玉米C_4光合酶基因转化超级杂交稻亲本及转基因水稻生物学特性研究
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摘要
光合作用是作物产量形成的生理基础,光合作用的强弱一定程度上决定了作物产量的高低。当前,我国超级稻育种正面临着产量“爬坡”的窘境,为进一步挖掘水稻产量潜力,通过转移和表达C4型光合酶基因来提高光合生产能力的高光效基因工程育种已成为水稻超高产育种研究的重要策略和主要内容。
为探讨以不同形式引入C4光合途径来改良主导恢复系光合特性的效果,本研究设置了3种不同光合酶基因互作表达形式,通过农杆菌介导法分别将C4型光合酶基因PEPC/PPDK和NADP-MDH/NADP-ME的双价cDNA转入超级杂交稻亲本湘恢299中,和通过转PEPC/PPDK和NADP-MDH/NADP-ME双价cDNA植株的杂交聚合形成四价转基因水稻为材料,较系统地研究了外源光合酶基因在湘恢299中的整合、转录与表达情况,分析了不同目的基因的表达对水稻光合生理、抗逆、品质和产量等基本生物学特性的影响,得出主要结论如下：
(1) PEPC、PPDK、NADP-MDH和NADP-ME基因能在水稻叶片中转录与表达,转基因植株PEPC的表达量为对照的1.3-5.9倍,NADP-ME的表达量为对照的1.4-6.8倍。且外源光合酶基因在转基因植株后代中可稳定遗传和高效表达。
(2)剔除选择标记的转目的基因植株平均分离频率为8.2%；PEPC、NADP-MDH酶活性与光合速率的相关系数为0.817和0.610,呈显著正相关。
(3)在水稻孕穗期、始穗期、齐穗期,转PEPC/PPDK基因恢复系平均光合速率比对照分别提高12.05%、8.14%、13.83；转基因杂交稻平均光合速率比对照组合分别提高3.59%、7.17%、9.51%。
(4)自始穗至成熟期,伴随着生育进程转基因株系叶绿素SPAD值呈下降趋势,对应于转基因水稻光合优势由强转弱的齐穗至乳熟期区段。
(5)转基因水稻CO2补偿点和光补偿点降低,平均光饱和点提高167μmol photos.m2.m-1,平均光饱和光合速率提高10.97%。在正常生长状态下,转光合酶基因水稻一定程度上表现为蒸腾速率、气孔导度和胞间CO2浓度的提高；气孔导度与光合速率呈显著正相关,与蒸腾速率和胞间CO2浓度呈一定的正相关；推测转基因水稻光合速率得以提高的可能机理在于增强了气孔导度。
(6)在干旱胁迫下,抽穗期转CO2光合酶基因水稻比对照具有较高的蒸腾速率、胞间CO2浓度和气孔导度,光合速率比对照提高75%-85%,水分利用效率比对照提高15%-85%。在高温胁迫下,苗期转C4光合酶基因水稻比对照具有较高的1级叶片比率。在干旱和高温胁迫下,转C4光合酶基因水稻均具有较高的PSⅡ最大光化学效率(Fv／Fm)和PSⅡ潜在活性(Fv／F0)。表明C4光合酶基因的表达对提高水稻的耐旱和耐高温能力有一定作用。
(7)转基因植株稻米垩白粒率、垩白面积和垩白度等都有不同程度地减少,部分株系达到显著差异水平。
(8)转基因恢复系和杂交稻普遍表现为株高增加,穗粒数和千粒重增多,但结实率降低；恢复系和杂交稻的理论单株产量增幅范围分别为-8.32%～19.10%和-19.34%～20.52%；转基因杂交稻测产结果显示,有3个组合产量增产幅度为7.45%、2.54%和1.13%。
综合看来,转PEPC/PPDK基因水稻在酶表达量、光合速率、气孔导度、耐热性、穗粒数和单株产量较有优势,转NADP-MDH/NADP-ME基因在抗旱能力方面略有优势,而四价转基因水稻多数性状上介于转PEPC/PPDK基因和转NADP-MDH/NADP-ME基因水稻之间。至于导致这种结果的可能机理以及生育后期叶绿素降解和光合能力衰退加快、结实率下降和整体产量不高等问题还有待深入研究。
It has been proved that photosynthesis is the physiological basis in crop yield formation, and to some extent, the intensity of photosynthesis determines the level of crop production. At present, China is facing the dilemma to further increase yield in super hybrid rice breeding. In order to promote rice yield potentiality, the genetic engineering by which C4 photosynthetic enzyme genes are introduced and expressed to increase the photosynthetic efficiency has become an important strategy and the main content in super high-yield rice breeding.
