从60Co诱变的粳稻中花11 M2代中发现了一个白化致死突变体，该突变体从发芽后至3叶期一直表现白化，3叶后逐渐死亡，根据随后的基因定位研究结果，将该白化突变体暂定名为albino lethal 4 (abl4)。与野生型相比，abl4突变体的叶绿素含量与类胡萝卜素含量极低，几乎难以检测到。叶绿素荧光分析结果表明，abl4叶片中的电子传递速率和实际光化学效率都为0，而最大光化学效率也极低，显示突变体没有光化学活性。abl4突变体中包括过氧化物酶、超氧化物歧化酶在内的抗氧化酶活性显著升高，过氧化氢酶（CAT）活性显著下降，脂质过氧化产物丙二醛含量显著提高，提示abl4突变体受到氧化胁迫。电子显微镜观察表明abl4不能形成完整的叶绿体，只有类似前质体结构。遗传分析表明，该突变表型受一对隐性核基因控制。利用abl4突变体与籼稻品种龙特甫B杂交获得的F2分离群体进行基因定位，首先将该基因定位于水稻第4条染色体上的SSR标记RM3785和RM303之间。随后，利用新开发的STS和dCAPS标记，进一步将ABL4 基因定位在RH4631和RH4633之间，物理距离约为201 kb。
An albino lethal mutant, temporarily designated as albino lethal 4(abl4), was obtained from 60Co γray radiation mutant pool of japonica rice Zhonghua 11. The seedling of abl4 showed albino phenotype from germination to 3leafstage, then died. The chlorophyll and carotenoid contents in abl4 mutant were too low to be detected. Moreover, the electron transport rate and the effective quantum yield of photosystem Ⅱ in abl4 were about zero. On the other hand, the potential photochemical efficiency of PSⅡ in abl4 was also at an undetectably low level, indicating that the activity of photochemical reaction was lost in the leaves of abl4. The activities of antioxidant enzymes induding superoxide dismutase and peroxidase were significantly increased, while catalase was decreased significantly. As results, the content of malondialdehyde was also increased significantly in abl4 compared with the wild type, suggesting that abl4 was strongly subjected to high oxidant stress. Transmission electron microscopy examination showed that abl4 plastids contained some empty vesicles without thylakoids. It was revealed that the phenotype of abl4 was controlled by a single recessive nuclear gene. An F2 population generated by crossing the abl4 mutant with the indica rice cultivar Longtefu B was used for gene mapping,ABL4 was preliminarily located between RM3785 and RM303 on rice chromosome 4 and further mapped to a 201 kb region.
\[1\]Abdallah F, Salamini F, Leister D. A prediction of the size and evolutionary origin of the proteome of chloroplasts of Arabidopsis. Trends Plant Sci, 2000, 5(4): 141142.
\[2\]Friso G, Giacomelli L, Ytterberg A J, et al. Indepth analysis of the thylakoid membrane proteome of Arabidopsis thaliana chloroplasts: new proteins, New functions, and a plastid proteome database. Plant Cell, 2004, 16(2): 478499.
\[3\]黄晓群, 赵海新, 董春林, 等. 水稻叶绿素合成缺陷突变体及其生物学研究进展. 西北植物学报, 2005, 25(8): 16851691.
\[4\]何冰, 刘玲珑, 张文伟, 等. 植物叶色突变体. 植物生理学通讯, 2006, 42(1)： 19.
\[5\]Iwata N, Omura T. Linkage studies in rice (Oryza sativa L) Some albino genes and their linkage relation with marker genes. Sci Bull Fac Agric Kyushu Univ, 1978, 33(1): 18.
\[6\]Iwata N, Satoh H, Omura T.Linkage analysis by use of trisomics in rice (Oryza sativa L.):IV. Linkage groups locating on chromosomes 2 and 10. Jpn J Breed, 1981, 31(0): 6667.
\[7\]赵海军, 吴殿星, 舒庆尧, 等. 携带白化转绿型叶色标记光温敏核不育系玉兔S的选育及其特征特性. 中国水稻科学, 2004, 18(6): 515521.
