小麦非叶器官的光合特性及其对籽粒产量的贡献
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摘要
以旗叶为对照,比较研究了小麦(Triticum aestivum)叶与非叶器官(穗、穗下节间、旗叶鞘)的光合生理以及对籽粒产量的贡献。结果表明:灌浆初期和中期,小麦穗下节间的叶绿素含量与旗叶差异不显著,但显著高于穗和旗叶鞘。灌浆后期,不同光合器官叶绿素含量由高到低依次为穗下节间、旗叶鞘、旗叶和穗。灌浆初期时叶片的群体光合速率占对照总群体光合的48.5%～50.0%,穗为29.5%～33.2%,穗下节间为12.2%～12.6%。灌浆中期,叶片的群体光合占比下降至42.9%～45.7%,穗提高至32.1%～34.4%。非叶光合器官中的RuBPCase和PEPCK活性在整个灌浆期都高于旗叶,且灌浆中期时穗部的PEPCK活性较灌浆初期时升高,而其他光合器官的PEPCK活性降低。穗下节间的ΦPSII显著大于旗叶,穗的Fv/Fm和ΦPSII均低于旗叶。穗部和叶片是籽粒产量的主要贡献者,其中,叶片的贡献为42.41%～44.97%,穗为17.77%～47.36%。研究结果将为小麦高产栽培的群体调控和品种选育提供理论依据。
The contribution of leaves and non-foliar photosynthetic organs(ear, peduncle and flag leaf sheath) to grain yield and the photosynthesis basis in wheat were studied with flag leaf as reference. The result showed that, no significant difference was found in chlorophyll content between flag leaves and peduncles during early and mid grain-filling stage, while chlorophyll content in ears and in flag leaf sheaths was remarkably lower than that in flag leaves and peduncles. At late grain-filling stage, the highest chlorophyll content was observed in peduncles, followed by flag leaf sheaths, flag leaves and ears. The canopy apparent photosynthetic(CAP) ratio to the total population(CK) of leaves, ears and peduncles were 48.5%–50.0%, 29.5%–33.2% and 12.2%–12.6% at early grain-filling stage. At mid grain-filling stage, the ratio of leaf CAP decreased to 42.9%–45.7%, ear CAP ratio increased to 32.1%–34.4%, compared with CK. RuBPCase activities and PEPCK activities in non-foliar photosynthetic organs were stronger than those in flag leaves during the whole grain-filling stage. PEPCK activities in ears enhanced from early to mid grain-filling stage, while PEPCK activities in other photosynthetic organs decreased. Peduncles performed higher ΦPSIⅡ value than flag leaves did, and the latter showed higher values in Fv/Fm value and ΦPSIⅡ than ears did. Leaves and ears were the dominating contributors to grain yield, with 42.41%–44.97% from leaves and 17.77%–47.36% from ears. The results will provide theoretical supports to cultivar screening and population control technologies of high-yielding wheat.
The contribution of leaves and non-foliar photosynthetic organs(ear, peduncle and flag leaf sheath) to grain yield and the photosynthesis basis in wheat were studied with flag leaf as reference. The result showed that, no significant difference was found in chlorophyll content between flag leaves and peduncles during early and mid grain-filling stage, while chlorophyll content in ears and in flag leaf sheaths was remarkably lower than that in flag leaves and peduncles. At late grain-filling stage, the highest chlorophyll content was observed in peduncles, followed by flag leaf sheaths, flag leaves and ears. The canopy apparent photosynthetic(CAP) ratio to the total population(CK) of leaves, ears and peduncles were 48.5%–50.0%, 29.5%–33.2% and 12.2%–12.6% at early grain-filling stage. At mid grain-filling stage, the ratio of leaf CAP decreased to 42.9%–45.7%, ear CAP ratio increased to 32.1%–34.4%, compared with CK. RuBPCase activities and PEPCK activities in non-foliar photosynthetic organs were stronger than those in flag leaves during the whole grain-filling stage. PEPCK activities in ears enhanced from early to mid grain-filling stage, while PEPCK activities in other photosynthetic organs decreased. Peduncles performed higher ΦPSIⅡ value than flag leaves did, and the latter showed higher values in Fv/Fm value and ΦPSIⅡ than ears did. Leaves and ears were the dominating contributors to grain yield, with 42.41%–44.97% from leaves and 17.77%–47.36% from ears. The results will provide theoretical supports to cultivar screening and population control technologies of high-yielding wheat.
