植物学报

QTL	Chr.	Interval (Mb)	Genes	Annotation	Reference
qHTSR1.1	1	1.90-1.95	Os01g0134800*	Glycosyl hydrolase family
Os01g0134900*	Glycosyl hydrolase family
Os01g0135000	Similar to 1,4-beta-D xylan xylanohydrolase
Os01g0135700*	OsCML16, calmodulin, calcium signaling during abiotic stress	Yang et al., 2018
Os01g0135800*	HSP20, heat shock protein
Os01g0135900*	HSP20, heat shock protein	Sarkar et al., 2020
Os01g0136000*	OsHSP16.9C, HSP17.5 heat shock protein	Sarkar et al., 2020
Os01g0136050	16.9 kDa heat shock protein
qHTSR4.1	4	15.72-15.92	Os04g0336600	Peptidase aspartic, catalytic domain containing protein
Os04g0337000*	Aspartyl protease domain containing protein
Os04g0337201*	SAD2, sensitive to ABA and drought2
Os04g0337300	SAD2, sensitive to ABA and drought2
Os04g0337500	Oxidoreductase
Os04g0337800*	IN2-2 protein
Os04g0338000*	Aldo/keto reductase family protein	Kim et al., 2019
qHTSR5.1	5	20.41-20.48	Os05g0417000*	DUF260 domain containing protein
Os05g0417100*	Deg Protease Protein, chloroplast development and maintenance of PSII function under high temperatures	Zheng et al., 2016
			Os05g0417800*	ulp1 protease family
qHTSR5.2	5	26.25-26.39	Os05g0418000*	GDP-dissociation inhibitor	Shad et al., 2023
Os05g0528500	OsWRKY58, WRKY transcription factor
Os05g0528600	OsYUCCA2, flavin monooxygenase-like enzyme, auxin biosynthesis	Yamamoto et al., 2007
Os05g0528900	Ribosomal protein
Os05g0529000*	OsCOLE1, regulation of intracellular auxin transport	Liu et al., 2016a
Os05g0529200	Enoyl-CoA hydratase/isomerase family protein
Os05g0529300	ER lumen protein retaining receptor
Os05g0529400*	Ubiquitin domain-containing protein
Os05g0529600*	ATP binding protein
Os05g0529700	Electron transporter/heat shock protein binding protein
Os05g0529900	Cell cycle control protein, OsHsfA4d; Heat stress transcription factor	Mittal et al., 2009
Os05g0530400*	Plant growth and balancing reactive oxygen species du- ring biotic and abiotic stress
Os05g0530500	Snf1 protein kinase; sucrose non-fermenting-1 related protein kinase 1	Kanegae et al., 2005
Os05g0530701	Leucine-rich repeat
Os05g0531000*	A member of S40 gene family, regulation of crosstalk among abiotic and biotic and developmental senescence
Os05g0531100	A member of S40 gene family, regulation of crosstalk among abiotic and biotic and developmental senescence
Os05g0531200	Pollen Ole e I allergen and extensin family protein precursor
Os05g0531400	Pollen Ole e I allergen and extensin family protein precursor
			Os05g0531500	Hydrolase

QTL	Chr.	Interval (Mb)	Genes	Annotation	Reference
qHTSR1.1	1	1.90-1.95	Os01g0134800*	Glycosyl hydrolase family
Os01g0134900*	Glycosyl hydrolase family
Os01g0135000	Similar to 1,4-beta-D xylan xylanohydrolase
Os01g0135700*	OsCML16, calmodulin, calcium signaling during abiotic stress	Yang et al., 2018
Os01g0135800*	HSP20, heat shock protein
Os01g0135900*	HSP20, heat shock protein	Sarkar et al., 2020
Os01g0136000*	OsHSP16.9C, HSP17.5 heat shock protein	Sarkar et al., 2020
Os01g0136050	16.9 kDa heat shock protein
qHTSR4.1	4	15.72-15.92	Os04g0336600	Peptidase aspartic, catalytic domain containing protein
Os04g0337000*	Aspartyl protease domain containing protein
Os04g0337201*	SAD2, sensitive to ABA and drought2
Os04g0337300	SAD2, sensitive to ABA and drought2
Os04g0337500	Oxidoreductase
Os04g0337800*	IN2-2 protein
Os04g0338000*	Aldo/keto reductase family protein	Kim et al., 2019
qHTSR5.1	5	20.41-20.48	Os05g0417000*	DUF260 domain containing protein
Os05g0417100*	Deg Protease Protein, chloroplast development and maintenance of PSII function under high temperatures	Zheng et al., 2016
			Os05g0417800*	ulp1 protease family
qHTSR5.2	5	26.25-26.39	Os05g0418000*	GDP-dissociation inhibitor	Shad et al., 2023
Os05g0528500	OsWRKY58, WRKY transcription factor
Os05g0528600	OsYUCCA2, flavin monooxygenase-like enzyme, auxin biosynthesis	Yamamoto et al., 2007
Os05g0528900	Ribosomal protein
Os05g0529000*	OsCOLE1, regulation of intracellular auxin transport	Liu et al., 2016a
Os05g0529200	Enoyl-CoA hydratase/isomerase family protein
Os05g0529300	ER lumen protein retaining receptor
Os05g0529400*	Ubiquitin domain-containing protein
Os05g0529600*	ATP binding protein
Os05g0529700	Electron transporter/heat shock protein binding protein
Os05g0529900	Cell cycle control protein, OsHsfA4d; Heat stress transcription factor	Mittal et al., 2009
Os05g0530400*	Plant growth and balancing reactive oxygen species du- ring biotic and abiotic stress
Os05g0530500	Snf1 protein kinase; sucrose non-fermenting-1 related protein kinase 1	Kanegae et al., 2005
Os05g0530701	Leucine-rich repeat
Os05g0531000*	A member of S40 gene family, regulation of crosstalk among abiotic and biotic and developmental senescence
Os05g0531100	A member of S40 gene family, regulation of crosstalk among abiotic and biotic and developmental senescence
Os05g0531200	Pollen Ole e I allergen and extensin family protein precursor
Os05g0531400	Pollen Ole e I allergen and extensin family protein precursor
			Os05g0531500	Hydrolase

