Characterization of photosynthesis, photoinhibition and the activities of C4 pathway enzymes in a superhigh-yield rice, Liangyoupeijiu

Characteristics of photosynthetic gas exchange, photoinhibition and C4 pathway enzyme activities in both flag leaves and lemma were compared between a superhigh-yield rice (Oryza sativa L.) hybrid, Liangyoupeijiu and a traditional rice hybrid, Shanyou63. Liangyoupeijiu had a similar light saturated assimilation rate (Asat) to Shanyou63, but a much higher apparent quantum yield (AQY), carboxylation efficiency (CE) and quantum yield of CO2 fixation (φCO2). Liangyoupeijiu also showed a higher resistance to photoinhibition and higher non-radiative energy dissipation associated with the xanthophyll cycle than Shanyou63 when subjected to strong light. In addition, Liangyoupeijiu had higher activities of the C4 pathway enzymes in both flag leaves and lemmas than Shanyou63. These results indicate that higher light and CO2 use efficiency, higher resistance to photoinhibition and C4 pathway in both flag leaf and lemma may contribute to the higher yield of the superhigh-yield rice hybrid, Liangyoupeijiu.     

超高产杂交稻不同生育期的光合色素含量、净光合速率和水分利用效率

 较为系统地研究了两个超高产杂交稻‘两优培九’、‘华安3号’和多年来大面积推广的常规杂交稻‘汕优63’不同生育期的光合色素含量、净光合速率和水分利用效率。结果表明,在苗期,3个杂交稻的单位叶面积的叶绿素（Chl）含量差别不大,类胡萝卜素（Car）的含量以‘汕优63’为最高。然而,随着发育阶段的推进,到分蘖期,尤其是抽穗期剑叶的单位叶面积Chl和Car含量,两个超高产杂交稻高于‘汕优63’。从苗期到抽穗期,超高产杂交稻‘两优培九’和‘华安3号’的净光合速率（Pn）都比‘汕优63’高,而在苗期的午间强光条件下和分蘖期的早晨以及抽穗期的早晚相对弱光条件下其Pn的差别尤为显著。在苗期,‘汕优63’的水分利用效率（WUE）略高于‘两优培九’和‘华安3号’;到分蘖期和抽穗期,在上午10时之前和下午14时以后的时间段,则是两个超高产杂交稻明显高于‘汕优63’。这些结果说明,超高产杂交稻‘两优培九’和‘华安3号’不仅有较高的Pn和较强的抗光抑制能力,而且还能充分利用早晨和傍晚较弱的光强进行光合作用,这些是它们之所以高产的重要生理基础。此外,超高产杂交稻在生长发育的中后期（分蘖期和抽穗期）具有较高的WUE,有利于节约农业用水。     

Midday photoinhibition of two newly developed super-rice hybrids

Super-rice hybrids are two-line hybrid rice cultivars with 15 to 20 % higher yields than the raditional three-line hybrid rice cultivars. Response of photosynthetic functions to midday photoinhibition was compared between seedlings of the traditional hybrid rice (Oryza sativa L.) Shanyou63 and two super-rice hybrids, Hua-an3 and Liangyoupeijiu. Under strong midday sunlight, in comparison with Shanyou63, the two super-rice hybrids were less photoinhibited, as indicated by the lower loss of the net photosynthetic rate (P _N), the quantum yield of photosystem 2 (Φ_PS2), and the maximum and effective quantum yield of PS2 photochemistry (F_v/F_m and F_v′/F_m′). They also had a much higher transpiration rate. Hence the super-rice hybrids could protect themselves against midday photoinhibition at the cost of water. The photoprotective de-epoxidized xanthophyll cycle components, antheraxanthin (A) and zeaxanthin (Z), were accumulated more in Hua-an3 and Liangyoupeijiu than in Shanyou63, but the size of xanthophyll cycle pool of the seedlings was not affected by midday photoinhibition. Compared to Shanyou63, the super-rice hybrids were better photoprotected under natural high irradiance stress and the accumulation of Z and A, not the size of the xanthophyll pool protected the rice hybrids against photoinhibition.     

