一个新的小麦黄化突变体研究
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摘要
本研究材料为西农1718高代品系自发黄化突变体,是较为稀有的种质材料,在研究小麦叶绿素合成代谢过程、叶绿体发育机制、光合膜与光合功能系统建成、叶绿体发育相关基因的表达、调控和克隆等方面具有非常重要的价值。本研究从形态特征、主要农艺性状、光合特性、遗传规律、叶绿体超微结构、色素蛋白及叶绿素生物合成主要中间产物等方面对该黄化突变体进行了研究,主要结果如下:
1.小麦黄化突变体自出苗即表现出黄色,随着发育黄化程度加深,极端黄化类型(金黄株)植株矮小,早衰死亡;黄-绿中间型茎、叶、穗呈不同程度的黄绿色,发育期较对照晚6～8d,苗相显弱,能结实,但结实率和粒重有所降低。花粉母细胞减数分裂观察发现,减数分裂行为正常,突变体染色体数目均为42条,即该黄化突变体未发生染色体数目和结构变异。花粉活性观察结果表明,黄化突变体花粉活力正常。
2.对突变体的叶绿素含量、光合速率及叶绿素荧光动力学参数进行比较分析。结果显示:突变体叶绿素含量(Chl)、类胡萝卜素(Caro)含量均显著低于突变亲本,金黄株、绿黄株、黄绿株叶绿素含量分别为突变亲本的17%、24%和58%,类胡萝卜素含量分别为突变体的53%、55%和83%,叶绿素含量降低幅度显著大于类胡萝卜素,表明该突变体为叶绿素缺乏突变体。金黄株净光合速率非常低,孕穗期、开花期仅为突变亲本的5.7%、2.4%;绿黄株净光合速率显著低于突变亲本,为突变亲本的57.7%、43.3%;而叶绿素含量仅为突变亲本一半的黄绿株,其净光合速率接近突变亲本,表明该黄化突变体叶绿素含量在一定范围内降低,单位叶绿素含量的光合效率提高。突变体Fo均显著低于突变亲本,金黄株、绿黄株的Fm,Fv, qP,qN均显著低于突变亲本,金黄株Fv/Fm、Fv/F0比值显著低于突变亲本,而绿黄株、黄绿株Fv/Fm、Fv/F0比值高于突变亲本,黄绿株Fm,Fv, qP,qN与突变亲本均无明显差异。结果表明,金黄株PSⅡ电子传递能力、QA的还原能力、PSII活性、PSII天线色素捕获光能效率及热耗散均大幅度降低,由此推测,金黄株光系统和叶绿体结构可能已遭到严重损伤;而黄绿株、绿黄株仍保持较高的PSII活性和光化学效率。
3.通过多年连续自交、与突变亲本回交以及与其它基因型进行正反交试验表明,突变体自交后代表现为,1金黄株:2黄-绿中间型:1绿株;突变体与突变亲本回交,后代表现为,1黄-绿中间型:1绿株。表明该小麦黄化突变体是由一对核基因控制的不完全显性遗传,其中,绿株和金黄株是纯合体,金黄株纯合致死,黄-绿中间型为杂合体。该黄化突变性状是能够稳定遗传的,突变性状的传递不仅可发生在亲子代之间,还可发生在不同基因型之间,且在不同遗传背景下,性状表现一致。
4.叶绿体观察发现:突变体叶绿体的数目、形态与突变亲本无明显差异;黄绿植株叶绿体超微结构与突变亲本无明显差异,都表现为类囊体膜系统发达,基质类囊体与基粒类囊体高度分化,构成基粒的垛叠层数较多,基粒数目也较多,且排列整齐,有少量嗜锇颗粒;绿黄株叶绿体结构存在一定缺陷,虽然能看到基粒片层与基质片层,但基粒数目明显少于突变亲本,且排列不规则,有淀粉粒及嗜锇颗粒;金黄株叶绿体结构存在严重缺陷,几乎所有的叶绿体内部均未见高度组织化的内膜系统,无基粒,而基质片层清晰可见,叶绿体内有淀粉粒和嗜锇颗粒,且嗜锇颗粒明显多于绿黄株、黄绿株及突变亲本。
5.利用SDS-尿素-聚丙烯酰胺凝胶电泳对突变体及突变亲本叶绿体类囊体色素蛋白组成进行比较研究,发现,突变体低分子量区多肽变化幅度大于高分子量区多肽,60～64kD多肽(主要为PSI反应中心蛋白)和42～54 kD多肽(主要为PSII反应中心天线蛋白),黄绿株与突变亲本无明显差别,而绿黄株和金黄株显著降低,且金黄株降低幅度大于绿黄株;33～35 kD多肽(主要为PSII反应中心蛋白),黄绿株、绿黄株表达量明显低于突变亲本,且绿黄株降低幅度大于黄绿株,23、27kD的多肽(主要为捕光色素蛋白),其变化最为显著,金黄株缺失,在黄绿株和绿黄株中,其降幅也极大;16～19kD多肽(主要为细胞色素b6f复合体和PSII光合放氧有关的多肽),其变化规律为,随黄化程度加深,其含量大幅度降低,黄绿株与突变亲本无明显差异,金黄株和绿黄株大幅度降低。
6.对叶绿素生物合成的一系列主要前体物质累积量进行测定分析,结果表明,金黄株、绿黄株和黄绿株中间产物ALA、PBG、UrogenIII、Coprogen和ProtoⅨ的含量均高于突变亲本,且随突变体黄化程度加深而递增,与叶绿素含量变化趋势相反,尤其ProtoⅨ累积更为显著,金黄株的含量达到突变亲本的210%,分离绿株与突变亲本无差别;而金黄株、绿黄株和黄绿株自ProtoⅨ以后的中间产物Mg-Proto、Pchlide均显著低于突变亲本,且随突变体黄化程度加深而递减,与叶绿素含量的变化一致,分离绿株与突变亲本无明显差异。由此判断,该黄化突变体叶绿素合成受阻,且受阻位点发生在ProtoⅨ到Mg-Proto部位。镁螯合酶催化了由ProtoⅨ到Mg-Proto,因此推断,该黄化突变体为镁螯合酶活性降低而导致叶绿素合成受阻。
