水稻黄叶转绿突变体818-6-8的目标基因定位与转录组分析

doi:10.16819/j.1001-7216.2026.241105

中国水稻科学 ›› 2026, Vol. 40 ›› Issue (3): 327-340.DOI: 10.16819/j.1001-7216.2026.241105

水稻黄叶转绿突变体818-6-8的目标基因定位与转录组分析

许扬¹^,²^,³, 王芳权¹^,², 李文奇¹^,², 陶亚军¹^,², 范方军¹^,², 陈智慧¹^,², 蒋彦婕¹^,², 朱建平¹^,², 李霞¹^,², 杨杰¹^,²^,³^,^*()

¹ 农业农村部淮河下游种质创制重点实验室/江苏省农业科学院粮食作物研究所/生物育种钟山实验室，南京 210014
² 扬州大学江苏省粮食作物现代化产业协同创新中心，江苏扬州 225009
³ 江苏大学生命科学学院，江苏镇江 212013

收稿日期:2024-11-08 修回日期:2024-12-27 出版日期:2026-05-10 发布日期:2026-05-13
通讯作者: *email: yangjie168@aliyun.com
基金资助:
江苏省重点研发计划资助项目(BE2022383);生物育种钟山实验室项目(ZSBBL-KY2023-04-1)

Gene Mapping and Transcriptome Analysis of a Green-revertible Yellow Leaf Mutant 818-6-8 in Rice

XU Yang¹^,²^,³, WANG Fangquan¹^,², LI Wenqi¹^,², TAO Yajun¹^,², FAN Fangjun¹^,², CHEN Zhihui¹^,², JIANG Yanjie¹^,², ZHU Jianping¹^,², LI Xia¹^,², YANG Jie¹^,²^,³^,^*()

¹ Key Laboratory of Germplasm Innovation in Downstream of Huaihe River, Ministry of Agriculture and Rural Affairs/Institute of Food Crops, Jiangsu Academy of Agricultural Sciences/Zhongshan Biological Breeding Laboratory, Nanjing 210014, China
² Jiangsu Co-innovation Center for Modern Production Technology of Grain Crops, Yangzhou University, Yangzhou 225009, China
³ Institute of Life Science, Jiangsu University, Zhenjiang 212013, China

Received:2024-11-08 Revised:2024-12-27 Online:2026-05-10 Published:2026-05-13

摘要/Abstract

摘要：

【目的】叶片是水稻最重要的光合作用器官。筛选和鉴定水稻叶色突变体，有利于探究叶绿体发育与光合作用的分子调控机制。【方法】在金粳818背景的基因编辑后代中获得一个黄叶转绿突变体818-6-8，并对其进行表型鉴定、遗传分析、基因定位和转录组分析。【结果】与野生型相比，突变体三叶期前叶片明显黄化，并伴随叶绿素含量显著降低；第4叶开始，突变体开始转绿并逐渐表现为与野生型无异。突变体中叶绿体数量较少、类囊体形态和基粒片层结构异常。818-6-8黄叶性状受1对隐性核基因控制。突变位点被定位于5号染色体2.29 Mb的区间内，分析并筛选得到20个候选基因。突变体中2090个基因表达量发生显著改变，其中1343个上调，747个下调；差异表达基因显著富集在叶绿体组织等生物学过程、叶绿体等细胞组分。突变体中多个叶绿素合成和叶绿体发育相关基因表达发生改变。【结论】本研究完成了突变体818-6-8的目标基因定位与转录组分析，为下一步目标基因克隆和叶绿体发育分子调控机理解析提供支持，为景观稻育种提供优异种质资源。

关键词: 水稻, 黄叶转绿突变体, 基因定位, 转录组分析

Abstract:

【Objective】Leaf is the most important photosynthetic organ in rice. Screening and identification of rice leaf color mutants are beneficial for exploring the molecular regulatory mechanisms of chloroplast development and photosynthesis. 【Method】A green-revertible yellow leaf mutant 818-6-8 was obtained from the gene-editing lines of Jingeng 818. Phenotypic identification, genetic analysis, gene mapping, and transcriptome analysis for 818-6-8 were performed. 【Results】Compared with the wild type, the mutant showed an obvious yellow-leaf phenotype until the three-leaf stage, accompanied by a significant decrease in chlorophyll content. As it grew, the mutant turned green and gradually became indistinguishable from the wild type at the four-leaf stage. Fewer chloroplasts, abnormal thylakoid morphology and stromal lamellas structure were observed in the mutant. The yellow-leaf trait of 818-6-8 was controlled by a pair of recessive nuclear alleles. The target mutation site was located within a 2.29 Mb interval on chromosome 5, and 20 annotated candidate genes were listed. The expression levels of 2090 genes in the mutant were significantly changed compared to the wild type, of which 1343 genes were up-regulated and 747 genes were down-regulated. Further, the differentially expressed genes were enriched in biological processes such as chloroplast organization and cellular components such as chloroplasts. The expression of genes associated with chlorophyll biosynthesis and chloroplast development of 818-6-8 changed significantly. 【Conclusion】These results support the next studies for cloning target genes and analyzing chloroplast-development molecular mechanisms, and provide a germplasm resource for ornamental rice breeding.

Key words: rice, green-revertible yellow leaf mutant, gene mapping, transcriptome analysis

许扬, 王芳权, 李文奇, 陶亚军, 范方军, 陈智慧, 蒋彦婕, 朱建平, 李霞, 杨杰. 水稻黄叶转绿突变体818-6-8的目标基因定位与转录组分析[J]. 中国水稻科学, 2026, 40(3): 327-340.

XU Yang, WANG Fangquan, LI Wenqi, TAO Yajun, FAN Fangjun, CHEN Zhihui, JIANG Yanjie, ZHU Jianping, LI Xia, YANG Jie. Gene Mapping and Transcriptome Analysis of a Green-revertible Yellow Leaf Mutant 818-6-8 in Rice[J]. Chinese Journal OF Rice Science, 2026, 40(3): 327-340.

图/表 12

表1 qRT-PCR引物序列

Table 1. Primer sequences for qRT-PCR

目标基因 Target gene	前引物 Forward primer (5′-3′)	后引物 Reverse primer (5′-3′)
OsCAO1	GATCCATACCCGATCGACAT	CGAGAGACATCCGGTAGAGC
OsDVR	AGCCCAGGTTCATCAAGGT	TGATCACCCTCTCGAAGAACT
OsYGL1	TGTTGGTGGGTCCTTGCTTT	ACTGAAGCCCCAGAGCTCTA
OsPORA	ATGGCTCTCCAAGTTCAG	TGGCTCACGCTAAGGAAC
OsPORB	CCGCAAGGAGGGAGCGGTG	CCTCTTGGTGCTAAGGCCG
OsCHLH	AACTGGATGAGCCAGAAGAGA	AAATGCAAAAGACTTGCGACT
OsCHLD	TCCTTTAGCTCACGGCCTTA	GTCCAACATCACCGCTCTTT
OsNYC1	TTTTGAGCGGTTCTTCTCAGA	CCTTTCACAACTCGCATCCT
OsNYC3	TCTATCTAGGTGCCAAAGGC	ATTCTGGCACCTGCTGTTTC
OsNYC4	CGTCTATGACCAACTCATGG	TGCGTCAGCTCTGTATTGCT
OsNOL	TCACAAGCCTTACGACCCAC	GCTCCCTCTCCATCATCTGC
OsHEMA	CGCTATTTCTGATGCTATGGGT	TCTTGGGTGATGATTGTTTGG
OsPAO	AAGCCTCCGATGTTACCGAA	CGAGGGTTTCCAGAATTTGA
OsRCCR1	GGATCGACGATTGATTTCATG	GTCGAGGCGTTCAGAAAGAT
Actin	TGGCATCTCTCAGCACATTCC	TGCACAATGGATGGGTCAGA
rpoA	CCATTCCCACAAGCAAAAAT	TCTTACCGCCTTCCGTAGAA
rpoB	TGGTACATATCCCTTATCTCAA	CTCCAGGACCCAAACAACTC
rbcL	CTTGGCAGCATTCCGAGTAA	ACAACGGGCTCGATGTGATA
rbcS	GCTTGGAGTTCAGCAAGGTC	AACGAAGGCATCAGGGTATG
PsaA	TTAGAAATCCGCCAATCCA	TGCTAGGCTCTACAACCATT
PsaD	CCGCTCCAAGTACAAGATCA	AAGAGCAGCCTGACAGATGA
PsbA	ACCCTCATTAGCAGATTCGT	GATTGTATTCCAGGCAGAGC
PsbD	AAGACAGATTCCGAGGGTGG	TGATTCGCTAGGGATTAAAGAG