Characterization of photosynthesis, photoinhibition and the activities of C4 pathway enzymes in a superhigh-yield rice, Liangyoupeijiu

Characteristics of photosynthetic gas exchange, photoinhibition and C₄ pathway enzyme activities in both flag leaves and lemma were compared between a superhigh-yield rice (Oryza sativa L.) hybrid, Liangyoupeijiu and a traditional rice hybrid, Shanyou63. Liangyoupeijiu had a similar light saturated assimilation rate (Asat) to Shanyou63, but a much higher apparent quantum yield (AQY), carboxylation efficiency (CE) and quantum yield of CO₂ fixation (ΦCO₂). Liangyoupeijiu also showed a higher resistance to photoinhibition and higher non-radiative energy dissipation associated with the xanthophyll cycle than Shanyou63 when subjected to strong light. In addition, Liangyoupeijiu had higher activities of the C₄ pathway enzymes in both flag leaves and lemmas than Shanyou63. These results indicate that higher light and CO₂ use efficiency, higher resistance to photoinhibition and C₄ pathway in both flag leaf and lemma may contribute to the higher yield of the superhigh-yield rice hybrid, Liangyoupeijiu.     

Relationship between leaf photosynthetic function at grain filling stage and yield in super high-yielding hybrid rice (Oryza sativa. L)

The characteristics of dry matter production before and after heading and the relationships between photosynthesis of flag leaves and dry matter accumulation in panicles were investigated on super high-yielding rice cv. Xieyou 9308 (the yield of up to 12 t/ha) with rice cv. Xieyou 63 as a control. The results showed that (i) the capacity of dry matter production before and after heading in Xieyou 9308, i.e. biomass and daily dry matter production, was remarkably higher than that in Xieyou 63, especially after heading; (ii) CO2 assimilation capacity in flag leaves in Xieyou 9308, namely Leaf Source Capacity (LSC), was also significantly higher than that in Xieyou 63, and the supply of photosynthate in leaves and the demand of grain filling were completely synchronous in Xieyou 9308, but photosynthetic function in flag leaves in Xieyou 63 declined sharply 20 days after heading and it was not enough to meet the demand of grain filling. These results confirmed that high efficient photosynthetic function in     

Relationship Between PSⅡExcitation Pressure and Content of Super High-Yielding Hybrid Rice

The changes in photochemical features of PhotosystemⅡ(PSⅡ) and contents of Rubisco large subunit (RLS) and small subunit (RSS) in flag leaf from 75DAS to 113DAS (from filling to harvesting stages) were investigated in two hybrid rices (Oryza sativa L.) cv. Liangyoupeijiu and cv. Shanyou 63 grown in the field. Liangyoupeijiu is a super high-yielding rice and Shanyou 63 has widely been planted in China in these years. The results indicate that soluble protein and chlorophyll in both cultivars degraded slowly at first and dramatically thereafter. The degradation speed of soluble protein in Shanyou 63 was faster than that in Liangyoupeijiu. Both Fv/Fm and qP decreased in parallel with leaf senescence, whereas qN fell at first and then rose. No significant change in excitation pressure (1-qP ) was found before 89DAS but a sharply increase in both cultivars after it occurred. Excitation pressure rose more rapidly in Shanyou 63 than that in Liangyoupeijiu. The changes of RLS and RSS content exhibited the same trend as that of soluble protein content. A better linear correlation between RLS, RSS degradation and elevation of (1-qP ) were shown in both cultivars. We suggest that the increase in PSⅡ excitation pressure possibly induced the quick senescence process in rice flag leaf. The high-yielding of Liangyoupeijiu may be due to its maintenance of stronger photosynthetic capacity, longer and more stable photosynthetic functional du-ration than that of Shanyou 63.     

水稻抽穗后剑叶衰老过程中光合关键酶的基因表达

协优 930 8、培矮 6 4s/ 9311和汕优 6 3三个超级高产水稻品种抽穗后 ,剑叶中光合速率减小、叶绿素含量下降、SOD活力降低和MDA含量上升的幅度不同 ,显示它们开花结实过程中叶片衰老的速度不同 ,以汕优 6 3衰老最快 ,协优 930 8最慢 ,培矮 6 4s/ 9311居中。Northern杂交和免疫定量研究表明 ,汕优 6 3和培矮6 4s/ 9311剑叶rbcS(Rubisco小亚基基因 )和rca(Ru bisco活化酶基因 )mRNA的表达随抽穗后天数的增加而下降 ,而协优 930 8在抽穗后第 10天才开始下降 ;Rubisco和Rubisco活化酶蛋白含量的下降趋势与rbcS和rcamRNA的变化趋势相似。因此 ,水稻抽穗后剑叶衰老过程中光合能力的下降与光合关键酶基因(rbcS和rca)表达下降密切相关     

Evaluation of genotypic variation in leaf photosynthetic rate and its associated factors by using rice diversity research set of germplasm

In order to evaluate genotypic variation, we measured leaf photosynthetic rate (Pn) and its associated factors for the rice diversity research set of germplasm (RDRS) selected from the Genebank in National Institute of Agrobiological Sciences (NIAS). Pn showed large genotypic variation from 11.9 to 32.1 micromol m(-2 )s(-1). The variation in stomatal conductance to CO2 (Gs) explained about 50% of that in Pn, while that in nitrogen concentration (N) in leaves explained about 35%. The genotype group which mainly consists of aus type indica tended to have higher Gs, and the genotype group which corresponds to japonica had a higher nitrogen concentration (N) in leaves. The relationships of Pn with Gs and N were not significantly different among genotype groups, suggesting photosynthetic efficiencies are similar among genotype groups.     

