水稻温敏型叶片白化转绿突变体tsa2的表型鉴定与基因定位

doi:10.3724/SP.J.1006.2019.82049

摘要/Abstract

摘要：

水稻温敏型叶色突变体是研究植物光合作用、叶绿体结构和功能以及温度影响叶绿体发育的理想材料。利用甲基磺酸乙酯(EMS)诱变籼型水稻(Oryza sativa L.)三系保持系西农1B, 从其后代中筛选到一个突变性状稳定遗传的温敏型叶片白化转绿突变体tsa2 (temperature-sensitive green-revertible albino 2)。与野生型相比, tsa2突变表型受温度影响, 22°C条件下萌发的野生型幼苗表型正常, 而tsa2幼苗完全白化, 且约40%白化苗死亡, 存活白化苗的光合色素含量、光合速率均显著降低, 成熟期主要农艺性状均显著变劣; 在28°C下萌发的tsa2幼苗叶片呈浅绿色并伴有白条纹, 其光合色素含量显著降低, 光合速率及主要农艺性状差异较小; 32°C下萌发的tsa2幼苗叶片无明显差异。透射电镜观察显示, 与野生型相比, tsa2在22°C下叶肉细胞中无叶绿体或存在异常发育叶绿体(尚未分化出基粒和基层), 在28°C下部分叶肉细胞含少量发育完整的叶绿体, 在32°C下叶肉细胞数量及形态均正常。实时荧光定量PCR (qRT-PCR)分析表明, 与野生型相比, tsa2突变体中部分光合色素代谢途径基因、叶绿体发育相关基因及光合作用相关基因的表达水平呈不同程度变化。遗传分析表明, tsa2突变表型受一对隐性核基因控制, TSA2被定位于第5染色体SSR标记S5-57和S5-119之间, 物理距离为718 kb。本研究为水稻遗传改良及研究温度影响叶绿体发育机制奠定了基础。

关键词: 水稻(Oryza sativa L.), 温敏, 白化转绿, 叶绿体超微结构, 基因定位

Abstract:

Temperature-sensitive leaf color mutants of rice are ideal materials in studies on photosynthesis, chloroplast structure and function, and chloroplast development. A temperature-sensitive green-revertible albino mutant (tsa2) with genetically stable mutational traits was screened out from the progeny of ethyl methane sulfonate (EMS) treated indica three-line maintainer line Xinong 1B. The wild type seedlings had normal phenotype at 22°C, while the mutant tsa2 had completely albino leaves and about 40% of albino seedlings died at the seedling stage; the photosynthetic pigment contents and photosynthetic rate of surviving albino seedlings decreased significantly, and the main agronomic traits were significantly lower than those of the wild type at maturity stage. When germinated at 28°C, tsa2 showed light-green leaves with white streaks and significantly lower photosynthetic pigment contents than the wild type, while a small difference of photosynthetic rate and main agronomic traits between the tsa2 and the wild type. No significant difference in leaves was observed between tsa2 and the wild type when seedlings germinated at 32°C. Transmission electron microscope observation revealed that the albino leaves of tsa2 demonstrated abnormal chloroplast development (without differentiated grana and granum lamella) or without chloroplast at 22°C and completely developed chloroplasts in partial mesophyll cells at 28°C, and normal number and morphology of mesophyll cells compared with wild type at 32°C. The analysis of qRT-PCR indicated that genes related to partial photosynthetic pigment metabolism pathways, chloroplast development and photosynthesis expressed in tsa2 to a varying degrees compared with these of the wild type. Genetic analysis suggested that mutational phenotype of tsa2 was controlled by a single recessive nuclear gene, TSA2, which was finally mapped between SSR markers S5-57 and S5-119 on chromosome 5, with a physical distance of 718 kb. These results lay a foundation for the research on genetic improvement and the mechanism explanation of chloroplast development affected by temperature in rice (Oryza sativa L.).

