CO_2浓度升高对冬小麦生长及氮素利用的效应
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摘要
本研究以冬小麦（Triticum aestivum L.）为供试材料，于2007-2010年连续3年利用开顶式气室（Open Top Chambers, OTCs）装置，通过土培和水培试验，系统研究了大气CO_2浓度升高对冬小麦光合、叶绿素荧光、光合源结构、根系形态和有机碳分泌、产量形成及氮素利用的影响。研究获得以下主要进展：
（1）在不施氮或低氮条件下CO_2浓度升高使冬小麦旗叶叶绿素a、叶绿素b和叶绿素总量减少，促进可溶性糖累积，导致光合适应；供氮充分时通过调节叶绿素，特别是叶绿素a而减缓或消除光合适应现象，而通过调节叶绿素b及可溶性糖累积对光合适应现象的调控作用相对有限，说明改善氮素营养可消除光合适应现象。
（2）CO_2浓度升高使冬小麦旗叶叶绿素基础荧光（Fo）增加1.4%~15.2%，最大荧光（Fm）、可变荧光（Fv）、最大光化学效率（Fv/Fm）以及潜在光化学活性（Fv/Fo）分别下降0.3%~9.3%、2.4%~12.0%、0.9%~4.7%和5.9%~22.7%；Fo值的增幅以及Fm、Fv、Fv/Fm和Fv/Fo的降幅随施氮水平提高而下降，表明CO_2浓度升高会导致PSII发生光损伤，使其电子传递、潜在活性和原初光能转换能力减弱，而改善氮素营养可减弱甚至消除CO_2浓度升高的这些负效应。
（3）CO_2浓度升高使冬小麦叶片、茎鞘及单茎总光合面积分别增加11.5%~37.3%、14.5%~18.1%和7.3%~38.3%。叶片和茎节长度增加是光合面积增加的主要原因。CO_2浓度与供氮水平互作只对少数光合器官形态产生影响；CO_2浓度升高对单茎花后干物质积累量、粒重叶比和粒数叶比在不施氮时影响不明显或导致下降，施氮后均使其显著增加。说明改善氮素营养可促进CO_2浓度升高对花后干物质生产的效应，并有利于提高单位叶源的库承载力。
（4）水培条件下，CO_2浓度升高使冬小麦幼苗根冠生物量、根长度、根面积、有机碳分泌速率及总量分别增加1.9%~42.8%、5.5%~27.7%、14.9%~92.9%、5.9%~71.4%、8.7%~11.8%和13.3%，叶片硝酸还原酶活性增加14.8%~33.5%，而对根冠比的影响缺乏规律性。在较低介质供氮水平下CO_2浓度升高对小麦幼苗根长和根面积以及有机碳分泌促进作用较大，表明当氮素营养不足时，CO_2浓度升高更有利于同化产物向根系的分配。
（5）CO_2浓度升高使冬小麦每盆籽粒重量平均增加48.3%~70.0%，增产原因主要是穗数和穗粒数增加的结果。CO_2浓度升高促进了花前不同器官干物质积累，使花前地上部营养器官贮存物质在花后向籽粒的转运量、转运率和贡献率明显增加。CO_2浓度升高对每盆籽粒重量、干物质积累及转运的影响，在不施氮时不显著，施氮后在大多数情况下达显著水平。说明施氮有助于CO_2浓度升高对每盆籽粒重量、花前物质积累和转运的正向效应。
（6）CO_2浓度升高使冬小麦开花期和成熟期植株氮素累积量分别增加15.0%~27.9%和13.9%~26.1%，使花前地上部营养器官氮素在花后向籽粒的转运量提高13.8%~77.4%；但对花前地上部氮素转运率及花后籽粒氮素累积贡献率的影响因品种而异。CO_2浓度升高也可提高氮素籽粒和干物质生产效率和氮素收获指数。不施氮时CO_2浓度升高对氮素累积、转运及籽粒中分配的影响不显著，施氮后显著增加。
（7）增加冬小麦中、后期追氮比例，更有利于发挥CO_2浓度升高对作物生长的促进效应。在不施氮条件下，CO_2浓度升高使冬小麦旗叶SPAD值和可溶性蛋白含量下降，导致旗叶出现明显光合适应；施氮时这些现象均未发生；与一次性底施相比，在施氮比例（底肥﹕拔节肥﹕孕穗肥）为5:5:0和5:3:2时提高了灌浆期旗叶光合速率。结果同时表明，分期施氮比全部底施更有利于增加CO_2浓度升高对小麦生育后期光合能力、花前物质积累与转运、籽粒产量以及氮素利用的促进效应。主要创新点：
（1）在介质供氮不足时，CO_2浓度升高会增加冬小麦叶片可溶性糖累积，导致PSII光损伤，叶绿素含量、PSII的电子传递、原初光化学效率和潜在活性降低，从而引起光合适应。改善氮素营养，特别增加中后施氮比例可减轻冬小麦光合适应，增强CO_2浓度升高对植株生长发育、籽粒产量形成和氮素利用的促进作用。
（2）CO_2浓度升高可促进冬小麦源器官和根系生长发育，增强花前碳氮物质积累和花后向籽粒的转运，增加籽粒产量和氮素累积量，提高氮素生产效率。
（3）综合分析研究结果认为，在介质供氮充分时，CO_2浓度升高导致花后非叶光合源面积增加及促进花前营养器官同化产物向籽粒转运是籽粒产量增加的主要原因。
The research was carried out with winter wheat（Triticum aestivum L.） as materialsduring the three growth seasons from2007to2010. Some pot and water culture experimentswere launched in four open top chambers (OTCs). The main aim of research was to assess theinfluence of CO_2concentration elevation on the photosynthesis, chlorophyll fluorescenceparameters, source components, root morphology and organic carbon secretion, yieldformation, nitrogen use efficency of winter wheat. The main results were as followings：
(1) CO_2enrichment reduced chlorophyll a, chlorophyll b and total chlorophyll contents,increased soluble sugar content in wheat flag leaf, and led to photosynthetic down-regulationunder no nitrogen application or low nitrogen level, but under ample nitrogen application, iteased off or removed photosythesitic down-regulation mainly by increasing chlorophyllcontent, especially chlorophyll a content, and limitedly by chlorophyll b and soluble sugarcontents. This showed nitrogen nutrition improvement could regulate leaf photosythesiticacclimation to some extent.
