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（２０     

超高产杂交稻两优培九的光合作用、光抑制和C4途径酶特性

比较研究了超高产杂交稻(Oryza sativa L.)两优培九和常规杂交稻汕优63剑叶的光合气体交换、光抑制, 以及其剑叶和稃片中的C₄酶活性. 发现两优培九和汕优63的光饱和同化速率(Asat)与汕优63相差无几, 但是, 其表观量子效率(AQY)、羧化效率(CE)和CO₂同化的量子效率(FCO₂)都远远高于汕优63. 强光条件下, 超高产杂交稻两优培九表现出比汕优63更强的抗光抑制能力, 并且具有较高的以叶黄素循环为量度的非辐射能量耗散. 此外, 两优培九的剑叶和稃片中的C₄途径酶活性均高于汕优63. 这些结果表明, 较高的光能和CO₂利用效率、较强的抗光抑制能力, 以及剑叶和稃片中C₄途径的较高表达可能是超高产杂交稻两优培九高产的重要保证.     

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。     

超高产杂交稻在强光胁迫及其恢复进程中的PSⅡ活性和叶黄素循环

研究了两系超高产杂交稻(Oryza sativa L.)"两优培九"和"华安3号"以及多年来大面积推广的三系杂交稻"汕优63"剑叶的PSⅡ活性和叶黄素循环对强光胁迫及其恢复进程的响应.结果表明,在2 000 μmol photons*m-2*s-1的强光胁迫下,3个杂交稻的PSⅡ光化学最大效率(Fv/Fm)、开放的PSⅡ反应中心捕获激发能效率(Fv′/Fm′)和PSⅡ的实际光化学转能效率(ΦPSⅡ)都随着光抑制进程而下降,其中以"汕优63"下降的幅度最大.光抑制过程中,杂交稻叶黄素循环的紫黄素(V)组分迅速下降,与此同时,脱环氧化组分环氧玉米黄素(A)和玉米黄素(Z)迅速积累,而超高产杂交稻"两优培九"和"华安3号"的A和Z的积累速度大大高于"汕优63".伴随A和Z的快速积累,它们的叶黄素循环的脱环氧化状态(DES)迅速上升,并在半小时左右基本达到最大值,其中"两优培九"和"华安3号"DES的上升速率仍然较"汕优63"高.光抑制处理结束后,转移到弱光(70 μmol photons*m-2*s-1)条件下恢复过程中,两个超高产杂交稻的Fv/Fm、Fv′/Fm′和ΦPSⅡ的恢复速率和恢复水平都高于"汕优63".同时,它们的叶黄素循环组分V、A、Z以及DES都逐渐恢复,但"两优培九"和"华安3号"的恢复速率和恢复水平仍然要优于"汕优63".以上结果说明,超高产杂交稻"两优培九"和"华安3号"较对照品种"汕优63"具有更强的抗光抑制及光保护能力,同时在光抑制结束后又能够更迅速地恢复光合功能,较强的抗光抑制能力和较高的恢复能力可能是超高产杂交稻高产的重要生理基础之一.     

在光氧化条件下不同类型高产稻的叶绿素荧光特性和膜脂过氧化特点

用不同类型高产稻(Oryza sativa L.)粳稻9516、具有粳型成分的两系法亚种间杂交稻培矮64/E32、两优培九(培矮64/9311)和籼型杂交稻X07S/紫恢100、冈优881、汕优63为材料,研究了孕穗期叶片在光氧化条件下的叶绿素荧光特性和膜脂过氧化表现.光氧化处理后,与籼型杂交稻比较,粳稻和具有粳型组分的亚种间杂交稻的PSⅡ原初光化学效率(Fv/Fm)、PSⅡ的线性电子传递的量子效率(ΦPSⅡ)和光化学猝灭系数(qP)下降的较少;超氧化物歧化酶(SOD)、过氧化氢酶(CAT)、过氧化物酶(POD)诱导的活性较高,活性氧 (O(-)/()2、H2O2)和丙二醛(MDA)的产生积累较少,叶绿素和蛋白质含量下降较少,表现出耐光氧化特性,这与在自然条件下生育后期叶绿素含量变化相一致.相关分析表明它们的耐光氧化特性与结实率密切相关,说明耐光氧化品种抗早衰,有利籽粒充实.这些结果启示我们:从超高产育种出发,兼顾杂种优势利用和抗早衰两方面考虑,在母本不育系中引入粳型成分是一个值得重视的育种策略.     

