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作物学报 ›› 2019, Vol. 45 ›› Issue (5): 662-675.doi: 10.3724/SP.J.1006.2019.82049

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

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

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()   

  1. 西南大学水稻研究所 / 西南大学农业科学研究院, 重庆 400715
  • 收稿日期:2018-10-10 接受日期:2019-01-12 出版日期:2019-05-12 网络出版日期:2019-02-12
  • 通讯作者: 王楠
  • 基金资助:
    本研究由国家自然科学基金项目(31771750);重庆市基础研究与前沿探索项目(cstc2018jcyjAX0424);重庆市研究生科研创新项目(CYS18084)

Phenotypic identification and gene mapping of temperature-sensitive green- revertible albino mutant tsa2 in rice (Oryza sativa L.)

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()   

  1. Rice Research Institute, Southwest University / Academy of Agricultural Sciences, Southwest University, Chongqing 400715, China
  • Received:2018-10-10 Accepted:2019-01-12 Published:2019-05-12 Published online:2019-02-12
  • Contact: Nan WANG
  • Supported by:
    This study was supported by the National Natural Science Foundation of China(31771750);Basic Research and Frontier Exploration Project in Chongqing(cstc2018jcyjAX0424);the Graduate Research and Innovation Project in Chongqing(CYS18084)

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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

图1

野生型(WT)和突变体tsa2的表型特征 A, B, C: 田间日均气温22°C时播种的野生型(WT)和突变体tsa2的苗期(A)、分蘖期(B)和成熟期(C)植株; D, E, F: 田间日均气温28°C时播种的野生型(WT)和突变体tsa2的苗期(D)、分蘖期(E)和成熟期(F)植株; G, H, I, J, K, L: 分别对应A, B, C, D, E, F中的叶片。图H, I, K和L中的1、2、3和4分别表示倒一叶、倒二叶、倒三叶和倒四叶。图A和D中, Bar = 2 cm; 图B和E中, Bar = 6 cm; 图C和F中, Bar = 13 cm; 图G~L中, Bar = 0.7 cm。"

图2

不同田间温度下播种的野生型(WT)和突变体tsa2的光合特性 A: 净光合速率(Pn); B: 气孔导度(Gs); C: 胞间CO2浓度(Ci); D: 蒸腾速率(Tr); **: 在0.01水平上差异显著。"

图3

不同田间温度下播种的野生型(WT)和突变体tsa2的农艺性状 A: 株高(cm); B: 穗长(cm); C: 分蘖数; D: 有效穗数; E: 每穗粒数; F: 每穗实粒数; G: 千粒重(g); H: 结实率(%); *: 在0.05水平上差异显著; **: 在0.01水平上差异显著。"

图4

不同温度下野生型(WT)和突变体tsa2幼苗表型及光合色素含量 A, B, C: 分别为22°C(A)、28°C(B)和32°C(C)下萌发的22 d野生型(WT)和突变体tsa2幼苗; D: 不同温度下萌发的野生型(WT)和突变体tsa2幼苗光合色素含量; **: 在0.01水平上差异显著。图A、B和C中, Bar = 2.5 cm。"

图5

不同温度下野生型(WT)及突变体tsa2叶肉细胞中叶绿体超微结构 A~D: 不同温度下萌发的22 d野生型(WT)(A)和突变体tsa2 (B, C, D)叶肉细胞结构; E, F, G, H: 分别为A、B、C、D中虚线区域的放大视野; I, J: 将22°C萌发的22 d突变体tsa2分别移至28°C (I)和32°C (J)下培养10 d的叶肉细胞结构; K, L: 将28°C萌发的22 d突变体tsa2分别移至22°C (K)和32°C (L)下培养10 d的叶肉细胞结构; M, N, O, P: 分别为I, J, K, L中虚线区域的放大视野。图A~D和I~L中, Bar=2 μm; 图E~H和M~P中, Bar=500 nm。"

图6

野生型(WT)及突变体tsa2叶片中光合色素代谢以及叶绿体发育和光合相关基因的表达分析 **: 在0.01水平上差异显著。"

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2

第5染色体上的部分多态性分子标记"

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

图7

TSA2基因在第5染色体上的分子定位 n = 285: 初步定位群体数目; n = 1150: 精细定位群体数目。"

表3

TSA2基因定位区间内蛋白质亚细胞定位预测"

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

表4

TSA2基因定位区间内部分注释基因"

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
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