引文
Arnon DI(1949).Copper enzyme in isolated chloroplasts polyphenoloxidase in Beta vulgaris.Plant Physiol,24(1):1-15
Ashcan G,Pfanz H(2003).Non-foliar photosynthesis-a strategy of additional carbon acquisition.Flora,198:81-97
Blanke MM,Ebert G(1992).Phosphoenolpyruvate carboxylase and carbon economy of apple seedlings.Exp Bot,43:965-968
Dong ST(1991).Study on relationship between canopy apparent photosynthesis and grain yield of high-yielding winter wheat.Acta Agron Sin,17(6):461-469(in Chinese with English abstract)[董树亭(1991).高产冬小麦群体光合能力与产量关系的研究.作物学报,17(6):461-469]
Dong ST,Hu CH,Gao RQ(1993).Rates of apparent photosynthesis,respiration and dry matter accumulation in maize canopies.Biol Planta,35(2):273-277
Du JY,Zhou XC,Yang LR(2004).Effect of damage of the photosynthesis organs in different wheat cultivars on grain yield per spike and their compensative effect.J Trit Crops,24(1):35-39(in Chinese with English abstract)[杜久元,周祥春,杨立荣(2004).不同小麦品种植株光合器官受损对单穗籽粒产量的影响及其补偿效应.麦类作物学报,24(1):35-39]
Feng B,Liu P,Li G,et al(2014).Effect of heat stress on the photosynthetic characteristics in flag leaves at the grain-filling stage of different heat-resistant winter wheat varieties.J Agron Crop Sci,4:143-155
Garrity DP,Sullivan CY,Watts DG(1984).Rapidly determining sorghum canopy photosynthetic rates with a mobile field chamber.Agron J,76(1):163-165
Gebbing T,Schnyder H(2001).13C Labeling kinetics of sucrose in glumes indicates significant refixation of respiration CO2 in the wheat ear.Aust J Plant Physiol,28:1047-1053
Gong YH,Lin N,Shi HQ,et al(2016).Canopy temperature characteristics and adaptability of stay-green wheat to low nitrogen and high temperature.J Northwest A&FUniv(Nat Sci Ed),44(9):49-55(in Chinese with English abstract)[龚月桦,林娜,石慧清等(2016).持绿型小麦冠温特性及其对低氮和高温的适应性.西北农林科技大学学报(自然科学版),44(9):49-55]
Guo WS,Feng CN,Yan LL,et al(1995).Analysis on sourcesink relationship after anthesis in wheat.Acta Agron Sin,21(3):334-340(in Chinese with English abstract)[郭文善,封超年,严六零等(1995).小麦开花后源库关系分析.作物学报,21(3):334-340]
Kong LA,Sun MZ,Xie Y,et al(2015).Photochemical and antioxidative responses of the glume and flag leaf to seasonal senescence in wheat.Front Plant Sci,6:358
Kong LA,Wang FH,Feng B,et al(2010).The structural and photosynthetic characteristics of the exposed peduncle of wheat(Triticum aestivum L.):an important photosynthetic source for grain-filling.BMC Plant Biol,10:141
Kong LA,Xie Y,Sun MZ,et al(2016).Comparison of the photosynthetic characteristics in the pericarp and flag leaves during wheat(Triticum aestivum L.)caryopsis development.Photosynthetic,54(1):40-46
Lilley RM,Walker DA(1974).An improved spectrophotometric assay for ribulosebisphosphate carboxylase.Biochem Biophys Acta,358:226-229
Maydup ML,Antonietta M,Guiamet JJ,et al(2010).The contribution of ear photosynthesis to grain filling in bread wheat(Triticum aestivum L.).Field Crops Res,119:48-58
Nedbal L,SoukupováJ,Kaftan D,et al(2000).Kinetic imaging of chlorophyll fluorescence using modulated light.Photosynthesis Res,66:3-12