(2) When air CO_2concentration was elevated, the average value of initialfluorescence(Fo) in wheat flag leaf increased1.4%~15.2%, and the average value ofmaximumu fluorescence(Fm), variable fluorescence(Fv), maximal photochemicalefficiency(Fv/Fm) and potential photochemical efficiency(Fv/Fo) reduced0.3%~9.3%,2.4%~12.0%,0.9%~4.7%and5.9%~22.7%, respectively. The ranges of Fo increase, and Fm,Fv, Fv/Fm and Fv/Fo decrease fell with the increase of nitrogen level. It showed elevated CO_2gave biger photo-damage to wheat PSII, and restrained its electron transport, potential activityand original light conversion ability. Nitrogen addication could weaken or eliminate thenegative effects.
(3) Compared of ambient CO_2concentration, CO_2enrichment increased thephotosynthesis areas of leaf, stem&sheath and whole stem by11.5%~37.3%,14.5%~18.1%and7.3%~38.3%, respectively. The expansion of photosynthesis area should mainly be due tothe increase of leaf and node lengths. The dry matter accumulation after anthesis, the ratios ofgrain number and weight to leaf area responded weakly or negatively to elevated CO_2under no nitrogen application, but significantly increased in the very great majority of cases undernitrogen addication. It showed that the positive effects of CO_2enrichment on dry matterproduction after anthesis and sink bearing capacity per source area could be enhanced byimproving nitrogen nutrition.
(4) The hydroponics results showed that compared with the background CO_2concentration, elevated CO_2increased root and shoot biomass, root length, root area, rootorganic carbon excretion rate and content, and leaf NR activity in wheat seedling by1.9%~42.8%,5.5%~27.7%,14.9%~92.9%,5.9%~71.4%,8.7%~11.8%,13.3%and14.8%~33.5%, but its effect on the root/shoot ratio lacked regularity. Elevated CO_2influnenced root length and area, organic carbon excretion more significantly under lownitrogen nutrition than under apmle nitrogen. It showed that elevated CO_2led to the moreassimilate product distributed to root when lacking nitrogen nutrition.
(5) CO_2enrichment significantly increased wheat grain weight per pot by48.3%~70.0%,mainly due to the improvement of spike number per pot and grain number per spike. ElevatedCO_2promoted the accumulation of dry matter in different organs before anthesis. The amountand rate of pre-anthesis dry matter in shoot transferring to grain after anthesis increased andthe contribution rate of pre-anthesis dry matter translocation rose with CO_2elevation. Theeffects of elevated CO_2on grain weight per pot, dry matter accumulation and translocationwere weak or not significant, but were mostly significant under nitrogen addication, whichindicated reasonable nitrogen application could improve the positive effects of elevated CO_2.