QTL	This study	Known QTLs or genes
qHTSR1.1	1	1.90-1.95	Nipponbare/Kasalath backcross inbred lines (BILs)	HT during grain filling	0.64-3.44	朱昌兰等, 2005
Teqin/Yuanjiang Common Wild Rice BILs	HT at flowering stage	0.38-2.18	奎丽梅等, 2008
N22/IR64 recombinant inbred lines (RILs)	Shoot length under heat stress	1.86-2.31	Kilasi et al., 2018
qHTSR1.2	1	39.82-39.86	Longdao 5/Zhongyouzao 8RILs	HTSR	37.6-39.08	刘进等, 2022a
Bala/Azucena RILs	HT at flowering stage	38.35-40.66	Jagadish et al., 2010
qHTSR2.2	2	24.58-24.72	255 core materials	HTSR	24.66	魏昭然, 2020
qHTSR4.1	4	15.71-15.92	996/4628 RILs	HT at flowering stage	15.74-18.82	Xiao et al., 2011; 盘毅等, 2011
Bala/Azucena RILs	HT at flowering stage	14.19-16.86	Jagadish et al., 2010
qHTSR 5.2	5	26.25-26.39	Zhongyouzao 8/Fengjin RILs	HT at flowering stage	3.08-26.85	张涛等, 2008
Longdao 5/Zhongyouzao 8 RILs	HTSR	25.78-26.28	刘进等, 2022a; 李林浩, 2023
R498/R3551 RILs	HT at flowering stage	26.69-27.69	陶磊, 2020
qHTSR9.1	9	9.42-9.46	Nipponbare/Kasalath bils	Thermo-tolerance of amylose content	5.27-21.85	朱昌兰等, 2006
qHTSR10	10	22.57-22.63	Bala/Azucena RILs	HT at flowering stage	20.84-23.45	Jagadish et al., 2010
233 core materials	HT during grain filling	22.62	杨飞, 2020
Sasanishiki/Habataki CSSLs	Pollen fertility under high temperature	19.89-23.01	Zhu et al., 2017

QTL	This study	Known QTLs or genes
qHTSR1.1	1	1.90-1.95	Nipponbare/Kasalath backcross inbred lines (BILs)	HT during grain filling	0.64-3.44	朱昌兰等, 2005
Teqin/Yuanjiang Common Wild Rice BILs	HT at flowering stage	0.38-2.18	奎丽梅等, 2008
N22/IR64 recombinant inbred lines (RILs)	Shoot length under heat stress	1.86-2.31	Kilasi et al., 2018
qHTSR1.2	1	39.82-39.86	Longdao 5/Zhongyouzao 8RILs	HTSR	37.6-39.08	刘进等, 2022a
Bala/Azucena RILs	HT at flowering stage	38.35-40.66	Jagadish et al., 2010
qHTSR2.2	2	24.58-24.72	255 core materials	HTSR	24.66	魏昭然, 2020
qHTSR4.1	4	15.71-15.92	996/4628 RILs	HT at flowering stage	15.74-18.82	Xiao et al., 2011; 盘毅等, 2011
Bala/Azucena RILs	HT at flowering stage	14.19-16.86	Jagadish et al., 2010
qHTSR 5.2	5	26.25-26.39	Zhongyouzao 8/Fengjin RILs	HT at flowering stage	3.08-26.85	张涛等, 2008
Longdao 5/Zhongyouzao 8 RILs	HTSR	25.78-26.28	刘进等, 2022a; 李林浩, 2023
R498/R3551 RILs	HT at flowering stage	26.69-27.69	陶磊, 2020
qHTSR9.1	9	9.42-9.46	Nipponbare/Kasalath bils	Thermo-tolerance of amylose content	5.27-21.85	朱昌兰等, 2006
qHTSR10	10	22.57-22.63	Bala/Azucena RILs	HT at flowering stage	20.84-23.45	Jagadish et al., 2010
233 core materials	HT during grain filling	22.62	杨飞, 2020
Sasanishiki/Habataki CSSLs	Pollen fertility under high temperature	19.89-23.01	Zhu et al., 2017

基于高密度Bin图谱的水稻苗期耐热性QTL定位

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赵凌 ¹ , 管菊 ¹ , 梁文化 ¹ , 张勇 ² , 路凯 ¹ , 赵春芳 ¹ , 李余生 ¹ , 张亚东 ¹^,^*

植物学报 | 研究论文 2025,60(3): 342-353

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植物学报 |研究论文 2025 , 60 (3) : 342 -353

基于高密度Bin图谱的水稻苗期耐热性QTL定位

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赵凌¹, 管菊¹, 梁文化¹, 张勇², 路凯¹, 赵春芳¹, 李余生¹, 张亚东¹^,^*

作者信息

¹江苏省农业科学院粮食作物研究所, 国家水稻改良中心南京分中心, 国家耐盐碱水稻技术创新中心华东中心, 江苏省优质水稻工程技术研究中心, 南京 210014

²江苏省农业科学院无锡分院生物育种研究室, 无锡 214000

通讯作者:

*张亚东, 男, 汉族, 江苏省靖江人, 研究员, 硕士生导师。2001年6月本科毕业于扬州大学农学院; 2003年6月硕士研究生毕业于扬州大学作物遗传育种专业; 2014年6月博士研究生毕业于南京农业大学作物遗传育种专业。2003.08-至今在江苏省农业科学院粮食作物研究所工作, 从事水稻遗传育种研究。其中2003.08-2006.07任研究实习员, 2006.08-2012.08任助理研究员, 2012.09-2018.09任副研究员, 2018.10-至今任研究员。2014.08- 2015.12先后在美国密西西比州立大学、德州农工大学做访问学者, 从事作物科学研究。E-mail: zhangyd@jaas.ac.cn

Mapping of QTLs for Heat Tolerance at the Seedling Stage in Rice Based on a High-density Bin Map

Zhao Ling¹, Guan Ju¹, Liang Wenhua¹, Zhang Yong², Lu Kai¹, Zhao Chunfang¹, Li Yusheng¹, Zhang Yadong¹^,^*

Affiliations

¹Jiangsu High Quality Rice Research & Development Center, East China Branch of National Center of Technology Innovation for Saline-Alkali Tolerant Rice, Nanjing Branch of China National Center for Rice Improvement, Institute of Food Crops, Jiangsu Academy of Agricultural Sciences, Nanjing 210014, China

²Biological Breeding Research Laboratory, Wuxi Branch of Jiangsu Academy of Agricultural Sciences, Wuxi 214000, China

Corresponding Author:

*E-mail: zhangyd@jaas.ac.cn

出版时间: 2025-05-10 doi: 10.11983/CBB24047

文章导航

摘要

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鉴定控制水稻(Oryza sativa)高温耐性的新位点和候选基因, 可为耐热遗传育种提供理论支撑, 具有重要的实践意义。利用粳稻(O. sativa subsp. japonica)品种TD70和籼稻(O. sativa subsp. indica)品种Kasalath衍生的重组自交系(RILs)群体为研究材料, 构建基于深度重测序的高密度Bin遗传图谱; 使用QTL IciMappingv软件基于完备复合区间作图法对水稻苗期高温胁迫下的幼苗存活率进行QTLs分析。共检测到26个控制苗期耐热性QTLs, 分布在除第3号染色体外的11条染色体上, LOD值为2.59-16.15, 其中4个QTLs的LOD值大于10, 7个QTLs与已知高温耐性QTLs的位置存在重叠或者部分重叠, 其主效QTL位点qHTSR5.2位于第5号染色体26.25-26.38 Mb区间, LOD值为16.15, 解释7.18%的表型贡献率。对4个主效QTLs区间进行基因功能注释和亲本间序列分析, 共发现27个注释有功能且在2个亲本间编码区存在非同义突变的基因。根据候选基因SNP的类型对RILs群体家系进行基因等位型分类和效应分析, 发现5个基因不同等位型的RILs群体家系高温处理后的幼苗存活率存在显著差异, 推测可能为候选基因, 可用于后续水稻高温耐性的分子机理研究。

关键词

耐热性 / 高密度Bin图谱 / QTL定位 / 水稻 / 苗期

Abstract

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INTRODUCTION: As the main grain crop, rice plays an important role in ensuring food security of China. Rise in global average temperature is detrimental to crop yield and heat stress is currently one of the major abiotic threats on rice production. There are significant variations in heat tolerance among different rice varieties. As a typical quantitative trait, heat tolerance of rice is controlled by multiple genes. Identification of new QTLs and genes related to heat tolerance is very important for the genetic research and the breeding of new heat-tolerant rice varieties.

RATIONALE:In recent years, many heat tolerant QTLs had been identified with different genetic populations and evaluation indicators at different growth stages. Most of those QTLs were mapped in large intervals due to the limited population sizes, simplified experimental designs and inaccurately controlled environments. The heat tolerance level identification in a population is very difficult for mature plants. Therefore, we developed a population of recombinant inbred lines (RILs) with 186 lines derived from japonica rice TD70 and indica rice Kasalath, which showed large variations in seedling survival rates under high temperature stress (HTSR). QTLs associated with HTSR were mapped by the high-density linkage Bin-map and candidate genes were identified.

RESULTS: Twenty-six QTLs related to the HTSR were mapped on 11 of the 12 chromosomes, with the exception of 3. The LOD values of single QTL ranged from 2.59-16.15, four of which with LOD values greater than 10. Seven QTLs were located within the same interval or adjacent to known heat tolerance QTLs. The major locus of qHTSR5.2 was located in the 26.25-26.38 Mb region of Chr. 5 with an LOD value of 12.07, which explained 7.18% of the total phenotypic variation in the HTSR. According to the annotation and sequence analysis of the genes located in the region of four major QTLs,we found that twenty-seven annotated genes with non-synonymous mutations in the coding regions between TD70 and Kasalath. Five of them were identified as potential candidate genes because the RILs sharing each of the distinct haplotypes of their parents for each gene exhibited significant different HTSR resistance level. Among them, three candidate genes encode heat shock proteins HSP20 or HSP17.5.

CONCLUSION: We detected 26 QTLs controlling seedling heat tolerance based on a high-density Bin map in a RIL population. Some of the QTLs were overlapped with known heat tolerance loci, indicating their strong effects on regulating heat tolerance of rice. Five candidate genes were identified through gene annotation, parental sequence comparison, effect analysis of heat tolerance between RILs with different haplotypes. The candidate genes identified in our study could be used for molecular mechanism research on high temperature tolerance of rice in the future.

Mapping of QTL for heat tolerance at seedling stage in rice based on a high-density Bin map. Heat tolerance of parents and RILs population during seedling stage. Location of QTLs contributing to heat tolerance at seedling stage.

Key words

heat tolerance / high-density Bin map / QTL mapping / rice / seedling stage

引用本文

赵凌, 管菊, 梁文化, 张勇, 路凯, 赵春芳, 李余生, 张亚东. 基于高密度Bin图谱的水稻苗期耐热性QTL定位. 植物学报, 2025 , 60 (3) : 342 -353 . DOI: 10.11983/CBB24047

Zhao Ling, Guan Ju, Liang Wenhua, Zhang Yong, Lu Kai, Zhao Chunfang, Li Yusheng, Zhang Yadong. Mapping of QTLs for Heat Tolerance at the Seedling Stage in Rice Based on a High-density Bin Map[J]. Chinese Bulletin of Botany, 2025 , 60 (3) : 342 -353 . DOI: 10.11983/CBB24047

正文

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水稻(Oryza sativa)是我国主要粮食作物, 对于保障粮食安全具有重要作用。水稻虽是喜温作物, 但当外界温度超过植株生长发育所需温度的上限时, 会受到高温胁迫, 其生长发育、产量和品质形成均会受到影响(杨军等, 2020; 宋有金等, 2021; 沈泓等, 2022)。近年来, 随着地球温室效应的加剧, 极端高温环境出现的频率以及强度显著增加, 对水稻、小麦(Triticum aestivum)和玉米(Zea mays)等主要粮食作物生产造成不同程度的危害。IPCC (2021)预计2016- 2035年全球地表平均温度将上升0.3-0.7°C。2022年, 世界范围内出现了超强高温, 长江中下游地区温度高达40°C以上且持续时间长达30天(郝立生等, 2022), 此时正值水稻成熟的关键期, 对水稻生产造成了巨大影响。国家统计局发布的粮食产量数据显示, 受南方地区高温和干旱影响, 2022年全国稻谷总产以及单产与2021年相比均有所下降(http://www.stats.gov.cn/sj/)。到2050年温室效应可能导致我国水稻单产降低10%- 19%, 高温热害已成为影响我国水稻生产的主要气象灾害(Chen and Chen, 2018)。

