棉花非叶绿色器官光合特性及对水分亏缺的适应机制
|
摘要
作物叶片通常被认为是主要光合器官，但许多作物的非叶绿色器官或组织，也能形成或含有叶绿素，并具有实际或潜在的光合能力，对产量形成具有一定的贡献。随着作物产量水平的提高，进一步增加单产的难度加大。重视作物非叶绿色器官的光合能力，提高作物整体光能利用效率，对作物生产水平的提高具有重要意义。本研究从棉花非叶绿色器官光合作用对产量形成的贡献、叶片和非叶绿色器官光合特性差异、水分亏缺条件下非叶绿色器官的光合生理适应性等3个方面入手，较系统的研究了棉花产量形成期各绿色器官的光合特性，揭示了非叶绿色器官光合作用对棉花产量形成的重要作用，明确了棉花不同非叶绿色器官光合作用及光保护机制，探讨了水分亏缺下棉花不同绿色器官光合作用的适应性。研究结果对进一步挖掘棉花光合物质生产潜力，提高单产奠定了理论基础，为棉花高产栽培及抗逆品种选育提供了理论依据。
1．研究了棉花不同生育时期叶片、主茎秆、苞叶和铃壳的光合面积、光合放氧能力和光合关键酶活性的变化，揭示了棉花各绿色器官光合作用对产量的相对贡献率。在棉花生育后期，叶片开始衰老，非叶绿色器官的光合面积增加至整体植株的38.2%；苞叶和铃壳的光合放氧能力和光合关键酶活性的下降幅度均低于叶片。结合各器官光合面积及其光合放氧能力，估算出各绿色器官光合对植株整体产量的相对贡献率，其中棉花生育后期茎杆和果实（铃壳和苞叶）对植株整体的相对贡献率分别达12.7%和23.7%；棉铃和茎杆对铃重的相对贡献率分别为24.1%和9%。因此，棉花生育后期非叶绿色器官的光合作用对其产量形成具有重要的作用。
2．研究了不同棉花品种（杂交种）非叶绿色器官的光合作用特性，阐明了杂交棉超高产形成的生理机理。研究表明，开花后5-15天，2个杂交棉新陆早43号和石杂2号叶片的光放氧能力较常规品种新陆早33号高，盛花期干物质积累速率快；随开花后天数的延长，石杂2号各绿色器官光合性能均表现出明显的下降趋势；3个不同品种（杂交种）间茎秆单位面积的光合放氧速率无显著差异，2个杂交棉生育后期茎秆具有较高的群体光合速率主要是由于株高及茎秆光合面积较高所致；与常规品种新陆早33号相比，2个杂交棉铃壳具有较高的群体光合速率，且在生育后期仍能维持较稳定的值，与杂交棉单株结铃较多、铃壳总光合面积较大有关，这是杂交棉新陆早43号和石杂2号生育后期仍能保持较高光合能力的重要原因。
3．由于棉铃具有较高的呼吸速率，在苞叶和棉铃之间形成一个高浓度CO_2的微环境。本研究揭示了棉花苞叶适应高浓度CO_2微环境的生理机制。测定结果表明，光照下棉铃（苞叶和铃）的胞间CO_2浓度是500-1300μmol·mol~(-1)。推理认为这种高浓度CO_2微环境早在110万年前四倍体棉花出现时就存在。因此，推测认为棉花苞叶具有适应高浓度CO_2微环境的能力，并从棉花叶片、苞叶的形态和生理等方面来检验所提出的这个假说。结果表明，与叶片相比，较低气孔导度的苞叶具有较高的瞬时水分利用效率；气体交换和蛋白组分分析均表明苞叶具有较高的J_(max)/V_(cmax)比率和Rieske FeS/Rubisco比值。这些结果均与理论上提出的植物适应高浓度CO_2的机制一致，这表明苞叶可作为研究植物适应高浓度CO_2的材料。
4．PSII的光失活可导致植物光化学效率下降，产量降低。植物体形成了多种保护机制来保护PSII免受光失活，然而有关非叶绿色器官光保护机制的研究较少。本文研究了棉花叶片、苞叶、主茎秆和铃壳的光保护机制，结果表明，苞叶具有较低的抗氧化酶活性，较高的ΔpH-叶黄素循环热耗散（Φ_(NPQ)）；主茎秆则倾向于通过依赖光和非依赖光的光化学耗散及较好的抗氧化酶体系；铃壳主要是通过其非常强的抗氧化酶体系和丰富的类胡萝卜素含量来进行热耗散，因为它具有较低的Φ_(NPQ)。因此，棉花各绿色器官具有不同的光保护机制。
5．本研究提出了一种采用P700氧化还原动力学原理非损伤测定棉花叶片的光失活速率常数（ki）和修复速率常数（k_r）的新方法。P700氧化还原动力学面积与气相氧电极所测得的单闪脉冲的放氧量呈线型相关，表明该方法可以精确测定功能性的PSII的相对含量。采用此方法，研究了棉花叶片不同轴面接触溶液（封闭气孔效应）条件下ki和k_r的变化规律。结果表明，ki与光强有关，远轴面朝向水溶液的棉花叶片ki略高些。黑暗中两种方式放置的棉花叶片k_r相差不大。当近轴面朝向水溶液时其k_r由光强决定。当远轴面朝向水溶液时，在中度光强下k_r达到峰值，而在高光强下k_r减小。当近轴面朝向水溶液时，其k_r在弱光下先增强，此后平缓增加，高光强下k_r显著增加。高光强下，近、远轴面分别朝向水溶液叶片之间的k_r存在很大差异，这可能是由于不同程度的氧胁迫导致的修复速率常数不一致。P700氧化还原动力学测定方法能快速、无损伤地测定棉花圆叶片的功能性PSII的相对含量，对研究棉花叶片光合能力差异及其内在机制具有重要意义，为未来通过P700氧化还原动力学方法深入分析棉花非叶绿色器官的光合能力差异及其内在机制提供了重要的技术支撑。
6．水分亏缺是导致作物产量降低的重要因素。研究了水分亏缺对棉花各绿色器官光合生理变化的影响机制，包括各绿色器官水分状况、光合速率、RuBPC活性及保护酶活性的变化等。水分亏缺严重降低了棉花各绿色器官的光合面积，但非叶绿色器官的下降幅度相对较小；随棉铃的发育，水分亏缺下棉花非叶绿色器官（苞叶和铃壳）的光合放氧速率、RuBPC和抗氧化酶活性的下降幅度均较叶片小，棉花非叶绿色器官的光合作用对棉株生物量积累的相对贡献率增加。
Although leaf was considered as the main photosynthetic organ, many parts of crop suchas non-foliar organs which contain chlorophyll can also performance photosynthesis,contributing to carbon acquisition and yield. This research was conducted from threeperspectives:1) changes in photosynthetic capacity of non-foliar organs in cotton and relativecontribution to yield during growth stages;2) differences in photosynthesis charactertics ofnon-foliar organs in cotton and3) the photosynthetic adaptations of green organs in cottonrespond to water stress. On the basis of the three points above, photosynthetic charactertics,relative contribution to its yield and adaptation respond to water stress of non-foliar organs incotton were studied to elucidate a) the changes in photosynthetic rate of green organs incotton during growth stages and the important photosynthetic contribution of non-foliar organto yield formation especially at later growth stage, b) differences in photosynthesischaractertic and photoprotection mechanism of non-foliar organ, and c) effects of waterdeficit on physiological process and the relative contribution of each organ to yield in cottonwas explored. This research may help to increase the potential of photosynthate production,thereby improving cotton yield. Additionally, the results obtained can provide theoreticalbasis for high-yielding cotton production and drought-tolerant cotton breeding. Below are themain results in this thesis:
1. The relative photosynthetic contribution of leaves, main stem, bracts and capsule wallsin cotton was revealed by measuring their time-course of surface area development, O2evolution capacity and photosynthetic enzyme activity. Because of the early senescence ofleaves, non-foliar organs increased their surface area up to38.2%of total at late growth stage.Bracts and capsule wall showed less ontogenetic decrease in O2evolution capacity per areaand photosynthetic enzyme activity than leaves at the late growth stage. The total capacity forO2evolution of stalks and bolls (bracts plus capsule wall) was12.7%and23.7%, respectively,as estimated by multiplying their surface area by their O2evolution capacity per area. We alsokept the bolls (from15days after anthesis) or main stem (at the early full bolling stage) indarkness for comparison with non-darkened controls. Darkening the bolls and main stemreduced the boll weight by24.1%and9%, respectively. Thus, we conclude that non-foliarorgans significantly contribute to the yield at the late growth stage.
2. The photosynthetic charactertics of non-foliar organs in super-high-yielding cottonduring growth stages were explored. Compared with traditional cotton Xinluzao33, thephotosynthetic rate of the main leaf of hybrid cotton Xinluzao43and Shiza2was higherduring5-15days after anthesis, which reason for faster dry matter accumulation rate ofXinluzao43and Shiza2cotton during early growth stage (Full Flowering). However, more decrease in photosynthetic rate of green organs of Shiza2during the growth stages, whichcaused its early senescence. There was no significance in photosynthetic rate of main stemamong these three cultivars. Thus, we suggested that the high height and larger surface area ofhybrid cotton result in their better canopy photosynthetic rate of stalks at later growth stage.The photosynthetic rate of capsule wall in hybrid cotton Xinluzao43and Shiza2was higherthan that in traditional cotton Xinluzao33, and maintain stable photosynthetic rate during latergrowth stage, company with their larger surface area which all contribution their highercanopy photosynthetic rate of fruits at later growth stage.
3. The rapid respiration rate of cotton (Gossypium hirsutum L.) fruits (bolls) produces aconcentrated CO_2micro-environment around the bolls and bracts. Elucidation of themechanisms by which plants adapt to elevated CO_2is needed. However, most studies of themechanisms investigated the response of plants adapted to current atmospheric CO_2. Weobserved that the intercellular CO_2concentration of a whole fruit (bract and boll) ranged from500to1300μmol mol1depending on the irradiance, even in ambient air. Arguably, this CO_2micro-environment has existed for at least1.1million years since the appearance of tetraploidcotton. Therefore, we hypothesized that the mechanisms by which cotton bracts have adaptedto elevated CO_2will indicate how plants will adapt to future increased atmospheric CO_2concentration. To test the hypothesis, the morphological and physiological traits of bracts andleaves of cotton were measured, including stomatal density, gas-exchange and proteincontents. Compared with leaves, bracts showed significantly smaller stomatal conductancewhich resulted in a significantly higher water use efficiency. Both gas-exchange and proteincontent showed a significantly greater RuBP regeneration/RuBP carboxylation capacity ratio(J_(max)/V_(cmax)) in bracts than in leaves. These results agree with the theoretical prediction thatadaptation of photosynthesis to elevated CO_2requires increased RuBP regeneration. Cottonbracts are readily-available material for studying adaption to elevated CO_2.
4. Photoinactivation of Photosystem II (PS II) during photosynthesis can lead to the lossof photochemical efficiency and decrease in crop yield. Plants have evolved variousphotoprotective strategies to ameliorate photoinactivation of PS II. Non-leaf organs of cottonalso contribute to carbon gain, but it is not clear how they photoprotect themselves. This studyinvestigated the photoprotective mechanisms in the leaf, bract, main stem and capsule wall ofcotton. Our results suggested that the bract mainly relies on high activities of antioxidativeenzymes and high ΔpH-and xanthophyll-regulated thermal dissipation (Φ_(NPQ)) forphotoprotection. The main stem preferentially dissipated its absorbed light energy vialight-regulated as well as light-independent non-photochemical quenching, aided by themoderately high activities of antioxidative enzymes. The capsule wall was less able to removereactive oxygen species due to lower activities of antioxidative enzymes, and less able todissipate energy via heat due to its lower Φ_(NPQ). Its main photoprotective mechanisms seem tobe (a) direct quenching of the energy by abundant carotenoids and (b) light-independentconstitutive thermal dissipation via Φf,D. The green organs of cotton have different ways touse or dissipate energy.
5. Here we report on a whole-tissue determination of the rate coefficient ofphotoinactivation ki, and that of repair k_rin cotton leaf discs. The P700kinetics area, directlyproportional to the oxygen yield per single-turnover, saturating flash, was used to obtain bothkiand k_r. Changes in kiand k_rof two orientations (abaxial/adaxial surface contacting water)were investigated in this study. The value of ki, directly proportional to irradiance, wasslightly higher when CO_2diffusion into the abaxial surface (richer in stomata) was blocked bycontact with water. The value of k_r, sizable in darkness, changed in the light depending onwhich surface was blocked by contact with water. When the abaxial surface was blocked, k_rfirst peaked at moderate irradiance and then decreased at high irradiance. When the adaxialsurface was blocked, k_rfirst increased at low irradiance, then plateaued, before increasingmarkedly at high irradiance. At the highest irradiance, k_rdiffered by an order of magnitudebetween the two orientations, attributable to different extents of oxidative stress affectingrepair. The P700kinetics area is a rapid, whole-tissue and in vivo measurement of relativeassay of functional PSII content, can help to explore the differences and underlyingmechanisms in photosynthetic capacity of leaf in cotton and also provide technical basis forexploring the differences and underlying mechanisms in photosynthetic capacity of variousgreen organs in cotton.
6. Water deficit is one of the most important causes of decreased yield in cultivatedplants. The physiological response of each organ to water deficit has not beencomprehensively studied in relation to the water status, photosynthetic rate, photosyntheticenzyme activity, antioxidant enzymes. Water deficit significantly decreased the surface area ofeach organ, but to a lesser extent in non-foliar organs. Non-foliar organs (bracts and capsulewall) showed less ontogenetic decrease in O2evolution capacity and in RuBPC activity (perdry weight) as well as better antioxidant systems than leaves at various days after anthesis. Addition ally, the relative contribution of biomass accumulation of non-foliar organs to thewhole cotton plant was estimated from the surface area, photosynthetic duration and the O2evolution capacity per area. Our result s showed that the relative contribution of biomassaccumulation of non-foliar organs increased under water deficit.