(6) Elevated CO_2increased plant nitrogen accumulation at anthesis and mature stages,the amount of pre-anthesis nitrogen in shoot transferring to grain after anthesis by15.0%~27.9%,13.9%~26.1%and13.8%~77.4%, respectively, but it had different effect onthe contribution rate of pre-anthesis nitrogen in shoot of two varieties. Compared with thebackground CO_2concentration, the high CO_2concentration treatments increased nitrogenefficiencies for grain and dry matter production, and nitrogen harvest index. The effects ofelevated CO_2were not significant under no nitrogen application, and all sigficant undernitrogen application.
(7) The effects of CO_2enrichment on wheat growth could be increased by applying morenitrogen at the middle or later stages. Elevated CO_2resulted in photosynthetic acclimation ofwheat flag leaf along with the decease of SPAD and soluble protein content under nonitrogen application, but the appearances were found under nitrogen addication; thephotosynthetic rate at grain-filling stage was enhanced by elevated CO_2under nitrogenapplications with ratios of5:5:0and5:3:2at before sowing, regreening and booting stage.Additionly, the length and area of the top three leaves at heading stage, photosynthesis after anthesis, grain yield, pre-anthesis dry matter accumulatio and translocation, and nitrogenusage were more easyly promoted by CO_2elevation under nitrogen spilt application thanunder nitrogen all-based application..
Main innovations:
(1) CO_2elevation would down-regulate winter wheat leaf photosythesis by inceasing leafsoluble sugar content and PSII light suppression, and reducing chlorophyll content, PSII lightchemistry efficiency and activity. By improvementing nitrogen nutrition, especiallyapplying more nitrogen at the middle and later stages, the leaf photosynthetic acclimationwould be relieved, and the positive effects of CO_2elevation on plant growth and development,yield formation and nitrogen usage would be strengthened.
(2) CO_2enrichment would promote the growth and development of source organs androot, pre-antheis dry matter and nitrogen accumulation in shoot, and translocation to grainafter anthesis, increase grain yield and nitrigen accumulation in grain, and improve thenitrogen production effciency.
3) The synthetical analyses showed that when nitrogen in medium was ample, thepositively effect of CO_2elevation on winter yield was mainly due to its accelerating thenon-leaf source areas and the translocation of dry matter restored in vegetative organs beforeanthesis to grain after anthesis.
（1）在不施氮或低氮条件下CO_2浓度升高使冬小麦旗叶叶绿素a、叶绿素b和叶绿素总量减少，促进可溶性糖累积，导致光合适应；供氮充分时通过调节叶绿素，特别是叶绿素a而减缓或消除光合适应现象，而通过调节叶绿素b及可溶性糖累积对光合适应现象的调控作用相对有限，说明改善氮素营养可消除光合适应现象。
（2）CO_2浓度升高使冬小麦旗叶叶绿素基础荧光（Fo）增加1.4%~15.2%，最大荧光（Fm）、可变荧光（Fv）、最大光化学效率（Fv/Fm）以及潜在光化学活性（Fv/Fo）分别下降0.3%~9.3%、2.4%~12.0%、0.9%~4.7%和5.9%~22.7%；Fo值的增幅以及Fm、Fv、Fv/Fm和Fv/Fo的降幅随施氮水平提高而下降，表明CO_2浓度升高会导致PSII发生光损伤，使其电子传递、潜在活性和原初光能转换能力减弱，而改善氮素营养可减弱甚至消除CO_2浓度升高的这些负效应。