Relationship between leaf photosynthetic function at grain filling stage and yield in super high-yielding hybrid rice (<Emphasis Type="Italic">Oryza sativa</Emphasis>. L)

The characteristics of dry matter production before and after heading and the relationships between photosynthesis of flag leaves and dry matter accumulation in panicles were investigated on super high-yielding rice cv. Xieyou 9308 (the yield of up to 12 t/ha) with rice cv. Xieyou 63 as a control. The results showed that (i) the capacity of dry matter production before and after heading in Xieyou 9308, i.e. biomass and daily dry matter production, was remarkably higher than that in Xieyou 63, especially after heading; (ii) CO₂ assimilation capacity in flag leaves in Xieyou 9308, namely Leaf Source Capacity (LSC), was also significantly higher than that in Xieyou 63, and the supply of photosynthate in leaves and the demand of grain filling were completely synchronous in Xieyou 9308, but photosynthetic function in flag leaves in Xieyou 63 declined sharply 20 days after heading and it was not enough to meet the demand of grain filling. These results confirmed that high efficient photosynthetic function in leaves after heading and its complete synchronization with grain filling are the key approaches to super high yield of rice.     

Photosynthetic efficiency and nitrogen distribution under different nitrogen management and relationship with physiological N-use efficiency in three rice genotypes

Nitrogen fertilization strategies were widely adopted to enhance grain production and improve nitrogen utilization in rice all over the world. For fertilization timing strategy, ear fertilization was usually employed in recent years. For fertilization amount strategy, nitrogen fertilization would continually increase to meet the demands of increasing people for food. However, under heavy ear fertilization as well as great nitrogen amount (NA), physiological N-use efficiency (PE, defined as grain production per unit nitrogen uptake by plants) decreased. Under three NA and two ratios of fertilization given during ear development period to total NA (ear fertilization distribution ratio, EFDR), net photosynthetic rate (Pn), Pn to nitrogen content per unit area (photosynthetic N-use efficiency, Pn/N), nitrogen accumulation in plant tissues and PE of three rice (Oryza sativaL.) genotypes, Jinyou 253, Liangyoupeijiu and Baguixiang were screened in the first and second seasons in 2002 so as to understand the fluctuation patterns of Pn/N and nitrogen distribution in leaf blades under great NA & EFDR and relationship with PE in rice. Results showed that under greater NA & EFDR, Pn in flag leaves at heading and plant nitrogen accumulation at maturity always increased and PE & Pn/N always decreased in spite of increased grain production. Rice distributed more nitrogen in leaf blade under greater NA and EFDR. PE indicated significantly (P<0.05) positive relationship with Pn/N and negative relationship with nitrogen distribution ratio in leaf blades at heading and maturity, and no association with Pn in two growing seasons. Results suggested that low PE in rice under great NA and heavy ear fertilization is associated to more nitrogen distribution in leaf blades and decreases in photosynthetic efficiency.     

水稻剑叶取向对其光合功能的影响

水稻的水平剑叶净光合速率 (Pn)和羧化效率(CE)显著高于直立剑叶 ,其胞间CO2 浓度 (Ci)显著低于直立剑叶 ,但两者的气孔导度 (Gs)没有明显差别。这表明剑叶取向对水稻叶片的光合能力有重要影响。水平剑叶的高Pn可能同其RuBP羧化酶含量和活性高有关。这可能是水平叶生长期间吸收光量较多的结果。     

PSⅡ Photochemistry and Xanthophyll Cycle in Two Superhigh yield Rice Hybrids,Liangyoupeijiu and Hua an 3 During Photoinhibition and Subsequent Restoration