Niu LY,Feng SW,Ru ZG,et al(2012).Rapid determination of single-stalk and population lodging resistance strengths and an assessment of the stem lodging wind speeds for winter wheat.Field Crops Res,139:1-8
Ru ZG,Feng SW,Li G(2015).High yield potential and effective ways of wheat varieties in Yellow and Huai river valley facultative winter wheat region.Sci Agric Sin,48(17):3388-3393(in Chinese with English abstract)[茹振钢,冯素伟,李淦(2015).黄淮麦区小麦品种的高产潜力与实现途径.中国农业科学,48(17):3388-3393]
Sanchez-Bragado R,Molero G,Reynolds MP,et al(2014).Relative contribution of shoot and ear photosynthesis to grain filling in wheat under good agronomical conditions assessed by differential organδ13C.J Exp Bot,65:5401-5413
Tambussi EA,Bort J,Guiamet JJ,et al(2007).The photosynthetic role of ears in C3 cereals:metabolism,water use efficiency and contribution to grain yield.Crit Rev Plant Sci,26:1
Tambussi EA,Nogués S,Araus,JL(2005).Ear of durum wheat under water stress:water relations and photosynthetic metabolism.Planta,221:446-458
Wang S,Yang JF,Han XR,et al(2008).Effect of fertilizer appliance on photosynthetic characteristic of spring maize.J Soil Fert China,(6):23-27(in Chinese with English abstract)[王帅,杨劲峰,韩晓日等(2008).不同施肥处理对旱作春玉米光合特性的影响.中国土壤与肥料,(6):23-27]
Wang YQ,Xi WX,Wang ZM,et al(2016).Contribution of ear photosynthesis to grain yield under rainfed and irrigation conditions for winter wheat cultivars released in the past 30 years in North China Plain.J Integ Agric,15(10):2247-2256
Wang ZM,Zhang YH,Zhang YP,et al(2004).Review on photosynthetic performance of ear organs in Triticeae crops.J Trit Crops,24(4):136-139(in Chinese with English abstract)[王志敏,张英华,张永平等(2004).麦类作物穗器官的光合作用研究进展.麦类作物学报,24(4):136-139]
Wei AL,Wang ZM(2001).A study on the contribution of different organs to grain weight in different genotype wheat.J Trit Crops,21(2):57-61(in Chinese with English abstract)[魏爱丽,王志敏(2001).小麦不同光合器官对穗粒重的作用及基因型差异研究.麦类作物学报,21(2):57-61]
Wei AL,Zhang YH,Huang Q,et al(2007).Dynamic photosynthetic rate and carbon assimilation enzyme activities of different green photosynthetic organs in different genotypes of wheat.Acta Agron Sin,33(9):1426-1431(in Chinese with English abstract)[魏爱丽,张英华,黄琴等(2007).小麦不同绿色器官光合速率与碳同化酶活性及其基因型差异研究.作物学报,33(9):1426-1431]
Wu YS,Xue H,Liu Y,et al(2010).Study on senescence and chlorophyll fluorescence traits of stay-green leaf in wheat.Agric Res Arid Areas,28(4):117-122(in Chinese with English abstract)[武永胜,薛辉,刘洋等(2010).持绿型小麦叶片衰老和叶绿素荧光特性的研究.干旱地区农业研究,28(4):117-122]
Xu KZ,Miao YN(1983).A study on physio-ecology photosynthesis in soybean III:The morphological and anatomical features of soybean leaves and photosynthetic rate.Soybean Sci,2(3):169-173(in Chinese with English abstract)[徐克章,苗以农(1983).大豆光合生理生态的研究-第3报:大豆叶形态解剖特征与光合作用速率.大豆科学,2(3):169-173]
Xu XL,Wang ZM,Zhang JP(2001).Effect of heat stress on photosynthetic characterisctic of different green organs in wheat.Acta Bot Sin,43(6):571-577(in English with Chinese abstract)[徐晓玲,王志敏,张俊平(2001).灌浆期热胁迫对小麦不同绿色器官光合性能的影响(英文).植物学报,43(6):571-577]
