空间环境诱变小麦叶绿素缺失突变体的研究
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摘要
本研究以空间环境诱变创制的三份小麦叶绿素缺失突变体Mt135、Mt6172和Mt18为试验材料,对其生物学特性、主要农艺性状、叶绿体超微结构、光合特性和遗传规律进行了初步研究,主要结果如下:
1、Mt135和Mt6172的表型特征主要表现为白化株、条纹株和绿株三种类型。其中白化株不能存活,苗期即死亡;条纹株正常成穗结实,但株高、株粒数、株粒重和千粒重显著降低。Mt18是一个受低温诱导的温度敏感型叶绿素突变体,叶色变化主要表现为绿-白-绿的变化过程,能够正常成穗结实,主要农艺性状都发生显著降低。
2、叶绿体超微结构分析表明,与原始亲本相比,Mt135条纹株绿色组织在苗期叶绿体数目减少,超微结构无显著差异;拔节期叶绿体数目与原始亲本无显著差异,基粒片层数目减少;抽穗期叶绿体数目和基粒片层数明显增多。Mt135条纹株白色组织和白化株叶绿体数目在苗期和拔节期减少,基粒类囊体数减少或消失,基质类囊体清晰可见,并在抽穗期出现空化叶绿体。Mt6172条纹株绿色组织仅在拔节期叶绿体数目少于原始亲本,叶绿体超微结构在三个时期均表现正常;Mt6172条纹株白色组织和白化株叶绿体存在不同类型的变异,叶绿体数目在三个时期均少于原始亲本。Mt18的叶绿体数目和结构在变白前期与原始亲本无显著差异,变白期突变体叶绿体基粒类囊体数减少甚至完全消失,复绿期突变体叶绿体数目少于原始亲本,大部分叶绿体恢复正常。
3、叶绿素荧光动力学参数分析表明,与原始亲本相比,Mt135条纹株绿色组织的光合效率在苗期和拔节期显著降低,抽穗期显著升高,而条纹株白色组织和完全白化株完全失去光合能力;Mt6172条纹株的光合效率苗期正常,拔节期和抽穗期显著降低,白化株依然保持着极低的光合能力;Mt18在变白期的光合效率显著降低,但变白前期和复绿期基本正常。另外,随着光照强度的增加,与原始亲本相比,Mt135条纹株绿色组织的光合效率在拔节期下降最显著,抽穗期显著升高,条纹株白色组织和白化株不能进行光合作用;Mt6172条纹株的光合电子传递速率在不同生育期均显著降低,而光能转化率和实际量子产量无显著变化,白化株光合效率极低;Mt18的光合效
4、初步遗传分析表明,Mt135和Mt18表现为细胞质遗传,而Mt6172是一个由核基因控制的隐性遗传突变材料。cDNA-AFLP初步分析显示,突变体与原始亲本均在分子水平上存在显著差异,扩增差异条带类型表现为增加、缺失、增强和减弱四种。
以上试验结果证实,空间环境可诱变创制不同遗传特点的小麦叶绿素缺失突变体。三份叶绿素缺失突变体的叶绿体超微结构存在不同程度的缺陷,光合作用受到较大的影响,光合特性与主要农艺性状发生了显著改变,为小麦光合作用基础研究提供了新材料。
Three chlorophyll-deficient wheat mutants, i.e., Mt135, Mt6172 and Mt18, which derived from spaceflight mutagenesis, were studied in terms of biological characteristics, main agronomic traits, chloroplast ultrastructure, photosynthesis and inherited model. The main results were as following:
1. The leaf color of the mutants Mt135 and Mt6172 showed three phenotypes: albino, stripe and green, in which the albino plants with entire albino leaves died during seedling stage, while the plant height, grains per plant, grain weight per plant and 1000-grain weight of plants with green-and-white striped leaves were lower than those of the wild types; Mt18 was a temperature-sensitive winter wheat chlorophyll mutant. The leaf color varied from green to albino and regreen with the temperature from low to normal during the whole growth stage, and the main agronomic traits were lower than those of the wild type.