图1 野生型金粳818和突变体818-6-8表型 A：野生型金粳818（WT）和突变体818-6-8幼苗期表型；B：野生型金粳818（WT）和突变体818-6-8的3叶期表型；C：野生型金粳818（WT）灌浆期植株，标尺=10 cm；D：突变体818-6-8灌浆期植株，标尺=10 cm。

Fig. 1. Phenotypes of the wild-type Jingeng 818 and the mutant 818-6-8 A, Phenotypes of wild type Jingeng 818 (WT) and mutant 818-6-8 at the seedling stage; B, Phenotypes of wild type Jingeng 818 (WT) and mutant 818-6-8 at the three-leaf stage; C, Wild type Jingeng 818 (WT) plant at the grain filling stage(bar=10 cm); D, Mutant 818-6-8 plant at the grain filling stage(bar=10 cm).

表2 野生型金粳818和818-6-8的叶绿素含量

Table 2. Chlorophyll content in Jingeng 818 and 818-6-8 mg/g

材料 Material	叶绿素a含量 Chlorophyll a content	叶绿素b含量 Chlorophyll b content	总叶绿素含量 Total chlorophyll content	类胡萝卜素含量 Carotenoid content
WT	2.209±0.228	0.953±0.139	3.162±0.284	0.565±0.067
818-6-8	2.185±0.206 ns	0.147±0.023**	2.332±0.218**	0.038±0.010**

表3 野生型金粳818和818-6-8的部分农艺性状比较

Table 3. Comparison of some agronomic traits between Jingeng 818 and 818-6-8

农艺性状 Agronomic trait	金粳818 Jingeng 818	818-6-8
株高Plant height(cm)	96.8 ± 2.3	96.7 ± 2.4 ns
每株分蘖数 Tiller number per plant	12.1 ± 2.6	12.0 ± 2.1 ns
抽穗期Heading date(d)	90.0 ± 1.0	89.0 ± 2.0 ns
每穗粒数 Number of grains per panicle	103.5 ± 5.8	106.0 ± 6.4 ns
结实率Seed-setting rate(%)	95.2 ± 0.5	94.7 ± 1.6 ns
千粒重1000-grain weight(g)	27.6 ± 1.4	27.9 ± 0.9 ns

图2 野生型金粳818和突变体818-6-8叶绿体超微结构 A：野生型叶绿体超微结构；B：A图中红框放大区域；C：突变体叶绿体超微结构；D：C图中红框放大区域。Cp：叶绿体；Thy：类囊体；OB：嗜锇体；Sg：淀粉粒。标尺= 2 μm。

Fig. 2. TEM images of chloroplast structure in the wild-type Jingeng 818 and the mutant 818-6-8 A, Chloroplast ultrastructure in the wild type; B, The magnified image of the selected area in A; C, Chloroplast ultrastructure in mutant; D, The magnified image of the selected area in C. Cp, Chloroplast; Thy, Thylakoid lamellae; OB, Osmiphilic body; Sg, Starch granule. Bars = 2 μm.