1．研究了棉花不同生育时期叶片、主茎秆、苞叶和铃壳的光合面积、光合放氧能力和光合关键酶活性的变化，揭示了棉花各绿色器官光合作用对产量的相对贡献率。在棉花生育后期，叶片开始衰老，非叶绿色器官的光合面积增加至整体植株的38.2%；苞叶和铃壳的光合放氧能力和光合关键酶活性的下降幅度均低于叶片。结合各器官光合面积及其光合放氧能力，估算出各绿色器官光合对植株整体产量的相对贡献率，其中棉花生育后期茎杆和果实（铃壳和苞叶）对植株整体的相对贡献率分别达12.7%和23.7%；棉铃和茎杆对铃重的相对贡献率分别为24.1%和9%。因此，棉花生育后期非叶绿色器官的光合作用对其产量形成具有重要的作用。
2．研究了不同棉花品种（杂交种）非叶绿色器官的光合作用特性，阐明了杂交棉超高产形成的生理机理。研究表明，开花后5-15天，2个杂交棉新陆早43号和石杂2号叶片的光放氧能力较常规品种新陆早33号高，盛花期干物质积累速率快；随开花后天数的延长，石杂2号各绿色器官光合性能均表现出明显的下降趋势；3个不同品种（杂交种）间茎秆单位面积的光合放氧速率无显著差异，2个杂交棉生育后期茎秆具有较高的群体光合速率主要是由于株高及茎秆光合面积较高所致；与常规品种新陆早33号相比，2个杂交棉铃壳具有较高的群体光合速率，且在生育后期仍能维持较稳定的值，与杂交棉单株结铃较多、铃壳总光合面积较大有关，这是杂交棉新陆早43号和石杂2号生育后期仍能保持较高光合能力的重要原因。
3．由于棉铃具有较高的呼吸速率，在苞叶和棉铃之间形成一个高浓度CO_2的微环境。本研究揭示了棉花苞叶适应高浓度CO_2微环境的生理机制。测定结果表明，光照下棉铃（苞叶和铃）的胞间CO_2浓度是500-1300μmol·mol~(-1)。推理认为这种高浓度CO_2微环境早在110万年前四倍体棉花出现时就存在。因此，推测认为棉花苞叶具有适应高浓度CO_2微环境的能力，并从棉花叶片、苞叶的形态和生理等方面来检验所提出的这个假说。结果表明，与叶片相比，较低气孔导度的苞叶具有较高的瞬时水分利用效率；气体交换和蛋白组分分析均表明苞叶具有较高的J_(max)/V_(cmax)比率和Rieske FeS/Rubisco比值。这些结果均与理论上提出的植物适应高浓度CO_2的机制一致，这表明苞叶可作为研究植物适应高浓度CO_2的材料。
4．PSII的光失活可导致植物光化学效率下降，产量降低。植物体形成了多种保护机制来保护PSII免受光失活，然而有关非叶绿色器官光保护机制的研究较少。本文研究了棉花叶片、苞叶、主茎秆和铃壳的光保护机制，结果表明，苞叶具有较低的抗氧化酶活性，较高的ΔpH-叶黄素循环热耗散（Φ_(NPQ)）；主茎秆则倾向于通过依赖光和非依赖光的光化学耗散及较好的抗氧化酶体系；铃壳主要是通过其非常强的抗氧化酶体系和丰富的类胡萝卜素含量来进行热耗散，因为它具有较低的Φ_(NPQ)。因此，棉花各绿色器官具有不同的光保护机制。
5．本研究提出了一种采用P700氧化还原动力学原理非损伤测定棉花叶片的光失活速率常数（ki）和修复速率常数（k_r）的新方法。P700氧化还原动力学面积与气相氧电极所测得的单闪脉冲的放氧量呈线型相关，表明该方法可以精确测定功能性的PSII的相对含量。采用此方法，研究了棉花叶片不同轴面接触溶液（封闭气孔效应）条件下ki和k_r的变化规律。结果表明，ki与光强有关，远轴面朝向水溶液的棉花叶片ki略高些。黑暗中两种方式放置的棉花叶片k_r相差不大。当近轴面朝向水溶液时其k_r由光强决定。当远轴面朝向水溶液时，在中度光强下k_r达到峰值，而在高光强下k_r减小。当近轴面朝向水溶液时，其k_r在弱光下先增强，此后平缓增加，高光强下k_r显著增加。高光强下，近、远轴面分别朝向水溶液叶片之间的k_r存在很大差异，这可能是由于不同程度的氧胁迫导致的修复速率常数不一致。P700氧化还原动力学测定方法能快速、无损伤地测定棉花圆叶片的功能性PSII的相对含量，对研究棉花叶片光合能力差异及其内在机制具有重要意义，为未来通过P700氧化还原动力学方法深入分析棉花非叶绿色器官的光合能力差异及其内在机制提供了重要的技术支撑。
6．水分亏缺是导致作物产量降低的重要因素。研究了水分亏缺对棉花各绿色器官光合生理变化的影响机制，包括各绿色器官水分状况、光合速率、RuBPC活性及保护酶活性的变化等。水分亏缺严重降低了棉花各绿色器官的光合面积，但非叶绿色器官的下降幅度相对较小；随棉铃的发育，水分亏缺下棉花非叶绿色器官（苞叶和铃壳）的光合放氧速率、RuBPC和抗氧化酶活性的下降幅度均较叶片小，棉花非叶绿色器官的光合作用对棉株生物量积累的相对贡献率增加。
Although leaf was considered as the main photosynthetic organ, many parts of crop suchas non-foliar organs which contain chlorophyll can also performance photosynthesis,contributing to carbon acquisition and yield. This research was conducted from threeperspectives:1) changes in photosynthetic capacity of non-foliar organs in cotton and relativecontribution to yield during growth stages;2) differences in photosynthesis charactertics ofnon-foliar organs in cotton and3) the photosynthetic adaptations of green organs in cottonrespond to water stress. On the basis of the three points above, photosynthetic charactertics,relative contribution to its yield and adaptation respond to water stress of non-foliar organs incotton were studied to elucidate a) the changes in photosynthetic rate of green organs incotton during growth stages and the important photosynthetic contribution of non-foliar organto yield formation especially at later growth stage, b) differences in photosynthesischaractertic and photoprotection mechanism of non-foliar organ, and c) effects of waterdeficit on physiological process and the relative contribution of each organ to yield in cottonwas explored. This research may help to increase the potential of photosynthate production,thereby improving cotton yield. Additionally, the results obtained can provide theoreticalbasis for high-yielding cotton production and drought-tolerant cotton breeding. Below are themain results in this thesis:
1. The relative photosynthetic contribution of leaves, main stem, bracts and capsule wallsin cotton was revealed by measuring their time-course of surface area development, O2evolution capacity and photosynthetic enzyme activity. Because of the early senescence ofleaves, non-foliar organs increased their surface area up to38.2%of total at late growth stage.Bracts and capsule wall showed less ontogenetic decrease in O2evolution capacity per areaand photosynthetic enzyme activity than leaves at the late growth stage. The total capacity forO2evolution of stalks and bolls (bracts plus capsule wall) was12.7%and23.7%, respectively,as estimated by multiplying their surface area by their O2evolution capacity per area. We alsokept the bolls (from15days after anthesis) or main stem (at the early full bolling stage) indarkness for comparison with non-darkened controls. Darkening the bolls and main stemreduced the boll weight by24.1%and9%, respectively. Thus, we conclude that non-foliarorgans significantly contribute to the yield at the late growth stage.
2. The photosynthetic charactertics of non-foliar organs in super-high-yielding cottonduring growth stages were explored. Compared with traditional cotton Xinluzao33, thephotosynthetic rate of the main leaf of hybrid cotton Xinluzao43and Shiza2was higherduring5-15days after anthesis, which reason for faster dry matter accumulation rate ofXinluzao43and Shiza2cotton during early growth stage (Full Flowering). However, more decrease in photosynthetic rate of green organs of Shiza2during the growth stages, whichcaused its early senescence. There was no significance in photosynthetic rate of main stemamong these three cultivars. Thus, we suggested that the high height and larger surface area ofhybrid cotton result in their better canopy photosynthetic rate of stalks at later growth stage.The photosynthetic rate of capsule wall in hybrid cotton Xinluzao43and Shiza2was higherthan that in traditional cotton Xinluzao33, and maintain stable photosynthetic rate during latergrowth stage, company with their larger surface area which all contribution their highercanopy photosynthetic rate of fruits at later growth stage.
3. The rapid respiration rate of cotton (Gossypium hirsutum L.) fruits (bolls) produces aconcentrated CO_2micro-environment around the bolls and bracts. Elucidation of themechanisms by which plants adapt to elevated CO_2is needed. However, most studies of themechanisms investigated the response of plants adapted to current atmospheric CO_2. Weobserved that the intercellular CO_2concentration of a whole fruit (bract and boll) ranged from500to1300μmol mol1depending on the irradiance, even in ambient air. Arguably, this CO_2micro-environment has existed for at least1.1million years since the appearance of tetraploidcotton. Therefore, we hypothesized that the mechanisms by which cotton bracts have adaptedto elevated CO_2will indicate how plants will adapt to future increased atmospheric CO_2concentration. To test the hypothesis, the morphological and physiological traits of bracts andleaves of cotton were measured, including stomatal density, gas-exchange and proteincontents. Compared with leaves, bracts showed significantly smaller stomatal conductancewhich resulted in a significantly higher water use efficiency. Both gas-exchange and proteincontent showed a significantly greater RuBP regeneration/RuBP carboxylation capacity ratio(J_(max)/V_(cmax)) in bracts than in leaves. These results agree with the theoretical prediction thatadaptation of photosynthesis to elevated CO_2requires increased RuBP regeneration. Cottonbracts are readily-available material for studying adaption to elevated CO_2.
4. Photoinactivation of Photosystem II (PS II) during photosynthesis can lead to the lossof photochemical efficiency and decrease in crop yield. Plants have evolved variousphotoprotective strategies to ameliorate photoinactivation of PS II. Non-leaf organs of cottonalso contribute to carbon gain, but it is not clear how they photoprotect themselves. This studyinvestigated the photoprotective mechanisms in the leaf, bract, main stem and capsule wall ofcotton. Our results suggested that the bract mainly relies on high activities of antioxidativeenzymes and high ΔpH-and xanthophyll-regulated thermal dissipation (Φ_(NPQ)) forphotoprotection. The main stem preferentially dissipated its absorbed light energy vialight-regulated as well as light-independent non-photochemical quenching, aided by themoderately high activities of antioxidative enzymes. The capsule wall was less able to removereactive oxygen species due to lower activities of antioxidative enzymes, and less able todissipate energy via heat due to its lower Φ_(NPQ). Its main photoprotective mechanisms seem tobe (a) direct quenching of the energy by abundant carotenoids and (b) light-independentconstitutive thermal dissipation via Φf,D. The green organs of cotton have different ways touse or dissipate energy.
5. Here we report on a whole-tissue determination of the rate coefficient ofphotoinactivation ki, and that of repair k_rin cotton leaf discs. The P700kinetics area, directlyproportional to the oxygen yield per single-turnover, saturating flash, was used to obtain bothkiand k_r. Changes in kiand k_rof two orientations (abaxial/adaxial surface contacting water)were investigated in this study. The value of ki, directly proportional to irradiance, wasslightly higher when CO_2diffusion into the abaxial surface (richer in stomata) was blocked bycontact with water. The value of k_r, sizable in darkness, changed in the light depending onwhich surface was blocked by contact with water. When the abaxial surface was blocked, k_rfirst peaked at moderate irradiance and then decreased at high irradiance. When the adaxialsurface was blocked, k_rfirst increased at low irradiance, then plateaued, before increasingmarkedly at high irradiance. At the highest irradiance, k_rdiffered by an order of magnitudebetween the two orientations, attributable to different extents of oxidative stress affectingrepair. The P700kinetics area is a rapid, whole-tissue and in vivo measurement of relativeassay of functional PSII content, can help to explore the differences and underlyingmechanisms in photosynthetic capacity of leaf in cotton and also provide technical basis forexploring the differences and underlying mechanisms in photosynthetic capacity of variousgreen organs in cotton.
6. Water deficit is one of the most important causes of decreased yield in cultivatedplants. The physiological response of each organ to water deficit has not beencomprehensively studied in relation to the water status, photosynthetic rate, photosyntheticenzyme activity, antioxidant enzymes. Water deficit significantly decreased the surface area ofeach organ, but to a lesser extent in non-foliar organs. Non-foliar organs (bracts and capsulewall) showed less ontogenetic decrease in O2evolution capacity and in RuBPC activity (perdry weight) as well as better antioxidant systems than leaves at various days after anthesis. Addition ally, the relative contribution of biomass accumulation of non-foliar organs to thewhole cotton plant was estimated from the surface area, photosynthetic duration and the O2evolution capacity per area. Our result s showed that the relative contribution of biomassaccumulation of non-foliar organs increased under water deficit.
引文
[1]杜明伟,罗宏海,张亚黎等.新疆超高产杂交棉的光合生产特征研究.中国农业科学,2009a,42(6):1952-1962
[2]杜明伟,冯国艺,姚炎帝等.杂交棉标杂A1和石杂2号超高产冠层特性及其与群体光合生产的关系.作物学报,2009b,35(6):1068-1077
[3]冯国艺.超高产棉花冠层结构形成机理及调控研究:石河子大学博士学位论文.石河子:石河子大学,2012
[4]李卫华,郝乃斌. C3植物中C4途径研究进展.植物学通报,1999,16(2):97-106
[5]李寒冰,胡玉熹,白克智等.水稻稃片和旗叶叶绿体超微结构低温荧光特性进行了比较研究.电子显微学报,2002,21(3):341-344
[6]李秀菊,职明星,石晓华等.小麦穗光合对不同花位籽粒及颖壳的影响.麦类作物学报,2006,26(5):146-148
[7]卢存福,贲桂英.高海拔地区植物的光合特性.植物学通报,1995,12(2):38-42
[8]施生锦,黄彬香,王志敏.小麦不同光合器官的光合性能研究.中国生态农业学报,2003,11(4):10-11
[9]薛丽华,章建新.大豆鼓粒期非叶光合器官与粒重的关系.大豆科学,2006,25(4):524-428.
[10]魏爱丽,王志敏.野生一粒麦与普通栽培小麦不同绿色器官光合特性与叶绿体结构特征.作物学报,2002,28(3):351-354
[11]魏爱丽,王志敏,翟志席等.土壤干旱对小麦旗叶和穗器官C4光合酶活性的影响.中国农业科学,2003,36(5):508-512
[12]魏爱丽,张英华,黄琴等.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,2007,33:(9)1426-1431
[13]张亚黎,冯国艺,胡渊渊等.棉花非叶器官光合能力的差异及与物质生产的关系.作物学报,2010,36(4):701-708
[14]张永平,王志敏,吴永成等.不同供水条件下小麦不同绿色器官的气孔特性研究.作物学报,2006,32(1):70-75
[15]中国农业科学院棉花研究所.中国棉花栽培学.上海:上海科学技术出版社,1983
[16] Abbad H, Jaafrai SE, Bort J, Araus JL. Comparison of flag leaf and ear photosynthesis with biomass and grain yield ofdurum wheat under various water conditions and genotypes. Agronomie,2004,24:19-28
[17] Ainsworth EA&Long SP. What have we learned from15years of free-air CO2enrichment (FACE)? A meta-analyticreview of the responses of photosynthesis, canopy properties and plant production to rising CO2. New Phytol,2005,165(2):351-371
[18] Anderson JM. Insights into the consequences of grana stacking of thylakoid membranes in vascular plants:a personalperspective. Aust J Plant Physiol,1999,26(7):625-639
[19] Aoyagi K&Bassham JA. Pyruvate orthophosphate dikinase of C3seeds and leaves as compared to the enzyme frommaize. Plant Physiol,1984,75:387-392
[20] Araus JL, Brown HR, Febrero A, Bort J, Serret MD. Ear photosynthesis, carbon isotope discrimination and thecontribution of respiratory CO2to differences in grain mass in durum wheat. Plant Cell Environ,1993,16(4):383-392
[21] Aro E-M, McCaffery S, Anderson JM. Photoinhibition and D1protein degradation in peas acclimated to differentgrowth irradiance. Plant Physiol,1993a,103:835-843
[22] Aro E-M, Virgin I, Andersson B. Photoinhibition of photosystem II. Inactivation, protein damage and turnover.Biochim Biophys Acta,1993b,1143(2):113-134
[23] Asada K. The water-water cycle in chloroplasts: scavenging of active oxygen and dissipation of excess photons. RevPlant Physiol Mol Biol,1999,50:601-639
[24] Aschan G&Pfanz H. Non-foliar photosynthesis-a strategy of additional carbon acquisition. Flora,2003,198(2):81-97.