不同水稻品种的耐热性存在明显差异, 鉴定耐热种质和发掘耐热基因/位点是培育耐热品种及应对高温危害的关键途径, 业已成为水稻分子生物学与遗传育种学研究的热点之一。水稻耐热性属于典型的数量性状, 不仅受多基因控制而且易受环境的影响, 存在明显的基因型与环境互作(杨军等, 2020)。近年来, 研究人员利用不同的遗传群体(F₂、F_2:3、BC、DH、RIL和CSSLs等)在不同生育时期(苗期、孕穗期、抽穗扬花期和灌浆期等), 采用不同的评价指标(花粉育性、产量/结实率、品质相关指标和抽穗开花时间等)在水稻12条染色体上定位到200多个耐热QTLs (Chen et al., 2021)。但多数鉴定的水稻耐热QTL区间较大, 且由于涉及不同的作图群体、实验设计、统计分析和环境, 这些QTL位点的贡献率和相关性有较大差异, 区间也不一致。李林浩(2023)对170个已发表的水稻耐热性相关QTL进行了元分析, 在12条染色体上共鉴定出35个QTL元, 每个Meta-QTL至少包含2个原始耐热性相关QTL, 其中Meta-QTL10.3区间包含的原始QTL数量最多, 为9个; Meta-QTL8.1和Meta- QTL9.3包含7个原始QTL。10多个耐热主效位点/基因(qHTCGR5、OsTT1、OsTT2、OsHTAS和TOGR1等)被精细定位或克隆, 为阐明水稻耐热性相关生理生化与分子遗传机制奠定了基础(Li et al., 2015; 胡时开和钱前, 2016; Liu et al., 2016b; Wang et al., 2016; 曹志斌等, 2020; Cao et al., 2020; Chen et al., 2021; 阚义和林鸿宣, 2022; Kan et al., 2022)。前人对水稻耐热性鉴定多是在孕穗期、抽穗期、开花期和灌浆期进行, 并以花粉育性、结实率、稻米品质或生理特性为指标进行评价(杨梯丰和刘斌, 2009)。鉴于成熟后植株较大, 利用遗传群体进行高温鉴定所需的温光条件很难控制, 限制了水稻高温耐性位点的定位和基因克隆相关研究进展。苗期耐热性鉴定所需条件容易达到和控制, 表型鉴定结果稳定性强(不受外界环境因素的影响), 进行QTL定位和候选基因克隆更加可靠。

水稻的热应激反应十分复杂, 涉及多个基因网络调控系统。目前, 对植物高温下参与生物合成与降解、调控和转运、能量和碳水化合物代谢及氧化还原等的基因和蛋白质研究的较为深入。植物在受高温胁迫后经过信号转导, 基因表达和转录产物的积累均发生了变化, 其中最重要的耐热反应是降低正常蛋白的合成, 加速热激蛋白(heat shock proteins, HSP)的转录和翻译(盘毅等, 2008; 栗振义等, 2016)。大量研究表明, 热激蛋白具有分子伴侣的作用, 其生成量与生物耐热性呈正相关, 热激蛋白的表达能稳定细胞膜系统、防止蛋白质变性和聚集及参与热胁迫信号转换等(Khan et al., 2019; Haq et al., 2019)。除HSP基因外, 目前已克隆的水稻耐热基因编码产物有26S蛋白酶体亚基、G蛋白γ亚基、细胞质膜定位的E3泛素连接酶和叶绿体前体蛋白等(阚义和林鸿宣, 2022)。尽管目前对水稻耐热性的遗传及分子机制研究有了一些进展, 但对于高温信号转导、耐热基因间的互作和HSP诱导高温抗性等方面的机理研究尚有欠缺。

解析水稻苗期耐热位点及相关基因, 对于揭示水稻耐热的分子机制及培育和应用高产水稻品种具有重要意义。Bin图谱是基于SNP标记的新一代遗传图谱, 由于SNP标记分布广泛、多样性高、数量大并且精确度高, 在复杂数量性状研究中具有明显的优势和广泛的应用前景(张亚东等, 2021; 赵凌等, 2022)。本研究以高温胁迫下幼苗存活率存在显著差异的粳稻TD70和籼稻Kasalath衍生的重组自交系(recombinant inbred line, RIL)为作图群体, 利用构建的密度连锁图谱进行苗期高温耐性QTL定位, 进一步鉴定候选基因, 以期为水稻耐热性遗传改良研究奠定基础。

1 材料与方法

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1.1 实验材料

以粳稻(Oryza sativa subsp. japonica)品系TD70 (天鹅谷///9520//72-496/苏御糯的后代)和籼稻(O. sativa subsp. indica)品种Kasalath的重组自交系群体为实验材料。2005年夏收取F₁代单株自交种子, 以单粒传法构建由186个家系组成的RILs。基因型调查为F₁₀世代, 表型调查为F₁₂世代。

1.2 材料种植及高温处理

2022年春季, 在江苏省农业科学院粮食作物研究所人工气候箱中种植RILs群体的186个家系及2个亲本。先用自来水浸种催芽, 7天后剔除长势较弱的幼苗, 移入剪掉底部的96孔板小孔中, 每系种植4行, 每行12株, 设置2次随机重复。实验材料在用基础盐培养基(PhytoTech LABS)配制的营养液(pH5.7-5.8)中培养至3叶1心后进行高温处理, 期间每3天更换1次营养液。幼苗培养和恢复培养期间人工气候室的光周期为16小时光照(28°C)/8小时黑暗(24°C), 相对湿度为60%。高温处理采用逐步升温的方式进行。先进行38°C 4小时、40°C 6小时高温预处理, 然后开始进行高温处理: 光周期为14小时光照(42°C)/10小时黑暗(38°C), 相对湿度为60%。高温处理5天后恢复培养7天, 然后调查高温幼苗存活率(high temperature seedling survival rate, HTSR)。幼苗存活率用高温胁迫下存活的幼苗数与处理幼苗数的百分比表示, 取2次重复的平均值进行统计分析。幼苗存活率经过反正弦转化后用于QTL定位。

1.3 QTL定位

利用186个家系及亲本的全基因组重测序数据, 构建基于SNP的高密度Bin遗传图谱, 图谱含有12 328个Bin标记, 标记间的平均物理距离为30.26 kb (张亚东等, 2021)。利用QTL IciMappingv软件3.4版基于完备区间作图法进行全基因组扫描(王建康, 2009; Meng et al., 2015), 以LOD=2.5为阈值检测控制苗期耐热性QTL, 进行QTL命名(McCouch et al., 1997)。

1.4 数据统计分析

使用Microsoft Excel 2016软件处理表型数据和作图、绘制频次分布图及进行方差分析。

2 结果与分析

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2.1 RIL群体及其亲本的苗期耐热性表型

在高温环境下, 对2个亲本与RILs群体家系进行幼苗存活率鉴定, 结果表明亲本TD70的苗期耐热性较强, 高温处理后的幼苗存活率为80.98%, 植株生长基本不受影响。而Kasalath在苗期对高温胁迫敏感, 幼苗存活率为12.54%, 高温处理后幼苗基本全部死亡。TD70的幼苗存活率极显著高于Kasalath (P<0.01)。RILs群体及其双亲苗期耐热性的表型变异见图1A。

RILs群体的幼苗存活率均值为32.76%, 最高的家系达98.1%, 最低的家系为1.2%, 群体中幼苗存活率存在明显的超亲现象。RILs群体中不同家系间的耐热性存在明显差异, 幼苗存活率呈现正态连续分布, 表明水稻苗期耐热性受数量性状调控(图1B)。

