水稻苗期低温失绿基因的精细定位和功能初步分析
摘要
水稻苗期低温失绿是一个重要的突变性状,研究该性状的遗传基础在理论上和应用上都具有重要的意义。本实验室前期研究发现,水稻品种Dular苗期叶色受温度影响。在早季低温条件下,Dular幼苗表现出白化失绿,叶绿体不能正常发育。光照培养箱低温试验表明,其失绿的临界温度为20~23℃。遗传分析表明,该低温失绿受隐性单基因cisc(t)控制,已将该基因定位在9号染色体两个SSR标记RM257与RM242之间。在此基础上,本研究对Dular幼苗低温敏感的时期和外源脱落酸(ABA)对幼苗低温敏感性的影响进行分析,对苗期低温失绿基因进行精细定位,并对候选基因进行初步的生物信息学分析和半定量RT-PCR分析,以期为基因的克隆、功能分析以及阐明植物耐冷分子机制奠定基础。主要结果如下:
(1)Dular幼苗在两叶期以前低温处理表现为白化或部分白化,三叶期以后低温处理则基本为绿色。持续低温试验表现为前2叶全部白化,第3叶长出时开始转绿,到5叶期大部分转绿。可见Dular幼苗对低温敏感主要在两叶期以前,而到三叶期时对低温已不敏感,具有一定的耐冷性。
(2)低温条件下施加外源ABA可以减弱Dular幼苗的白化程度,提高其耐冷性。随着ABA浓度的升高,幼苗的白化程度逐渐降低。当ABA浓度大于0.4μM时,幼苗恢复绿色,但植株矮化,生长受到抑制。由此推测控制幼苗低温失绿的基因很可能是与ABA途径有交叉的冷响应途径中的一个关键基因。
(3)在前期定位的基础上开发出新的SSR标记和InDel标记,利用Lemont/Dular、Ⅱ-32B/Dular的两个F2群体中的800个低温白化单株进行基因定位,将该基因定位在标记InDel13-2与InDel11之间的88 kb范围内。
(4)在88kb定位区间内共有13个基因。有2个基因编码的蛋白存在叶绿体前导肽,与叶绿体有关,故可作为候选基因。另外,对3个基因(2个为bZIP和bHLH类型转录因子,1个为编码磷脂酰肌醇的PHIN基因)在低温处理下进行RT-PCR半定量分析,结果显示:随着处理时间的延长,bZIP和PHIN表达量先增强后减弱,而bHLH持续增强,但差异不太明显。说明这3个基因均与冷胁迫相关,都是可能的候选基因。
Cold-induced seedling chlorosis is an important mutant character in rice. Research of the genetic basis of this character is meaningful in both theory and application. Previous studies of our laboratory have found that the seedling leaf colour of rice cultivar Dular is affected by temperature. Under the low temperature condition in early season, Dular showed albino seedlings, in which chloroplasts could not develop normally. Experiments in light incubator revealed that the critical temperature of seedling chlorosis is 20~23℃. Genetic analysis showed that the albino trait was controlled by a single recessive gene cisc(t), which was mapped in a region between SSR markers RM257 and RM242 on chromosome 9. On this basis, this study was to identify the sensative period of Dular seedlings to low temperature, analyze the effect of abscisic acid (ABA) on the low temperature sensitivity of Dular seedlings, fine map the cold-induced seedling chlorosis gene, and preliminarily analyze candidate genes with bioinformatics methods and semi-quantitative RT-PCR, aiming at facilitating the cloning and functional analysis of the gene and elucidating the molecular mechanism of plant cold tolerance. The main results are as follows:
(1) Dular seedlings showed albino or partial albino when treated at low temperature before two-leaf stage, but were basically green when treated after three-leaf stage. In sustained low temperature test, Dular seedlings showed entire albino for the first two leaves, but the third leaf began to exhibit a little green when it came out, and most leaves turned green at five-leaf stage. This result indicates that the low temperature sensative time of Dular seedlings is before the three-leaf stage.
(2) Under low-temperature condition, exogenous ABA could reduce the albino extent and increase the low temperature resistance of Dular seedlings, and the albino extent decreased with the increase of ABA concentration. When the ABA concentration was was higher than 0.4μM, Dular seedlings recovered green but became dwarf, indicating that their growth was suppressed. Based on these results, we speculated that the cold-induced seedling chlorosis gene is likely to be a key gene in a cold response pathway cross-talking with the ABA dependent pathway.
(3) Based on the result of primary mapping, new SSR and InDel markers were developed and fine maping of the target gene was performed using 800 cold-induced albino plants from two F2 populations of Lemont/Dular andⅡ-32B/Dular. The target gene was mapped in a region of 88 kb between markers InDel13-2 and InDel11.