[25] Aschan G, Pfanz H, Vodnik D, Bati F. Photosynthetic performance of vegetative and reproductive structures of greenhellebore (Helleborus viridis L. agg.). Photosynthetica,2005,43(1):55-64
[26] Aschan G&Pfanz H. Why snowdrop (Galanthus nivalis L.) tepals have green marks? Flora,2006,8(23):623-632
[27] Ashraf M&Foolad MR. Roles of glycine betaine and proline in improving plant abiotic stress resistance. Environ ExpBot,2007,59(2):206-216
[28] Athar H&Ashraf M. Photosynthesis under drought stress. In: Pessarakli, M.(Ed.), Handbook Photosynthesis, seconded. CRC Press, New York, USA,2005,795-810
[29] Bachmann KM, Ebbert V, Adams WW III, Verhoven AS, Logan BA, Demmig-Adams B. Effects of lincomycin on PSIIefficiency, non-photochemical quenching, D1protein and xanthophylls cycle during photoinhibition and recovery. FuncPlant Biol,2004,31:803-813
[30] Bajji M, Lutts S, Kinet JM. Water deficit effects on solute contribution to osmotic adjustment as a function of leafageing in three durum wheat (Triticum durum Desf.) cultivars performing differently in arid conditions. Plant Sci,2001,160(4):669-681
[31] Barlow EWR, Lee JW, Rana Munns Smart MG. Water relations of the developing wheat grain. Aust J Plant Physiol,1980,7(5):519-525
[32] Bates LS, Waldren RP, Teare JD. Rapid determination of proline for water stress studies.Plant Soil,1973,39(1):205-207
[33] Ben Ahmed C, Ben Rouina B, Athar HR, Boukhris M. Olive tree (Oleaeuropaea L. cv. Chemlali) under salt stress:water relations and ions contents. Pak J Bot,2006,38(5):1477-1484
[34] Ben Ahmed C, Ben Rouina B, Boukhris M. Effects of water deficit on olive trees cv. Chemlali under field conditions inarid region in Tunisia. Sci Hort,2007,113(3):267-277
[35] Ben Ahmed C, Ben Rouina B, Sensoy S, Boukhris M, Ben Abdallah F. Changes in gas exchange, proline accumulationand antioxidative enzyme activities in three olive cultivarsunder contrastingwater availability regimes. Environ Exp Bot,2009,67(2):345-352
[36] Bettarini I., Vaccari FP, Miglietta F. Elevated CO2concentrations and stomatal density-observations from17plantspecies growing in a CO2spring in central Italy. Global Change Biology,1998,4(1):17-22
[37] Bhatt JG. Transport of radioactivity in relation to bracts in the cotton plant. Ann Bot,1988,62(6):571-573
[38] Birkhold KB, Koch KE, Darnell RL. Carbon and nitrogen economy of developing rabbiteye blueberry fruit. J Am SocHortic Sci,1992,117(1):139-145
[39] Biswall B. Carotenoid catabolism during leaf senescence and its control by light. J Photochem Photobiol B,1995,30(1):3-14
[40] Bj rkman O. Carboxydismutase activity in shade-adapted and sun-adapted species of higher plants. PhysiologiaPlantarum,1968,21(1):1-10
[41] Bj rkman O. Response to different quantum flux densities. In Lange OL, Nobel PS, Osmond CB, Ziegler H, eds.Physiological Plant Ecology I. Responses to the Physical Environment, Encyclopedia of plant physiology, new series.Springer-Verlag: Berlin,1981,57-107
[42] Black CC&Vines HM. Alternative plant photosynthetic CO2fixation cycles.In DW Newman,KG Wilson,eds,Modelsin Plant Physioligy and Biochemistry,Vol1.CRC Press.BocaRaton, FL,1987,57-61
[43] Blanke MM&Lenz F. Fruit photosynthesis a review. Plant Cell Environ,1989,12(1):31-46
[44] Blanke MM&Ebert G. Phosphoenolpyruvate carboxylase and carbon economy of apple seedlings. J Exp Bot,1992,43(7):965-968
[45] Blum A. Photosynthesis and transpiration in leaves and ears of wheat and barley varieties. J Exp Bot,1985,36(3):432-440
[46] Blum A. The effects of heat stress on wheat leaf and ear photosynthesis. J Exp Bot,1986,37(174):111-118
[47] Blum A, Sinmena B, Mayer J, Golan G and Shpiler L (1994) Stem reserve mobilization supports wheat-grain fillingunder heat stress. Australian Journal of Plant Physiology,21(6):771-781
[48] Bondada BR, Oosterhuis DM, Wullschleger SD, Kim KS, Harris WM. Anatomical considerations related tophotosynthesis in cotton leaves, bracts, and the capsule wall. J Exp Bot,1994,45(1):111-118
[49] Bondada BR, Oosterhuis DM, Murphy JB, Kim KS. Effect of water stress on the epicuticular wax composition andultrastructure of cotton leaf, bract, and boll. Environ Exp Bot,1996,36(1):61-69
[50] Bondada BR&Oosterhuis DM. Comparative epidermal ultrastructure of cotton (Gossypium hirsutum L.) leaf, bractand capsule wall. Ann Bot,2000,86(6):1143-1152
[51] Bondada BR&Oosterhuis DM. Morphometic analysis of chloroplasts of cotton leaf and fruiting organs.Biol Plantarum,2003,47(2):281-284
[52] Bort J, Febrero A, Amaro T, Araus JL. Role of awns in ear water-use efficiency and grain weight in barley. Agronomie,1994,2:133-139
[53] Bort J, Brown RH, Araus JL. Refixation of respiratory CO2in the ears of C3-cereals. J Exp.Bot,1996,47(10):1567-1575.
[54] Boyer JS. Plant productivity and environment. Sci,1982,218(4571):443-448
[55] Bravd BA, Palgi A, Lurie S. Changing ribulose diphosphate carboxylase/oxygenase activity in ripening tomato fruit.Plant Physiol,1977,60:309-312
[56] Brown KJ. Translocation of carbohydrates in cotton: Movement to the fruiting bodies.Ann Bot,1968,32(4):703-713
[57] Caley CY, Duffus CM, Jeffcoat B. Photosynthesis in the pericarp of developing wheat grains. J Exp Bot,1990,41(3):303-307
[58] Caldwell MM, Osmond CB, Nott DL. C4pathway photosynthesis at low temperature in cold-tolerant Atriplex species.Plant Physiol,1977,60:157-164
[59] Camp PJ, Huber SC, Burke JJ, Moreland DE. Biochemical changes that occur during senescence of wheat leaves. I.Basis for the reduction of photosynthesis. Plant Physiol,1982,70:1641-1646
[60] Chapin III FS. The mineral nutrition of wild plants. Annu Rev Ecol Syst,1980,11:233-260
[61] Chaves MM. Effects of water deficits on carbon assimilation. J Exp Bot,1991,42(1):1-16
[62] Chaves MM, Maroco JP, Pereira JS. Understanding plant responses to drought-from genes to the whole plant. FunctPlant Biol,2003,30(3):239-264
[63] Chow WS&Aro E-M. Photoinactivation and mechanisms of recovery. In: Wydrzynski T and Satoh K (eds)Photosystem II: the light-driven water: plastoquinone oxidoreductase. Advances in photosynthesis and respiration, vol22Springer, Dordrecht,2005,627-648
[64] Chow WS&Hope AB. Electron fluxes through Photosystem I in cucumber leaf discs probed by far-red light.Photosynth Res,2004,81:77-89
[65] Chow WS, Hope AB, Anderson JM. Oxygen per flash from leaf disks quantifies photosystem II. Biochim Biophys Acta,1989,973(1):105-108
[66] Chow WS, Fan D-Y, Oguchi R, Jia H, Losciale P, Park Y-I, He J, quist G, Shen Y-G, Anderson JM. Quantifying andmonitoring functional Photosystem II and the stoichiometry of the two photosystems in leaf segments: Approaches andapproximations. Photosynth Res,2012,113(1):63-74
[67] Claussen W. Proline as a measure of stress in tomato plants. Plant Sci,2005,168(1):241-248
[68] Cockfield SD. Relative availability of nitrogen in host plants of invertebrate herbivores: three possible nutritional andphysiological definitions. Oecologia,1988,77:91-94
[69] Condon AG, Richards RA,Farquhar GD. Carbon isotope discrimination is positively correlated with grain yield and drymatter production in filed-grown wheat. Crop Sci,1987,996-1001
[70] Constable GA&Rawson HM. Carbon production and utilization in cotton: Inferences from a carbon budget. Aust JPlant Physiol,1980,7(5):539-553
[71] Cornic G. Drought stress and high light effects on leaf photosynthesis. In Photoinhibition of Photosynthesis: FromMolecular Mechanisms to the Field. Eds. N R Baker and J R Bowyer. Oxford BIOS Scientific Publishers.1994,297-313
[72] Cornic G. Drought stress inhibits photosynthesis by decreasing stomatal aperture-not by affecting ATP systhesis. TrendsPlant Sci,2000,5:187-188
[73] Crookston RK, O'Toole J, Ozbun JL. Characterization of the bean pod as a photosynthetic organ. Crop Sci,1974,14:708-712
[74] Cruz-Aguado JA, Rodés R, Pérez IP, Dorado M. Morphological characteristics and yield components associated withaccumulation and loss of dry mass in the internodes of wheat. Field Crops Research,2000,66(2):129-139
[75] Demmig-Adams B&Adams III WW. Photoprotection and other responses of plants to high light stress. Ann Rev PlantPhysiol PlantMol Biol,1992,43(1):599-626
[76] Drake BG, Gonzalez-Meler MA, Long SP. More efficient plants-a consequence of rising atmospheric CO2. Annu RevPlant Physiol Plant Mol Bio,1997, l48:609-639
[77] Duffus CM&Cochrane MP. Formation of the barley grain-morphology, physiology, and biochemistry. In: MacGregorAW, Bhatty RS (eds) Barley: chemistry and technology. American Association of Cereal Chemists, St. Paul,1993,31-72
[78] Ehdaie B, Alloush GA, Waines JG. Genotypic variation in linear rate of grain growth and contribution of stem reservesto grian yield in wheat. Field Crops Research,2008,106(1):34-43
[79] Eamus D. The interaction of rising CO2and temperatures with water use efficiency. Plant Cell Environ,1991,14:843-582
[80] Ehsanzadeh P, Mahmoudieh R, Fareed N. Photosynthetic contribution of the inflorescence and adjacent green tissue todiverse environmental conditions. J Biol Sci,2007,7(2):263-269
[81] Elmore CD. Contributions of the capsule wall and bracts to the developing cotton fruits. Crop Sci1973,13:751-752
[82] Elstner EF. Metabolism of activated oxygen species. In: Davies DD (ed.):The biochemistry of Plants, Biochemistry ofMetabolism. Academic Press, San Diego.1987, Vol11.253-315
[83] Endress PK. Floral phyllotaxis and floral evolution. Botanische Jahrbücher für Systematik,1987,108:417-438
[84] Endress PK. Evolutionary diversification of the flowers in Angiosperms. Am J Bot,2011,98(3):370-396
[85] Evans LT, Bingham J, Jackson P, Sutherland J. Effect of awns and drought on the supply of photosynthate and itsdistribution within wheat ears. Ann Appl Biol,1972,70(1):67-76
[86] Evans JR. Photosynthesis and nitrogen relationships in leaves of C3plants. Oecologia,1989,78(1):9-19
[87] Evans JR, Jakobsen I, gren E. Photosynthetic light-response curves2. Gradients of light-absorption andphotosynthetic capacity. Planta,1993,189:191-200
[88] Evans JR. Potential errors in electron transport rates calculated from chlorophyll fluorescence as revealed by amultilayer leaf model. Plant Cell Physiol,2009,50(4):698-706
[89] Ewart AJ. On assimilatory inhibition in plants. J Linn Soc,1896,31(217):364-461
[90] Farquhar GD, von Caemmere S, Berry JA. A biochemical model of photosynthetic CO2assimilation in leaves of C3species. Planta,1980,149(1):78-90
[91] Field C, Mooney H. The photosynthesis-nitrogen relationship in wild plants. In On the Economy of Plant Form andFunction. Ed. G.T. Givnish. Cambridge University Press,1986,25-55
[92] Foyer CH, Descourvieres P, Kunert KJ. Protection against oxygen radicals: an important defence mechanism studied intransgenic plants. Plant Cell Environ,1994,17:507-523
[93] Fu J&Huang B. Involvement of antioxidants and lipid peroxidation in the adaptation of two cool-season grasses tolocalized drought stress. Environ Exp Bot,2001,45(2):105-114
[94] Gebbing T, Schnyder H.13C Clabelling kinetics of sucrose in glumes indicates significant refixation of respiratory CO2in the wheat ear. Aust J Plant Physiol,2001,28(10):1047-1053
[95] Genty B, Briantais JM, Baker NR. The relationship between the quantum yield of photosynthetic electron transport andquenching of chlorophyll fluorescence. Biochim Biophys Acta,1989,990(1):87–92
[96] Grossman A, Bartlett S, Chua N-H. Energy-dependent uptake of cytoplasmically synthesized polypeptide bychloroplasts. Nature,1980,285:625-628
[97] Gunderson CA&Wullschleger SD. Photosynthetic acclimation in trees to rising atmospheric CO2: a broaderperspective. Photosynth Res,1994,39(3):369-388
[98] Gupta AS, Webb RP, Holaday AS and Allen RD. Overexpression of superoxide dismutase protects plants fromoxidative stress (Induction of ascobate peroxidase in superoxide dismutase-overpressing plants). Plant Physiol,1993,103:1067-1073
[99] Hattenschwiler S, Miglietta F, Raschi A, Korner C. Thirty years of in situ tree growth under elevated CO2: a model forfuture forest responses? Global Change Biol,1997,3:463-471
[100] He J&Chow WS. The rate coefficient of repair of photosystem II after photoinactivation. Physiol Plant,2003,118(2):297-304
[101] Hong SS&Xu DQ. Light-induced increase in initial chlorophyll fluorescence Folevel and the reversible inactivation ofPSII reaction centers in soybean leaves. Photosynth Res,1999,61(3):269-280
[102] Hedley CL, Harvey DM, Keely RJ. Role of PEP carboxylase during seed development in Pisum sativum. Nature,1975,258:352-354
[103] Hemmers H and Gulz PG. Chemistry and morphology of epicuticular waxes from leaves of five Euphorbia species. ZNatuforsch,1986,41(5-6):521-525
[104] Hendrickson L, Furbank RT, Chow WS. A simple alternative approach to assessing the fate of absorbed light energyusing chlorophyll fluorescence. Photosynth Res,2004,82:73-81
[105] Hikosaka K&Hirose T. Leaf and canopy photosynthesis of C3plants at elevated CO2in relation to optimal partitioningof nitrogen among photosynthetic components: theoretical prediction. Ecol Model,1998,106(2-3):247-259
[106] Hikosaka K. Nitrogen partitioning in the photosynthetic apparatus of Plantago asiatica leaves grown under differenttemperature and light conditions: similarities and differences between temperature and light acclimation. Plant CellPhysiol,2005,46(8):1283-1290
[107] Hodges DM, DeLong JM, Forney CF, Prange RK. Improving the thiobarbituric acid-reactive-substances assay forestimating lipid peroxidation in plant tissues containing anthocyanin and other interfering compounds. Planta,1999,207(4):604-611
[108] Hormaetxe K, Becerril JM, Fleck I, Pinto M, Garcia-Plazaola JI. Functional role of red (retro)-carotenoids as passivelight filters in theleaves of Buxus sempervirens L.: increased protection of photosynthetic tissues? J Exp Bot,2005,56(420):2629-2636
[109] Horton P. Prospects for crop improvement through the genetic manipulation of photosynthesis: morphological andbiochemical aspects of light capture. J Exp Bot,2000,51(suppl1):475-485
[110] Imaizumi N, Usuda H, Nakamoto H and Ishihara K. Changes in the rate of photosynthesis during grain filling and theenzymatic activities associated with the photosynthetic carbon metabolism in rice panicles. Plant Cell Physiol,1990,31(6):835-843.