To explore the effects on improving photosynthetic characteristics by introducing C4 photosynthetic enzyme genes in different interactive forms,3 expression modules were set up, i.e., by Agrobacterium-mediated method, the maize cDNA of PEPC/PPDK and that of NADP-MDH/NADP-ME were transformed into Xianghui299 (R299), a super hybrid rice parental line, respectively, and through hybridization of two transgenic lines harbored with cDNA of PEPC/PPDK and that of NADP-MDH/NADP-ME, a new transgenic line combined with all four genes above was acquired. On this basis, the integration, transcription and expression of foreign C4 photosynthetic enzyme genes were systematically studied, and effects of the expression of different target genes were analyzed on the biological characteristics such as photosynthesis, stress tolerance, grain quality and yield. The major results were shown as below:
(1) It was found that PEPC, PPDK, NADP-MDH and NADP-ME genes could be integrated, transcribed and expressed in rice leaves. The enzyme activity of PEPC and ME in transgenic rice plants were 1.3 to 5.9 and 1.4 to 6.8 times of those in wild type plants, respectively. The foreign C4 photosynthetic enzyme genes could be stably inherited and effectively expressed in progenies.
(2) The average segregated percentage of the transgenic plants with target genes but selectable marker gene free (SMF) was 8.2%. Several selectable marker-free transgenic homozygous lines of T3 generation were obtained. The correlation coefficients of PEPC and NADP-MDH to Pn (photosynthetic rate) were 0.817 and 0.610, both of which are significant at the 5% level.
(3) In the booting, heading, full heading stages, the average Pn of PEPC/PPDK transgenic restorer lines was higher than that of the control by 12.05%,8.14%,13.83%, respectively, while the average Pn of PEPC/PPDK transgenic rice hybrids was higher than that of the control by 3.59%,7.17%,9.51%, respectively.
(4) From heading stage to mature stage, the SPAD value of chlorophyll in transgenic leaves declined gradually along with the reproductive process. The occurrence of turning point of chlorophyll SPAD (10DAF) was roughly consistent to the full heading stage, after which Pn declined dramatically.
(5) Compared with the control, the transgenic restorer lines with PEPC/PPDK, NADP-MDH/NADP-ME or all four genes had a little bit lower CO2 compensation point and light compensation point but a higher light saturation point by 167μmol photos.m 2.m-1 and a higher light saturated photosynthetic rate by 10.97%. Under normal conditions, the transgenic plants displayed enhanced transpiration rate (VPD), stomatal conductance (Gs) and intercellular CO2 concentration (Ci) to some extent. Gs was positively and significantly correlated to Pn, and positively but not significantly correlated to VPD and Ci. It could be inferred that the enhanced stomatal conductance contributed a lot to its higher Pn.
(6) Under drought stress, in heading stage the transgenic rice showed higher transpiration rate, intercellular CO2 concentration and stomatal conductance than the control; and compared with the control, the Pn and WUE (water use efficiency) of the transgenic rice were increased by 75%-85% and 15%-85%, respectively. Under high temperature stress, in seedling stage the transgenic rice showed higher ratio of 1st grade leaves. Under both drought and high temperature stress conditions, the transgenic rice had higher maximal photochemical efficiency of PSII (Fv/Fm) and higher PSII potential activity (Fv/F0). Therefore, it could be inferred that expression of the foreign C4 photosynthetic genes served a useful function for resistance to drought and high temperature.
(7) The transgenic rice carrying C4 photosynthetic enzyme genes decreased in chalky rice percentage, chalky area and chalkiness to a different extent, and in some cases, the differences reached a statistically significant level between the transgenic rice and the control.
(8) The transgenic restorer lines and their hybrids showed an increase in plant height, the number of spikelets per panicle and grain weight but a decrease in seed setting rate. Assessment of yield capacity test of the transgenic hybrids showed that three had a higher yield than the control hybrids by 7.45%,2.54% and 1.13%.
On the whole, for PEPC/PPDK transgenic rice, it was superior to the others in enzyme expression, Pn, Gs, high temperature resistance, the number of spikelets per panicle and grain yield per plant. The NADP-MDH/NADP-ME transgenic rice had an advantage on drought resistance over the others. When it comes to the transgenic rice carrying all four genes, most traits had values between the PEPC/PPDK transgenic rice and the NADP-MDH/NADP-ME transgenic rice.The possible mechanism leading to this result and the problems such as faster degradation of chlorophyll and sharper decline of photosynthetic capacity in later growth stages, and lower spikelet fertility and yield advantage should be further studied.