自然条件下不同高产稻生育后期剑叶叶绿素荧光和膜脂过氧化的表现

以籼型(Oryza sativa L.)杂交组合汕优63为对照,以中粳9516、两系亚种间杂交组合培矮64S/E32、培矮64S/9311、亚种间三系杂交稻冈优881和两系杂交组合X07S/紫恢100为材料,研究其在生育后期(抽穗-成熟)自然条件下剑叶的叶绿素衰减、CO2交换、叶绿素荧光参数和膜脂过氧化表现.结果表明: 水稻在生育后期伴随叶绿素衰减,其叶内的原初光化学效率Fv/Fm、PSⅡ非环式电子传递效率ΦPSⅡ、电子流传递速率ETR都有相应地下降,这种光能转化的障碍使多余的光能传递给PSⅡ的还原侧,产生O(-)/(*)2累积,发生膜脂过氧化和MDA的积累,引起光合色素及光合膜的破坏,发生光氧化早衰.这种现象在品种间有明显差异,耐光氧化的粳稻9516,其叶内的 Fv/Fm、ΦPSⅡ、ETR、qP下降较少,具有较稳定的光能转化能力,不易早衰,具有较高的结实率;而对光氧化敏感的籼稻汕优63其叶内的Fv/Fm、ΦPSⅡ、ETR,光化学猝灭参数qP下降较多,易发生膜脂过氧化,导致叶片早衰,影响水稻灌浆结实和产量;而二系的和三系的杂交稻的耐光氧化特性和早衰表现居于中间.从水稻超高产育种的角度出发,在目前株型良好的基础上,兼顾杂种优势和防止早衰两方面考虑,在母本中利用粳型或带有粳型基因的不育系是育种上一个值得重视的策略.     

超级杂交稻两优培九及其亲本的光氧化特性

通过比较超级杂交稻两优培九和其亲本的光能利用与活性氧代谢的差异,为培育耐光氧化的杂交稻提供选亲配组的生理依据．使用TPS-光合仪和FMS2荧光仪(Hansateeh,UK)分别测定了人工光氧化处理后水稻叶片的光合速率与叶绿素荧光,同时测定了叶绿素、蛋白质和丙二醛的含量。以及超氧化物歧化酶和过氧化物酶的活性．结果表明,两优培九的光合速率低于母本2．4％,高于父本23％;光氧化处理8d后,与母本相比。两优培九的叶绿素、蛋白质、原初光化学效率、超氧化物歧化酶和过氧化物酶分别高于母本33％、15％、30％、32％和100％;光化学猝灭系数和丙二醛分别低于母本9％和50％。与父本相比,则差异不大．超级杂交稻两优培九在光氧化条件下光能利用率较高,在耐光氧化特性上具有超亲的光合生理特性．     

在光氧化条件下不同类型高产稻的叶绿素荧光特性和膜脂过氧化特点

用不同类型高产稻 (OryzasativaL .)粳稻 95 16、具有粳型成分的两系法亚种间杂交稻培矮 6 4/E32、两优培九(培矮 6 4/ 9311)和籼型杂交稻X0 7S/紫恢 10 0、冈优 881、汕优 6 3为材料 ,研究了孕穗期叶片在光氧化条件下的叶绿素荧光特性和膜脂过氧化表现。光氧化处理后 ,与籼型杂交稻比较 ,粳稻和具有粳型组分的亚种间杂交稻的PSⅡ原初光化学效率 (Fv/Fm)、PSⅡ的线性电子传递的量子效率 (ΦPSⅡ)和光化学猝灭系数 (qP)下降的较少 ;超氧化物歧化酶 (SOD)、过氧化氢酶 (CAT)、过氧化物酶 (POD)诱导的活性较高 ,活性氧 (O-·2 、H2 O2 )和丙二醛 (MDA)的产生积累较少 ,叶绿素和蛋白质含量下降较少 ,表现出耐光氧化特性 ,这与在自然条件下生育后期叶绿素含量变化相一致。相关分析表明它们的耐光氧化特性与结实率密切相关 ,说明耐光氧化品种抗早衰 ,有利籽粒充实。这些结果启示我们 :从超高产育种出发 ,兼顾杂种优势利用和抗早衰两方面考虑 ,在母本不育系中引入粳型成分是一个值得重视的育种策略。     