[111] Ishihara K, Kiyota E, Imaizumi N. On the contribution of panicle photosynthesis to grain yield in rice plants. Jpn JCrop Sci,1991,60(Extra Issue1):122-123
[112] Johnson RR&Moss DN. Effect of water stress on14CO2fixation and translocation in wheat during grain filling. CropSci,1976,16:697-701
[113] Johnson DA, Richards RA, and Turner NC. Yield, water, relations, gas exchange, and surface reflectances of nearisogenic wheat lines differing in glaucousness. Crop Sci,1983,23(2):318-325
[114] Jones MB, Brown JC, Raschi A, Miglietta F. The effects on Arbutus unedo L. of long-term exposure to elevated CO2.Global Change Biology,1995,1(4):295-302
[115] Jordan WR, Shouse PJ, Blum A, Miller FR, Monk RL. Environmental physiology of sorghum. II. Epicuticular wax loadand cuticular transpiration. Crop Sci,1984,24(6):1168-1173
[116] Kato MC, Hikosaka K, Hirotsu N, Makino A, Hirose T. The excess light energy that is neither utilized in photosynthesisnor dissipated by photoprotective mechanisms determines the rate of photoinactivation in Photosystem II. Plant CellPhysiol,2003,44(3):318-325
[117] Kikuta SB&Richter H. Graphical evaluation and partitioning of turgor responses to drought in leaves of durum wheat.Planta,1986,168:36-42
[118] Kimball BA, Kobayashi K, Bindi M. Responses of agricultural crops to free-air CO2enrichment. Adv Agron,2002,77:293-368
[119] Kingston-Smith AH, Harbinson J, Williams J, Foyer CH. Effect of chilling on carbon assimilation, enzyme activation,and photosynthetic electron transport in the absence of photoinhibition in maize leaves. Plant Physiol,1997,114(3):1039-1046
[120] Knoppik D, Selinger H, Ziegler-J ns A. Differences between the flag leaf and the ear of a spring wheat cultivar(Triticum aestivum cv. Arkas) with respect to the CO2response of assimilation, respiration and stomatal conductance.Physiol Plant,1986,68(3):451-457
[121] Kocheva K, Lambrev P, Georgiev G, Goltsev V, Karabaliev M. Evaluation of chlorophyll fluorescence and membraneinjury in the leaves of barley cultivars under osmotic stress. Bioelectrochemistry,2004,63(1-2):121-124
[122] Kou J, Oguchi R, Fan D-Y, Chow WS. The time course of photoinactivation of photosystem II in leaves revisited.Photosynth Res,2012,113(1-3):157-164
[123] Lawlor DW, Kontturi M, Young AT. Photosynthesis by flag leaves of wheat in relation to protein, ribulose bisphosphatecarboxylase activity and nitrogen supply. J Exp Bot,1989,40(1):43-52
[124] Lawlor DW. The effects of water deficit on photosynthesis. In Environment and Plant Metabolism. Flexibility andAcclimation, Ed. N Smirnoff. Oxford: BIOS Scientific Publisher.1995, pp.129-160
[125] Lawlor DW. Limitation to photosynthesis in water-stressed leaves: stomata vs. metabolism and the role of ATP. AnnBot,2002,89(7):871-885
[126] Lawlor DW&Cornic G. Photosynthetic carbon assimilation and associated metabolism in relation to water deficits inhigher plants. Plant Cell Environ,2002,25(2):275-294
[127] Lee H-Y, Hong Y-N, Chow WS. Photoinactivation of photosystem II complexes and photoprotection by non-functionalneighbours in Capsicum annuum L. leaves. Planta,2001,212(3):332-342
[128] Levizou E, Petropoulou Y, Manetas Y. Carotenoid composition of peridermal twigs does fully conform to a shadeacclimation hypothesis. Photosynthetica,2004,42(4):591-596
[129] Li XJ, Wang HG, Li HB, Zhang LY, Teng NJ, Lin QQ, Wang Jian, Kuang TY, Li ZS, Li B, Zhang AM and Lin JX. Awnplay a dominant role in carbohydrate production during the grain filling stages in wheat (Triticum aestivum).Physiol Plantarum,2006,127(4):701-709
[130] Lichtenthaler HK. Chlorophylls and carotenoids: pigments of photosynthetic biomembranes. Methods Enzymol,1987,148:350-382
[131] Lizana C, Wentworth M, Martínez JP, Villegas D, Meneses R, Murchie EH, Pastenes C, Lercari B, Vernieri P, Horton P,Pinto M. Differential adaptation of two varieties of common bean to abiotic stress. I. Effects of drought on yield andphotosynthesis. J Exp Bot,2006,57(3):685-697
[132] Long SP, Ainsworth EA, Rogers A, Ort DR. Rising atmospheric carbon dioxide: plants face the future. Annu Rev PlantBiol,2004,55:591-628
[133] Losciale P, Oguchi R, Hendrickson L, Hope AB, Corelli-Grappadelli L, Chow WS. A rapid, whole-tissue determinationof the functional fraction of Photosystem II after photoinhibition of leaves based on flash-induced P700redox kinetics.Physiol Plant,2008,132(1):23-32
[134] Matto AK, Hoffman-Falk H, Marder JB, Edelman M. Regulation of protein metabolism: Coupling of photosyntheticelectron transport to in vivo degradation of the rapidly metabolized32-kilodalton protein of the chloroplast membranes.Proc Natl Acad Sci USA,1984,81(5):1380-1384
[135] Makino A, Mae T, Ohira K. Photosynthesis and ribulose-1,5-bisphosphate carboxylase in rice leaves. Plant Physiol,1983,73(4):1002-1007
[136] Makino A&Osmond B. Effects of nitrogen nutrition on nitrogen partitioning between chloroplasts and mitochondria inpea and wheat. Plant Physiol,1991,96(2):355-362
[137] Makino A, Harada M, Kaneko K, Mae T, Shimada T, Yamamoto N. Whole-plant growth and N allocation in transgenicrice plants with decreased content of ribulose-1,5-bisphosphate carboxylase under different CO2partial pressures. AustJ Plant Physiol,2000,27(1):1-12
[138] Martinez DE, Luquez VM, Bartoli CG, Guizmét JJ. Persistence of photosynthetic components and photochemical effectin ears of water-stressed wheat(Triticum aestivum). Physiol Plantarum,2003,119(4):519-525
[139] Maydup ML, Anotonietta M, Guiamet JJ, Graciano C, López, Tambussi EA. The contribution of ear photosynthesis tograin filling in bread wheat (Triticum aestivum L.). Field Crop Research.,2010,119(1):48-58
[140] McKee IF&Woodward FI. The effect of growth at elevated CO2concentrations on photosynthesis in wheat. Plant CellEnviron,1994,17(7):853-859
[141] Meehl GA, Stocker TF, Collins WD, Friedlingstein P, Gaye AT, Gregory JM, Kitoh A, Knutti R, Murphy JM, Noda A,Raper SCB, Watterson IG, Weaver AJ, Zhao Z-C. Global Climate Projections. In Solomon S, Qin D, Manning M, ChenZ, Marquis M, Averyt KB, Tignor M, Miller HL, eds. Climate Change2007: The Physical Science Basis. Contributionof Working Group I to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change. CambridgeUniversity Press: Cambridge and New York,2007,747-845
[142] Medlyn BE. The optimal allocation of nitrogen within the C3photosynthetic system at elevated CO2. Aust J PlantPhysiol,1996,23(5):593-603
[143] Medrano H, Escalona JM, Bota J, Gulías J and Flexas J. Regulation of photosynthesis of C3plants in response toprogressive drought: stomatal conductance as a reference parameter. Ann Bot,2002,89(7):895-905
[144] Melis A. Photosystem-II damage and repair cycle in chloroplasts: what modulates the rate of photodamage in vivo?Trends Plant Sci,1999,4(4):130-135
[145] Miglietta F&Raschi A. Studying the effect of elevated CO2in the open in a naturally enriched environment in CentralItaly.1993, Vegetatio104/105:391-400
[146] Mooney HA&Gulmon SL. Environmental and evolutionary constraints on the photosynthetic characteristics of higherplants. In: Solbrig OT, Jain S, Johnson GB, Raven PH, eds. Topics in plant population biology. New York, NY, USA:Columbia University Press,1979,316-337
[147] Morgan JM. Osmotic adjustment in the spikelets and leaves of wheat. J Exp Bot,1980,31(2):655-665
[148] Morgan JM. Osmoregulation and water-stressed in higher plants. Annu Rev Plant Physiol,1984,35(1):299-319.
[149] Mulchi CL, Slaughter L, Saleem M, Lee EH, Pausch R, Rowland R. Growth and physiological characteristics ofsoybean in open top chambers in response to ozone and increased atmospheric CO2. Agr Ecosyst Environ,1992,38(1-2):107-118
[150] Müller P, Li XP, Niyogi KK. Non-photochemical quenching. A response to excess light energy. Plant Physiol,2001,125(4):1558-1566
[151] Nakano H, Makino A, Mae T. The effect of elevated partial pressures of CO2on the relationship betweenphotosynthetic capacity and N content in rice leaves. Plant Physiol,1997,115(1):191-198
[152] Nayyar H. Accumulation of osmolytes and osmotic adjustment in water-stressed wheat (Triticum aestivum) and maize(Zea mays) as affected by calcium and its antagonists. Environ Exp Bot,2003,50(3):253-264
[153] Nie GY, Long SP, Garcia RL, Kimball BA, Lamorte RL, Pinter PJ, Wall GW, Webber AN. Effects of free-air CO2enrichment on the development of the photosynthetic apparatus in wheat, as indicated by changes in leaf protein. PlantCell Environ,1995,18(8):855-864
[154] Nishimyama Y, Yamamoto H, Allakhverdiev SI, Inaba M, Yokota A, Murata N. Oxidative stress inhibits the repair ofphotodamage to the photosynthetic machinery. EMBO J,2001,20(20):5587-5594
[155] Nishimyama Y, Allakhverdiev SI, Murata N. Protein synthesis is the primary target of reactive oxygen species in thephotosystem II. Physiol Plant,2011,142(1):35-46
[156] Oguchi R, Terashima I, Chow WS. The involvement of dual mechanisms of photoinactivation of photosystem II inCapsicum annuum L. plants. Plant Cell Physiol,2009,50(10):1815-1825
[157] Oguchi R, Terashima I, Kou J, Chow WS. Operation of dual mechanisms that both lead to photoinactivation ofPhotosystem II in leaves by visible light. Physiol Plant,2011a,142(1):47-55
[158] Oguchi R, Douwstra P, Fujita T, Chow WS, Terashima I. Intra-leaf gradients of photoinhibition induced by differentcolor lights: Implications for the dual mechanisms of photoinhibition and for the application of conventionalchlorophyll fluorometers. New Phytol,2011b,191(1):146-159
[159] Ohad I, Berg A, Berkowicz SM, Kaplan A, Keren N. Photoinactivation of photosystem II: is there more than one way toskin a cat? Physiol Plant,2011,142(1):79-86
[160] Onoda Y, Hikosaka K, Hirose T. Seasonal change in the balance between capacities of rubp carboxylation and rubpregeneration affects CO2response of photosynthesis in Polygonum cuspidatum. J Expl Bot,2005,56(412):755-763.