2. Chloroplast ultrastructure observation showed that the chloroplast number in green tissue of Mt135 striped plants was less than that of the wild type, but no significant difference in the ultrastructure. The similar number of chloroplast but less number of granum-thylakoids in Mt135 striped plants was observed at the elongation stage. At the heading stage, the number of chloroplast and granum-thylakoids were increased significantly. Both at seedling and elongation stage, the chloroplast number of white tissue in Mt135 striped plants and albino plants was less than that of the wild type, and the number of granum-thylakoids reduced or completely disappeared, but the strom-thylakoid was obviously visible, and some chloroplasts without internal ultrastructure were found at heading stage. Mt6172 had less chloroplast in the green tissue at the elongation stage, but had normal ultrastructure at the three stages. There were different chloroplast variation of the white tissue in Mt6172 striped plants and albino plants, and the chloroplast number was less than that of the wild type among the three growth stages. There was no significant difference between Mt18 and the wild type during prior albino stage; at the albino stage, the number of granum-thylakoids and grana lamellae became fewer or completely disappeared, but the strom-thylakoid was obviously visible; after returning green, structure of most chloroplasts recovered into normal, but number of chloroplast was still lower than that of the wild type.
3. The analysis of chlorophyll fluorescence kinetic parameters between the mutants and their wild type revealed that the photosynthetic efficiency of the green tissue in Mt135 striped plants reduced significantly at both seedling and elongation stages, and then increased at heading stage, while the white tissue of Mt135 striped plants and albino plants lost photosynthetic capacity completely; the photosynthetic efficiency of Mt6172 striped plants was normal at seedling stage, and was reduced significantly at elongation and heading stages, and albino plants still maintained very low photosynthetic capacity; the photosynthetic efficiency of Mt18 was normal at both prior albino and re-green stages and lower than that of the wild type at albino stage. In addition, with the increase of light intensity, the photosynthetic efficiency of the green tissue in Mt135 striped plants was lowest at elongation stage, and then increased significantly at heading stage. However, the white tissue of Mt135 striped plants and albino plants lost photosynthetic capacity completely; Mt6172 striped plants was significantly lower than that of wild type at different growth stages, while there was no significant differences in qP and Y(Ⅱ) between mutant and wild type, and albino plants still maintained very low photosynthetic capacity; the photosynthetic efficiency of Mt18 was significantly lower than that of the wild type only at albino stage.
4. Preliminary genetic analysis showed that the inheritance model of Mt135 and Mt6172 was cytoplasmic, while Mt6172 a recessive nuclear. cDNA-AFLP analysis revealed significant differences between the chlorophyll-deficient mutants and their wild types at the molecular level. The type of different bands among the three mutants showed increment, deletion, strongth and weakness.
It was concluded that different chlorophyll-deficient wheat mutants could be induced by spaceflight environment, with the characteristics of destructed chloroplast ultrastructure, deeply affected photosynthesis as well as decreased main agronomic traits. These chlorophyll-deficient wheat mutants could be new materials for basic study of wheat photosynthesis.
1、Mt135和Mt6172的表型特征主要表现为白化株、条纹株和绿株三种类型。其中白化株不能存活,苗期即死亡;条纹株正常成穗结实,但株高、株粒数、株粒重和千粒重显著降低。Mt18是一个受低温诱导的温度敏感型叶绿素突变体,叶色变化主要表现为绿-白-绿的变化过程,能够正常成穗结实,主要农艺性状都发生显著降低。
2、叶绿体超微结构分析表明,与原始亲本相比,Mt135条纹株绿色组织在苗期叶绿体数目减少,超微结构无显著差异;拔节期叶绿体数目与原始亲本无显著差异,基粒片层数目减少;抽穗期叶绿体数目和基粒片层数明显增多。Mt135条纹株白色组织和白化株叶绿体数目在苗期和拔节期减少,基粒类囊体数减少或消失,基质类囊体清晰可见,并在抽穗期出现空化叶绿体。Mt6172条纹株绿色组织仅在拔节期叶绿体数目少于原始亲本,叶绿体超微结构在三个时期均表现正常;Mt6172条纹株白色组织和白化株叶绿体存在不同类型的变异,叶绿体数目在三个时期均少于原始亲本。Mt18的叶绿体数目和结构在变白前期与原始亲本无显著差异,变白期突变体叶绿体基粒类囊体数减少甚至完全消失,复绿期突变体叶绿体数目少于原始亲本,大部分叶绿体恢复正常。
3、叶绿素荧光动力学参数分析表明,与原始亲本相比,Mt135条纹株绿色组织的光合效率在苗期和拔节期显著降低,抽穗期显著升高,而条纹株白色组织和完全白化株完全失去光合能力;Mt6172条纹株的光合效率苗期正常,拔节期和抽穗期显著降低,白化株依然保持着极低的光合能力;Mt18在变白期的光合效率显著降低,但变白前期和复绿期基本正常。另外,随着光照强度的增加,与原始亲本相比,Mt135条纹株绿色组织的光合效率在拔节期下降最显著,抽穗期显著升高,条纹株白色组织和白化株不能进行光合作用;Mt6172条纹株的光合电子传递速率在不同生育期均显著降低,而光能转化率和实际量子产量无显著变化,白化株光合效率极低;Mt18的光合效
4、初步遗传分析表明,Mt135和Mt18表现为细胞质遗传,而Mt6172是一个由核基因控制的隐性遗传突变材料。cDNA-AFLP初步分析显示,突变体与原始亲本均在分子水平上存在显著差异,扩增差异条带类型表现为增加、缺失、增强和减弱四种。
以上试验结果证实,空间环境可诱变创制不同遗传特点的小麦叶绿素缺失突变体。三份叶绿素缺失突变体的叶绿体超微结构存在不同程度的缺陷,光合作用受到较大的影响,光合特性与主要农艺性状发生了显著改变,为小麦光合作用基础研究提供了新材料。