（3）CO_2浓度升高使冬小麦叶片、茎鞘及单茎总光合面积分别增加11.5%~37.3%、14.5%~18.1%和7.3%~38.3%。叶片和茎节长度增加是光合面积增加的主要原因。CO_2浓度与供氮水平互作只对少数光合器官形态产生影响；CO_2浓度升高对单茎花后干物质积累量、粒重叶比和粒数叶比在不施氮时影响不明显或导致下降，施氮后均使其显著增加。说明改善氮素营养可促进CO_2浓度升高对花后干物质生产的效应，并有利于提高单位叶源的库承载力。
（4）水培条件下，CO_2浓度升高使冬小麦幼苗根冠生物量、根长度、根面积、有机碳分泌速率及总量分别增加1.9%~42.8%、5.5%~27.7%、14.9%~92.9%、5.9%~71.4%、8.7%~11.8%和13.3%，叶片硝酸还原酶活性增加14.8%~33.5%，而对根冠比的影响缺乏规律性。在较低介质供氮水平下CO_2浓度升高对小麦幼苗根长和根面积以及有机碳分泌促进作用较大，表明当氮素营养不足时，CO_2浓度升高更有利于同化产物向根系的分配。
（5）CO_2浓度升高使冬小麦每盆籽粒重量平均增加48.3%~70.0%，增产原因主要是穗数和穗粒数增加的结果。CO_2浓度升高促进了花前不同器官干物质积累，使花前地上部营养器官贮存物质在花后向籽粒的转运量、转运率和贡献率明显增加。CO_2浓度升高对每盆籽粒重量、干物质积累及转运的影响，在不施氮时不显著，施氮后在大多数情况下达显著水平。说明施氮有助于CO_2浓度升高对每盆籽粒重量、花前物质积累和转运的正向效应。
（6）CO_2浓度升高使冬小麦开花期和成熟期植株氮素累积量分别增加15.0%~27.9%和13.9%~26.1%，使花前地上部营养器官氮素在花后向籽粒的转运量提高13.8%~77.4%；但对花前地上部氮素转运率及花后籽粒氮素累积贡献率的影响因品种而异。CO_2浓度升高也可提高氮素籽粒和干物质生产效率和氮素收获指数。不施氮时CO_2浓度升高对氮素累积、转运及籽粒中分配的影响不显著，施氮后显著增加。
（7）增加冬小麦中、后期追氮比例，更有利于发挥CO_2浓度升高对作物生长的促进效应。在不施氮条件下，CO_2浓度升高使冬小麦旗叶SPAD值和可溶性蛋白含量下降，导致旗叶出现明显光合适应；施氮时这些现象均未发生；与一次性底施相比，在施氮比例（底肥﹕拔节肥﹕孕穗肥）为5:5:0和5:3:2时提高了灌浆期旗叶光合速率。结果同时表明，分期施氮比全部底施更有利于增加CO_2浓度升高对小麦生育后期光合能力、花前物质积累与转运、籽粒产量以及氮素利用的促进效应。主要创新点：
（1）在介质供氮不足时，CO_2浓度升高会增加冬小麦叶片可溶性糖累积，导致PSII光损伤，叶绿素含量、PSII的电子传递、原初光化学效率和潜在活性降低，从而引起光合适应。改善氮素营养，特别增加中后施氮比例可减轻冬小麦光合适应，增强CO_2浓度升高对植株生长发育、籽粒产量形成和氮素利用的促进作用。
（2）CO_2浓度升高可促进冬小麦源器官和根系生长发育，增强花前碳氮物质积累和花后向籽粒的转运，增加籽粒产量和氮素累积量，提高氮素生产效率。
（3）综合分析研究结果认为，在介质供氮充分时，CO_2浓度升高导致花后非叶光合源面积增加及促进花前营养器官同化产物向籽粒转运是籽粒产量增加的主要原因。
The research was carried out with winter wheat（Triticum aestivum L.） as materialsduring the three growth seasons from2007to2010. Some pot and water culture experimentswere launched in four open top chambers (OTCs). The main aim of research was to assess theinfluence of CO_2concentration elevation on the photosynthesis, chlorophyll fluorescenceparameters, source components, root morphology and organic carbon secretion, yieldformation, nitrogen use efficency of winter wheat. The main results were as followings：
(1) CO_2enrichment reduced chlorophyll a, chlorophyll b and total chlorophyll contents,increased soluble sugar content in wheat flag leaf, and led to photosynthetic down-regulationunder no nitrogen application or low nitrogen level, but under ample nitrogen application, iteased off or removed photosythesitic down-regulation mainly by increasing chlorophyllcontent, especially chlorophyll a content, and limitedly by chlorophyll b and soluble sugarcontents. This showed nitrogen nutrition improvement could regulate leaf photosythesiticacclimation to some extent.
(2) When air CO_2concentration was elevated, the average value of initialfluorescence(Fo) in wheat flag leaf increased1.4%~15.2%, and the average value ofmaximumu fluorescence(Fm), variable fluorescence(Fv), maximal photochemicalefficiency(Fv/Fm) and potential photochemical efficiency(Fv/Fo) reduced0.3%~9.3%,2.4%~12.0%,0.9%~4.7%and5.9%~22.7%, respectively. The ranges of Fo increase, and Fm,Fv, Fv/Fm and Fv/Fo decrease fell with the increase of nitrogen level. It showed elevated CO_2gave biger photo-damage to wheat PSII, and restrained its electron transport, potential activityand original light conversion ability. Nitrogen addication could weaken or eliminate thenegative effects.