1.A novel spontaneous aurea mutation of wheat (Triticum aestivum L.) was first described in this paper. It was find from a winter wheat culture genotype (Xinong 1718) in 1999-2000. When grown in the greenhouse and in the field, the mutant plants could be clearly distinguished from the green ones at any stage from seedling to near maturity. When the mutant plant selfed, three types offsprings couled be seen: green plant, intermediate of yellow-greenplants and yellow plant(aurea). The aurea plants died at the seedling stage for plants grown in soil, and the intermediate plants are viable but less vigorous and later development than the green plant. The mutant causes accumulation of chlorophyll reduce, resulting in a yellow color in the plant in different degreed. There were no significient differences in some agronomy traits between wild-type and yellownish-green plants, while the greenish-yellow mutant plants performed worse compared to the wild type in the field, with decreases in seed production of up to 70%.
2.The chlorophyll content and the fluorescence parameters (Fo,Fm,Fv,Fv /Fm, qP,qN )were measured to evaluate the mutant. The result showed that the mutant wheat were shown to have a greatly diminished rate of chlorophyll accumulation compared to the wild type, and the ratio of Chl a / Chl b in the mutated plants were smaller than that in the wild type, and it tended to decrease as the etiolation degree increased. The net photosynthesis rate during booting stage and blooming stage in the golden types were only 5.7% and 2.4%, respectively, of that in the wilt type; while in the greenish-yellow type, it was about one-half (57.7% and 43.3%) of that in the wild type; and in the yellowish-green type, they were close to that in the wild type. The value of Fo in mutated plants were decreased significantly. The values of Fm,Fv,qP,qN in aurea type and greenish-yellow type, were decreased significantly. The values of Fm,Fv,qP,qN in yellowish-green type, have no significant difference with wild type. The value of Fv /Fm,Fv/F0 were decreased significantly in aurea type and in the greenish-yellow and yellowish-green plants, they were higher than that of the wild type.
3.The aurea mutant wheat were analyzed genetically after inbreds and crosses with normal green plants. Selfing the M1 plant(chlorina) results in 1:2:1 segregation ratio of aurea : chlorina : green plants. The progeny of M1 green plants were all green. The test cross between a chlorina plant and a normal plant (as male or female) gave a 1 normal green: l yellow-green ratio. The result can be explained on the assumption that the aurea is homozygous for Au and the chlorina heterozygous for Au. It is concluded that the semi-dominant mutant is controlled by a single nucleus gene.This is the genetic constitution of the segregants being Au/Au, normal, Au/au, yellow-green, au/au, aurea.