Xue X,Xu HM,Wu HY,et al(2017).Research progress of cyclic electron transport in plant photosynthesis.Plant Physiol J,53(2):145-158(in Chinese with English abstract)[薛娴,许会敏,吴鸿洋等(2017).植物光合作用循环电子传递的研究进展.植物生理学报,53(2):145-158]
Zhang HF,Xu WG,Wang HW,et al(2014).Pyramiding expression of maize genes encoding phosphoenolpyruvate carboxylase(PEPC)and pyruvate orthophosphate dikinase(PPDK)synergistically improve the photosynthetic characteristics of transgenic wheat.Protoplasm,251(5):1163-1173
Zhang YP,Wang ZM,Huang Q,et al(2008).Changes of chloroplast ultramicrostructure and function of different green organs in wheat under limited irrigation.Acta Agron Sin,34(7):1213-1219(in Chinese with English abstract)[张永平,王志敏,黄琴等(2008).不同水分供给对小麦叶与非叶器官叶绿体结构和功能的影响.作物学报,34(7):1213-1219]
Zheng PY,Li XY(1986).Preliminary observation on the morphology of the photosynthetic cells of barley leaf sheath at different positions on the main stem.Acta Agron Sin,12(3):183-187(in Chinese with English abstract)[郑丕尧,李小云(1986).大麦不同叶位叶鞘同化细胞形态结构的初步观察.作物学报,12(3):183-187]
Zheng PY,Li YM(1993).Comparison between the cell morphology and photosynthetic performance of photosynthetic organs of festucoin crops and panicoid crops.Sci Agric Sin,(4):29-35(in Chinese with English abstract)[郑丕尧,李雁鸣(1993).麦类和黍类作物光合器官细胞形态和光合性能的比较.中国农业科学,(4):29-35]
Zhou BW,Serret MD,Elazab A(2016).Wheat ear carbon assimilation and nitrogen remobilization contribute significantly to grain yield.J Integr Plant Biol,58(11):914-926
Zhou B,Sanz-Sáez,á,Elezab A(2014).Physiological traits contributed to the recent increase in yield potential of winter wheat from Henan Province,China.J Integr Plant Biol,56(5):492-504
Ashcan G,Pfanz H(2003).Non-foliar photosynthesis-a strategy of additional carbon acquisition.Flora,198:81-97
Blanke MM,Ebert G(1992).Phosphoenolpyruvate carboxylase and carbon economy of apple seedlings.Exp Bot,43:965-968
Dong ST(1991).Study on relationship between canopy apparent photosynthesis and grain yield of high-yielding winter wheat.Acta Agron Sin,17(6):461-469(in Chinese with English abstract)[董树亭(1991).高产冬小麦群体光合能力与产量关系的研究.作物学报,17(6):461-469]
Dong ST,Hu CH,Gao RQ(1993).Rates of apparent photosynthesis,respiration and dry matter accumulation in maize canopies.Biol Planta,35(2):273-277
Du JY,Zhou XC,Yang LR(2004).Effect of damage of the photosynthesis organs in different wheat cultivars on grain yield per spike and their compensative effect.J Trit Crops,24(1):35-39(in Chinese with English abstract)[杜久元,周祥春,杨立荣(2004).不同小麦品种植株光合器官受损对单穗籽粒产量的影响及其补偿效应.麦类作物学报,24(1):35-39]
Feng B,Liu P,Li G,et al(2014).Effect of heat stress on the photosynthetic characteristics in flag leaves at the grain-filling stage of different heat-resistant winter wheat varieties.J Agron Crop Sci,4:143-155
Garrity DP,Sullivan CY,Watts DG(1984).Rapidly determining sorghum canopy photosynthetic rates with a mobile field chamber.Agron J,76(1):163-165
Gebbing T,Schnyder H(2001).13C Labeling kinetics of sucrose in glumes indicates significant refixation of respiration CO2 in the wheat ear.Aust J Plant Physiol,28:1047-1053