(3) Compared of ambient CO_2concentration, CO_2enrichment increased thephotosynthesis areas of leaf, stem&sheath and whole stem by11.5%~37.3%,14.5%~18.1%and7.3%~38.3%, respectively. The expansion of photosynthesis area should mainly be due tothe increase of leaf and node lengths. The dry matter accumulation after anthesis, the ratios ofgrain number and weight to leaf area responded weakly or negatively to elevated CO_2under no nitrogen application, but significantly increased in the very great majority of cases undernitrogen addication. It showed that the positive effects of CO_2enrichment on dry matterproduction after anthesis and sink bearing capacity per source area could be enhanced byimproving nitrogen nutrition.
(4) The hydroponics results showed that compared with the background CO_2concentration, elevated CO_2increased root and shoot biomass, root length, root area, rootorganic carbon excretion rate and content, and leaf NR activity in wheat seedling by1.9%~42.8%,5.5%~27.7%,14.9%~92.9%,5.9%~71.4%,8.7%~11.8%,13.3%and14.8%~33.5%, but its effect on the root/shoot ratio lacked regularity. Elevated CO_2influnenced root length and area, organic carbon excretion more significantly under lownitrogen nutrition than under apmle nitrogen. It showed that elevated CO_2led to the moreassimilate product distributed to root when lacking nitrogen nutrition.
(5) CO_2enrichment significantly increased wheat grain weight per pot by48.3%~70.0%,mainly due to the improvement of spike number per pot and grain number per spike. ElevatedCO_2promoted the accumulation of dry matter in different organs before anthesis. The amountand rate of pre-anthesis dry matter in shoot transferring to grain after anthesis increased andthe contribution rate of pre-anthesis dry matter translocation rose with CO_2elevation. Theeffects of elevated CO_2on grain weight per pot, dry matter accumulation and translocationwere weak or not significant, but were mostly significant under nitrogen addication, whichindicated reasonable nitrogen application could improve the positive effects of elevated CO_2.
(6) Elevated CO_2increased plant nitrogen accumulation at anthesis and mature stages,the amount of pre-anthesis nitrogen in shoot transferring to grain after anthesis by15.0%~27.9%,13.9%~26.1%and13.8%~77.4%, respectively, but it had different effect onthe contribution rate of pre-anthesis nitrogen in shoot of two varieties. Compared with thebackground CO_2concentration, the high CO_2concentration treatments increased nitrogenefficiencies for grain and dry matter production, and nitrogen harvest index. The effects ofelevated CO_2were not significant under no nitrogen application, and all sigficant undernitrogen application.
(7) The effects of CO_2enrichment on wheat growth could be increased by applying morenitrogen at the middle or later stages. Elevated CO_2resulted in photosynthetic acclimation ofwheat flag leaf along with the decease of SPAD and soluble protein content under nonitrogen application, but the appearances were found under nitrogen addication; thephotosynthetic rate at grain-filling stage was enhanced by elevated CO_2under nitrogenapplications with ratios of5:5:0and5:3:2at before sowing, regreening and booting stage.Additionly, the length and area of the top three leaves at heading stage, photosynthesis after anthesis, grain yield, pre-anthesis dry matter accumulatio and translocation, and nitrogenusage were more easyly promoted by CO_2elevation under nitrogen spilt application thanunder nitrogen all-based application..
Main innovations:
(1) CO_2elevation would down-regulate winter wheat leaf photosythesis by inceasing leafsoluble sugar content and PSII light suppression, and reducing chlorophyll content, PSII lightchemistry efficiency and activity. By improvementing nitrogen nutrition, especiallyapplying more nitrogen at the middle and later stages, the leaf photosynthetic acclimationwould be relieved, and the positive effects of CO_2elevation on plant growth and development,yield formation and nitrogen usage would be strengthened.
(2) CO_2enrichment would promote the growth and development of source organs androot, pre-antheis dry matter and nitrogen accumulation in shoot, and translocation to grainafter anthesis, increase grain yield and nitrigen accumulation in grain, and improve thenitrogen production effciency.
3) The synthetical analyses showed that when nitrogen in medium was ample, thepositively effect of CO_2elevation on winter yield was mainly due to its accelerating thenon-leaf source areas and the translocation of dry matter restored in vegetative organs beforeanthesis to grain after anthesis.
引文
白莉萍,仝乘风,林而达,卢志光,饶敏杰.2004.大气CO2浓度增加对冬小麦品质性状的影响.自然科学进展,14(1):111~115
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