4.In order to better understand the mechanism of aurea mutant, comparative study on chloroplast ultrastructure was conducted between aurea mutant and its wild type(Xinong1718) under the electromicroscope. Using morphometrical measurements, the mutants chloroplast numbers, shape, size and the distribution in cell were similar with those of the parent of mutant. In contrast, the ultrastructure of chloroplasts among three phenotype types and the parent of mutant were markedly differences: in the yellowish-green mutant, a well-organized thylakoid system, differentiated into grana and intergrana thylakoids, resembled the parent of mutant; in the greenish-yellow chloroplasts, their thylakoid systems appeared to be disorganized, and the grana contained fewer lamellae, the number of grana per chloroplast section was reduced; in the golden type, the development of chloroplast was the most bad among the mutants, no grana but the stroma thylakoid were clearly, accumulation of osmiophilic droplet and starch granules were more than those of other phenotype types and wild type. The thylakoid morphology is more nearly wild type in appearance.
5. The precursors of tetrapyrrole synthesis,δ-aminolevulinic acid (ALA), porphobilinogen (PBG), uroporphyrinogen(Urogen), Coprogen, ProtoⅨ, Mg-Proto and Pchlide were over accumulated in mutant wheat, while protoporphyrin IX(MgPP and MgPme were investigated. The result showed that the content of ALA、PBG、UrogenIII、Coprogen and ProtoⅨw ere all over to the wild type; in contrast, Mg-Proto and Pchlide were all decreased than wild type. This result indicated Coprogen and protoporphyrin accumulated in the mutant, and the rate of conversion of ProtoⅨto Mg- Proto was very low than that of wild type. It is concluded that, the mutation brings about a restriction in the rate of conversion of protoporphyrin to magnesium protoporphyrin; and in the mutant, the enzymes converting protoporphyrin into Mg- Proto are most likely absent or damaged. Mg-Chelatase catalyses this step.
6. The pigment protein complexes are separated as shown by fully denaturing SDS-PAGE electrophoresis. Eleven chlorophyll-protein complexes of wheat were resolved in wild type. As compared with mutant parent wheat, in all type mutant, there were no significant changes in the composition of 60~64 kD polypeptide, which are mainly reaction center chlorophyll-protein complexes of PS I and 42~54 kD polypeptide, which are mainly PSII reaction center antenna protein. In contrast, a deficiency in 23~35 kD polypeptides, which are mainly light harvesting chlorophyll protein (LHCP) complex and reaction center chlorophyll-protein complexes of PSII, had been noted in all kinds of mutant wheat, particularly the two major polypeptides of LHCII, 23kD and 27kD polypeptides, were greatly diminished in amount in greenish-yellow and yellowish-green plants, and it had been absent in aurea plant. These data led to the suggestion that the LHCP complexes were more sensitivity to chlorophyll-deficiency than the other chlorophyll-protein complexes.