Gong YH,Lin N,Shi HQ,et al(2016).Canopy temperature characteristics and adaptability of stay-green wheat to low nitrogen and high temperature.J Northwest A&FUniv(Nat Sci Ed),44(9):49-55(in Chinese with English abstract)[龚月桦,林娜,石慧清等(2016).持绿型小麦冠温特性及其对低氮和高温的适应性.西北农林科技大学学报(自然科学版),44(9):49-55]
Guo WS,Feng CN,Yan LL,et al(1995).Analysis on sourcesink relationship after anthesis in wheat.Acta Agron Sin,21(3):334-340(in Chinese with English abstract)[郭文善,封超年,严六零等(1995).小麦开花后源库关系分析.作物学报,21(3):334-340]
Kong LA,Sun MZ,Xie Y,et al(2015).Photochemical and antioxidative responses of the glume and flag leaf to seasonal senescence in wheat.Front Plant Sci,6:358
Kong LA,Wang FH,Feng B,et al(2010).The structural and photosynthetic characteristics of the exposed peduncle of wheat(Triticum aestivum L.):an important photosynthetic source for grain-filling.BMC Plant Biol,10:141
Kong LA,Xie Y,Sun MZ,et al(2016).Comparison of the photosynthetic characteristics in the pericarp and flag leaves during wheat(Triticum aestivum L.)caryopsis development.Photosynthetic,54(1):40-46
Lilley RM,Walker DA(1974).An improved spectrophotometric assay for ribulosebisphosphate carboxylase.Biochem Biophys Acta,358:226-229
Maydup ML,Antonietta M,Guiamet JJ,et al(2010).The contribution of ear photosynthesis to grain filling in bread wheat(Triticum aestivum L.).Field Crops Res,119:48-58
Nedbal L,SoukupováJ,Kaftan D,et al(2000).Kinetic imaging of chlorophyll fluorescence using modulated light.Photosynthesis Res,66:3-12
Niu LY,Feng SW,Ru ZG,et al(2012).Rapid determination of single-stalk and population lodging resistance strengths and an assessment of the stem lodging wind speeds for winter wheat.Field Crops Res,139:1-8
Ru ZG,Feng SW,Li G(2015).High yield potential and effective ways of wheat varieties in Yellow and Huai river valley facultative winter wheat region.Sci Agric Sin,48(17):3388-3393(in Chinese with English abstract)[茹振钢,冯素伟,李淦(2015).黄淮麦区小麦品种的高产潜力与实现途径.中国农业科学,48(17):3388-3393]
Sanchez-Bragado R,Molero G,Reynolds MP,et al(2014).Relative contribution of shoot and ear photosynthesis to grain filling in wheat under good agronomical conditions assessed by differential organδ13C.J Exp Bot,65:5401-5413
Tambussi EA,Bort J,Guiamet JJ,et al(2007).The photosynthetic role of ears in C3 cereals:metabolism,water use efficiency and contribution to grain yield.Crit Rev Plant Sci,26:1
Tambussi EA,Nogués S,Araus,JL(2005).Ear of durum wheat under water stress:water relations and photosynthetic metabolism.Planta,221:446-458
Wang S,Yang JF,Han XR,et al(2008).Effect of fertilizer appliance on photosynthetic characteristic of spring maize.J Soil Fert China,(6):23-27(in Chinese with English abstract)[王帅,杨劲峰,韩晓日等(2008).不同施肥处理对旱作春玉米光合特性的影响.中国土壤与肥料,(6):23-27]
Wang YQ,Xi WX,Wang ZM,et al(2016).Contribution of ear photosynthesis to grain yield under rainfed and irrigation conditions for winter wheat cultivars released in the past 30 years in North China Plain.J Integ Agric,15(10):2247-2256
Wang ZM,Zhang YH,Zhang YP,et al(2004).Review on photosynthetic performance of ear organs in Triticeae crops.J Trit Crops,24(4):136-139(in Chinese with English abstract)[王志敏,张英华,张永平等(2004).麦类作物穗器官的光合作用研究进展.麦类作物学报,24(4):136-139]