(4) There were 13 genes in the 88-kb region. Two genes contained chloroplast-targeted leading peptide in the proteins encoded by them and therefore could be regarded as candidate genes. In addition, semi-quantitative RT-PCR analysis of three genes (two encoding bZIP and bHLH type transcription factors and one encoding phosphatidylinositol PHIN) under low temperature treatment showed that the expression of both bZIP and PHIN increased at beginning and decreased afterwards, while bHLH expression consistently increased, but the difference was not very significant. This suggests that these three genes are all related to cold stress and therefore could be possible candidate genes.
(1)Dular幼苗在两叶期以前低温处理表现为白化或部分白化,三叶期以后低温处理则基本为绿色。持续低温试验表现为前2叶全部白化,第3叶长出时开始转绿,到5叶期大部分转绿。可见Dular幼苗对低温敏感主要在两叶期以前,而到三叶期时对低温已不敏感,具有一定的耐冷性。
(2)低温条件下施加外源ABA可以减弱Dular幼苗的白化程度,提高其耐冷性。随着ABA浓度的升高,幼苗的白化程度逐渐降低。当ABA浓度大于0.4μM时,幼苗恢复绿色,但植株矮化,生长受到抑制。由此推测控制幼苗低温失绿的基因很可能是与ABA途径有交叉的冷响应途径中的一个关键基因。
(3)在前期定位的基础上开发出新的SSR标记和InDel标记,利用Lemont/Dular、Ⅱ-32B/Dular的两个F2群体中的800个低温白化单株进行基因定位,将该基因定位在标记InDel13-2与InDel11之间的88 kb范围内。
(4)在88kb定位区间内共有13个基因。有2个基因编码的蛋白存在叶绿体前导肽,与叶绿体有关,故可作为候选基因。另外,对3个基因(2个为bZIP和bHLH类型转录因子,1个为编码磷脂酰肌醇的PHIN基因)在低温处理下进行RT-PCR半定量分析,结果显示:随着处理时间的延长,bZIP和PHIN表达量先增强后减弱,而bHLH持续增强,但差异不太明显。说明这3个基因均与冷胁迫相关,都是可能的候选基因。
Cold-induced seedling chlorosis is an important mutant character in rice. Research of the genetic basis of this character is meaningful in both theory and application. Previous studies of our laboratory have found that the seedling leaf colour of rice cultivar Dular is affected by temperature. Under the low temperature condition in early season, Dular showed albino seedlings, in which chloroplasts could not develop normally. Experiments in light incubator revealed that the critical temperature of seedling chlorosis is 20~23℃. Genetic analysis showed that the albino trait was controlled by a single recessive gene cisc(t), which was mapped in a region between SSR markers RM257 and RM242 on chromosome 9. On this basis, this study was to identify the sensative period of Dular seedlings to low temperature, analyze the effect of abscisic acid (ABA) on the low temperature sensitivity of Dular seedlings, fine map the cold-induced seedling chlorosis gene, and preliminarily analyze candidate genes with bioinformatics methods and semi-quantitative RT-PCR, aiming at facilitating the cloning and functional analysis of the gene and elucidating the molecular mechanism of plant cold tolerance. The main results are as follows:
(1) Dular seedlings showed albino or partial albino when treated at low temperature before two-leaf stage, but were basically green when treated after three-leaf stage. In sustained low temperature test, Dular seedlings showed entire albino for the first two leaves, but the third leaf began to exhibit a little green when it came out, and most leaves turned green at five-leaf stage. This result indicates that the low temperature sensative time of Dular seedlings is before the three-leaf stage.
(2) Under low-temperature condition, exogenous ABA could reduce the albino extent and increase the low temperature resistance of Dular seedlings, and the albino extent decreased with the increase of ABA concentration. When the ABA concentration was was higher than 0.4μM, Dular seedlings recovered green but became dwarf, indicating that their growth was suppressed. Based on these results, we speculated that the cold-induced seedling chlorosis gene is likely to be a key gene in a cold response pathway cross-talking with the ABA dependent pathway.
(3) Based on the result of primary mapping, new SSR and InDel markers were developed and fine maping of the target gene was performed using 800 cold-induced albino plants from two F2 populations of Lemont/Dular andⅡ-32B/Dular. The target gene was mapped in a region of 88 kb between markers InDel13-2 and InDel11.
(4) There were 13 genes in the 88-kb region. Two genes contained chloroplast-targeted leading peptide in the proteins encoded by them and therefore could be regarded as candidate genes. In addition, semi-quantitative RT-PCR analysis of three genes (two encoding bZIP and bHLH type transcription factors and one encoding phosphatidylinositol PHIN) under low temperature treatment showed that the expression of both bZIP and PHIN increased at beginning and decreased afterwards, while bHLH expression consistently increased, but the difference was not very significant. This suggests that these three genes are all related to cold stress and therefore could be possible candidate genes.
引文
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