2.2 苗期耐热性QTL定位

利用RILs群体共检测到26个控制苗期耐热性QTLs, 分布在除第3号染色体外的11条染色体上(图2)。其中在第10号染色体上只定位到1个控制苗期耐热性的QTL位点, 在第1、2、8、9、11和12号染色体上均定位到2个QTL位点, 在第4、5和6号染色体上定位到3个QTL位点, 在第7号染色体上定位到4个QTL位点。26个QTLs的LOD值介于2.59-16.15之间, 贡献率为1.88%-7.18%, 加性效应值为-5.99%-1.85%。其中qHTSR2.1、qHTSR2.2、qHTSR4.1、qHTSR5.2、qHTSR8.1、qHTSR8.2和qHTSR12.1被定位在大于0.1 Mb的区间内。

LOD值最大的位点qHTSR5.2位于第5号染色体RBN5335-RBN5336区间, 物理位置为26 253 196- 26 385 406 bp, LOD值为16.15, 表型贡献率为7.18%, 增效等位基因来自Kasalath, 可降低高温胁迫下幼苗存活率5.99%。除qHTSR5.2外, 还检测到3个LOD值大于10的QTL, 分别为qHTSR1.1、qHTSR4.1和qHTSR5.1。qHTSR1.1位于第1号染色体距顶端约1.9 Mb处, LOD值为11.18, 其表型贡献率为4.79%,来自Kasalath的增效等位基因可使幼苗存活率降低3.53%。qHTSR4.1位于第4号染色体15.71- 15.92 Mb区间, LOD值为10.25, 表型贡献率为4.81%, 来自Kasalath的增效等位基因可以使幼苗存活率降低3.38%。qHTSR5.1位于第5号染色体顶端20.41- 20.48 Mb区间, LOD值为12.07, 表型贡献率为4.86%, 来自Kasalath的增效等位基因可使幼苗存活率降低3.78%。

2.3 候选基因的筛选

利用水稻注释数据库(Rice Annotation Project Database, https://rapdb.dna.affrc.go.jp)对检测到的影响苗期高温存活率的4个主效QTL, qHTSR1.1、qHTSR4.1、qHTSR5.1和qHTSR5.2区间内的基因进行功能注释(表1)。

qHTSR1.1位于第1号染色体1.90-1.95 Mb区间, LOD值为11.8, 表型贡献率为4.79%, 该区间有8个注释有功能的基因。序列分析发现, 除基因Os01g0135000和Os01g0136050外, 其它6个基因的编码区在2个亲本间均存在非同义突变。其中Os01g0135700编码钙调蛋白基因OsCML16, 与水稻非生物胁迫过程中的钙信号转导有关, Os01g0135800、Os01g0135900和Os01g0136000均编码热激蛋白(Yang et al., 2018; Sarkar et al., 2020)。根据这4个基因的SNP类型对RILs群体家系进行等位型分类, SNP位点核苷酸为亲本TD70型的命名为HapT, 为Kasalath型的命名为HapK。通过方差分析检测HapT和HapK等位型对群体家系高温幼苗存活率的效应, 结果发现其中3个编码热激蛋白的基因(Os01g0135800、Os01g0135900和Os01g0136000)不同等位型RILs家系间高温幼苗存活率存在显著差异(P=0.0367、0.0156、0.0218), 推测为qHTSR1.1可能的候选基因(图4)。Os01g0135800以及Os01g0135900编码HSP20热激蛋白, 其中Os01g0135800 (LOC_Os01g04340)全长为807 bp, 编码区存在2处错义突变和4处移码突变; Os01g0135900 (LOC_Os01g04350)全长为839 bp, 编码区在TD70与Kasalath间存在4处错义突变和1处移码突变。Os01g0136000 (LOC_Os01g04360)编码HSP17.5热激蛋白, 全长为542 bp, 注释为水稻OsHSP16.9C基因(Zhang et al., 2016), 其DNA序列在TD70与Kasalath间仅存在1处错义突变(A/C), 导致编码的氨基酸由Asn变成Lys (图3)。

qHTSR4.1位于第4号染色体15.72-15.92 Mb区间, 该区间内有7个注释有功能的基因, 其中4个基因的编码区在2个亲本间存在非同义突变, 分别编码天冬氨酰蛋白酶结构域蛋白、SAD2、IN2-2蛋白和氧化还原酶。但是4个基因的不同等位型RILs家系间的HTSR差异不显著。

qHTSR5.1位于第5号染色体20.41-20.48 Mb区间, LOD值为12.07, 表型贡献率为4.86%, 区间内有4个注释有功能的基因, 编码区在2个亲本间均存在非同义突变, 分别编码DUF 260结构域蛋白质、Deg蛋白酶、ulp1蛋白酶家族成员以及GDP解离抑制剂蛋白。进一步分析发现, Os05g0417100和Os05g0417800基因不同等位型的RILs家系间高温幼苗存活率存在显著差异(P=0.044 7、0.041 6), 推测它们为该位点可能的候选基因(图4)。Os05g0417100 (LOC_Os05g34460)基因(TCM5)全长1 845 bp, 编码615 aa的Deg蛋白酶, 具有在高温下维持叶绿体发育和PSII功能的作用(Zheng et al., 2016), 其DNA序列在2个亲本间仅存在1处错义突变(A/G), 导致编码的氨基酸由Lys变成Arg。Os05g0417800 (LOC_Os05g34520)基因全长2 289 bp, 编码ulp1蛋白酶家族成员, 其DNA序列在2个亲本间存在8处错义突变、7处移码突变和1处同义突变(图3)。

qHTSR5.2位于第5号染色体26.25-26.39 Mb区间内, 该区间有18个注释有功能的基因。其中基因Os05g0529000、Os05g0529400、Os05g0529600、Os05g0530400和Os05g0531000的编码区在2个亲本间存在非同义突变, 进一步分析发现这5个基因不同等位型RILs家系间HTSR均差异不显著。

通过对4个主效QTL中的基因注释、亲本间序列比较、等位型和效应分析, 最终确定其中2个主效QTL的候选基因。qHTSR1.1位点有3个候选基因, Os01g0135800、Os01g0135900和Os01g0136000, 分别编码热激蛋白HSP20和HSP17.5, 其中Os01g0136000为已克隆的OsHSP16.9C基因。qHTSR5.1位点有2个候选基因, Os05g0417100编码Deg蛋白酶, 在高温下维持叶绿体的发育和PSII的功能; Os05g0417800编码ulp1蛋白酶家族成员。