\[8\]夏九成, 王玉平, 马炳田, 等. 水稻(Oryza sativa L.)苗期低温白化突变体al12的超微结构与基因定位. 遗传学报, 2006, 33(12): 11121119.
\[9\]陈涛, 张亚东, 赵凌, 等.水稻白化转绿突变基因gra(t)的精细定位与候选基因分析.遗传学报, 2009, 36(2): 117123.
\[10\]余庆波, 江华, 米华玲, 等. 水稻白化突变体alb21 生理特性和基因定位. 上海师范大学学报:自然科学版, 2005, 34(1): 7075.
\[11\]孙萌萌, 余庆波, 张慧琦, 等.控制水稻叶绿体发育基因OsALB23的定位.植物生理与分子生物学报,2006, 32(4): 433437.
\[12\]Jung K H, Hur J, Ryu C H, et al. Characterization of a rice chlorophylldeficient mutant using the TDNA genetrap system. Plant Cell Physiol, 2003, 44(5): 463472.
\[13\]Su N, Hu M L, Wu D X, et al. Disruption of a rice pentatricopeptide repeat protein causes a seedlingspecific albino phenotype and its utilization to enhance seed purity in hybrid rice production. Plant Physiol, 2012, 159(1): 227238.
\[14\]Gothandam K M, Kim E S, Cho H, et al. OsPPR1, a pentatricopeptide repeat protein of rice is essential for the chloroplast biogenesis. Plant Mol Biol, 2005, 58(3): 421433.
\[15\]Wellburn A R. The spectral determination of chlorophyll a and b, as well as total carotenoids, using various solvents with spectrophotometers of different resolution. Plant Physiol, 1994, 144: 307313.
\[16\]Rogers S O, Bendich A J. Extraction of DNA from plant tissues. Plant Mol Biol Manual, 1988, A6: 110.
\[17\]McCouch S R, Teytelman L, Xu Y B, et al. Development and mapping of 2240 new SSR markers for rice (Oryza sativa L.). DNA Res, 2002, 9(6): 199207.
\[18\]Panaud O, Chen X, McCouch S R. Development microsatellite markers characterization of simple sequence length polymorphism (SSLPs) in rice (Oryza sativa L). Mol Gen Genet, 1996, 252(5): 597607.
\[19\]Michelmore R W, Paran I, Kesseli R V. Identification of markers linked to diseaseresistance genes by bulked segregation analysis: A rapid method to detect markers in specific genomic regions by using segregation population. Proc Natl Acad Sci USA, 1991, 88(21): 98289832.
\[20\]Stenbaek A, Jensen P E. Redox regulation of chlorophyll biosynthesis. Phytochemistry, 2010, 71 (8/9): 853859.
\[21\]李育红, 王宝和, 戴正元, 等. 一个水稻新型叶色突变体的形态结构与遗传定位.中国水稻科学, 2011, 25(6): 587593.
\[22\]奉保华, 杨杨, 施勇烽, 等. 水稻淡褐斑叶突变体lbsl1的遗传分析与基因定位.中国水稻科学, 2012, 26(3): 297301.
\[23\]Zhang J X, Kirkham M B. Droughtstressinduced changes in activities of superoxide dismutase, catalase, and peroxidase in wheat species. Plant Cell Physiol, 35(5): 785791.
\[24\]Shim I S, Momose Y, Yamamoto A, et al. Inhibition of catalase activity by oxidative stress and its relationship to salicylic acid accumulation in plants. Plant Growth Regul, 2003, 39(3): 285292.
\[25\]金怡, 刘合芹, 汪得凯, 等.一个水稻苗期白条纹叶抽穗期白穗突变体的遗传分析和基因定位. 中国水稻科学, 2011, 25(6): 461466.
\[26\]汪庆, 汪得凯, 陶跃之. 一个新的水稻半矮化小穗突变体的遗传分析与基因定位. 中国水稻科学, 2011, 25(6): 677680.