1.小麦黄化突变体自出苗即表现出黄色,随着发育黄化程度加深,极端黄化类型(金黄株)植株矮小,早衰死亡;黄-绿中间型茎、叶、穗呈不同程度的黄绿色,发育期较对照晚6～8d,苗相显弱,能结实,但结实率和粒重有所降低。花粉母细胞减数分裂观察发现,减数分裂行为正常,突变体染色体数目均为42条,即该黄化突变体未发生染色体数目和结构变异。花粉活性观察结果表明,黄化突变体花粉活力正常。
2.对突变体的叶绿素含量、光合速率及叶绿素荧光动力学参数进行比较分析。结果显示:突变体叶绿素含量(Chl)、类胡萝卜素(Caro)含量均显著低于突变亲本,金黄株、绿黄株、黄绿株叶绿素含量分别为突变亲本的17%、24%和58%,类胡萝卜素含量分别为突变体的53%、55%和83%,叶绿素含量降低幅度显著大于类胡萝卜素,表明该突变体为叶绿素缺乏突变体。金黄株净光合速率非常低,孕穗期、开花期仅为突变亲本的5.7%、2.4%;绿黄株净光合速率显著低于突变亲本,为突变亲本的57.7%、43.3%;而叶绿素含量仅为突变亲本一半的黄绿株,其净光合速率接近突变亲本,表明该黄化突变体叶绿素含量在一定范围内降低,单位叶绿素含量的光合效率提高。突变体Fo均显著低于突变亲本,金黄株、绿黄株的Fm,Fv, qP,qN均显著低于突变亲本,金黄株Fv/Fm、Fv/F0比值显著低于突变亲本,而绿黄株、黄绿株Fv/Fm、Fv/F0比值高于突变亲本,黄绿株Fm,Fv, qP,qN与突变亲本均无明显差异。结果表明,金黄株PSⅡ电子传递能力、QA的还原能力、PSII活性、PSII天线色素捕获光能效率及热耗散均大幅度降低,由此推测,金黄株光系统和叶绿体结构可能已遭到严重损伤;而黄绿株、绿黄株仍保持较高的PSII活性和光化学效率。
3.通过多年连续自交、与突变亲本回交以及与其它基因型进行正反交试验表明,突变体自交后代表现为,1金黄株:2黄-绿中间型:1绿株;突变体与突变亲本回交,后代表现为,1黄-绿中间型:1绿株。表明该小麦黄化突变体是由一对核基因控制的不完全显性遗传,其中,绿株和金黄株是纯合体,金黄株纯合致死,黄-绿中间型为杂合体。该黄化突变性状是能够稳定遗传的,突变性状的传递不仅可发生在亲子代之间,还可发生在不同基因型之间,且在不同遗传背景下,性状表现一致。
4.叶绿体观察发现:突变体叶绿体的数目、形态与突变亲本无明显差异;黄绿植株叶绿体超微结构与突变亲本无明显差异,都表现为类囊体膜系统发达,基质类囊体与基粒类囊体高度分化,构成基粒的垛叠层数较多,基粒数目也较多,且排列整齐,有少量嗜锇颗粒;绿黄株叶绿体结构存在一定缺陷,虽然能看到基粒片层与基质片层,但基粒数目明显少于突变亲本,且排列不规则,有淀粉粒及嗜锇颗粒;金黄株叶绿体结构存在严重缺陷,几乎所有的叶绿体内部均未见高度组织化的内膜系统,无基粒,而基质片层清晰可见,叶绿体内有淀粉粒和嗜锇颗粒,且嗜锇颗粒明显多于绿黄株、黄绿株及突变亲本。
5.利用SDS-尿素-聚丙烯酰胺凝胶电泳对突变体及突变亲本叶绿体类囊体色素蛋白组成进行比较研究,发现,突变体低分子量区多肽变化幅度大于高分子量区多肽,60～64kD多肽(主要为PSI反应中心蛋白)和42～54 kD多肽(主要为PSII反应中心天线蛋白),黄绿株与突变亲本无明显差别,而绿黄株和金黄株显著降低,且金黄株降低幅度大于绿黄株;33～35 kD多肽(主要为PSII反应中心蛋白),黄绿株、绿黄株表达量明显低于突变亲本,且绿黄株降低幅度大于黄绿株,23、27kD的多肽(主要为捕光色素蛋白),其变化最为显著,金黄株缺失,在黄绿株和绿黄株中,其降幅也极大;16～19kD多肽(主要为细胞色素b6f复合体和PSII光合放氧有关的多肽),其变化规律为,随黄化程度加深,其含量大幅度降低,黄绿株与突变亲本无明显差异,金黄株和绿黄株大幅度降低。
6.对叶绿素生物合成的一系列主要前体物质累积量进行测定分析,结果表明,金黄株、绿黄株和黄绿株中间产物ALA、PBG、UrogenIII、Coprogen和ProtoⅨ的含量均高于突变亲本,且随突变体黄化程度加深而递增,与叶绿素含量变化趋势相反,尤其ProtoⅨ累积更为显著,金黄株的含量达到突变亲本的210%,分离绿株与突变亲本无差别;而金黄株、绿黄株和黄绿株自ProtoⅨ以后的中间产物Mg-Proto、Pchlide均显著低于突变亲本,且随突变体黄化程度加深而递减,与叶绿素含量的变化一致,分离绿株与突变亲本无明显差异。由此判断,该黄化突变体叶绿素合成受阻,且受阻位点发生在ProtoⅨ到Mg-Proto部位。镁螯合酶催化了由ProtoⅨ到Mg-Proto,因此推断,该黄化突变体为镁螯合酶活性降低而导致叶绿素合成受阻。
1.A novel spontaneous aurea mutation of wheat (Triticum aestivum L.) was first described in this paper. It was find from a winter wheat culture genotype (Xinong 1718) in 1999-2000. When grown in the greenhouse and in the field, the mutant plants could be clearly distinguished from the green ones at any stage from seedling to near maturity. When the mutant plant selfed, three types offsprings couled be seen: green plant, intermediate of yellow-greenplants and yellow plant(aurea). The aurea plants died at the seedling stage for plants grown in soil, and the intermediate plants are viable but less vigorous and later development than the green plant. The mutant causes accumulation of chlorophyll reduce, resulting in a yellow color in the plant in different degreed. There were no significient differences in some agronomy traits between wild-type and yellownish-green plants, while the greenish-yellow mutant plants performed worse compared to the wild type in the field, with decreases in seed production of up to 70%.