超高产杂交稻剑叶衰老过程中PSII功能的变化

对田间自然条件下培矮64S/E32和对照汕优63的剑叶衰老过程中PSII光化学效率、光能耗散和叶黄素循环组分的动态变化进行了研究.结果表明,水稻剑叶衰老过程中PSⅡ光化学效率(F/Fm)、PSII量子产量(φPSII)、光化学猝灭系数(qP)和光合速率(Pn)均降低,但非辐射热耗散(NPQ)、叶黄素循环库和中午时的脱环氧化状态则升高.移栽后55-75 d两品种剑叶的φPSII和qP变化相近,此后培矮64S/E32比汕优63下降缓慢.在生育后期的衰老过程中培矮64S/E32剑叶的PSII功能比汕优63下降慢,并具有更高的光能利用效率.     

超级杂交稻两优培九及其亲本低温强光适应特性

以超级杂交稻两优培九(培矮64S/9311)和母本培矮64S(PA64S)以及父本中籼9311为材料,以叶绿素荧光参数和活性氧代谢等指标研究生育后期亲本和后代对低温强光的适应特性。结果显示:低温强光处理后,与父母本相比,两优培九的叶绿素、PSⅡ原初光化学效率(Fv/Fm)和光化学猝灭系数(qP)和实际PSⅡ光能捕获的效率(φPSⅡ)的降低较少,说明在低温强光下,超级杂交稻两优培九吸收的光能较多地转化为化学能;同时两优培九的内源活性氧清除酶系如超氧化物歧化酶和过氧化物酶诱导的活性高,表现耐低温强光的特性,从而能有效清除叶片内的活性氧,因而膜脂过氧化较轻。试验结果表明:在生育后期超级杂交稻两优培九对低温强光适应的特性具有超双亲偏母本的现象。     

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.     

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.     

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 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.     

超高产水稻组合‘培矮64S/E32’的耐光氧化特性及其机理

甲基紫精（MV）介导的光氧化导致两个水稻品种‘汕优63’(目前我国大面积推广的杂交水稻)和‘培矮64S/E32’（新培育的超高产杂交水稻）细胞电解质渗漏率都增加,前者增加的量大于后者,显示‘培矮64S/E32’的细胞膜系统受光氧化的伤害小。光氧化条件下,‘培矮64S/E32’仍能维持较高的光合放氧能力（Amax、Φi）、PSⅡ活性（Fv/Fm、ΦPSⅡ）和叶绿素荧光猝灭系数（qP、NPQ）,而且光氧化引起‘培矮64S/E32’这些参数的下降幅度也小。另外,光氧化导致了两个水稻品种抗氧化酶SOD和APX活性增加,‘培矮64S/E32’增加的幅度约为‘汕优63’的3倍。结果表明超高产水稻具有更强的耐光氧化能力。     

Photosynthetic light and CO2 utilization and C4 traits of two novel super-rice hybrids

Characteristics of photosynthetic light and CO2 use efficiency from seedling to heading stage, and C4 pathway enzyme activities in both flag leaves and lemma were compared between two newly developed super-rice hybrids (Oryza sativa L.), Liangyoupeijiu and Hua-an 3, and a traditional rice hybrid, Shanyou 63. At seedling and tillering stages, Liangyoupeijiu and Hua-an 3 had higher net photosynthetic rates (Pn) and light saturated assimilation rates (Asat) than did Shanyou 63, at both normal (360 micromol mol(-1)) and doubled (720 micromol mol(-1)) CO2 concentrations. At the heading stage, the flag leaves of all three rice hybrids had similar Pn and Asat. However, the two super-rice hybrids had higher apparent quantum yield (AQY) and carboxylation efficiency (CE) during all three typical developmental stages, and higher quantum yield of CO2 fixation (PhiCO2) at the tillering and heading stages. In addition, Liangyoupeijiu showed significantly higher activities of the C(4) pathway enzymes in both flag leaves and lemmas than did Shanyou 63. As a result, flag leaves of the two super-rice hybrids had higher Pn at morning, noontime and late afternoon during the daily cycle. Since most of the grain yield of rice comes from the photosynthesis of flag leaves, the similar Asat and much higher AQY, CE and PhiCO2 at heading stage of the two super-rice hybrids indicates that higher photosynthetic efficiency rather than higher photosynthetic capacity may be the primary factor contributing to their higher grain yields.