Wei AL,Wang ZM(2001).A study on the contribution of different organs to grain weight in different genotype wheat.J Trit Crops,21(2):57-61(in Chinese with English abstract)[魏爱丽,王志敏(2001).小麦不同光合器官对穗粒重的作用及基因型差异研究.麦类作物学报,21(2):57-61]
Wei AL,Zhang YH,Huang Q,et al(2007).Dynamic photosynthetic rate and carbon assimilation enzyme activities of different green photosynthetic organs in different genotypes of wheat.Acta Agron Sin,33(9):1426-1431(in Chinese with English abstract)[魏爱丽,张英华,黄琴等(2007).小麦不同绿色器官光合速率与碳同化酶活性及其基因型差异研究.作物学报,33(9):1426-1431]
Wu YS,Xue H,Liu Y,et al(2010).Study on senescence and chlorophyll fluorescence traits of stay-green leaf in wheat.Agric Res Arid Areas,28(4):117-122(in Chinese with English abstract)[武永胜,薛辉,刘洋等(2010).持绿型小麦叶片衰老和叶绿素荧光特性的研究.干旱地区农业研究,28(4):117-122]
Xu KZ,Miao YN(1983).A study on physio-ecology photosynthesis in soybean III:The morphological and anatomical features of soybean leaves and photosynthetic rate.Soybean Sci,2(3):169-173(in Chinese with English abstract)[徐克章,苗以农(1983).大豆光合生理生态的研究-第3报:大豆叶形态解剖特征与光合作用速率.大豆科学,2(3):169-173]
Xu XL,Wang ZM,Zhang JP(2001).Effect of heat stress on photosynthetic characterisctic of different green organs in wheat.Acta Bot Sin,43(6):571-577(in English with Chinese abstract)[徐晓玲,王志敏,张俊平(2001).灌浆期热胁迫对小麦不同绿色器官光合性能的影响(英文).植物学报,43(6):571-577]
Xue X,Xu HM,Wu HY,et al(2017).Research progress of cyclic electron transport in plant photosynthesis.Plant Physiol J,53(2):145-158(in Chinese with English abstract)[薛娴,许会敏,吴鸿洋等(2017).植物光合作用循环电子传递的研究进展.植物生理学报,53(2):145-158]
Zhang HF,Xu WG,Wang HW,et al(2014).Pyramiding expression of maize genes encoding phosphoenolpyruvate carboxylase(PEPC)and pyruvate orthophosphate dikinase(PPDK)synergistically improve the photosynthetic characteristics of transgenic wheat.Protoplasm,251(5):1163-1173
Zhang YP,Wang ZM,Huang Q,et al(2008).Changes of chloroplast ultramicrostructure and function of different green organs in wheat under limited irrigation.Acta Agron Sin,34(7):1213-1219(in Chinese with English abstract)[张永平,王志敏,黄琴等(2008).不同水分供给对小麦叶与非叶器官叶绿体结构和功能的影响.作物学报,34(7):1213-1219]
Zheng PY,Li XY(1986).Preliminary observation on the morphology of the photosynthetic cells of barley leaf sheath at different positions on the main stem.Acta Agron Sin,12(3):183-187(in Chinese with English abstract)[郑丕尧,李小云(1986).大麦不同叶位叶鞘同化细胞形态结构的初步观察.作物学报,12(3):183-187]
Zheng PY,Li YM(1993).Comparison between the cell morphology and photosynthetic performance of photosynthetic organs of festucoin crops and panicoid crops.Sci Agric Sin,(4):29-35(in Chinese with English abstract)[郑丕尧,李雁鸣(1993).麦类和黍类作物光合器官细胞形态和光合性能的比较.中国农业科学,(4):29-35]
Zhou BW,Serret MD,Elazab A(2016).Wheat ear carbon assimilation and nitrogen remobilization contribute significantly to grain yield.J Integr Plant Biol,58(11):914-926
Zhou B,Sanz-Sáez,á,Elezab A(2014).Physiological traits contributed to the recent increase in yield potential of winter wheat from Henan Province,China.J Integr Plant Biol,56(5):492-504
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