2.The chlorophyll content and the fluorescence parameters (Fo,Fm,Fv,Fv /Fm, qP,qN )were measured to evaluate the mutant. The result showed that the mutant wheat were shown to have a greatly diminished rate of chlorophyll accumulation compared to the wild type, and the ratio of Chl a / Chl b in the mutated plants were smaller than that in the wild type, and it tended to decrease as the etiolation degree increased. The net photosynthesis rate during booting stage and blooming stage in the golden types were only 5.7% and 2.4%, respectively, of that in the wilt type; while in the greenish-yellow type, it was about one-half (57.7% and 43.3%) of that in the wild type; and in the yellowish-green type, they were close to that in the wild type. The value of Fo in mutated plants were decreased significantly. The values of Fm,Fv,qP,qN in aurea type and greenish-yellow type, were decreased significantly. The values of Fm,Fv,qP,qN in yellowish-green type, have no significant difference with wild type. The value of Fv /Fm,Fv/F0 were decreased significantly in aurea type and in the greenish-yellow and yellowish-green plants, they were higher than that of the wild type.
3.The aurea mutant wheat were analyzed genetically after inbreds and crosses with normal green plants. Selfing the M1 plant(chlorina) results in 1:2:1 segregation ratio of aurea : chlorina : green plants. The progeny of M1 green plants were all green. The test cross between a chlorina plant and a normal plant (as male or female) gave a 1 normal green: l yellow-green ratio. The result can be explained on the assumption that the aurea is homozygous for Au and the chlorina heterozygous for Au. It is concluded that the semi-dominant mutant is controlled by a single nucleus gene.This is the genetic constitution of the segregants being Au/Au, normal, Au/au, yellow-green, au/au, aurea.
4.In order to better understand the mechanism of aurea mutant, comparative study on chloroplast ultrastructure was conducted between aurea mutant and its wild type(Xinong1718) under the electromicroscope. Using morphometrical measurements, the mutants chloroplast numbers, shape, size and the distribution in cell were similar with those of the parent of mutant. In contrast, the ultrastructure of chloroplasts among three phenotype types and the parent of mutant were markedly differences: in the yellowish-green mutant, a well-organized thylakoid system, differentiated into grana and intergrana thylakoids, resembled the parent of mutant; in the greenish-yellow chloroplasts, their thylakoid systems appeared to be disorganized, and the grana contained fewer lamellae, the number of grana per chloroplast section was reduced; in the golden type, the development of chloroplast was the most bad among the mutants, no grana but the stroma thylakoid were clearly, accumulation of osmiophilic droplet and starch granules were more than those of other phenotype types and wild type. The thylakoid morphology is more nearly wild type in appearance.
5. The precursors of tetrapyrrole synthesis,δ-aminolevulinic acid (ALA), porphobilinogen (PBG), uroporphyrinogen(Urogen), Coprogen, ProtoⅨ, Mg-Proto and Pchlide were over accumulated in mutant wheat, while protoporphyrin IX(MgPP and MgPme were investigated. The result showed that the content of ALA、PBG、UrogenIII、Coprogen and ProtoⅨw ere all over to the wild type; in contrast, Mg-Proto and Pchlide were all decreased than wild type. This result indicated Coprogen and protoporphyrin accumulated in the mutant, and the rate of conversion of ProtoⅨto Mg- Proto was very low than that of wild type. It is concluded that, the mutation brings about a restriction in the rate of conversion of protoporphyrin to magnesium protoporphyrin; and in the mutant, the enzymes converting protoporphyrin into Mg- Proto are most likely absent or damaged. Mg-Chelatase catalyses this step.
6. The pigment protein complexes are separated as shown by fully denaturing SDS-PAGE electrophoresis. Eleven chlorophyll-protein complexes of wheat were resolved in wild type. As compared with mutant parent wheat, in all type mutant, there were no significant changes in the composition of 60~64 kD polypeptide, which are mainly reaction center chlorophyll-protein complexes of PS I and 42~54 kD polypeptide, which are mainly PSII reaction center antenna protein. In contrast, a deficiency in 23~35 kD polypeptides, which are mainly light harvesting chlorophyll protein (LHCP) complex and reaction center chlorophyll-protein complexes of PSII, had been noted in all kinds of mutant wheat, particularly the two major polypeptides of LHCII, 23kD and 27kD polypeptides, were greatly diminished in amount in greenish-yellow and yellowish-green plants, and it had been absent in aurea plant. These data led to the suggestion that the LHCP complexes were more sensitivity to chlorophyll-deficiency than the other chlorophyll-protein complexes.
引文
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