[161] Onoda Y, Hirose T, Hikosaka K. Effect of elevated CO2levels on leaf starch, nitrogen and photosynthesis of plantsgrowing at three natural CO2springs in Japan. Ecol Res,2007,22(3):475-484
[162] Onoda Y, Hirose T, Hikosaka K. Does leaf phootosynthesis adapt to CO2-enriched environments? An experiment onplants originating from three natural CO2springs. New Phytol,2009,182(3):698-709
[163] Oosterhuis DM, Chipamaunga J, Bate GC. Nitrogen uptake of field-grown cotton. I. Distribution in plant componentsin relation to fertilization and yield. Expl Agric,1983,19(1):91-101
[164] Paoletti E, Nourrisson G, Garrec JP, Raschi A. Modifications of the leaf surface structures of Quercus ilex L. in open,naturally CO2-enriched environments. Plant Cell Environ,1998,21(10):1071-1075
[165] Pérez-Alfocea F, Estan MT, Caro M, Guerrier G. Osmotic adjustment in Lycopersicon esculentum and L. pennelliiunder NaCl and polyethylene glycol6000iso-osmotic stresses. Physiol Plant,1993,87(4):493-498
[166] Pheloung PC&Siddique KHM. Contribution of stem dry matter to grain yield in wheat cultivars. Aust J Plant Physiol,1991,18(1):53-64
[167] Pic E, Tyssender de la Serve B, Tardieu F, Turc O. Leaf senescence induced by mild water deficiet follows the samesequence of macroscopic, bichemical and molecular events as monocarpic senescence in pea. Plant Physiol,2002,128(1):236-246
[168] Pinheiro HA, DaMatta FM, Chaves ARM, Fontes EPB, Loureiro ME. Drought tolerance in relation to protectionagainst oxidative stress in clones of Coffea canephora subjected to long-term drought. Plant Sci,2004,167(6):1307-1314
[169] Polle A, Otter T, Seifert F. Apoplastic peroxidases and lignification in needles of Norway spruce (Picea abies L.). PlantPhysiol,1994,106(1):53-60
[170] Poole I, Weyers JDB, Lawson T, Raven JA. Variations in stomatal density and index: implications for palaeoclimaticreconstructions. Plant Cell Environ,1996,19(6):705-12
[171] Porra RJ, Thompson WA, Kriedemann PE. Determination of accurate extinction coefficients and simultaneousequations for assaying chlorophylls a and b extracted with four different solvents: verification of the concentration ofchlorophyll standards by atomic absorption spectroscopy. BBA-Bioenergetics,1989,975(3):384-394
[172] Powles SB. Photoinhibition of photosynthesis induced by visible light. Annu Rev Plant Physiol,1984,35(1):15-44
[173] Prásil O, Adir N, Ohad I. Dynamics of photosystem II: mechanism of photoinhibition and recovery processes. In:Barber J (ed) Topics in Photosyntheses, Vol.11, Elsevier, Amsterdam,1992, pp.295-348
[174] Prestidge RA&McNeill S. The role of nitrogen in the ecology of grassland Auchenorryncha. In Lee JA, McNeil S andRorison IH (eds) Nitrogen as an Ecological Factor. Blackwell Scientific Publishers, Oxford,1983, pp257-281
[175] Price GD, von Caemmerer S, Evans JR, Siebke K, Anderson JM, Badger MR. Photosynthesis is strongly reduced byantisense suppression of chloroplastic cytochrome bf complex in transgenic tobacco. Aust J Plant Physiol,1998,25(4):445-452
[176] Pugnaire FI, Haase P, Incoll LD, Clark SC. Response of the tussock grass Stipa tenacissima to watering in a semi-aridenvironment. Funct Ecol,1996,10:265-274
[177] Read SM&Northcote DH. Minimization of variation in the response to different proteins of the Coomassie blue G dyebinding assay for protein. Anal Biochem,1981,116(1):53-64
[178] Reddy AR, Chaitanya KV, Jutur PP, Sumithra K. Differential antioxidative responses to water stress among fivemulberry (Morus alba L.) cultivars. Environ Exp Bot,2004,52(1):33-42
[179] Specht JE, Hume DJ, Kumudini SV. Soybean yield potential-a genetic and physiological perspective. Crop Science,1999,39(6):1560-1570
[180] Rodermel S, Haley J, Jiang CZ, Tsai CH, Bogorad L. A mechanism for intergenomic integration: abundance of ribulosebisphosphate carboxylase small-subunit protein influences the translation of the large-subunit mRNA. Proceedings ofthe National Academy of Sciences of the United States of America,1996,93(9):3881-3885
[181] Royer DL. Stomatal density and stomatal index as indicators of paleoatmospheric CO2concentration. Rev PalaeobotPalyno,2001,114(1-2):1-28
[182] Sage RF. Acclimation of photosynthesis to increasing atmospheric CO2: the gas exchange perspective. Photosynthe Res,1994,39(3):351-368
[183] Salvador RJ&Pearce RB. Husk removal and its effects on maize grain yield.Crop Sci,1988,28(6):961-964
[184] Sánchez-Díaz M, García JL, Antolín MC, Araus JL. Effects of soil drought and atmospheric humidity on yield, gasexchange and stable carbon composition of barley. Photosynthetica,2002,40(3):415-421
[185] Sayre RT, Kennedy RA, Pringnitz DJ. Photosynthetic enzyme activities and localization in Mollugo verticillatapopulations differing in the levels of C3and C4cycle operation. Plant Physiol,1979,64(2):293-299
[186] Scarascia-Mugnozza G, De Angelis P, Matteucci G, Valentini R. Long-term exposure to elevated [CO2] in a naturalQuercus ilex L. community: net photosynthesis and photochemical efficiency of PSII at different levels of water stressPlant Cell Environ,1996,19(6):643-654
[187] Schuman GE, Stanley MA, Knudsen D. Automated Total Nitrogen Analysis of Soil and Plant Samples. Soil Sci Soc AmJ,1972,37(3):480-481
[188] Serraj R&Sinclair TR. Osmolyte accumulation: can it really help increase crop yield under drought conditions? PlantCell Environ,2002,25(2):333-341
[189] Sinclair TR&Ludlow MM. Who taught plants thermodynamics? The unfulfilled potential of plant water potential.Aust. J Plant Physiol,1985,12(3):213-217
[190] Singal HR, Sheoran IS, Singh R. In vitro enzyme activites and products of14CO2assimilation in flag leaf and ear partsof wheat (Triticum aestivum L.).Photosynthe Res,1986,8(2):113-122
[191] Smirnoff N. Tansley Review No.52. The role of active oxygen in the response of plants to water deficit and desiccation.New Phytol,1993,125:27-58
[192] Sofo A, Dichio B, Xiloyannis C, Masia A. Lipoxygenase activity and proline accumulation in leaves and roots of olivetree in response to drought stress. Physiol Plant,2004,121(1):58-65
[193] Specht JE, Hume DJ, Kumudini SV. Soybean yield potential-a genetic and physiological perspective. Crop Science.,1999,39(6):1560-1570
[194] Spence RD, Wu H, Sharpe PJH, Clark KG. Water-stressed effects on guard cell anatomy and the mechanical advantageof the epidermal cells. Plant Cell Environ,1986,9(3):197-202
[195] Stitt M. Rising carbon dioxide levels and their potential significance for carbon flow in photosynthetic cells. Plant, CellEnviron,1991,14(8):741-762
[196] Suzuki Y, Makino A. Availability of Rubisco small subunit up-regulates the transcript levels of large subunit forstoichiometric assembly of its holoenzyme in rice. Plant Physiol,2012,160(1):533-540
[197] Tambussi EA, Nogués S, Araus JL. Ear of durum wheat under water stress water relations and photosynthesismetabolism. Planta,2005,221(3):446-458
[198] Tambussi EA, Bort J, Guiamet JJ, Nogues S, Araus JL. The photosynthetic role of ears in C3cereals: metabolism,water use efficiency and contribution to grain yield. Crit Rev Plant Sci,2007,26(1):1-16
[199] Taniguchi M, Kuroda H, Satoh K. ATP-dependent protein synthesis in isolated pea chloroplasts: Evidence foraccumulation of a translation intermediate of the D1protein. FEBS Lett,1993,317(1-2):57-61
[200] Terashima I, Fujita T, Inoue T, Chow WS, Oguchi R. Green light drive leaf photosynthesis more efficiently than redlight in strong white light: Revisiting the enigmatic question of why leaves are green. Plant Cell Physiol,2009,50(4):674-697
[201] Tyystj rvi E, Aro E-M. The rate constant of photoinhibition, measured in lincomycin-treated leaves, is directlyproportional to light intensity. Proc Natl Acad Sci USA,1996,93(5):2213-2218
[202] Tezara W, Mitchell V, Driscoll SP, Lawlor DW. Effects of water deficit and its interaction with CO2supply on thebiochemistry and physiology of photosynthesis in sunflower. J Exp Bot,2002,53(375):1781-1791
[203] Ting IP&Osmond CB. Photosynthetic phosphoenolpyruvate carboxylases characteristics of alloenzymes from leavesof C3and C4plants. Plant Physiol,1973,51(3):439-447
[204] Tognetti R, Giovannelli A, Longobucco A, Miglietta F, Raschi A. Water relations of oak species growing in the naturalCO2spring of Rapolano (central Italy). Ann For Sci,1996,53(2-3):475-485
[205] Tsuyama M, Shibata M, Kobayashi Y. Leaf factors affecting the relationship between chlorophyll fluorescence and therate of photosynthetic electron transport as determined from CO2uptake. J Plant Physiol,2003,160(10):1131-1139
[206] Van Gorkom HJ, Schelvis JPM. Kok’s oxygen clock: What makes it tick? The structure of P680and consrquences of itsoxidizing power. Photosynth Res,1993,38(3):297-301
[207] Van Iersel MW&Oosterhuis DM. Diurnal water relations of expanding and full-sized cotton fruits and subtendingleaves. Plant, Cell Environ,1995,18(7):807-812
[208] Van Iersel MW&Oosterhuis DM. Drought effects on the water relations of cotton fruits, bracts, and leaves duringontogeny. Environ Exp Bot,1996,36(1):51-59
[209] Vass I. Role of charge recombination processes in photodamage and photoprotection of the photosystem II complex.Physiol Plant,2011,142(1):6-16
[210] von Balthazar M&Endress PK. Floral bract function, flowering process and breeding systems of Sarcandra andChloranthus (Chloranthaceae). Plant Syst Evol,1999,218(3-4):161-178
[211] von Caemmerer S&Farquhar GD. Some relationships between the biochemistry of photosynthesis and the gasexchange of leaves. Planta,1981,153(4):376-387
[212] von Caemmerer S, Evans JR, Hudson GS, Andrews TJ. The kinetics of ribulose-1,5-bisphosphatecarboxylase/oxygenase in vivo inferred from measurements of photosynthesis in leaves of transgenic tobacco. Planta,1994,195(1):88-97
[213] Walcroft AS, Whitehead D, Silvester WB, Kelliher FM. The response of photosyntheticmodel parameters totemperature and nitrogen content in Pinus radiata D. Don. Plant Cell Environ,1997,20(11):1338-1348
[214] Wang ZM, Wei AL, Zheng DM. Photosynthetic characteristics of non leaf organs of winter wheat cultvars differeing inear type and their relationship with grain mass per ear. Photosynthetica,2001,39(2):239-244
[215] Wardlaw IF. Interaction between drought and chronic high temperature during kernel filling in wheat in a controlledenvrironment. Ann Bot,2002,90(4):469-476
[216] Warren CR&Adams MA. Phosphorus affects growth and partitioning of nitrogen to Rubisco in Pinus pinaster. TreePhysiol,2002,22(1):11-19
[217] Warren CR&Adams MA. Evergreen trees do not maximize instantaneous photosynthesis. Trends Plant Sci,2004,9(6):270-274
[218] Webber AN, Nie GY, Long SP. Acclimation of photosynthetic proteins to rising atmospheric CO2. Photosynth Res,1994,39(3):413-425
[219] Weete JD, Leek G.L, Peterson CM, Currie HE and Branch WD. Lipid and surface wax synthesis in water-stressedcotton leaves. Plant Physiol,1978,62(5):675-677
[220] Wendel JF. New World tetraploid cottons contain Old-World cytoplasm. Proc Natl Acad Sci USA,1989,86(11):4132-4136
[221] Wendel JF&Albert VA. Phylogenetics of the cotton genus (Gossypium): Character-state weighted parsimony analysisof chloroplast-DNA restriction site data and its systematic and biogeographic implications. Systematic Botany,1992,17:115-143
[222] Willmer CM. Morphology and anatomy of stomata. In Stomata. London: Longman,1983
[223] Woodson JD&Chory J. Coordination of gene expression between organellar and nuclear genomes. Nat Rev Genet,2008,9(5):383-395
[224] Wullschleger SD, Oosterhuis DM, Rutherford SA. Importance of bracts in the carbon economy of cotton. ArkansasFarm Res,1990,39(3):4
[225] Wullschleger SD&Oosterhuis DM. Photosynthetic and respiratory activity of fruiting forms within the cottoncanopy. Plant Physiol,1990a,94(2),463-469
[226] Wullschleger SD&Oosterhuis DM. Photosynthetic carbon production and use by developing cotton leaves and bolls.Crop Sci,1990b,30(6):1259-1264
[227] Wullschleger SD&Oosterhuis DM. Photosynthesis, transpiration, and water-use efficiency of cotton leaves and fruit.Photosynthetica,1991,25:505-515
[228] Wullschleger SD, Oosterhuis DM, Hurren RG, Hanson PJ. Evidence for light-dependent recycling of respired carbondioxide by the cotton fruit. Plant Physiol,1991,97(2):574-579
[229] Xu HL, Ishii R, Yamagishi T, Kumura A. Effects of water deficit on photosynthesis in wheat plants. III. Effect onnon-stomatal mediated photosynthesis and RuBP carboxylase content in different plant parts. Jpn J Crop Sci,1990,59(1):153-157
[230] Xu HL, Gauthier L, Desjardins Y, Gosselin A. Photosynthesis in leaves, fruits, stem and petioles of greenhouse-growntomato plants. Photosynthetica,1997,33(1):113-123
[231] Xu XL&Ishii R. Wheat cultivar differences in photosynthetic response to low soil water potentials:1. Marintenance ofphotosynthesis and leaf water potential. Jpn.J Crop Sci,1996,65:509-517
[232] Xu ZZ&Zhou GS. Responses of leaf stomatal density to water status and its relationship with photosynthesis in agrass. J Exp Bot,2008,59(12):3317-3325
[233] Xu ZZ, Zhou GS, Wang YL, Han GX, Li YJ. Changes in chlorophyll fluorescence in maize plants with imposed rapiddehydration at different leaf ages. J Plant Growth Regulat,2008,27(1):83-92
[234] Yabuta Y, Motoki T, Yoshimura K, Takeda T, Ishikawa T and Shigeoka S. Thylakoid membrane-bound ascorbateperoxidase is a limiting factor of antioxidative systems under photo-oxidative stress. Plant J,2002,32(6):915-925
[235] Yang JC, Zhang JH, Wang ZQ, Zhu QS, Liu LJ. Activities of starch hydrolytic enzymes and sucrose-phosphatesynthase in the stems of rice subjected to water stress during grain filling. J Exp Bot,2001,52(364):2169-2179
[236] Yamori W, Takahashi S, Makino A, Price GD, Badger MR, von Caemmerer S. The roles of ATP synthase and theCytochrome b6/f complexes in limiting chloroplast electron transport and determining photosynthetic capacity. PlantPhysiol,2011,155(2):956-962
[237] Zhang YH, Wang ZM, Shu HW. Phosphoenolpyruvate carboxylase activity in ear organs is related to proteinconcentration in grains of winter wheat. J Cereal Sci,2008,47(2):386-391
[238] Zhang YP, Zhang YH, Wang ZM, Wang ZJ. Characteristics of canopy structure and contributions of non-leaf organs toyield in winter wheat under different irrigated conditions. Field Crop Res,2011,123(3):187-195
[239] Zhao DL&Oosterhuis DM. Photosynthetic capacity and carbon contribution of leaves and bracts to developing floralbuds in cotton. Photosynthetica,1999,36(1-2):279-290
[240] Zhu XG, Ort DR, Whitmarsh J, Long SP. The slow reversibility of photosystem Ⅱ thermal energy dissipation ontransfer from high to low light may cause large losses in carbon gain by crop canopies: a theoretical analysis. J Exp Bot,2004,55(400):1167-1175
[241] Ziegler J ns A. Gas-exchange of ears of cereals in response to carbon dioxide and light. I. Relative contributions ofparts of the ears of wheat, oat, and barley to the gas exchange of the whole organ. Planta,1989,178(2):84-91
[2]杜明伟,冯国艺,姚炎帝等.杂交棉标杂A1和石杂2号超高产冠层特性及其与群体光合生产的关系.作物学报,2009b,35(6):1068-1077
[3]冯国艺.超高产棉花冠层结构形成机理及调控研究:石河子大学博士学位论文.石河子:石河子大学,2012
[4]李卫华,郝乃斌. C3植物中C4途径研究进展.植物学通报,1999,16(2):97-106
[5]李寒冰,胡玉熹,白克智等.水稻稃片和旗叶叶绿体超微结构低温荧光特性进行了比较研究.电子显微学报,2002,21(3):341-344
[6]李秀菊,职明星,石晓华等.小麦穗光合对不同花位籽粒及颖壳的影响.麦类作物学报,2006,26(5):146-148
[7]卢存福,贲桂英.高海拔地区植物的光合特性.植物学通报,1995,12(2):38-42
[8]施生锦,黄彬香,王志敏.小麦不同光合器官的光合性能研究.中国生态农业学报,2003,11(4):10-11
[9]薛丽华,章建新.大豆鼓粒期非叶光合器官与粒重的关系.大豆科学,2006,25(4):524-428.