为探讨以不同形式引入C4光合途径来改良主导恢复系光合特性的效果,本研究设置了3种不同光合酶基因互作表达形式,通过农杆菌介导法分别将C4型光合酶基因PEPC/PPDK和NADP-MDH/NADP-ME的双价cDNA转入超级杂交稻亲本湘恢299中,和通过转PEPC/PPDK和NADP-MDH/NADP-ME双价cDNA植株的杂交聚合形成四价转基因水稻为材料,较系统地研究了外源光合酶基因在湘恢299中的整合、转录与表达情况,分析了不同目的基因的表达对水稻光合生理、抗逆、品质和产量等基本生物学特性的影响,得出主要结论如下：
(1) PEPC、PPDK、NADP-MDH和NADP-ME基因能在水稻叶片中转录与表达,转基因植株PEPC的表达量为对照的1.3-5.9倍,NADP-ME的表达量为对照的1.4-6.8倍。且外源光合酶基因在转基因植株后代中可稳定遗传和高效表达。
(2)剔除选择标记的转目的基因植株平均分离频率为8.2%；PEPC、NADP-MDH酶活性与光合速率的相关系数为0.817和0.610,呈显著正相关。
(3)在水稻孕穗期、始穗期、齐穗期,转PEPC/PPDK基因恢复系平均光合速率比对照分别提高12.05%、8.14%、13.83；转基因杂交稻平均光合速率比对照组合分别提高3.59%、7.17%、9.51%。
(4)自始穗至成熟期,伴随着生育进程转基因株系叶绿素SPAD值呈下降趋势,对应于转基因水稻光合优势由强转弱的齐穗至乳熟期区段。
(5)转基因水稻CO2补偿点和光补偿点降低,平均光饱和点提高167μmol photos.m2.m-1,平均光饱和光合速率提高10.97%。在正常生长状态下,转光合酶基因水稻一定程度上表现为蒸腾速率、气孔导度和胞间CO2浓度的提高；气孔导度与光合速率呈显著正相关,与蒸腾速率和胞间CO2浓度呈一定的正相关；推测转基因水稻光合速率得以提高的可能机理在于增强了气孔导度。
(6)在干旱胁迫下,抽穗期转CO2光合酶基因水稻比对照具有较高的蒸腾速率、胞间CO2浓度和气孔导度,光合速率比对照提高75%-85%,水分利用效率比对照提高15%-85%。在高温胁迫下,苗期转C4光合酶基因水稻比对照具有较高的1级叶片比率。在干旱和高温胁迫下,转C4光合酶基因水稻均具有较高的PSⅡ最大光化学效率(Fv／Fm)和PSⅡ潜在活性(Fv／F0)。表明C4光合酶基因的表达对提高水稻的耐旱和耐高温能力有一定作用。
(7)转基因植株稻米垩白粒率、垩白面积和垩白度等都有不同程度地减少,部分株系达到显著差异水平。
(8)转基因恢复系和杂交稻普遍表现为株高增加,穗粒数和千粒重增多,但结实率降低；恢复系和杂交稻的理论单株产量增幅范围分别为-8.32%～19.10%和-19.34%～20.52%；转基因杂交稻测产结果显示,有3个组合产量增产幅度为7.45%、2.54%和1.13%。
综合看来,转PEPC/PPDK基因水稻在酶表达量、光合速率、气孔导度、耐热性、穗粒数和单株产量较有优势,转NADP-MDH/NADP-ME基因在抗旱能力方面略有优势,而四价转基因水稻多数性状上介于转PEPC/PPDK基因和转NADP-MDH/NADP-ME基因水稻之间。至于导致这种结果的可能机理以及生育后期叶绿素降解和光合能力衰退加快、结实率下降和整体产量不高等问题还有待深入研究。
It has been proved that photosynthesis is the physiological basis in crop yield formation, and to some extent, the intensity of photosynthesis determines the level of crop production. At present, China is facing the dilemma to further increase yield in super hybrid rice breeding. In order to promote rice yield potentiality, the genetic engineering by which C4 photosynthetic enzyme genes are introduced and expressed to increase the photosynthetic efficiency has become an important strategy and the main content in super high-yield rice breeding.