表4 818-6-8和正常绿色品种杂交F2的叶色分离

Table 4. Segregation of leaf color in F2 populations from the crosses between 818-6-8 and green leaf varieties

杂交组合 Material	总株数 Total number of plants	绿叶植株数 No. of green leaf plants	黄叶植株数 No. of yellow leaf plants	χ²
金粳818/818-6-8 Jingeng 818/818-6-8	883	679	204	2.73
818-6-8/金粳818 818-6-8/Jingeng 818	858	657	201	1.05
818-6-8/9311	1089	836	253	1.72
818-6-8/蜀恢881 818-6-8/Shuhui 881	1207	876	331	3.65

图3 Delta SNP-index和SNP-index在染色体上的分布 A：Delta拟合图；B：显性性状样品的SNP-index拟合图；C：隐性性状样品的SNP-index拟合图。图中横坐标表示染色体的位置；散点表示计算得到delta SNP-index和SNP-index值；黑色曲线表示相应的拟合值；红色虚线表示Loess拟合值为99%的阈值线。

Fig. 3. Distribution of delta SNP-index and SNP-index on chromosomes A, The delta fitting plot; B, The SNP-index fitting plot for dominant trait samples; C, The SNP-index fitting plot for recessive trait samples. The X axis represents the position of chromosomes; The scattered points represent the values of delta SNP-index and SNP-index; The black curves represent the corresponding fitting value; The red dashed lines represent the threshold line with a Loess fitting value of 99%.

表5 候选基因

Table 5. Candidate gene

基因编号 Gene ID	基因注释 Gene annotation	显著差异表达 Significantly differential expression
LOC_Os05g18150	BKRF1编码类EBNA-1蛋白BKRF1 Encodes an EBNA-1-like protein	否No
LOC_Os05g18294	含CRAL-TRIO结构域的蛋白质C589.09，线粒体亚型X4 C589.09, SEC14细胞质因子家族蛋白，推测，表达 protein containing CRAL-TRIO domain, mitochondrial isoform X4, SEC14 cytosolic factor family protein, putative, expressed	否No
LOC_Os05g18470	随机Slug蛋白, 含CRAL/TRIO结构域蛋白，表达5 Random Slug protein 5, Protein containing CRAL/TRIO domain, expressed	否No
LOC_Os05g18660	未鉴定蛋白LOC4338253, OsFBDUF24-F-box和含DUF结构域蛋白，表达 Uncharacterized protein LOC4338253, OsFBDUF24-F-box and DUF domain-containing protein, expressed	否No
LOC_Os05g18670	S型阴离子通道SLAH4-like, C₄二羧酸转运蛋白/含有苹果酸转运蛋白结构域蛋白，表达 S-type anion channel SLAH4-like, C₄-dicarboxylate transporter/malate transporter domain-containing protein, expressed	否No
LOC_Os05g18750	50S核糖体蛋白L25, 核糖体蛋白L25，推测，表达 50S ribosomal protein L25, Ribosomal protein L25, putative, expressed	否No
LOC_Os05g18790	Os05g0270800 甲酰基转移酶 Formyltransferase	否No
LOC_Os05g18850*	水稻DICER相似蛋白基因 OsDCL1c Rice DICER-like protein gene OsDCL1c	升高 Increased
LOC_Os05g18950	含五肽重复序列的蛋白At1g10330, 五萜肽，推测，表达 Protein containing pentatricopeptide repeat At1g10330, Pentatricopeptide, putative, expressed	否 No
LOC_Os05g19150	Os05g0273800 水解酶，含有α/β折叠家族结构域蛋白，表达 Hydrolase, alpha/beta fold family domain-containing protein, expressed	否 No
LOC_Os05g19370	60S核糖体蛋白L15, 60S核糖体蛋白L15，推测表达 60S ribosomal protein L15, 60S ribosomal protein L15, putative, expressed	否 No
LOC_Os05g19380	含五肽重复序列的蛋白质At2g18940，叶绿体, 五萜肽，推测，表达 Protein containing pentatricopeptide repeat At2g18940, chloroplast, Pentatricopeptide, putative, expressed	否 No
LOC_Os05g19390*	三角状五肽重复蛋白OsPPR939; OsPPR920 Pentatricopeptide repeat protein OsPPR939; OsPPR920	否 No
LOC_Os05g19480	过氧化物酶体生物发生蛋白2, 过氧化物酶体组装蛋白，推测，表达 Peroxisome biogenesis protein 2, Peroxisome assembly protein, putative, expressed	否 No
LOC_Os05g19954	tRNA假尿苷合酶A1, tRNA假尿苷合酶家族蛋白，推测表达 tRNA pseudouridine synthase A1, tRNA pseudouridine synthase family protein, putative, expressed	否 No
LOC_Os05g20100	可能的甘油-3-磷酸酰基转移酶3, 甘油-3-磷酸酰基转移酶，推测，表达 Probable glycerol-3-phosphate acyltransferase 3, Glycerol-3-phosphate acyltransferase, putative, expressed	否 No
LOC_Os05g17604	可能的LRR受体样丝氨酸/苏氨酸蛋白激酶At1g56130亚型X1, SHR5受体样激酶，推测表达 Probable LRR receptor-like serine/threonine-protein kinase At1g56130 isoform X1 SHR5 receptor-like kinase, putative, expressed	升高 Increased
LOC_Os05g18274	未知蛋白, 表达蛋白 Unknown protein, Expressed protein	升高 Increased
LOC_Os05g19010	Os05g0272300 表达蛋白, 阿魏酰辅酶A木质素转移酶 Expressed protein	升高 Increased
LOC_Os05g19910*	OsFMT; OsAT5 Feruloyl-CoA: monolignol transferase OsFMT; OsAT5	降低Decreased