[10]魏爱丽,王志敏.野生一粒麦与普通栽培小麦不同绿色器官光合特性与叶绿体结构特征.作物学报,2002,28(3):351-354
[11]魏爱丽,王志敏,翟志席等.土壤干旱对小麦旗叶和穗器官C4光合酶活性的影响.中国农业科学,2003,36(5):508-512
[12]魏爱丽,张英华,黄琴等.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,2007,33:(9)1426-1431
[13]张亚黎,冯国艺,胡渊渊等.棉花非叶器官光合能力的差异及与物质生产的关系.作物学报,2010,36(4):701-708
[14]张永平,王志敏,吴永成等.不同供水条件下小麦不同绿色器官的气孔特性研究.作物学报,2006,32(1):70-75
[15]中国农业科学院棉花研究所.中国棉花栽培学.上海:上海科学技术出版社,1983
[16] Abbad H, Jaafrai SE, Bort J, Araus JL. Comparison of flag leaf and ear photosynthesis with biomass and grain yield ofdurum wheat under various water conditions and genotypes. Agronomie,2004,24:19-28
[17] Ainsworth EA&Long SP. What have we learned from15years of free-air CO2enrichment (FACE)? A meta-analyticreview of the responses of photosynthesis, canopy properties and plant production to rising CO2. New Phytol,2005,165(2):351-371
[18] Anderson JM. Insights into the consequences of grana stacking of thylakoid membranes in vascular plants:a personalperspective. Aust J Plant Physiol,1999,26(7):625-639
[19] Aoyagi K&Bassham JA. Pyruvate orthophosphate dikinase of C3seeds and leaves as compared to the enzyme frommaize. Plant Physiol,1984,75:387-392
[20] Araus JL, Brown HR, Febrero A, Bort J, Serret MD. Ear photosynthesis, carbon isotope discrimination and thecontribution of respiratory CO2to differences in grain mass in durum wheat. Plant Cell Environ,1993,16(4):383-392
[21] Aro E-M, McCaffery S, Anderson JM. Photoinhibition and D1protein degradation in peas acclimated to differentgrowth irradiance. Plant Physiol,1993a,103:835-843
[22] Aro E-M, Virgin I, Andersson B. Photoinhibition of photosystem II. Inactivation, protein damage and turnover.Biochim Biophys Acta,1993b,1143(2):113-134
[23] Asada K. The water-water cycle in chloroplasts: scavenging of active oxygen and dissipation of excess photons. RevPlant Physiol Mol Biol,1999,50:601-639
[24] Aschan G&Pfanz H. Non-foliar photosynthesis-a strategy of additional carbon acquisition. Flora,2003,198(2):81-97.
[25] Aschan G, Pfanz H, Vodnik D, Bati F. Photosynthetic performance of vegetative and reproductive structures of greenhellebore (Helleborus viridis L. agg.). Photosynthetica,2005,43(1):55-64
[26] Aschan G&Pfanz H. Why snowdrop (Galanthus nivalis L.) tepals have green marks? Flora,2006,8(23):623-632
[27] Ashraf M&Foolad MR. Roles of glycine betaine and proline in improving plant abiotic stress resistance. Environ ExpBot,2007,59(2):206-216
[28] Athar H&Ashraf M. Photosynthesis under drought stress. In: Pessarakli, M.(Ed.), Handbook Photosynthesis, seconded. CRC Press, New York, USA,2005,795-810
[29] Bachmann KM, Ebbert V, Adams WW III, Verhoven AS, Logan BA, Demmig-Adams B. Effects of lincomycin on PSIIefficiency, non-photochemical quenching, D1protein and xanthophylls cycle during photoinhibition and recovery. FuncPlant Biol,2004,31:803-813
[30] Bajji M, Lutts S, Kinet JM. Water deficit effects on solute contribution to osmotic adjustment as a function of leafageing in three durum wheat (Triticum durum Desf.) cultivars performing differently in arid conditions. Plant Sci,2001,160(4):669-681
[31] Barlow EWR, Lee JW, Rana Munns Smart MG. Water relations of the developing wheat grain. Aust J Plant Physiol,1980,7(5):519-525
[32] Bates LS, Waldren RP, Teare JD. Rapid determination of proline for water stress studies.Plant Soil,1973,39(1):205-207
[33] Ben Ahmed C, Ben Rouina B, Athar HR, Boukhris M. Olive tree (Oleaeuropaea L. cv. Chemlali) under salt stress:water relations and ions contents. Pak J Bot,2006,38(5):1477-1484
[34] Ben Ahmed C, Ben Rouina B, Boukhris M. Effects of water deficit on olive trees cv. Chemlali under field conditions inarid region in Tunisia. Sci Hort,2007,113(3):267-277
[35] Ben Ahmed C, Ben Rouina B, Sensoy S, Boukhris M, Ben Abdallah F. Changes in gas exchange, proline accumulationand antioxidative enzyme activities in three olive cultivarsunder contrastingwater availability regimes. Environ Exp Bot,2009,67(2):345-352
[36] Bettarini I., Vaccari FP, Miglietta F. Elevated CO2concentrations and stomatal density-observations from17plantspecies growing in a CO2spring in central Italy. Global Change Biology,1998,4(1):17-22
[37] Bhatt JG. Transport of radioactivity in relation to bracts in the cotton plant. Ann Bot,1988,62(6):571-573
[38] Birkhold KB, Koch KE, Darnell RL. Carbon and nitrogen economy of developing rabbiteye blueberry fruit. J Am SocHortic Sci,1992,117(1):139-145
[39] Biswall B. Carotenoid catabolism during leaf senescence and its control by light. J Photochem Photobiol B,1995,30(1):3-14
[40] Bj rkman O. Carboxydismutase activity in shade-adapted and sun-adapted species of higher plants. PhysiologiaPlantarum,1968,21(1):1-10
[41] Bj rkman O. Response to different quantum flux densities. In Lange OL, Nobel PS, Osmond CB, Ziegler H, eds.Physiological Plant Ecology I. Responses to the Physical Environment, Encyclopedia of plant physiology, new series.Springer-Verlag: Berlin,1981,57-107
[42] Black CC&Vines HM. Alternative plant photosynthetic CO2fixation cycles.In DW Newman,KG Wilson,eds,Modelsin Plant Physioligy and Biochemistry,Vol1.CRC Press.BocaRaton, FL,1987,57-61
[43] Blanke MM&Lenz F. Fruit photosynthesis a review. Plant Cell Environ,1989,12(1):31-46
[44] Blanke MM&Ebert G. Phosphoenolpyruvate carboxylase and carbon economy of apple seedlings. J Exp Bot,1992,43(7):965-968
[45] Blum A. Photosynthesis and transpiration in leaves and ears of wheat and barley varieties. J Exp Bot,1985,36(3):432-440
[46] Blum A. The effects of heat stress on wheat leaf and ear photosynthesis. J Exp Bot,1986,37(174):111-118
[47] Blum A, Sinmena B, Mayer J, Golan G and Shpiler L (1994) Stem reserve mobilization supports wheat-grain fillingunder heat stress. Australian Journal of Plant Physiology,21(6):771-781
[48] Bondada BR, Oosterhuis DM, Wullschleger SD, Kim KS, Harris WM. Anatomical considerations related tophotosynthesis in cotton leaves, bracts, and the capsule wall. J Exp Bot,1994,45(1):111-118
[49] Bondada BR, Oosterhuis DM, Murphy JB, Kim KS. Effect of water stress on the epicuticular wax composition andultrastructure of cotton leaf, bract, and boll. Environ Exp Bot,1996,36(1):61-69
[50] Bondada BR&Oosterhuis DM. Comparative epidermal ultrastructure of cotton (Gossypium hirsutum L.) leaf, bractand capsule wall. Ann Bot,2000,86(6):1143-1152
[51] Bondada BR&Oosterhuis DM. Morphometic analysis of chloroplasts of cotton leaf and fruiting organs.Biol Plantarum,2003,47(2):281-284
[52] Bort J, Febrero A, Amaro T, Araus JL. Role of awns in ear water-use efficiency and grain weight in barley. Agronomie,1994,2:133-139
[53] Bort J, Brown RH, Araus JL. Refixation of respiratory CO2in the ears of C3-cereals. J Exp.Bot,1996,47(10):1567-1575.
[54] Boyer JS. Plant productivity and environment. Sci,1982,218(4571):443-448
[55] Bravd BA, Palgi A, Lurie S. Changing ribulose diphosphate carboxylase/oxygenase activity in ripening tomato fruit.Plant Physiol,1977,60:309-312
[56] Brown KJ. Translocation of carbohydrates in cotton: Movement to the fruiting bodies.Ann Bot,1968,32(4):703-713
[57] Caley CY, Duffus CM, Jeffcoat B. Photosynthesis in the pericarp of developing wheat grains. J Exp Bot,1990,41(3):303-307
[58] Caldwell MM, Osmond CB, Nott DL. C4pathway photosynthesis at low temperature in cold-tolerant Atriplex species.Plant Physiol,1977,60:157-164
[59] Camp PJ, Huber SC, Burke JJ, Moreland DE. Biochemical changes that occur during senescence of wheat leaves. I.Basis for the reduction of photosynthesis. Plant Physiol,1982,70:1641-1646
[60] Chapin III FS. The mineral nutrition of wild plants. Annu Rev Ecol Syst,1980,11:233-260
[61] Chaves MM. Effects of water deficits on carbon assimilation. J Exp Bot,1991,42(1):1-16
[62] Chaves MM, Maroco JP, Pereira JS. Understanding plant responses to drought-from genes to the whole plant. FunctPlant Biol,2003,30(3):239-264
[63] Chow WS&Aro E-M. Photoinactivation and mechanisms of recovery. In: Wydrzynski T and Satoh K (eds)Photosystem II: the light-driven water: plastoquinone oxidoreductase. Advances in photosynthesis and respiration, vol22Springer, Dordrecht,2005,627-648
[64] Chow WS&Hope AB. Electron fluxes through Photosystem I in cucumber leaf discs probed by far-red light.Photosynth Res,2004,81:77-89
[65] Chow WS, Hope AB, Anderson JM. Oxygen per flash from leaf disks quantifies photosystem II. Biochim Biophys Acta,1989,973(1):105-108
[66] Chow WS, Fan D-Y, Oguchi R, Jia H, Losciale P, Park Y-I, He J, quist G, Shen Y-G, Anderson JM. Quantifying andmonitoring functional Photosystem II and the stoichiometry of the two photosystems in leaf segments: Approaches andapproximations. Photosynth Res,2012,113(1):63-74
[67] Claussen W. Proline as a measure of stress in tomato plants. Plant Sci,2005,168(1):241-248
[68] Cockfield SD. Relative availability of nitrogen in host plants of invertebrate herbivores: three possible nutritional andphysiological definitions. Oecologia,1988,77:91-94
[69] Condon AG, Richards RA,Farquhar GD. Carbon isotope discrimination is positively correlated with grain yield and drymatter production in filed-grown wheat. Crop Sci,1987,996-1001
[70] Constable GA&Rawson HM. Carbon production and utilization in cotton: Inferences from a carbon budget. Aust JPlant Physiol,1980,7(5):539-553
[71] Cornic G. Drought stress and high light effects on leaf photosynthesis. In Photoinhibition of Photosynthesis: FromMolecular Mechanisms to the Field. Eds. N R Baker and J R Bowyer. Oxford BIOS Scientific Publishers.1994,297-313
[72] Cornic G. Drought stress inhibits photosynthesis by decreasing stomatal aperture-not by affecting ATP systhesis. TrendsPlant Sci,2000,5:187-188
[73] Crookston RK, O'Toole J, Ozbun JL. Characterization of the bean pod as a photosynthetic organ. Crop Sci,1974,14:708-712
[74] Cruz-Aguado JA, Rodés R, Pérez IP, Dorado M. Morphological characteristics and yield components associated withaccumulation and loss of dry mass in the internodes of wheat. Field Crops Research,2000,66(2):129-139
[75] Demmig-Adams B&Adams III WW. Photoprotection and other responses of plants to high light stress. Ann Rev PlantPhysiol PlantMol Biol,1992,43(1):599-626
[76] Drake BG, Gonzalez-Meler MA, Long SP. More efficient plants-a consequence of rising atmospheric CO2. Annu RevPlant Physiol Plant Mol Bio,1997, l48:609-639
[77] Duffus CM&Cochrane MP. Formation of the barley grain-morphology, physiology, and biochemistry. In: MacGregorAW, Bhatty RS (eds) Barley: chemistry and technology. American Association of Cereal Chemists, St. Paul,1993,31-72
[78] Ehdaie B, Alloush GA, Waines JG. Genotypic variation in linear rate of grain growth and contribution of stem reservesto grian yield in wheat. Field Crops Research,2008,106(1):34-43
[79] Eamus D. The interaction of rising CO2and temperatures with water use efficiency. Plant Cell Environ,1991,14:843-582
[80] Ehsanzadeh P, Mahmoudieh R, Fareed N. Photosynthetic contribution of the inflorescence and adjacent green tissue todiverse environmental conditions. J Biol Sci,2007,7(2):263-269
[81] Elmore CD. Contributions of the capsule wall and bracts to the developing cotton fruits. Crop Sci1973,13:751-752
[82] Elstner EF. Metabolism of activated oxygen species. In: Davies DD (ed.):The biochemistry of Plants, Biochemistry ofMetabolism. Academic Press, San Diego.1987, Vol11.253-315
[83] Endress PK. Floral phyllotaxis and floral evolution. Botanische Jahrbücher für Systematik,1987,108:417-438
[84] Endress PK. Evolutionary diversification of the flowers in Angiosperms. Am J Bot,2011,98(3):370-396
[85] Evans LT, Bingham J, Jackson P, Sutherland J. Effect of awns and drought on the supply of photosynthate and itsdistribution within wheat ears. Ann Appl Biol,1972,70(1):67-76
[86] Evans JR. Photosynthesis and nitrogen relationships in leaves of C3plants. Oecologia,1989,78(1):9-19
[87] Evans JR, Jakobsen I, gren E. Photosynthetic light-response curves2. Gradients of light-absorption andphotosynthetic capacity. Planta,1993,189:191-200
[88] Evans JR. Potential errors in electron transport rates calculated from chlorophyll fluorescence as revealed by amultilayer leaf model. Plant Cell Physiol,2009,50(4):698-706
[89] Ewart AJ. On assimilatory inhibition in plants. J Linn Soc,1896,31(217):364-461
[90] Farquhar GD, von Caemmere S, Berry JA. A biochemical model of photosynthetic CO2assimilation in leaves of C3species. Planta,1980,149(1):78-90
[91] Field C, Mooney H. The photosynthesis-nitrogen relationship in wild plants. In On the Economy of Plant Form andFunction. Ed. G.T. Givnish. Cambridge University Press,1986,25-55
[92] Foyer CH, Descourvieres P, Kunert KJ. Protection against oxygen radicals: an important defence mechanism studied intransgenic plants. Plant Cell Environ,1994,17:507-523
[93] Fu J&Huang B. Involvement of antioxidants and lipid peroxidation in the adaptation of two cool-season grasses tolocalized drought stress. Environ Exp Bot,2001,45(2):105-114
[94] Gebbing T, Schnyder H.13C Clabelling kinetics of sucrose in glumes indicates significant refixation of respiratory CO2in the wheat ear. Aust J Plant Physiol,2001,28(10):1047-1053
[95] Genty B, Briantais JM, Baker NR. The relationship between the quantum yield of photosynthetic electron transport andquenching of chlorophyll fluorescence. Biochim Biophys Acta,1989,990(1):87–92
[96] Grossman A, Bartlett S, Chua N-H. Energy-dependent uptake of cytoplasmically synthesized polypeptide bychloroplasts. Nature,1980,285:625-628
[97] Gunderson CA&Wullschleger SD. Photosynthetic acclimation in trees to rising atmospheric CO2: a broaderperspective. Photosynth Res,1994,39(3):369-388
[98] Gupta AS, Webb RP, Holaday AS and Allen RD. Overexpression of superoxide dismutase protects plants fromoxidative stress (Induction of ascobate peroxidase in superoxide dismutase-overpressing plants). Plant Physiol,1993,103:1067-1073
[99] Hattenschwiler S, Miglietta F, Raschi A, Korner C. Thirty years of in situ tree growth under elevated CO2: a model forfuture forest responses? Global Change Biol,1997,3:463-471
[100] He J&Chow WS. The rate coefficient of repair of photosystem II after photoinactivation. Physiol Plant,2003,118(2):297-304
[101] Hong SS&Xu DQ. Light-induced increase in initial chlorophyll fluorescence Folevel and the reversible inactivation ofPSII reaction centers in soybean leaves. Photosynth Res,1999,61(3):269-280
[102] Hedley CL, Harvey DM, Keely RJ. Role of PEP carboxylase during seed development in Pisum sativum. Nature,1975,258:352-354
[103] Hemmers H and Gulz PG. Chemistry and morphology of epicuticular waxes from leaves of five Euphorbia species. ZNatuforsch,1986,41(5-6):521-525
[104] Hendrickson L, Furbank RT, Chow WS. A simple alternative approach to assessing the fate of absorbed light energyusing chlorophyll fluorescence. Photosynth Res,2004,82:73-81
[105] Hikosaka K&Hirose T. Leaf and canopy photosynthesis of C3plants at elevated CO2in relation to optimal partitioningof nitrogen among photosynthetic components: theoretical prediction. Ecol Model,1998,106(2-3):247-259
[106] Hikosaka K. Nitrogen partitioning in the photosynthetic apparatus of Plantago asiatica leaves grown under differenttemperature and light conditions: similarities and differences between temperature and light acclimation. Plant CellPhysiol,2005,46(8):1283-1290
[107] Hodges DM, DeLong JM, Forney CF, Prange RK. Improving the thiobarbituric acid-reactive-substances assay forestimating lipid peroxidation in plant tissues containing anthocyanin and other interfering compounds. Planta,1999,207(4):604-611
[108] Hormaetxe K, Becerril JM, Fleck I, Pinto M, Garcia-Plazaola JI. Functional role of red (retro)-carotenoids as passivelight filters in theleaves of Buxus sempervirens L.: increased protection of photosynthetic tissues? J Exp Bot,2005,56(420):2629-2636
[109] Horton P. Prospects for crop improvement through the genetic manipulation of photosynthesis: morphological andbiochemical aspects of light capture. J Exp Bot,2000,51(suppl1):475-485
[110] Imaizumi N, Usuda H, Nakamoto H and Ishihara K. Changes in the rate of photosynthesis during grain filling and theenzymatic activities associated with the photosynthetic carbon metabolism in rice panicles. Plant Cell Physiol,1990,31(6):835-843.
