色素含量和光合作用的影响[J].中国农学通报,2012,28(6):47-52.GAO J,HAO X Y,JU H,et al.Effect of elevated[CO2]on photosynthetic pigment contents and photosynthesis of Summer Soybean[J].Chinese Agricultural Science Bulletin,2012,28(6):47-52.
[8]袁蕊,聂磊云,等.大气CO2浓度升高对辣椒光合作用及相关生理特性的影响[J].生态学杂志,2017,36(12):3 510-3 516.YUAN R,NIE L Y,et al.Effect of elevated[CO2]on photosynthesis and physiological characteristics of pepper(Capsicum annuumL.)[J].Chinese Journal of Ecology,2017,36(12):3510-3 516.
[9]YELLE S,BEESON J R C,TRUDEL M J.Acclimation of two tomato species to high atmospheric CO2[J].Plant Physiology,1989,90(4):1 473-1 477.
[10]刘晓萌,于凌飞,黄耀,等.CO2浓度升高下粳稻叶片光合作用对光强变化的响应[J].生态学杂志,2018,37(4):1 051-1 057.LIU X M,YU L F,HUANG Y,et al.Response of photosynthesis in leaves of Japonicarice to light intensities at elevated CO2concentration[J].Chinese Journal of Ecology,2018,37(4):1 051-1 057.
[11]韩燕青,刘鑫,胡维平,等.CO2浓度升高对苦草(Vallisneria natans)叶绿素荧光特性的影响[J].植物研究,2017,37(1):45-51.HAN Y Q,LIU X,HU W P,et al.Effects of CO2enrichment on chlorophyll fluorescence characteristics of Vallisneria natans[J].Bulletin of Botanical Research,2017,37(1):45-51.
[12]PORRAS M E.,LORENZO P,MEDRANO E.et al.Photosynthetic acclimation to elevated CO2concentration in a sweet pepper(Capsicum annuum)crop under Mediterranean greenhouse conditions:influence of the nitrogen source and salinity[J].Functional Plant Biology,2017,44(6):573-586.
[13]汪燕平.清以来宁夏枸杞作为地道药材的形成史[J].史林,2017,(3):67-76+220.WANG Y P.Formation history of Lycium barbarum as a genuine medicinal material since Qing dynasty[J].Historical Review,2017,(3):67-76+220.
[14]吕海洋.枸杞多酚的结构鉴定及抗氧化活性评价[D].北京:中国农业科学院,2016.
[15]叶子飘.光合作用对光响应新模型及其应用[J].生物数学学报,2008,23(4):710-716.YE Z P.New model of photosynthesis response to light and its application[J].Journal of Biomathematics,2008,23(4):710-716.
[16]叶子飘,于强.光合作用对胞间和大气CO2响应曲线的比较[J].生态学杂志,2009,28(11):2 233-2 238.YE Z P,YU Q.A comparison of response curves of photosynthesis to intercellular and air CO2concentrations[J].Chinese Journal of Ecology,2009,28(11):2 233-2 238.
[17]张韫,李响,张帆,等.CO2升高对红松幼苗水分生理特征与土壤含水率变化的影响[J].北京林业大学学报,2015,37(1):37-41.ZHANG W,LI X,ZHANG F,et al.Effects of elevated CO2on physiological characteristics of water in Pinus koraiensis seedlings and soil moisture in a controlled environment[J].Journal of Beijing Forestry University,2015,37(1):37-41.
[18]XU Z Z,JIANG Y L,ZHOU G S.Elevated-CO2response of stomata and its dependence on environmental factors[J].Frontiers in Plant Science,2016,7:657.
[19]居辉,姜帅,李靖涛,等.北方冬麦区CO2浓度增高与氮肥互作对冬小麦生理特性和产量的影响[J].中国农业科学,2015,48(24):4 948-4 956.JU H,JIANG S,LI J T,et al.Interactive effects of elevated CO2 and nitrogen on the physiology and yield of Winter Wheat in North Winter Wheat Region of China[J].Scientia Agricultura Sinica,2015,48(24):4 948-4 956.
[20]NGUYEN C T,HIROYUKI S,ETSUSHI K.Effects of elevated CO2 concentration on growth and photosynthesis of Chinese yam under different temperature regimes[J].Plant Production Science,2017,20:2 227-2 236.
[21]JEFFERSON R S,ANGELICA E P,et al.Photosynthetic acclimation to elevated CO2combined with partial rootzone drying results in improved water use efficiency,drought tolerance and leaf carbon balance of grapevines(Vitis labrusca)[J].Environmental and Experimental Botany,2017,134:82-95.
[22]候晶东,曹兵,宋丽华.CO2浓度倍增对宁夏枸杞光合特性的影响[J].南京林业大学学报:自然科学版,2012,36(5):71-76.HOU J D,CAO B,SONG L H.Effect of doubled CO2concentration on photosynthesis characteristics of Lycium barbarum seeding[J].Journal of Nanjing Forestry University(Natural Science Edition)2012,36(5):71-76.
[23]CAWAS B.E,MIMI H-S,JUNTARO N.CO2sensing and CO2regulation of stomatal conductance:advances and open questions[J].Trends in Plant Science,2016,21:16-30.