Three chlorophyll-deficient wheat mutants, i.e., Mt135, Mt6172 and Mt18, which derived from spaceflight mutagenesis, were studied in terms of biological characteristics, main agronomic traits, chloroplast ultrastructure, photosynthesis and inherited model. The main results were as following:
1. The leaf color of the mutants Mt135 and Mt6172 showed three phenotypes: albino, stripe and green, in which the albino plants with entire albino leaves died during seedling stage, while the plant height, grains per plant, grain weight per plant and 1000-grain weight of plants with green-and-white striped leaves were lower than those of the wild types; Mt18 was a temperature-sensitive winter wheat chlorophyll mutant. The leaf color varied from green to albino and regreen with the temperature from low to normal during the whole growth stage, and the main agronomic traits were lower than those of the wild type.
2. Chloroplast ultrastructure observation showed that the chloroplast number in green tissue of Mt135 striped plants was less than that of the wild type, but no significant difference in the ultrastructure. The similar number of chloroplast but less number of granum-thylakoids in Mt135 striped plants was observed at the elongation stage. At the heading stage, the number of chloroplast and granum-thylakoids were increased significantly. Both at seedling and elongation stage, the chloroplast number of white tissue in Mt135 striped plants and albino plants was less than that of the wild type, and the number of granum-thylakoids reduced or completely disappeared, but the strom-thylakoid was obviously visible, and some chloroplasts without internal ultrastructure were found at heading stage. Mt6172 had less chloroplast in the green tissue at the elongation stage, but had normal ultrastructure at the three stages. There were different chloroplast variation of the white tissue in Mt6172 striped plants and albino plants, and the chloroplast number was less than that of the wild type among the three growth stages. There was no significant difference between Mt18 and the wild type during prior albino stage; at the albino stage, the number of granum-thylakoids and grana lamellae became fewer or completely disappeared, but the strom-thylakoid was obviously visible; after returning green, structure of most chloroplasts recovered into normal, but number of chloroplast was still lower than that of the wild type.
3. The analysis of chlorophyll fluorescence kinetic parameters between the mutants and their wild type revealed that the photosynthetic efficiency of the green tissue in Mt135 striped plants reduced significantly at both seedling and elongation stages, and then increased at heading stage, while the white tissue of Mt135 striped plants and albino plants lost photosynthetic capacity completely; the photosynthetic efficiency of Mt6172 striped plants was normal at seedling stage, and was reduced significantly at elongation and heading stages, and albino plants still maintained very low photosynthetic capacity; the photosynthetic efficiency of Mt18 was normal at both prior albino and re-green stages and lower than that of the wild type at albino stage. In addition, with the increase of light intensity, the photosynthetic efficiency of the green tissue in Mt135 striped plants was lowest at elongation stage, and then increased significantly at heading stage. However, the white tissue of Mt135 striped plants and albino plants lost photosynthetic capacity completely; Mt6172 striped plants was significantly lower than that of wild type at different growth stages, while there was no significant differences in qP and Y(Ⅱ) between mutant and wild type, and albino plants still maintained very low photosynthetic capacity; the photosynthetic efficiency of Mt18 was significantly lower than that of the wild type only at albino stage.
4. Preliminary genetic analysis showed that the inheritance model of Mt135 and Mt6172 was cytoplasmic, while Mt6172 a recessive nuclear. cDNA-AFLP analysis revealed significant differences between the chlorophyll-deficient mutants and their wild types at the molecular level. The type of different bands among the three mutants showed increment, deletion, strongth and weakness.
It was concluded that different chlorophyll-deficient wheat mutants could be induced by spaceflight environment, with the characteristics of destructed chloroplast ultrastructure, deeply affected photosynthesis as well as decreased main agronomic traits. These chlorophyll-deficient wheat mutants could be new materials for basic study of wheat photosynthesis.
引文
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