PSⅡ photochemistry and xanthophyll cycle during photoinhibition (exposed to strong light of 2 000 μmol photons·m-2·s-1) and the subsequent restoration were compared between two superhigh yield rice hybrids (Liangyoupeijiu and Hua an 3, the newly developed rice hybrids from two parental lines) and the traditional rice hybrid Shanyou 63 developed from three parental lines. The results showed that the maximal efficiency of PSⅡ photochemistry (Fv/Fm), the efficiency of excitation energy capture by open PSⅡ centers (Fv′/Fm′), and the yield of PSⅡ electron transport (ΦPSⅡ) of the three rice hybrids decreased during photoinhibition. However, a greater decrease in Fv/Fm, Fv′/Fm′, and ΦPSⅡ was observed in Shanyou 63 than in Liangyoupeijiu and Hua an 3. At the same time, the components of xanthophyll cycle, antherxanthin (A) and zeathanxin (Z) increased rapidly while violaxanthin (V) decreased considerably. Both the rate of accumulation and the amount of A and Z in the two superhigh yield rice hybrids were higher than that in Shanyou 63. The de epoxidation state (DES) of xanthophyll cycle increased rapidly with the fast accumulation of A and Z, and reached the maximal level after first 30 min during photoinhibition. Of the three hybrids, the increasing rate of DES in Liangyoupeijiu and Hua an 3 was higher than that in Shanyou 63. After photoinhibition treatment, the plant materials were transferred to a dim light (70 μmol photons·m-2·s-1) for restoration. During restoration, both chlorophyll fluorescence parameters and xanthophyll cycle relaxed gradually, but the rate and level of restoration in the two superhigh yield rice hybrids were higher than those in Shanyou 63. Our results suggest that Liangyoupeijiu and Hua an 3 had higher resistance to photoinhibition and higher capacity of non radiative energy dissipation associated with xanthophyll cycle, as well as higher rate of restoration after photoinhibition, than Shanyou 63 when subjected to strong light.     

水稻叶片光合作用对开放式空气CO2浓度增高(FACE)的响应与适应

利用便携式光合气体分析系统 (LI 6 4 0 0 ) ,比较测定了高CO2 浓度 (FACE ,free airCO2 enrich ment)和普通空气CO2 浓度下生长的水稻叶片的净光合速率、水分利用率、表观量子效率和RuBP羧化效率等光合参数 .在各自生长CO2 浓度 (380vs 5 80 μmol·mol-1)下测定时 ,高CO2 浓度 (5 80 μmol·mol-1)下生长的水稻叶片的净光合速率、碳同化的表观量子效率和水分利用率明显高于普通空气 (380 μmol·mol-1)下生长的水稻叶片 .但是 ,随着FACE处理时间的延长 ,高CO2 浓度对净光合速率的促进作用逐渐减小 .在相同CO2 浓度下测定时 ,FACE条件下生长的水稻叶片净光合速率和羧化效率明显比普通空气下生长的对照低 .尽管高CO2 浓度下生长的水稻叶片的气孔导度明显低于普通空气中生长的水稻叶片 ,但两者胞间CO2 浓度差异不显著 ,因此高CO2 浓度下生长的水稻叶片光合下调似乎不是由气孔导度降低造成的 .     

超高产杂交稻光合特性的研究

比较了超高产杂交稻(Oryza sativa L.)X07S/紫恢100和两优培九与多年来大面积推广的杂交稻汕优63的光合功能和抗光胁迫能力.结果表明,超高产杂交稻X07S/紫恢100和两优培九的净光合速率(Pn)分别比汕优63高9.1%和11.9%,而其蒸腾速率(Tr)分别比汕优63低37.46%和31.42%,此外,其水分利用效率(WUE)分别比汕优63高出74.2%和63.5%;经强光(2 000μmol photons@m-2@s-1)处理2 h之后,X07S/紫恢100和两优培九的光系统Ⅱ光化学量子效率分别上升28.3%和37.0%,荧光光化学猝灭系数分别升高46.2%和18.0%,而汕优63的这两项参数值却变化很小;同时,X07S/紫恢100和两优培九的非光化学猝灭系数均下降50%左右,而汕优63则上升近50%,表明超高产杂交稻较高的光合能力、水分利用效率以及较强的抗光抑制能力可能是其高产的生理基础.     

Relationship between leaf photosynthetic function at grain filling stage and yield in super high-yielding hybrid rice ( Oryza sativa . L)

The characteristics of dry matter production before and after heading and the relationships between photosynthesis of flag leaves and dry matter accumulation in panicles were investigated on super high-yielding rice cv. Xieyou 9308 (the yield of up to 12 t/ha) with rice cv. Xieyou 63 as a control. The results showed that (i) the capacity of dry matter production before and after heading in Xieyou 9308, i.e. biomass and daily dry matter production, was remarkably higher than that in Xieyou 63, especially after heading; (ii) CO₂ assimilation capacity in flag leaves in Xieyou 9308, namely Leaf Source Capacity (LSC), was also significantly higher than that in Xieyou 63, and the supply of photosynthate in leaves and the demand of grain filling were completely synchronous in Xieyou 9308, but photosynthetic function in flag leaves in Xieyou 63 declined sharply 20 days after heading and it was not enough to meet the demand of grain filling. These results confirmed that high efficient photosynthetic function in leaves after heading and its complete synchronization with grain filling are the key approaches to super high yield of rice.     