3 讨论

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3.1 水稻耐热性相关QTL比较

高温胁迫是影响我国乃至全球水稻生产的主要非生物逆境因素之一, 因此耐热性一直是水稻遗传和分子生物学研究的重要课题。但由于对遗传群体进行耐热性鉴定需要特定的设施条件, 特别是在成熟期进行高温耐性鉴定较为困难。目前已报道的水稻耐热性相关QTL仅有200多个, 涉及利用不同的遗传群体, 在不同的生育时期鉴定不同的性状。

本研究利用籼粳交RILs群体构建高密度Bin遗传图谱, 检测到26个控制苗期耐热性的QTLs, 通过遗传和物理图谱对比(http:/www.gramene.org), 发现一些位点与已知耐热性位点存在区间重叠, 如qHTSR1.1、qHTSR2.2、qHTSR5.2、qHTSR9.1和qHTSR10。本研究检测到的qHTSR5.2位于第5号染色体26.25-26.39 Mb区间。张涛等(2008)利用中优早8号/丰锦的重组自交系群体在第5号染色体3.08-26.85 Mb区间定位到控制水稻抽穗扬花期耐热性QTLs, 其LOD值为4.55。刘进等(2022a)利用龙稻5号/中优早8号的RILs群体, 在第5号染色体25.78-26.28 Mb区间检测到控制高温幼苗存活率的位点, 覆盖了我们检测到的qHTSR5.2。陶磊(2020)利用R498/R3551的RILs群体在第5号染色体26.698-27.69 Mb区间检测到控制开花期高温耐性的位点, 位于本研究定位到的qHTSR5.2附近。李林浩(2023)对已发表的水稻耐热性相关QTL进行元分析, 也认为第5号染色体24.26- 27.34 Mb区间内存在1个QTL元, 覆盖了本研究的qHTSR5.2 (表2)。

本研究在第1号染色体上定位到的qHTSR1.1位于约1.9 Mb, 与朱昌兰等(2005)定位到的灌浆期耐热性主效位点qHR1、奎丽梅等(2008)定位到的抽穗期开花期耐热性位点qHT1以及Kilasi等(2018)定位到的高温苗期根长位点位于同一区间; 位于约39.82 Mb的qHTSR1.2与利用Bala/Azucena RILs、龙稻5号/中优早8号RILs分别定位到的开花期耐热性位点qtl_1.1及苗期耐热性位点qHTG1.2位于同一区间(Jagadish et al., 2010; 刘进等, 2022a)。位于第4号染色体的位点qHTSR4.1则与Jagadish等(2010)、Xiao等(2011)及盘毅等(2011)定位到的开花期耐热性QTL位于同一区间(表2)。

此外, 本研究发现的qHTSR2.2与魏昭然(2020)利用255份籼稻核心材料通过全基因组关联分析定位到的苗期高温耐性位点qSR_ind2-3, qHTSR9.1与朱昌兰等(2006)鉴定的直链淀粉含量高温耐性位点qHTAC9-1所在的染色体区间相近; qHTSR10与利用Bala/Azucena RILs、233份核心材料和Sasanishiki/ Habataki CSSLs分别检测到的开花期耐热性、灌浆期耐热性及高温下小穗育性位点在同一区间(Jagadish et al., 2010; Zhu et al., 2017; 杨飞, 2020)。

这些QTL在不同群体中利用不同的耐热性指标被鉴定出来, 说明其在调控水稻耐热性方面具有较强的效应, 可能有较大的应用前景, 尚需进一步研究。

3.2 苗期耐热性候选基因的筛选

植物HSP种类众多, 根据分子量大小分为HSP100 (100 kDa以上)、HSP90 (83-90 kDa)、HSP70 (66-78 kDa)、HSP60 (60 kDa)和小热激蛋白sHSPs (15-42 kDa) 5大家族(栗振义等, 2016)。大部分HSP在受到高温胁迫时会大量表达, 参与胁迫应答, 使植株表现出更强的耐热性, 及耐盐和干旱等能力。不同家族的HSP具有不同的生物学功能。多数HSP100蛋白具有ATP酶结构, 为发挥分子伴侣作用提供能量。HSP90底物多为信号转导蛋白, 如类固醇激素受体或者信号激酶(Murthy and Ravishankar, 2016)。HSP70的保守性最强, 主要在植物的叶绿体、线粒体和内质网等细胞器中行使功能, 部分参与辅助多肽折叠及蛋白转运, 引导错误折叠蛋白降解(Bauer et al., 2018)。HSP60是热休克依赖型热激蛋白, 主要通过参与合成蛋白的折叠与转运进程发挥作用, 促进新蛋白的正确形成(Jacob et al., 2017)。sHSPs分子量低, 具有分子伴侣活性, 可与蛋白形成稳定的复合体, 进而更易接近ATP依赖性HSPs, 最终使蛋白质正确折叠与构建, 以便后续加工(Zhang et al., 2015)。

sHSPs是植物中含量最丰富且分布最广泛的热激蛋白, 具有保守的ACD结构域和可变的两端序列区域, 与耐热性密切相关, 近年来备受关注。最早发现植物sHSPs受到高温胁迫诱导表达(俞佳虹等, 2017)。随着研究的不断深入, 发现干旱、低温和病原菌等生物和非生物胁迫均能诱导sHSPs的表达量升高几十至几百倍, 且表达强度与胁迫强度和持续时间相关, 在受到胁迫后一段时间表达量很稳定, 可能对胁迫后修复起重要作用(Siddique et al., 2008)。刘进等(2022b)以周南稻/赣早籼58号的RILs为研究材料, 构建高密度Bin遗传图谱, 筛选出8个苗期耐热性调控的可能目标基因, 其中包括3个调控胁迫耐性的转录因子以及1个HSP40型的sHSPs基因Os02g0804500, 编码含有DNAJ结构域蛋白。Sarkar等(2009)发现19个sHSPs基因在高温胁迫下明显上调表达。超表达HSP17.7基因可以提高水稻的耐热性和抗旱性(Murakami et al., 2004)。42°C高温特异性诱导HSP26大量表达, 推测HSP26能保护叶绿体避免氧化胁迫对其造成破坏(Lee et al., 2000)。郭虹霞等(2019)克隆了2个水稻sHSPs基因Os16.9A和Os16.9B, 它们的启动子均含有CAAT-box、TATA-box、GATA-box及与热激等胁迫应答有关的顺式作用元件; Os16.9A的启动子还包含响应冷和干旱应答的顺式作用元件。