表5 候选基因

Table 5. Candidate gene

基因编号 Gene ID	基因注释 Gene annotation	显著差异表达 Significantly differential expression
LOC_Os05g18150	BKRF1编码类EBNA-1蛋白BKRF1 Encodes an EBNA-1-like protein	否No
LOC_Os05g18294	含CRAL-TRIO结构域的蛋白质C589.09，线粒体亚型X4 C589.09, SEC14细胞质因子家族蛋白，推测，表达 protein containing CRAL-TRIO domain, mitochondrial isoform X4, SEC14 cytosolic factor family protein, putative, expressed	否No
LOC_Os05g18470	随机Slug蛋白, 含CRAL/TRIO结构域蛋白，表达5 Random Slug protein 5, Protein containing CRAL/TRIO domain, expressed	否No
LOC_Os05g18660	未鉴定蛋白LOC4338253, OsFBDUF24-F-box和含DUF结构域蛋白，表达 Uncharacterized protein LOC4338253, OsFBDUF24-F-box and DUF domain-containing protein, expressed	否No
LOC_Os05g18670	S型阴离子通道SLAH4-like, C₄二羧酸转运蛋白/含有苹果酸转运蛋白结构域蛋白，表达 S-type anion channel SLAH4-like, C₄-dicarboxylate transporter/malate transporter domain-containing protein, expressed	否No
LOC_Os05g18750	50S核糖体蛋白L25, 核糖体蛋白L25，推测，表达 50S ribosomal protein L25, Ribosomal protein L25, putative, expressed	否No
LOC_Os05g18790	Os05g0270800 甲酰基转移酶 Formyltransferase	否No
LOC_Os05g18850*	水稻DICER相似蛋白基因 OsDCL1c Rice DICER-like protein gene OsDCL1c	升高 Increased
LOC_Os05g18950	含五肽重复序列的蛋白At1g10330, 五萜肽，推测，表达 Protein containing pentatricopeptide repeat At1g10330, Pentatricopeptide, putative, expressed	否 No
LOC_Os05g19150	Os05g0273800 水解酶，含有α/β折叠家族结构域蛋白，表达 Hydrolase, alpha/beta fold family domain-containing protein, expressed	否 No
LOC_Os05g19370	60S核糖体蛋白L15, 60S核糖体蛋白L15，推测表达 60S ribosomal protein L15, 60S ribosomal protein L15, putative, expressed	否 No
LOC_Os05g19380	含五肽重复序列的蛋白质At2g18940，叶绿体, 五萜肽，推测，表达 Protein containing pentatricopeptide repeat At2g18940, chloroplast, Pentatricopeptide, putative, expressed	否 No