[111] Ishihara K, Kiyota E, Imaizumi N. On the contribution of panicle photosynthesis to grain yield in rice plants. Jpn JCrop Sci,1991,60(Extra Issue1):122-123
[112] Johnson RR&Moss DN. Effect of water stress on14CO2fixation and translocation in wheat during grain filling. CropSci,1976,16:697-701
[113] Johnson DA, Richards RA, and Turner NC. Yield, water, relations, gas exchange, and surface reflectances of nearisogenic wheat lines differing in glaucousness. Crop Sci,1983,23(2):318-325
[114] Jones MB, Brown JC, Raschi A, Miglietta F. The effects on Arbutus unedo L. of long-term exposure to elevated CO2.Global Change Biology,1995,1(4):295-302
[115] Jordan WR, Shouse PJ, Blum A, Miller FR, Monk RL. Environmental physiology of sorghum. II. Epicuticular wax loadand cuticular transpiration. Crop Sci,1984,24(6):1168-1173
[116] Kato MC, Hikosaka K, Hirotsu N, Makino A, Hirose T. The excess light energy that is neither utilized in photosynthesisnor dissipated by photoprotective mechanisms determines the rate of photoinactivation in Photosystem II. Plant CellPhysiol,2003,44(3):318-325
[117] Kikuta SB&Richter H. Graphical evaluation and partitioning of turgor responses to drought in leaves of durum wheat.Planta,1986,168:36-42
[118] Kimball BA, Kobayashi K, Bindi M. Responses of agricultural crops to free-air CO2enrichment. Adv Agron,2002,77:293-368
[119] Kingston-Smith AH, Harbinson J, Williams J, Foyer CH. Effect of chilling on carbon assimilation, enzyme activation,and photosynthetic electron transport in the absence of photoinhibition in maize leaves. Plant Physiol,1997,114(3):1039-1046
[120] Knoppik D, Selinger H, Ziegler-J ns A. Differences between the flag leaf and the ear of a spring wheat cultivar(Triticum aestivum cv. Arkas) with respect to the CO2response of assimilation, respiration and stomatal conductance.Physiol Plant,1986,68(3):451-457
[121] Kocheva K, Lambrev P, Georgiev G, Goltsev V, Karabaliev M. Evaluation of chlorophyll fluorescence and membraneinjury in the leaves of barley cultivars under osmotic stress. Bioelectrochemistry,2004,63(1-2):121-124
[122] Kou J, Oguchi R, Fan D-Y, Chow WS. The time course of photoinactivation of photosystem II in leaves revisited.Photosynth Res,2012,113(1-3):157-164
[123] Lawlor DW, Kontturi M, Young AT. Photosynthesis by flag leaves of wheat in relation to protein, ribulose bisphosphatecarboxylase activity and nitrogen supply. J Exp Bot,1989,40(1):43-52
[124] Lawlor DW. The effects of water deficit on photosynthesis. In Environment and Plant Metabolism. Flexibility andAcclimation, Ed. N Smirnoff. Oxford: BIOS Scientific Publisher.1995, pp.129-160
[125] Lawlor DW. Limitation to photosynthesis in water-stressed leaves: stomata vs. metabolism and the role of ATP. AnnBot,2002,89(7):871-885
[126] Lawlor DW&Cornic G. Photosynthetic carbon assimilation and associated metabolism in relation to water deficits inhigher plants. Plant Cell Environ,2002,25(2):275-294
[127] Lee H-Y, Hong Y-N, Chow WS. Photoinactivation of photosystem II complexes and photoprotection by non-functionalneighbours in Capsicum annuum L. leaves. Planta,2001,212(3):332-342
[128] Levizou E, Petropoulou Y, Manetas Y. Carotenoid composition of peridermal twigs does fully conform to a shadeacclimation hypothesis. Photosynthetica,2004,42(4):591-596
[129] Li XJ, Wang HG, Li HB, Zhang LY, Teng NJ, Lin QQ, Wang Jian, Kuang TY, Li ZS, Li B, Zhang AM and Lin JX. Awnplay a dominant role in carbohydrate production during the grain filling stages in wheat (Triticum aestivum).Physiol Plantarum,2006,127(4):701-709
[130] Lichtenthaler HK. Chlorophylls and carotenoids: pigments of photosynthetic biomembranes. Methods Enzymol,1987,148:350-382
[131] Lizana C, Wentworth M, Martínez JP, Villegas D, Meneses R, Murchie EH, Pastenes C, Lercari B, Vernieri P, Horton P,Pinto M. Differential adaptation of two varieties of common bean to abiotic stress. I. Effects of drought on yield andphotosynthesis. J Exp Bot,2006,57(3):685-697
[132] Long SP, Ainsworth EA, Rogers A, Ort DR. Rising atmospheric carbon dioxide: plants face the future. Annu Rev PlantBiol,2004,55:591-628
[133] Losciale P, Oguchi R, Hendrickson L, Hope AB, Corelli-Grappadelli L, Chow WS. A rapid, whole-tissue determinationof the functional fraction of Photosystem II after photoinhibition of leaves based on flash-induced P700redox kinetics.Physiol Plant,2008,132(1):23-32
[134] Matto AK, Hoffman-Falk H, Marder JB, Edelman M. Regulation of protein metabolism: Coupling of photosyntheticelectron transport to in vivo degradation of the rapidly metabolized32-kilodalton protein of the chloroplast membranes.Proc Natl Acad Sci USA,1984,81(5):1380-1384
[135] Makino A, Mae T, Ohira K. Photosynthesis and ribulose-1,5-bisphosphate carboxylase in rice leaves. Plant Physiol,1983,73(4):1002-1007
[136] Makino A&Osmond B. Effects of nitrogen nutrition on nitrogen partitioning between chloroplasts and mitochondria inpea and wheat. Plant Physiol,1991,96(2):355-362
[137] Makino A, Harada M, Kaneko K, Mae T, Shimada T, Yamamoto N. Whole-plant growth and N allocation in transgenicrice plants with decreased content of ribulose-1,5-bisphosphate carboxylase under different CO2partial pressures. AustJ Plant Physiol,2000,27(1):1-12
[138] Martinez DE, Luquez VM, Bartoli CG, Guizmét JJ. Persistence of photosynthetic components and photochemical effectin ears of water-stressed wheat(Triticum aestivum). Physiol Plantarum,2003,119(4):519-525
[139] Maydup ML, Anotonietta M, Guiamet JJ, Graciano C, López, Tambussi EA. The contribution of ear photosynthesis tograin filling in bread wheat (Triticum aestivum L.). Field Crop Research.,2010,119(1):48-58
[140] McKee IF&Woodward FI. The effect of growth at elevated CO2concentrations on photosynthesis in wheat. Plant CellEnviron,1994,17(7):853-859
[141] Meehl GA, Stocker TF, Collins WD, Friedlingstein P, Gaye AT, Gregory JM, Kitoh A, Knutti R, Murphy JM, Noda A,Raper SCB, Watterson IG, Weaver AJ, Zhao Z-C. Global Climate Projections. In Solomon S, Qin D, Manning M, ChenZ, Marquis M, Averyt KB, Tignor M, Miller HL, eds. Climate Change2007: The Physical Science Basis. Contributionof Working Group I to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change. CambridgeUniversity Press: Cambridge and New York,2007,747-845
[142] Medlyn BE. The optimal allocation of nitrogen within the C3photosynthetic system at elevated CO2. Aust J PlantPhysiol,1996,23(5):593-603
[143] Medrano H, Escalona JM, Bota J, Gulías J and Flexas J. Regulation of photosynthesis of C3plants in response toprogressive drought: stomatal conductance as a reference parameter. Ann Bot,2002,89(7):895-905
[144] Melis A. Photosystem-II damage and repair cycle in chloroplasts: what modulates the rate of photodamage in vivo?Trends Plant Sci,1999,4(4):130-135
[145] Miglietta F&Raschi A. Studying the effect of elevated CO2in the open in a naturally enriched environment in CentralItaly.1993, Vegetatio104/105:391-400
[146] Mooney HA&Gulmon SL. Environmental and evolutionary constraints on the photosynthetic characteristics of higherplants. In: Solbrig OT, Jain S, Johnson GB, Raven PH, eds. Topics in plant population biology. New York, NY, USA:Columbia University Press,1979,316-337
[147] Morgan JM. Osmotic adjustment in the spikelets and leaves of wheat. J Exp Bot,1980,31(2):655-665
[148] Morgan JM. Osmoregulation and water-stressed in higher plants. Annu Rev Plant Physiol,1984,35(1):299-319.
[149] Mulchi CL, Slaughter L, Saleem M, Lee EH, Pausch R, Rowland R. Growth and physiological characteristics ofsoybean in open top chambers in response to ozone and increased atmospheric CO2. Agr Ecosyst Environ,1992,38(1-2):107-118
[150] Müller P, Li XP, Niyogi KK. Non-photochemical quenching. A response to excess light energy. Plant Physiol,2001,125(4):1558-1566
[151] Nakano H, Makino A, Mae T. The effect of elevated partial pressures of CO2on the relationship betweenphotosynthetic capacity and N content in rice leaves. Plant Physiol,1997,115(1):191-198
[152] Nayyar H. Accumulation of osmolytes and osmotic adjustment in water-stressed wheat (Triticum aestivum) and maize(Zea mays) as affected by calcium and its antagonists. Environ Exp Bot,2003,50(3):253-264
[153] Nie GY, Long SP, Garcia RL, Kimball BA, Lamorte RL, Pinter PJ, Wall GW, Webber AN. Effects of free-air CO2enrichment on the development of the photosynthetic apparatus in wheat, as indicated by changes in leaf protein. PlantCell Environ,1995,18(8):855-864
[154] Nishimyama Y, Yamamoto H, Allakhverdiev SI, Inaba M, Yokota A, Murata N. Oxidative stress inhibits the repair ofphotodamage to the photosynthetic machinery. EMBO J,2001,20(20):5587-5594
[155] Nishimyama Y, Allakhverdiev SI, Murata N. Protein synthesis is the primary target of reactive oxygen species in thephotosystem II. Physiol Plant,2011,142(1):35-46
[156] Oguchi R, Terashima I, Chow WS. The involvement of dual mechanisms of photoinactivation of photosystem II inCapsicum annuum L. plants. Plant Cell Physiol,2009,50(10):1815-1825
[157] Oguchi R, Terashima I, Kou J, Chow WS. Operation of dual mechanisms that both lead to photoinactivation ofPhotosystem II in leaves by visible light. Physiol Plant,2011a,142(1):47-55
[158] Oguchi R, Douwstra P, Fujita T, Chow WS, Terashima I. Intra-leaf gradients of photoinhibition induced by differentcolor lights: Implications for the dual mechanisms of photoinhibition and for the application of conventionalchlorophyll fluorometers. New Phytol,2011b,191(1):146-159
[159] Ohad I, Berg A, Berkowicz SM, Kaplan A, Keren N. Photoinactivation of photosystem II: is there more than one way toskin a cat? Physiol Plant,2011,142(1):79-86
[160] Onoda Y, Hikosaka K, Hirose T. Seasonal change in the balance between capacities of rubp carboxylation and rubpregeneration affects CO2response of photosynthesis in Polygonum cuspidatum. J Expl Bot,2005,56(412):755-763.