本研究通过对4个苗期耐热主效QTL位点中的基因注释、亲本间序列比较、等位型和效应分析, 确定了5个候选基因。其中, Os01g0135800、Os01g0135900和Os01g0136000分别编码热激蛋白HSP20和HSP17.5, 属于sHSP家族。通过ENSEMBL数据库(http://plants.ensembl.org)查询, 发现Os01g0135800、Os01g0135900和Os01g0136000均具有蛋白质复合物聚合和蛋白质折叠功能, 能响应高温、盐、过氧化氢和活性氧胁迫。Os01g0136000是已克隆的OsHSP16.9C基因, 编码HSP17.5热激蛋白。OsHSP16.9C具有极强的组织特异性, 在穗中表达水平最高, 节中次之, 在根、茎、幼叶和老叶中表达水平较低。45°C热处理后幼苗和幼穗中的OsHSP16.9C表达量均急剧上升, 并分别在0.5和1小时达到峰值, 随后逐步降低(Zhang et al., 2016)。Os01g0135800和Os01g0135900的具体生物学功能有待进一步探索。

qHTSR5.1位点有2个候选基因, Os05g0417100是水稻TCM5基因, 编码Deg蛋白酶, 高温下维持叶绿体发育和PSII的功能。TCM5在茎和叶等绿色组织中高表达, 并在高温下3叶期幼苗中高表达(Zheng et al., 2016)。预测Os05g0417800编码ulp1蛋白酶家族成员, 但具体生物学功能尚未见报道。

作者贡献声明

收起

赵凌: 构思并撰写论文初稿; 管菊: 配制培养基、高温处理; 梁文化: 遗传群体田间种植; 张勇: 协助高温处理; 路凯: 协助统计耐热性表型数据; 赵春芳: 遗传群体分子图谱构建; 李余生: 遗传群体田间种植; 张亚东: 遗传群体构建、构思和修改文章。

基金

收起

江苏省种业振兴“揭榜挂帅”项目(No.JBGS(2021)040)

参考文献

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doi: 10.11983/CBB24047

接收时间：2024-03-28
首发时间：2026-10-20
出版时间：2025-05-10

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收稿日期：2024-03-28
录用日期：2024-05-27

基金

江苏省种业振兴“揭榜挂帅”项目(No.JBGS(2021)040)

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²江苏省农业科学院无锡分院生物育种研究室, 无锡 214000

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2种不同金属材料的力学参数

科 Family	属数 Number of genus	种数 Number of species	占总种数比例 Percentage of total species (%)	属 Genus	种数 Number of species	占总种数比例 Percentage of total species (%)
鹅膏菌科Amanitaceae	2	11	5.26	鹅膏菌属 Amanita	10	4.78
小菇科 Mycenaceae	2	12	5.74	丝盖伞属 Inocybe	5	2.39
多孔菌科 Polyporaceae	8	14	6.70	蜡蘑属 Laccaria	5	2.39
红菇科 Russulaceae	3	23	11.00	小皮伞属 Marasmius	6	2.87
				小菇属 Mycena	11	5.26
				光柄菇属 Pluteus	5	2.39
				红菇属 Russula	17	8.13
				栓菌属 Trametes	5	2.39

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TxT

QTL	Chr.	Interval (Mb)	Genes	Annotation	Reference
qHTSR1.1	1	1.90-1.95	Os01g0134800*	Glycosyl hydrolase family
Os01g0134900*	Glycosyl hydrolase family
Os01g0135000	Similar to 1,4-beta-D xylan xylanohydrolase
Os01g0135700*	OsCML16, calmodulin, calcium signaling during abiotic stress	Yang et al., 2018
Os01g0135800*	HSP20, heat shock protein
Os01g0135900*	HSP20, heat shock protein	Sarkar et al., 2020
Os01g0136000*	OsHSP16.9C, HSP17.5 heat shock protein	Sarkar et al., 2020
Os01g0136050	16.9 kDa heat shock protein
qHTSR4.1	4	15.72-15.92	Os04g0336600	Peptidase aspartic, catalytic domain containing protein
Os04g0337000*	Aspartyl protease domain containing protein
Os04g0337201*	SAD2, sensitive to ABA and drought2
Os04g0337300	SAD2, sensitive to ABA and drought2
Os04g0337500	Oxidoreductase
Os04g0337800*	IN2-2 protein
Os04g0338000*	Aldo/keto reductase family protein	Kim et al., 2019
qHTSR5.1	5	20.41-20.48	Os05g0417000*	DUF260 domain containing protein
Os05g0417100*	Deg Protease Protein, chloroplast development and maintenance of PSII function under high temperatures	Zheng et al., 2016
			Os05g0417800*	ulp1 protease family
qHTSR5.2	5	26.25-26.39	Os05g0418000*	GDP-dissociation inhibitor	Shad et al., 2023
Os05g0528500	OsWRKY58, WRKY transcription factor
Os05g0528600	OsYUCCA2, flavin monooxygenase-like enzyme, auxin biosynthesis	Yamamoto et al., 2007
Os05g0528900	Ribosomal protein
Os05g0529000*	OsCOLE1, regulation of intracellular auxin transport	Liu et al., 2016a
Os05g0529200	Enoyl-CoA hydratase/isomerase family protein
Os05g0529300	ER lumen protein retaining receptor
Os05g0529400*	Ubiquitin domain-containing protein
Os05g0529600*	ATP binding protein
Os05g0529700	Electron transporter/heat shock protein binding protein
Os05g0529900	Cell cycle control protein, OsHsfA4d; Heat stress transcription factor	Mittal et al., 2009
Os05g0530400*	Plant growth and balancing reactive oxygen species du- ring biotic and abiotic stress
Os05g0530500	Snf1 protein kinase; sucrose non-fermenting-1 related protein kinase 1	Kanegae et al., 2005
Os05g0530701	Leucine-rich repeat
Os05g0531000*	A member of S40 gene family, regulation of crosstalk among abiotic and biotic and developmental senescence
Os05g0531100	A member of S40 gene family, regulation of crosstalk among abiotic and biotic and developmental senescence
Os05g0531200	Pollen Ole e I allergen and extensin family protein precursor
Os05g0531400	Pollen Ole e I allergen and extensin family protein precursor
			Os05g0531500	Hydrolase

QTL

Chr.