Key words: rice (Oryza sativa L.), temperature-sensitive, green-revertible albino, chloroplast ultrastructure, gene mapping

尚丽娜,陈新龙,米胜南,委刚,王玲,张雅怡,雷霆,林永鑫,黄兰杰,朱美丹,王楠. 水稻温敏型叶片白化转绿突变体tsa2的表型鉴定与基因定位[J]. 作物学报, 2019, 45(5): 662-675.

Li-Na SHANG,Xin-Long CHEN,Sheng-Nan MI,Gang WEI,Ling WANG,Ya-Yi ZHANG,Ting LEI,Yong-Xin LIN,Lan-Jie HUANG,Mei-Dan ZHU,Nan WANG. Phenotypic identification and gene mapping of temperature-sensitive green- revertible albino mutant tsa2 in rice (Oryza sativa L.)[J]. Acta Agronomica Sinica, 2019, 45(5): 662-675.

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链接本文: https://zwxb.chinacrops.org/CN/10.3724/SP.J.1006.2019.82049

https://zwxb.chinacrops.org/CN/Y2019/V45/I5/662

图/表 11

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表1

光合色素代谢以及叶绿体发育和光合相关基因的定量引物"

引物 Primer	正向序列 Forward sequence (5'-3')	反向序列 Reverse sequence (5'-3')
UBIQUITIN 5	ACCACTTCGACCGCCACTACT	ACGCCTAAGCCTGCTGGTT
OsCAO1	GACACCTTCATCTGGGCTTCAA	CGAGAGACATCCGGTAGAGC
OsCHLI	GTTCGAGCCTGGTTTGCTTGC	CTCTCCACGGTGTTCCATCCTG
OsCHLa/b	GGTGGTCAAGGTGTCATTGTCAT	ACGCGACTGGATCTTTGGAGAG
OsCHLM	CCATCCATTGGTCTCCTTATGACA	GTAGCCTACTTACCATCAATGAGTC
OsCHLD	GCTTGCAGAAAGCTACACAAGC	AGGCCGTGAGCTAAAGGAGA
OsDVR	CAGGTCGAGACCGTCAAGAAC	ATGACCTGGATCGGCACCTTG
OsHEMD	TGGAAGGCTGCTGGAAATCCTAAG	TCCTTGGAAGCTCTGAGGCCAA
OsPSY1	GCCTCAAGCAGGCCTATCATC	GTGATGTGCGAGGCATTTGGTC
OsPSY2	GACAAATTCTGCGTGCCAGGTT	GACAGCAGCTTCTTTGCCTTGTT
OsPSY3	TTCAGACAGGCCGAAGAAGGC	GTAGGCCCTCTTGGTGAAGTTGT
OsPSAA	TGGGGTTGATCCTAAGGAGATACCA	CCTCCGCGAAAACTAAGAAATTCTG
OsPSAB	TGGCAGGGCAACGTTTCACAATT	CAGTCGCCCAAACAAGATGTCCAAAT
OsPSAC	GTGTACGAGCTTGTCCAACAGATGTAT	GCAGGCGGATTCGCATCTCTTAC
OsPSAD	AGCAGGTGTTCGAGATGCCCAC	ATTGGGGAAGACGCGGTAGAACT
OsPSAE	AGGCACCAAGGTGAAGATCCTGAG	AGGGCGTAGTTGTTGGTCGACAC
OsPSAF	TCAAGAAGCTCCAGTCGTCGCTC	AGGCCGAACTTGCCGTAGTTCTC
OsPSAH	GTCGCCGTACAACCCTCTCCAG	CTTCTTGATCGGAAGCAGGTCCG
OsPSAL	TCTGAGAAGCCAACGTACCAGGTG	AGGTTGGAGAGGTACCAGGCGAC