To explore the effects on improving photosynthetic characteristics by introducing C4 photosynthetic enzyme genes in different interactive forms,3 expression modules were set up, i.e., by Agrobacterium-mediated method, the maize cDNA of PEPC/PPDK and that of NADP-MDH/NADP-ME were transformed into Xianghui299 (R299), a super hybrid rice parental line, respectively, and through hybridization of two transgenic lines harbored with cDNA of PEPC/PPDK and that of NADP-MDH/NADP-ME, a new transgenic line combined with all four genes above was acquired. On this basis, the integration, transcription and expression of foreign C4 photosynthetic enzyme genes were systematically studied, and effects of the expression of different target genes were analyzed on the biological characteristics such as photosynthesis, stress tolerance, grain quality and yield. The major results were shown as below:
(1) It was found that PEPC, PPDK, NADP-MDH and NADP-ME genes could be integrated, transcribed and expressed in rice leaves. The enzyme activity of PEPC and ME in transgenic rice plants were 1.3 to 5.9 and 1.4 to 6.8 times of those in wild type plants, respectively. The foreign C4 photosynthetic enzyme genes could be stably inherited and effectively expressed in progenies.
(2) The average segregated percentage of the transgenic plants with target genes but selectable marker gene free (SMF) was 8.2%. Several selectable marker-free transgenic homozygous lines of T3 generation were obtained. The correlation coefficients of PEPC and NADP-MDH to Pn (photosynthetic rate) were 0.817 and 0.610, both of which are significant at the 5% level.
(3) In the booting, heading, full heading stages, the average Pn of PEPC/PPDK transgenic restorer lines was higher than that of the control by 12.05%,8.14%,13.83%, respectively, while the average Pn of PEPC/PPDK transgenic rice hybrids was higher than that of the control by 3.59%,7.17%,9.51%, respectively.
(4) From heading stage to mature stage, the SPAD value of chlorophyll in transgenic leaves declined gradually along with the reproductive process. The occurrence of turning point of chlorophyll SPAD (10DAF) was roughly consistent to the full heading stage, after which Pn declined dramatically.
(5) Compared with the control, the transgenic restorer lines with PEPC/PPDK, NADP-MDH/NADP-ME or all four genes had a little bit lower CO2 compensation point and light compensation point but a higher light saturation point by 167μmol photos.m 2.m-1 and a higher light saturated photosynthetic rate by 10.97%. Under normal conditions, the transgenic plants displayed enhanced transpiration rate (VPD), stomatal conductance (Gs) and intercellular CO2 concentration (Ci) to some extent. Gs was positively and significantly correlated to Pn, and positively but not significantly correlated to VPD and Ci. It could be inferred that the enhanced stomatal conductance contributed a lot to its higher Pn.
(6) Under drought stress, in heading stage the transgenic rice showed higher transpiration rate, intercellular CO2 concentration and stomatal conductance than the control; and compared with the control, the Pn and WUE (water use efficiency) of the transgenic rice were increased by 75%-85% and 15%-85%, respectively. Under high temperature stress, in seedling stage the transgenic rice showed higher ratio of 1st grade leaves. Under both drought and high temperature stress conditions, the transgenic rice had higher maximal photochemical efficiency of PSII (Fv/Fm) and higher PSII potential activity (Fv/F0). Therefore, it could be inferred that expression of the foreign C4 photosynthetic genes served a useful function for resistance to drought and high temperature.
(7) The transgenic rice carrying C4 photosynthetic enzyme genes decreased in chalky rice percentage, chalky area and chalkiness to a different extent, and in some cases, the differences reached a statistically significant level between the transgenic rice and the control.
(8) The transgenic restorer lines and their hybrids showed an increase in plant height, the number of spikelets per panicle and grain weight but a decrease in seed setting rate. Assessment of yield capacity test of the transgenic hybrids showed that three had a higher yield than the control hybrids by 7.45%,2.54% and 1.13%.
On the whole, for PEPC/PPDK transgenic rice, it was superior to the others in enzyme expression, Pn, Gs, high temperature resistance, the number of spikelets per panicle and grain yield per plant. The NADP-MDH/NADP-ME transgenic rice had an advantage on drought resistance over the others. When it comes to the transgenic rice carrying all four genes, most traits had values between the PEPC/PPDK transgenic rice and the NADP-MDH/NADP-ME transgenic rice.The possible mechanism leading to this result and the problems such as faster degradation of chlorophyll and sharper decline of photosynthetic capacity in later growth stages, and lower spikelet fertility and yield advantage should be further studied.
引文
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