Interval (Mb)

Genes

Annotation

Reference

qHTSR1.1

1.90-1.95

Os01g0134800*

Glycosyl hydrolase family

Os01g0134900*

Glycosyl hydrolase family

Os01g0135000

Similar to 1,4-beta-D xylan xylanohydrolase

Os01g0135700*

OsCML16, calmodulin, calcium signaling during abiotic stress

Yang et al., 2018

Os01g0135800*

HSP20, heat shock protein

Os01g0135900*

HSP20, heat shock protein

Sarkar et al., 2020

Os01g0136000*

OsHSP16.9C, HSP17.5 heat shock protein

Sarkar et al., 2020

Os01g0136050

16.9 kDa heat shock protein

qHTSR4.1

15.72-15.92

Os04g0336600

Peptidase aspartic, catalytic domain containing protein

Os04g0337000*

Aspartyl protease domain containing protein

Os04g0337201*

SAD2, sensitive to ABA and drought2

Os04g0337300

SAD2, sensitive to ABA and drought2

Os04g0337500

Oxidoreductase

Os04g0337800*

IN2-2 protein

Os04g0338000*

Aldo/keto reductase family protein

Kim et al., 2019

qHTSR5.1

20.41-20.48

Os05g0417000*

DUF260 domain containing protein

Os05g0417100*

Deg Protease Protein, chloroplast development and maintenance of PSII function under high temperatures

Zheng et al., 2016

Os05g0417800*

ulp1 protease family

qHTSR5.2

26.25-26.39

Os05g0418000*

GDP-dissociation inhibitor

Shad et al., 2023

Os05g0528500

OsWRKY58, WRKY transcription factor

Os05g0528600

OsYUCCA2, flavin monooxygenase-like enzyme, auxin biosynthesis

Yamamoto et al., 2007

Os05g0528900

Ribosomal protein

Os05g0529000*

OsCOLE1, regulation of intracellular auxin transport

Liu et al., 2016a

Os05g0529200

Enoyl-CoA hydratase/isomerase family protein

Os05g0529300

ER lumen protein retaining receptor

Os05g0529400*

Ubiquitin domain-containing protein

Os05g0529600*

ATP binding protein

Os05g0529700

Electron transporter/heat shock protein binding protein

Os05g0529900

Cell cycle control protein, OsHsfA4d; Heat stress transcription factor

Mittal et al., 2009

Os05g0530400*

Plant growth and balancing reactive oxygen species du- ring biotic and abiotic stress

Os05g0530500

Snf1 protein kinase; sucrose non-fermenting-1 related protein kinase 1

Kanegae et al., 2005

Os05g0530701

Leucine-rich repeat

Os05g0531000*

A member of S40 gene family, regulation of crosstalk among abiotic and biotic and developmental senescence

Os05g0531100

A member of S40 gene family, regulation of crosstalk among abiotic and biotic and developmental senescence

Os05g0531200

Pollen Ole e I allergen and extensin family protein precursor

Os05g0531400

Pollen Ole e I allergen and extensin family protein precursor

Os05g0531500

Hydrolase

QTL	This study	Known QTLs or genes
qHTSR1.1	1	1.90-1.95	Nipponbare/Kasalath backcross inbred lines (BILs)	HT during grain filling	0.64-3.44	朱昌兰等, 2005
Teqin/Yuanjiang Common Wild Rice BILs	HT at flowering stage	0.38-2.18	奎丽梅等, 2008
N22/IR64 recombinant inbred lines (RILs)	Shoot length under heat stress	1.86-2.31	Kilasi et al., 2018
qHTSR1.2	1	39.82-39.86	Longdao 5/Zhongyouzao 8RILs	HTSR	37.6-39.08	刘进等, 2022a
Bala/Azucena RILs	HT at flowering stage	38.35-40.66	Jagadish et al., 2010
qHTSR2.2	2	24.58-24.72	255 core materials	HTSR	24.66	魏昭然, 2020
qHTSR4.1	4	15.71-15.92	996/4628 RILs	HT at flowering stage	15.74-18.82	Xiao et al., 2011; 盘毅等, 2011
Bala/Azucena RILs	HT at flowering stage	14.19-16.86	Jagadish et al., 2010
qHTSR 5.2	5	26.25-26.39	Zhongyouzao 8/Fengjin RILs	HT at flowering stage	3.08-26.85	张涛等, 2008
Longdao 5/Zhongyouzao 8 RILs	HTSR	25.78-26.28	刘进等, 2022a; 李林浩, 2023
R498/R3551 RILs	HT at flowering stage	26.69-27.69	陶磊, 2020
qHTSR9.1	9	9.42-9.46	Nipponbare/Kasalath bils	Thermo-tolerance of amylose content	5.27-21.85	朱昌兰等, 2006
qHTSR10	10	22.57-22.63	Bala/Azucena RILs	HT at flowering stage	20.84-23.45	Jagadish et al., 2010
233 core materials	HT during grain filling	22.62	杨飞, 2020
Sasanishiki/Habataki CSSLs	Pollen fertility under high temperature	19.89-23.01	Zhu et al., 2017

QTL

This study

Known QTLs or genes

Chr.

Position (Mb)

Population

Traits

Position (Mb)

Reference

qHTSR1.1

1.90-1.95

Nipponbare/Kasalath backcross inbred lines (BILs)

HT during grain filling

0.64-3.44

朱昌兰等, 2005

Teqin/Yuanjiang Common Wild Rice BILs

HT at flowering stage

0.38-2.18

奎丽梅等, 2008

N22/IR64 recombinant inbred lines (RILs)

Shoot length under heat stress

1.86-2.31

Kilasi et al., 2018

qHTSR1.2

39.82-39.86

Longdao 5/Zhongyouzao 8RILs

HTSR

37.6-39.08

刘进等, 2022a

Bala/Azucena RILs

HT at flowering stage

38.35-40.66

Jagadish et al., 2010

qHTSR2.2

24.58-24.72

255 core materials

HTSR

24.66

魏昭然, 2020

qHTSR4.1

15.71-15.92

996/4628 RILs

HT at flowering stage

15.74-18.82

Xiao et al., 2011; 盘毅等, 2011

Bala/Azucena RILs

HT at flowering stage

14.19-16.86

Jagadish et al., 2010

qHTSR 5.2

26.25-26.39

Zhongyouzao 8/Fengjin RILs

HT at flowering stage

3.08-26.85

张涛等, 2008

Longdao 5/Zhongyouzao 8 RILs

HTSR

25.78-26.28

刘进等, 2022a; 李林浩, 2023

R498/R3551 RILs

HT at flowering stage

26.69-27.69

陶磊, 2020

qHTSR9.1

9.42-9.46

Nipponbare/Kasalath bils

Thermo-tolerance of amylose content

5.27-21.85

朱昌兰等, 2006

qHTSR10

22.57-22.63

Bala/Azucena RILs

HT at flowering stage

20.84-23.45

Jagadish et al., 2010

233 core materials

HT during grain filling

22.62

杨飞, 2020

Sasanishiki/Habataki CSSLs

Pollen fertility under high temperature

19.89-23.01

Zhu et al., 2017