LOC_Os05g19390*	三角状五肽重复蛋白OsPPR939; OsPPR920 Pentatricopeptide repeat protein OsPPR939; OsPPR920	否 No
LOC_Os05g19480	过氧化物酶体生物发生蛋白2, 过氧化物酶体组装蛋白，推测，表达 Peroxisome biogenesis protein 2, Peroxisome assembly protein, putative, expressed	否 No
LOC_Os05g19954	tRNA假尿苷合酶A1, tRNA假尿苷合酶家族蛋白，推测表达 tRNA pseudouridine synthase A1, tRNA pseudouridine synthase family protein, putative, expressed	否 No
LOC_Os05g20100	可能的甘油-3-磷酸酰基转移酶3, 甘油-3-磷酸酰基转移酶，推测，表达 Probable glycerol-3-phosphate acyltransferase 3, Glycerol-3-phosphate acyltransferase, putative, expressed	否 No
LOC_Os05g17604	可能的LRR受体样丝氨酸/苏氨酸蛋白激酶At1g56130亚型X1, SHR5受体样激酶，推测表达 Probable LRR receptor-like serine/threonine-protein kinase At1g56130 isoform X1 SHR5 receptor-like kinase, putative, expressed	升高 Increased
LOC_Os05g18274	未知蛋白, 表达蛋白 Unknown protein, Expressed protein	升高 Increased
LOC_Os05g19010	Os05g0272300 表达蛋白, 阿魏酰辅酶A木质素转移酶 Expressed protein	升高 Increased
LOC_Os05g19910*	OsFMT; OsAT5 Feruloyl-CoA: monolignol transferase OsFMT; OsAT5	降低Decreased

图4 突变体818-6-8与金粳818叶片中差异表达基因火山图红点表示显著上调的基因，绿点表示显著下调的基因，灰点表示表达量无显著差异的基因。

Fig. 4. Volcano map of DEG, in leaves of Jingeng 818 and 818-6-8 Red dots indicate significant up-regulated genes; the green dots indicate significant down-regulated genes; Gray dots indicate the genes with no significant difference in expression levels.

图5 突变体818-6-8与金粳818叶片中差异表达基因GO功能分类 1 ：叶绿体组织；2：异戊烯基二磷酸生物合成，甲基赤藓糖醇4-磷酸途径；3：叶绿体靶向蛋白；4：质体转录；5：tRNA代谢；6：核糖体核糖核酸；7：类囊体膜组织；8：氧化还原；9：DNA模板转录正向调控；10：蛋白质再折叠；11：叶绿体；12：叶绿体基质；13：叶绿体膜；14：叶绿体类囊体膜；15：细胞质；16：叶绿体类囊体腔；17：谷氨酰tRNA（Gln）氨基转移酶复合物；18：质体大核糖体亚基；19：类囊体膜的组分；20：叶绿体拟核；21：铁离子结合；22：血红素结合；23：rRNA结合；24：假尿苷合酶活性；25：氧化还原酶活性；26：多糖结合；27：单加氧酶活性；28：谷氨酰胺tRNA合酶（谷氨酰胺水解）活性；29：金属离子结合；30：氧化还原酶活性。