OsPSBA	GCGGTTCCCTATTCAGTGCTATG	TAACCATGAGCGGCCACAATATT
OsPSBB	TAGTTTCTGGTTGGGCTGGCTC	CTCCAACCACCCCACGAATTG
OsPSBD	GTAGGATGGTCTGGCCTATTGCTT	ACTATTGGCAGGGGTGGAAACTG
OsPSBE	TGTCTGGAAGCACGGGAGAACGT	GTTTGGCCGAGGACTTCCAAACAC
OsPSBF	CTATTTTTACAGTGCGATGGCTGG	TATCGTTGGATGAACTGCATTGCT
OsPSBO	TCGAGGAGAAGGACGGAATCGAC	CTTTGGGTCGAGGAAGGACGAAC
OsPSBY	TGCGGTCAGACTGAGAGCTTGAG	AAGGGAAGAGAGGATTAGGAACAGGATT
OsPETA	GCAGCAAGGTTATGAAAACCCAC	AACAGCACCCACATTCAACCCT
OsPETB	TTCAGACCTCGCAACCAGACTG	AACAAAAGGCAAGGGTTCTTCGA
OsPETC	CTCAAGGGTGACCCGACGTACCT	AGGGGCAGATGAACTTGTTCTCG
OsPETD	GCGTGGCCCAACGATCTTTTAT	TCCAGAGGAGTTGCAAACGGAT
OsPETH	GTTCTGGGATCCAGGGTAGCAAC	ACTGACATGCAGAGCACCTTGTTC
OsATPA	TGAATCTCCTGCTCCGGGTATAAT	TGCTGTTTTGCCGGTTTGTCT
OsATPB	TCGCAATTCTTGGGTTGGATGA	CAACATACTTTCCCGGAGAACCG
OsATPG	AGGTGGAGCTCCTCTACTCCAAGT	TCAGCTTCCCTTCCTTGGTGGT
OsNDHA	CGAGCTGCCGCTCAATCTATTAG	AGGCTGACGCCAAAGATTCCATC
OsNDHB	TAACAGCTACTCTAGGGGGAATGTT	CTTGCCCCACCCATGAGTAAAT
OsRBCS2	ACTCCAAGAGCTCGCAGACC	GCCTGTAGTTGGCACCCAGA
OsLHCA2	CCTGGTGGTCTGTGGTTTGACC	CCTGGAACCAAGCTCCCATGACG
OsLHCA3	GCTCAGGCTCTTCTCTCTGGGAG	CTCAGGGACTGCTTGGATGCGA
OsLHCA4	CCCTTCTCCCACCTACCTCAACG	TCAACCCGATCTTCGTCAGCACC
OsLHCA5	TCACCTCGACGGCACATTACCT	CTGCTTCAAACCACACTGGCAG
OsLHCB1.3	AAGTGAAGTGGGGGACCGTAGC	TCTCGTCGCACTAAACCCATCTTC
OsLHCB2.1	CACGATCGAGATGGTGCCAAC	CGGCCTGCGGCTTACATTAA
OsLHCB3	GACTTCAAGGAGCCCGTGTGG	CCATGAGGACGACCTGGAATCC
OsLHCB4.1	TCTTTCGCGCGCAATTCAAAC	TGCAAGTCGCCATTAACCACC
OsLHCB5	GATGAGCTCGCCAAGTGGTACG	CTGAGGCCAAAAGGGTCGTAGC

表1

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标记 Marker	正向序列 Forward sequence (5'-3')	反向序列 Reverse sequence (5'-3')
RM405	TATGCTTTCTGTCAGCTTCC	CTGCTGTGAAAGAGTTGACG
ZTQ43	TGCAGAGACAAGGAAGCGG	TCCTGATCGTTGAGCAGGC
S5-20	TCAGGTCTTACGACGGTATGG	GGACACACTAGAATCTACGCACG
S5-57	GAAGCTATTAGCCGGGATCG	GCCAAGGCAAAGCTCTCTT
S5-119	AGCGCAGATATGCTTTCCAA	TATGGCAGTGCAGAGGTGA
RM18053	GAGACCAGAGGGAGACAAAGA	CTTAGGTCTCCCGACAGTCACG