[24]RUBEN V,PILAR P,RAFAEL M-C,et al.Improved responses to elevated CO2in durum wheat at a low nitrate supply associated with the up-regulation of photosynthetic genes and the activation of nitrate assimilation[J].Plant Science,2017,260:119-128.
[25]陆志峰.钾素营养对冬油菜叶片光合作用的影响机制研究[D].武汉:华中农业大学,2017.
[26]ZHU X C,SONG F B,LIU S Q,et al.Arbuscular mycorrhiza improve growth,nitrogen uptake,and nitrogen use efficiency in wheat grown under elevated CO2[J].Mycorrhiza,2016,26(2):133-140.
[27]LI D X,LIU H L,QIAO Y Z,et al.Effects of elevated CO2on the growth,seed yield,and water use efficiency of soybean(Glycine max(L.)Merr.)under drought stress[J].Agricultural Water Management,2013,129:105-112.
[28]MICHAEL T,DANANJALI G,NAOKI H,et al.Effects of elevated carbon dioxide on photosynthesis and carbon partitioning:aperspective on root sugar sensing and hormonal crosstalk[J].Plant Physiology,2017,8:578.
[29]曹宏杰,倪红伟.大气CO2升高对土壤碳循环影响的研究进展[J].生态环境学报,2013,22(11):1 846-1 852.CAO H J,NI H W.Research progress on the effects of elevated CO2concentration on carbon cycling[J].Ecology and Environmental Sciences,2013,22(11):1 846-1 852.
[30]CAO B,DANG Q L,YU X G,et al.Effects of CO2and nitrogen on morphological and biomass traits of white birch(Betula papyrifer)seedlings[J].Forest Ecology and Management,2008,254:217-224.
[2]IPCC.Climate Change 2013:The physical science basis.Contribution of working group I to the fifth assessment report of the intergovernmental panel on climate change.Cambridge:Cambridge University Press,2013.
[3]种培芳,姬江丽,李毅,等.红砂(Reaumuria soongorica)对大气CO2浓度升高及降水变化的光合生理响应[J].中国沙漠,2017,37(4):714-723.ZHONG P F,JI J L,LI Y,et al.Photosynthetic physiology responses to elevated CO2concentration and changing precipitation in desert plant Reaumuria soongorica[J].Journal of Desert Research,2017,37(4):714-723.
[4]徐明怡,倪红伟.CO2浓度升高对植物光合作用的影响[J].国土与自然资源研究,2016,(2):83-86.XU M Y,NI H W.Effects of elevated CO2concentration on plant photosynthesis[J].Territory&Natural Resources Study,2016,(2):83-86.
[5]JIANG Y L,XU Z Z,ZHOU G S.Elevated CO2can modify the response to a water status gradient in a steppe grass:from cell organelles to photosynthetic capacity to plant growth[J].BMC Plant Biology,2016,16:157.
[6]WANG L,FENG Z Z,et al.Effects of elevated atmospheric CO2on physiology and yield of wheat(Triticum aestivum L.):A meta-analytic test of current hypotheses[J].Agriculture,Ecosystems and Environment 2013,178:57-63.
[7]高霁,郝兴宇,居煇,等.自由大气CO2浓度升高对夏大豆光合色素含量和光合作用的影响[J].中国农学通报,2012,28(6):47-52.GAO J,HAO X Y,JU H,et al.Effect of elevated[CO2]on photosynthetic pigment contents and photosynthesis of Summer Soybean[J].Chinese Agricultural Science Bulletin,2012,28(6):47-52.
[8]袁蕊,聂磊云,等.大气CO2浓度升高对辣椒光合作用及相关生理特性的影响[J].生态学杂志,2017,36(12):3 510-3 516.YUAN R,NIE L Y,et al.Effect of elevated[CO2]on photosynthesis and physiological characteristics of pepper(Capsicum annuumL.)[J].Chinese Journal of Ecology,2017,36(12):3510-3 516.
[9]YELLE S,BEESON J R C,TRUDEL M J.Acclimation of two tomato species to high atmospheric CO2[J].Plant Physiology,1989,90(4):1 473-1 477.
[10]刘晓萌,于凌飞,黄耀,等.CO2浓度升高下粳稻叶片光合作用对光强变化的响应[J].生态学杂志,2018,37(4):1 051-1 057.LIU X M,YU L F,HUANG Y,et al.Response of photosynthesis in leaves of Japonicarice to light intensities at elevated CO2concentration[J].Chinese Journal of Ecology,2018,37(4):1 051-1 057.
[11]韩燕青,刘鑫,胡维平,等.CO2浓度升高对苦草(Vallisneria natans)叶绿素荧光特性的影响[J].植物研究,2017,37(1):45-51.HAN Y Q,LIU X,HU W P,et al.Effects of CO2enrichment on chlorophyll fluorescence characteristics of Vallisneria natans[J].Bulletin of Botanical Research,2017,37(1):45-51.