Fig. 5. GO functional classification of differentially expressed genes in leaves of Jingeng 818 and 818-6-81, Chloroplast organization; 2, Isopentenyl diphosphate biosynthetic process, methylerythritol 4-phosphate pathway; 3, Protein targeting to chloroplast; 4, Plastid transcription; 5, tRNA metabolic process; 6, rRNA processing; 7, Thylakoid membrane organization; 8, Oxidation-reduction process; 9, Positive regulation of transcription, DNA-templated; 10, Protein refolding; 11, Chloroplast; 12, Chloroplast stroma; 13, Chloroplast envelope; 14, Chloroplast thylakoid membrane; 15, Cytoplasm; 16, Chloroplast thylakoid lumen; 17, Glutamyl-tRNA(Gln) amidotransferase complex; 18, Plastid large ribosomal subunit; 19, Integral component of thylakoid membrane; 20, Chloroplast nucleoid; 21, Iron ion binding; 22, Heme binding; 23, rRNA binding; 24, Pseudouridine synthase activity; 25, Oxidoreductase activity, acting on paired donors; 26, Polysaccharide binding; 27, Monooxygenase activity; 28, Glutaminyl-tRNA synthase (glutamine-hydrolyzing) activity; 29, Metal ion binding; 30, Oxidoreductase activity, acting on paired donors.

图6 突变体818-6-8与金粳818叶片中差异表达基因KEGG通路富集分析 1 ：二萜生物合成；2：淀粉和蔗糖代谢；3：苯丙素生物合成；4：苯丙氨酸代谢；5：油菜素内酯生物合成；6：嘧啶代谢；7：甜菜素生物合成；8：氨基糖和核苷酸糖代谢；9：谷胱甘肽代谢；10：色氨酸代谢；11：单萜类生物合成；12：氰基氨基酸代谢；13：牛磺酸和次牛磺酸代谢；14：核糖体；15：氨基酰基tRNA生物合成；16：RNA降解；17：碱基切除修复；18：真核生物核糖体生物合成；19：非同源末端连接；20：植物MAPK信号通路。

Fig. 6. KEGG pathway enrichment analysis of differentially expressed genes in leaves of Jingeng 818 and 818-6-81, Diterpenoid biosynthesis; 2, Starch and sucrose metabolism; 3, Phenylpropanoid biosynthesis; 4, Phenylalanine metabolism; 5, Brassinosteroid biosynthesis; 6, Pyrimidine metabolism; 7, Betalain biosynthesis; 8, Amino sugar and nucleotide sugar metabolism; 9, Glutathione metabolism; 10, Tryptophan metabolism; 11, Monoterpenoid biosynthesis; 12, Cyanoamino acid metabolism; 13, Taurine and hypotaurine metabolism; 14, Ribosome; 15, Aminoacyl-tRNA biosynthesis; 16, RNA degradation; 17, Base excision repair; 18, Ribosome biogenesis in eukaryotes; 19, Non-homologous end-joining; 20, MAPK signaling pathway-plant.

图7 野生型金粳818和突变体818-6-8叶绿素合成、叶绿体发育及光合作用相关基因的表达量 A：叶绿素合成相关基因的表达量；B：叶绿体发育及光合作用相关基因的表达量。**表示差异极显著水平（P < 0.01）；*表示差异显著水平（P < 0.05）；ns表示无显著性差异。

Fig. 7. Expression levels of chlorophyll synthesis, chloroplast development and photosynthesis related genes in the wild-type Jingeng 818 and its mutant 818-6-8 A, The expression levels of genes related to chlorophyll synthesis; B, The expression levels of genes related to chloroplast development and photosynthesis. ** indicate significant differences at 0.01 level; * indicate differences at 0.05 level; ns indicate no significant difference.

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水稻黄叶转绿突变体818-6-8的目标基因定位与转录组分析

Gene Mapping and Transcriptome Analysis of a Green-revertible Yellow Leaf Mutant 818-6-8 in Rice

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