基因登录号 Accession number	蛋白质亚细胞定位预测 Protein subcellular localization prediction (https://www.genscript.com/)		基因登录号 Accession number		蛋白质亚细胞定位预测 Protein subcellular localization prediction (https://www.genscript.com/)
LOC_Os05g09200	chlo: 7, nucl: 3, mito: 3		LOC_Os05g09640		chlo: 10, nucl: 1, cyto: 1, mito: 1
LOC_Os05g09240	chlo: 7, mito: 6		LOC_Os05g09704		chlo: 9.5, chlo_mito: 5.5, vacu: 2, plas: 1
LOC_Os05g09280	E.R.: 4.5, cyto: 4, E.R._plas: 4, plas: 2.5, mito: 2		LOC_Os05g09708		chlo: 8, cyto: 2, nucl: 1, mito: 1, extr: 1
LOC_Os05g09290	chlo: 14		LOC_Os05g09724		chlo: 8, vacu: 2, nucl: 1, plas: 1, extr: 1
LOC_Os05g09370	chlo: 12, nucl: 1		LOC_Os05g09740		chlo: 11, vacu: 2
基因登录号 Accession number	蛋白质亚细胞定位预测 Protein subcellular localization prediction (https://www.genscript.com/)	基因登录号 Accession number		蛋白质亚细胞定位预测 Protein subcellular localization prediction (https://www.genscript.com/)
LOC_Os05g09400	chlo: 12, mito: 2	LOC_Os05g10210		chlo: 4, cyto: 2.5, vacu: 2, cyto_nucl: 2, mito: 1
LOC_Os05g09430	chlo: 9, mito: 3, extr: 1	LOC_Os05g10290		chlo: 9, cyto: 2, mito: 1, extr: 1
LOC_Os05g09450	chlo: 7, extr: 3, nucl: 2, cyto: 1	LOC_Os05g10300		chlo: 3, nucl: 3, E.R.: 3, vacu: 2, mito: 1, plas: 1
LOC_Os05g09480	chlo: 10, mito: 2, nucl: 1	LOC_Os05g10330		chlo: 6, cyto: 2, vacu: 2, E.R.: 2, nucl: 1
LOC_Os05g09490	chlo: 9, cyto: 5	LOC_Os05g10350		chlo: 8, cyto: 3, nucl: 1.5, nucl_plas: 1.5
LOC_Os05g09510	chlo: 12, nucl: 1	LOC_Os05g10380		chlo: 12, mito: 2
LOC_Os05g09540	chlo: 9, mito: 4	LOC_Os05g10420		cyto: 7, chlo: 6
LOC_Os05g09550	chlo: 11, nucl: 1, plas: 1	LOC_Os05g10430		chlo: 6, nucl: 5, extr: 1, E.R.: 1
LOC_Os05g09590	chlo: 5, mito: 5, nucl: 4	LOC_Os05g10550		chlo: 6, nucl: 5, cyto: 3

基因登录号 Accession number	基因注释 Gene annotation
LOC_Os05g09060	collagen alpha-2, putative, expressed
LOC_Os05g09360	glycine-rich protein, putative, expressed
LOC_Os05g09410	histidine-containing phosphotransfer protein 4, putative, expressed
LOC_Os05g09440	malic enzyme
LOC_Os05g09500	cytosolic hexokinase
LOC_Os05g09520	calmodulin binding protein
LOC_Os05g09600	GA11916-PA, putative, expressed
LOC_Os05g09620	SCC3, putative, expressed
LOC_Os05g09630	homeobox domain containing protein, expressed
LOC_Os05g09650	ubiquinone biosynthesis protein CoQ₄, putative, expressed
LOC_Os05g09660	HAD superfamily phosphatase, putative, expressed
LOC_Os05g09680	acid phosphatase family, putative, expressed
LOC_Os05g09732	acid phosphatase family, putative, expressed
LOC_Os05g10310	acid phosphatase, putative, expressed
LOC_Os05g10370	acid phosphatase, putative, expressed