超高产杂交稻剑叶中C4途径酶活性和稳定碳同位素分异作用的变化

以超高产杂交水稻(Oryza sativa L.)"培矮64S/E32"和多年来大面积推广的杂交稻"汕优63"为材料,研究孕穗后剑叶中C4途径酶和对稳定碳同位素分异作用的变化.结果表明,籽粒灌浆期(移栽后68～75 d)的两个品种剑叶中NADP-MDH活性最高,随后下降;超高产杂交水稻"培矮64S/E32"的NADP-MDH的活性明显高于"汕优63";PEPCase和NADP-ME活性在黄熟期之前的叶片中持续上升.不同生育期的叶片与籽粒的△1aC值相近(19.49‰～19.82‰),在成熟期时较高.超高产水稻"培矮64S/E32"叶片的平均△13C值比"汕优63"高0.43‰.     

水稻亚种间杂种F1 光合特性研究

用绵恢725、蜀恢527和蜀恢881三个籼型恢复系、1个美国稻Lemont和1个爪哇稻香大粒作母本, 与1个日本特早熟粳稻Kitaake杂交, 研究了5个杂种F1及其亲本的光合生理表现。结果表明, 在高光通量密度(Photosynthetic flux density, PFD)条件下, 5个杂种F1净光合速率(Pn)明显高于双亲或双亲之一, 推测亲本与杂种F1之间不同的Pn同叶片中Rubsico活性有关。杂种F1的比较中, 在表观量子效率(j)、羧化效率(CE)、CO2补偿点(G)等方面, 籼粳亚种间杂种F1(绵恢725/Kitaake、蜀恢527/Kitaake、蜀恢881/Kitaake)对2个亚种内杂种F1香大粒/Kitaake(粳爪交)、Lemont/Kitaake(不同生态型的粳粳交, 美国稻属于特殊粳稻)具有明显的优势, 而蜀恢881含有粳型血缘, 蜀恢881/Kitaake也比典型籼粳亚种间杂种F1 绵恢725/Kitaake、蜀恢527/Kitaake优势稍逊一筹。5个杂种F1因为具有不同的遗传差异而表现出不同的光合优势, 在这方面, 典型籼粳亚种间杂交蜀恢527/Kitaake、绵恢725/Kitaake要优于其他杂交种, 说明亲本间遗传差异越大, 其杂种F1的光合优势越强。     

油菜素内酯对低氧胁迫下黄瓜幼苗叶片线粒体、叶绿体超微结构及光合的影响

在水培条件下,研究24-表油菜素内酯(EBR)对低氧胁迫下黄瓜幼苗叶片叶绿体和线粒体超微结构及光合的影响.结果表明:与正常通气条件相比,低氧胁迫下表观量子效率(AQY)和羧化效率(CE)显著降低,而光补偿点(LCP)、暗呼吸速率(Rd)和CO2补偿点(CCP)显著升高;低氧胁迫并添加油菜素内酯处理下,CE与低氧胁迫处理相比显著提高29.4％,而LCP和Rd分别显著下降15.0％和14.4％.光响应Pn-PPFD曲线和CO2响应Pn-Ci曲线表明,低氧胁迫下净光合速率(Pn)增幅降低,而添加油菜素内酯有利于Pn增幅的提高.低氧胁迫下叶绿体和线粒体结构受到伤害,而油菜素内酯可以缓解低氧胁迫对黄瓜幼苗叶绿体和线粒体超微结构的不良影响,使叶片维持较好的光合性能.     

低温对杂交水稻乳熟期剑叶光合作用和光合产物运输的影响

为了解寒露风引起晚稻减产的生理原因,本试验以两个不同抗冷力的杂交水稻组合为材料,研究了低温对水稻乳熟期剑叶光合作用与果糖１,６－二磷酸酯酶（ＦＢＰａｓｅ）活性的影响。结果表明：低温引起离体剑叶光合效率与ＦＢＰａｓｅ 活性下降,不抗冷的“汕优６３”及其亲本的光合效率与ＦＢＰａｓｅ 活性比抗冷的“秀优５７”及其亲本下降幅度较大。乳熟期是籽粒灌浆的高峰期,自然低温（寒露风）造成晚造水稻减产与乳熟期连体剑叶的光合作用下降和ＦＢＰａｓｅ 对低温的适应和调节能力有关