[12]PORRAS M E.,LORENZO P,MEDRANO E.et al.Photosynthetic acclimation to elevated CO2concentration in a sweet pepper(Capsicum annuum)crop under Mediterranean greenhouse conditions:influence of the nitrogen source and salinity[J].Functional Plant Biology,2017,44(6):573-586.
[13]汪燕平.清以来宁夏枸杞作为地道药材的形成史[J].史林,2017,(3):67-76+220.WANG Y P.Formation history of Lycium barbarum as a genuine medicinal material since Qing dynasty[J].Historical Review,2017,(3):67-76+220.
[14]吕海洋.枸杞多酚的结构鉴定及抗氧化活性评价[D].北京:中国农业科学院,2016.
[15]叶子飘.光合作用对光响应新模型及其应用[J].生物数学学报,2008,23(4):710-716.YE Z P.New model of photosynthesis response to light and its application[J].Journal of Biomathematics,2008,23(4):710-716.
[16]叶子飘,于强.光合作用对胞间和大气CO2响应曲线的比较[J].生态学杂志,2009,28(11):2 233-2 238.YE Z P,YU Q.A comparison of response curves of photosynthesis to intercellular and air CO2concentrations[J].Chinese Journal of Ecology,2009,28(11):2 233-2 238.
[17]张韫,李响,张帆,等.CO2升高对红松幼苗水分生理特征与土壤含水率变化的影响[J].北京林业大学学报,2015,37(1):37-41.ZHANG W,LI X,ZHANG F,et al.Effects of elevated CO2on physiological characteristics of water in Pinus koraiensis seedlings and soil moisture in a controlled environment[J].Journal of Beijing Forestry University,2015,37(1):37-41.
[18]XU Z Z,JIANG Y L,ZHOU G S.Elevated-CO2response of stomata and its dependence on environmental factors[J].Frontiers in Plant Science,2016,7:657.
[19]居辉,姜帅,李靖涛,等.北方冬麦区CO2浓度增高与氮肥互作对冬小麦生理特性和产量的影响[J].中国农业科学,2015,48(24):4 948-4 956.JU H,JIANG S,LI J T,et al.Interactive effects of elevated CO2 and nitrogen on the physiology and yield of Winter Wheat in North Winter Wheat Region of China[J].Scientia Agricultura Sinica,2015,48(24):4 948-4 956.
[20]NGUYEN C T,HIROYUKI S,ETSUSHI K.Effects of elevated CO2 concentration on growth and photosynthesis of Chinese yam under different temperature regimes[J].Plant Production Science,2017,20:2 227-2 236.
[21]JEFFERSON R S,ANGELICA E P,et al.Photosynthetic acclimation to elevated CO2combined with partial rootzone drying results in improved water use efficiency,drought tolerance and leaf carbon balance of grapevines(Vitis labrusca)[J].Environmental and Experimental Botany,2017,134:82-95.
[22]候晶东,曹兵,宋丽华.CO2浓度倍增对宁夏枸杞光合特性的影响[J].南京林业大学学报:自然科学版,2012,36(5):71-76.HOU J D,CAO B,SONG L H.Effect of doubled CO2concentration on photosynthesis characteristics of Lycium barbarum seeding[J].Journal of Nanjing Forestry University(Natural Science Edition)2012,36(5):71-76.
[23]CAWAS B.E,MIMI H-S,JUNTARO N.CO2sensing and CO2regulation of stomatal conductance:advances and open questions[J].Trends in Plant Science,2016,21:16-30.
[24]RUBEN V,PILAR P,RAFAEL M-C,et al.Improved responses to elevated CO2in durum wheat at a low nitrate supply associated with the up-regulation of photosynthetic genes and the activation of nitrate assimilation[J].Plant Science,2017,260:119-128.
[25]陆志峰.钾素营养对冬油菜叶片光合作用的影响机制研究[D].武汉:华中农业大学,2017.
[26]ZHU X C,SONG F B,LIU S Q,et al.Arbuscular mycorrhiza improve growth,nitrogen uptake,and nitrogen use efficiency in wheat grown under elevated CO2[J].Mycorrhiza,2016,26(2):133-140.
[27]LI D X,LIU H L,QIAO Y Z,et al.Effects of elevated CO2on the growth,seed yield,and water use efficiency of soybean(Glycine max(L.)Merr.)under drought stress[J].Agricultural Water Management,2013,129:105-112.
[28]MICHAEL T,DANANJALI G,NAOKI H,et al.Effects of elevated carbon dioxide on photosynthesis and carbon partitioning:aperspective on root sugar sensing and hormonal crosstalk[J].Plant Physiology,2017,8:578.
[29]曹宏杰,倪红伟.大气CO2升高对土壤碳循环影响的研究进展[J].生态环境学报,2013,22(11):1 846-1 852.CAO H J,NI H W.Research progress on the effects of elevated CO2concentration on carbon cycling[J].Ecology and Environmental Sciences,2013,22(11):1 846-1 852.
[30]CAO B,DANG Q L,YU X G,et al.Effects of CO2and nitrogen on morphological and biomass traits of white birch(Betula papyrifer)seedlings[J].Forest Ecology and Management,2008,254:217-224.