[161] Onoda Y, Hirose T, Hikosaka K. Effect of elevated CO2levels on leaf starch, nitrogen and photosynthesis of plantsgrowing at three natural CO2springs in Japan. Ecol Res,2007,22(3):475-484
[162] Onoda Y, Hirose T, Hikosaka K. Does leaf phootosynthesis adapt to CO2-enriched environments? An experiment onplants originating from three natural CO2springs. New Phytol,2009,182(3):698-709
[163] Oosterhuis DM, Chipamaunga J, Bate GC. Nitrogen uptake of field-grown cotton. I. Distribution in plant componentsin relation to fertilization and yield. Expl Agric,1983,19(1):91-101
[164] Paoletti E, Nourrisson G, Garrec JP, Raschi A. Modifications of the leaf surface structures of Quercus ilex L. in open,naturally CO2-enriched environments. Plant Cell Environ,1998,21(10):1071-1075
[165] Pérez-Alfocea F, Estan MT, Caro M, Guerrier G. Osmotic adjustment in Lycopersicon esculentum and L. pennelliiunder NaCl and polyethylene glycol6000iso-osmotic stresses. Physiol Plant,1993,87(4):493-498
[166] Pheloung PC&Siddique KHM. Contribution of stem dry matter to grain yield in wheat cultivars. Aust J Plant Physiol,1991,18(1):53-64
[167] Pic E, Tyssender de la Serve B, Tardieu F, Turc O. Leaf senescence induced by mild water deficiet follows the samesequence of macroscopic, bichemical and molecular events as monocarpic senescence in pea. Plant Physiol,2002,128(1):236-246
[168] Pinheiro HA, DaMatta FM, Chaves ARM, Fontes EPB, Loureiro ME. Drought tolerance in relation to protectionagainst oxidative stress in clones of Coffea canephora subjected to long-term drought. Plant Sci,2004,167(6):1307-1314
[169] Polle A, Otter T, Seifert F. Apoplastic peroxidases and lignification in needles of Norway spruce (Picea abies L.). PlantPhysiol,1994,106(1):53-60
[170] Poole I, Weyers JDB, Lawson T, Raven JA. Variations in stomatal density and index: implications for palaeoclimaticreconstructions. Plant Cell Environ,1996,19(6):705-12
[171] Porra RJ, Thompson WA, Kriedemann PE. Determination of accurate extinction coefficients and simultaneousequations for assaying chlorophylls a and b extracted with four different solvents: verification of the concentration ofchlorophyll standards by atomic absorption spectroscopy. BBA-Bioenergetics,1989,975(3):384-394
[172] Powles SB. Photoinhibition of photosynthesis induced by visible light. Annu Rev Plant Physiol,1984,35(1):15-44
[173] Prásil O, Adir N, Ohad I. Dynamics of photosystem II: mechanism of photoinhibition and recovery processes. In:Barber J (ed) Topics in Photosyntheses, Vol.11, Elsevier, Amsterdam,1992, pp.295-348
[174] Prestidge RA&McNeill S. The role of nitrogen in the ecology of grassland Auchenorryncha. In Lee JA, McNeil S andRorison IH (eds) Nitrogen as an Ecological Factor. Blackwell Scientific Publishers, Oxford,1983, pp257-281
[175] Price GD, von Caemmerer S, Evans JR, Siebke K, Anderson JM, Badger MR. Photosynthesis is strongly reduced byantisense suppression of chloroplastic cytochrome bf complex in transgenic tobacco. Aust J Plant Physiol,1998,25(4):445-452
[176] Pugnaire FI, Haase P, Incoll LD, Clark SC. Response of the tussock grass Stipa tenacissima to watering in a semi-aridenvironment. Funct Ecol,1996,10:265-274
[177] Read SM&Northcote DH. Minimization of variation in the response to different proteins of the Coomassie blue G dyebinding assay for protein. Anal Biochem,1981,116(1):53-64
[178] Reddy AR, Chaitanya KV, Jutur PP, Sumithra K. Differential antioxidative responses to water stress among fivemulberry (Morus alba L.) cultivars. Environ Exp Bot,2004,52(1):33-42
[179] Specht JE, Hume DJ, Kumudini SV. Soybean yield potential-a genetic and physiological perspective. Crop Science,1999,39(6):1560-1570
[180] Rodermel S, Haley J, Jiang CZ, Tsai CH, Bogorad L. A mechanism for intergenomic integration: abundance of ribulosebisphosphate carboxylase small-subunit protein influences the translation of the large-subunit mRNA. Proceedings ofthe National Academy of Sciences of the United States of America,1996,93(9):3881-3885
[181] Royer DL. Stomatal density and stomatal index as indicators of paleoatmospheric CO2concentration. Rev PalaeobotPalyno,2001,114(1-2):1-28
[182] Sage RF. Acclimation of photosynthesis to increasing atmospheric CO2: the gas exchange perspective. Photosynthe Res,1994,39(3):351-368
[183] Salvador RJ&Pearce RB. Husk removal and its effects on maize grain yield.Crop Sci,1988,28(6):961-964
[184] Sánchez-Díaz M, García JL, Antolín MC, Araus JL. Effects of soil drought and atmospheric humidity on yield, gasexchange and stable carbon composition of barley. Photosynthetica,2002,40(3):415-421
[185] Sayre RT, Kennedy RA, Pringnitz DJ. Photosynthetic enzyme activities and localization in Mollugo verticillatapopulations differing in the levels of C3and C4cycle operation. Plant Physiol,1979,64(2):293-299
[186] Scarascia-Mugnozza G, De Angelis P, Matteucci G, Valentini R. Long-term exposure to elevated [CO2] in a naturalQuercus ilex L. community: net photosynthesis and photochemical efficiency of PSII at different levels of water stressPlant Cell Environ,1996,19(6):643-654
[187] Schuman GE, Stanley MA, Knudsen D. Automated Total Nitrogen Analysis of Soil and Plant Samples. Soil Sci Soc AmJ,1972,37(3):480-481
[188] Serraj R&Sinclair TR. Osmolyte accumulation: can it really help increase crop yield under drought conditions? PlantCell Environ,2002,25(2):333-341
[189] Sinclair TR&Ludlow MM. Who taught plants thermodynamics? The unfulfilled potential of plant water potential.Aust. J Plant Physiol,1985,12(3):213-217
[190] Singal HR, Sheoran IS, Singh R. In vitro enzyme activites and products of14CO2assimilation in flag leaf and ear partsof wheat (Triticum aestivum L.).Photosynthe Res,1986,8(2):113-122
[191] Smirnoff N. Tansley Review No.52. The role of active oxygen in the response of plants to water deficit and desiccation.New Phytol,1993,125:27-58
[192] Sofo A, Dichio B, Xiloyannis C, Masia A. Lipoxygenase activity and proline accumulation in leaves and roots of olivetree in response to drought stress. Physiol Plant,2004,121(1):58-65
[193] Specht JE, Hume DJ, Kumudini SV. Soybean yield potential-a genetic and physiological perspective. Crop Science.,1999,39(6):1560-1570
[194] Spence RD, Wu H, Sharpe PJH, Clark KG. Water-stressed effects on guard cell anatomy and the mechanical advantageof the epidermal cells. Plant Cell Environ,1986,9(3):197-202
[195] Stitt M. Rising carbon dioxide levels and their potential significance for carbon flow in photosynthetic cells. Plant, CellEnviron,1991,14(8):741-762
[196] Suzuki Y, Makino A. Availability of Rubisco small subunit up-regulates the transcript levels of large subunit forstoichiometric assembly of its holoenzyme in rice. Plant Physiol,2012,160(1):533-540
[197] Tambussi EA, Nogués S, Araus JL. Ear of durum wheat under water stress water relations and photosynthesismetabolism. Planta,2005,221(3):446-458
[198] Tambussi EA, Bort J, Guiamet JJ, Nogues S, Araus JL. The photosynthetic role of ears in C3cereals: metabolism,water use efficiency and contribution to grain yield. Crit Rev Plant Sci,2007,26(1):1-16
[199] Taniguchi M, Kuroda H, Satoh K. ATP-dependent protein synthesis in isolated pea chloroplasts: Evidence foraccumulation of a translation intermediate of the D1protein. FEBS Lett,1993,317(1-2):57-61
[200] Terashima I, Fujita T, Inoue T, Chow WS, Oguchi R. Green light drive leaf photosynthesis more efficiently than redlight in strong white light: Revisiting the enigmatic question of why leaves are green. Plant Cell Physiol,2009,50(4):674-697
[201] Tyystj rvi E, Aro E-M. The rate constant of photoinhibition, measured in lincomycin-treated leaves, is directlyproportional to light intensity. Proc Natl Acad Sci USA,1996,93(5):2213-2218
[202] Tezara W, Mitchell V, Driscoll SP, Lawlor DW. Effects of water deficit and its interaction with CO2supply on thebiochemistry and physiology of photosynthesis in sunflower. J Exp Bot,2002,53(375):1781-1791
[203] Ting IP&Osmond CB. Photosynthetic phosphoenolpyruvate carboxylases characteristics of alloenzymes from leavesof C3and C4plants. Plant Physiol,1973,51(3):439-447
[204] Tognetti R, Giovannelli A, Longobucco A, Miglietta F, Raschi A. Water relations of oak species growing in the naturalCO2spring of Rapolano (central Italy). Ann For Sci,1996,53(2-3):475-485
[205] Tsuyama M, Shibata M, Kobayashi Y. Leaf factors affecting the relationship between chlorophyll fluorescence and therate of photosynthetic electron transport as determined from CO2uptake. J Plant Physiol,2003,160(10):1131-1139
[206] Van Gorkom HJ, Schelvis JPM. Kok’s oxygen clock: What makes it tick? The structure of P680and consrquences of itsoxidizing power. Photosynth Res,1993,38(3):297-301
[207] Van Iersel MW&Oosterhuis DM. Diurnal water relations of expanding and full-sized cotton fruits and subtendingleaves. Plant, Cell Environ,1995,18(7):807-812
[208] Van Iersel MW&Oosterhuis DM. Drought effects on the water relations of cotton fruits, bracts, and leaves duringontogeny. Environ Exp Bot,1996,36(1):51-59
[209] Vass I. Role of charge recombination processes in photodamage and photoprotection of the photosystem II complex.Physiol Plant,2011,142(1):6-16
[210] von Balthazar M&Endress PK. Floral bract function, flowering process and breeding systems of Sarcandra andChloranthus (Chloranthaceae). Plant Syst Evol,1999,218(3-4):161-178
[211] von Caemmerer S&Farquhar GD. Some relationships between the biochemistry of photosynthesis and the gasexchange of leaves. Planta,1981,153(4):376-387
[212] von Caemmerer S, Evans JR, Hudson GS, Andrews TJ. The kinetics of ribulose-1,5-bisphosphatecarboxylase/oxygenase in vivo inferred from measurements of photosynthesis in leaves of transgenic tobacco. Planta,1994,195(1):88-97
[213] Walcroft AS, Whitehead D, Silvester WB, Kelliher FM. The response of photosyntheticmodel parameters totemperature and nitrogen content in Pinus radiata D. Don. Plant Cell Environ,1997,20(11):1338-1348
[214] Wang ZM, Wei AL, Zheng DM. Photosynthetic characteristics of non leaf organs of winter wheat cultvars differeing inear type and their relationship with grain mass per ear. Photosynthetica,2001,39(2):239-244
[215] Wardlaw IF. Interaction between drought and chronic high temperature during kernel filling in wheat in a controlledenvrironment. Ann Bot,2002,90(4):469-476
[216] Warren CR&Adams MA. Phosphorus affects growth and partitioning of nitrogen to Rubisco in Pinus pinaster. TreePhysiol,2002,22(1):11-19
[217] Warren CR&Adams MA. Evergreen trees do not maximize instantaneous photosynthesis. Trends Plant Sci,2004,9(6):270-274
[218] Webber AN, Nie GY, Long SP. Acclimation of photosynthetic proteins to rising atmospheric CO2. Photosynth Res,1994,39(3):413-425
[219] Weete JD, Leek G.L, Peterson CM, Currie HE and Branch WD. Lipid and surface wax synthesis in water-stressedcotton leaves. Plant Physiol,1978,62(5):675-677
[220] Wendel JF. New World tetraploid cottons contain Old-World cytoplasm. Proc Natl Acad Sci USA,1989,86(11):4132-4136
[221] Wendel JF&Albert VA. Phylogenetics of the cotton genus (Gossypium): Character-state weighted parsimony analysisof chloroplast-DNA restriction site data and its systematic and biogeographic implications. Systematic Botany,1992,17:115-143
[222] Willmer CM. Morphology and anatomy of stomata. In Stomata. London: Longman,1983
[223] Woodson JD&Chory J. Coordination of gene expression between organellar and nuclear genomes. Nat Rev Genet,2008,9(5):383-395
[224] Wullschleger SD, Oosterhuis DM, Rutherford SA. Importance of bracts in the carbon economy of cotton. ArkansasFarm Res,1990,39(3):4
[225] Wullschleger SD&Oosterhuis DM. Photosynthetic and respiratory activity of fruiting forms within the cottoncanopy. Plant Physiol,1990a,94(2),463-469
[226] Wullschleger SD&Oosterhuis DM. Photosynthetic carbon production and use by developing cotton leaves and bolls.Crop Sci,1990b,30(6):1259-1264
[227] Wullschleger SD&Oosterhuis DM. Photosynthesis, transpiration, and water-use efficiency of cotton leaves and fruit.Photosynthetica,1991,25:505-515
[228] Wullschleger SD, Oosterhuis DM, Hurren RG, Hanson PJ. Evidence for light-dependent recycling of respired carbondioxide by the cotton fruit. Plant Physiol,1991,97(2):574-579
[229] Xu HL, Ishii R, Yamagishi T, Kumura A. Effects of water deficit on photosynthesis in wheat plants. III. Effect onnon-stomatal mediated photosynthesis and RuBP carboxylase content in different plant parts. Jpn J Crop Sci,1990,59(1):153-157
[230] Xu HL, Gauthier L, Desjardins Y, Gosselin A. Photosynthesis in leaves, fruits, stem and petioles of greenhouse-growntomato plants. Photosynthetica,1997,33(1):113-123
[231] Xu XL&Ishii R. Wheat cultivar differences in photosynthetic response to low soil water potentials:1. Marintenance ofphotosynthesis and leaf water potential. Jpn.J Crop Sci,1996,65:509-517
[232] Xu ZZ&Zhou GS. Responses of leaf stomatal density to water status and its relationship with photosynthesis in agrass. J Exp Bot,2008,59(12):3317-3325
[233] Xu ZZ, Zhou GS, Wang YL, Han GX, Li YJ. Changes in chlorophyll fluorescence in maize plants with imposed rapiddehydration at different leaf ages. J Plant Growth Regulat,2008,27(1):83-92
[234] Yabuta Y, Motoki T, Yoshimura K, Takeda T, Ishikawa T and Shigeoka S. Thylakoid membrane-bound ascorbateperoxidase is a limiting factor of antioxidative systems under photo-oxidative stress. Plant J,2002,32(6):915-925
[235] Yang JC, Zhang JH, Wang ZQ, Zhu QS, Liu LJ. Activities of starch hydrolytic enzymes and sucrose-phosphatesynthase in the stems of rice subjected to water stress during grain filling. J Exp Bot,2001,52(364):2169-2179
[236] Yamori W, Takahashi S, Makino A, Price GD, Badger MR, von Caemmerer S. The roles of ATP synthase and theCytochrome b6/f complexes in limiting chloroplast electron transport and determining photosynthetic capacity. PlantPhysiol,2011,155(2):956-962
[237] Zhang YH, Wang ZM, Shu HW. Phosphoenolpyruvate carboxylase activity in ear organs is related to proteinconcentration in grains of winter wheat. J Cereal Sci,2008,47(2):386-391
[238] Zhang YP, Zhang YH, Wang ZM, Wang ZJ. Characteristics of canopy structure and contributions of non-leaf organs toyield in winter wheat under different irrigated conditions. Field Crop Res,2011,123(3):187-195
[239] Zhao DL&Oosterhuis DM. Photosynthetic capacity and carbon contribution of leaves and bracts to developing floralbuds in cotton. Photosynthetica,1999,36(1-2):279-290
[240] Zhu XG, Ort DR, Whitmarsh J, Long SP. The slow reversibility of photosystem Ⅱ thermal energy dissipation ontransfer from high to low light may cause large losses in carbon gain by crop canopies: a theoretical analysis. J Exp Bot,2004,55(400):1167-1175
[241] Ziegler J ns A. Gas-exchange of ears of cereals in response to carbon dioxide and light. I. Relative contributions ofparts of the ears of wheat, oat, and barley to the gas exchange of the whole organ. Planta,1989,178(2):84-91