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水稻耐低温逆境研究：分子生理机制及育种展望

本站小编 Free考研考试/2022-01-01
刘次桃1, 王威1, 毛毕刚2, 储成才,1 1. 中国科学院遗传与发育生物学研究所,植物基因组学国家重点实验室,北京 100101
2. 湖南杂交水稻研究中心,杂交水稻国家重点实验室,长沙 410125

Cold stress tolerance in rice: physiological changes, molecular mechanism, and future prospects

Citao Liu1, Wei Wang1, Bigang Mao2, Chengcai Chu,1 1. State Key Laboratory of Plant Genomics, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, China
2. State Key Laboratory of Hybrid Rice, China National Hybrid Rice Research and Development Center, Changsha 410125, China

通讯作者: 储成才，博士，研究员，研究方向：资源高效利用的分子基础和作物品种的分子设计。E-mail: ccchu@genetics.ac.cn

第一联系人: 作者简介: 刘次桃,博士,研究方向：水稻耐受非生物逆境的基因克隆及功能分析。E-mail: liucitao2007@sina.com
编委: 张红生
收稿日期:2018-01-8修回日期:2018-02-13网络出版日期:2018-03-20
基金资助: 国家自然科学基金青年基金项目(编号：31501283)资助.


Editorial board:
Received:2018-01-8Revised:2018-02-13Online:2018-03-20
Fund supported: [Supported by the National Natural Science Foundation of China (No. 31501283)].


摘要
低温严重影响水稻的地理分布、生长发育及产量。水稻在低温逆境下会产生一系列的生理及代谢变化,如：叶绿素荧光的改变,电解质渗漏增加,活性氧、丙二醛、蔗糖、脂质过氧化物、脯氨酸等代谢物含量升高,植物内源激素ABA和GA的改变等。了解水稻在低温逆境下的生理代谢变化及低温应答分子机理对水稻耐低温性状的遗传改良具有重要的意义。本文系统总结了水稻在低温逆境下的生理代谢变化、已定位和克隆的耐低温基因/QTL,以及水稻应答低温逆境信号转导机制的最新研究进展,以期为水稻的耐低温育种提供参考。
关键词： 水稻;低温胁迫;耐低温相关基因;数量性状;信号传导

Abstract
Low temperature is a major factor affecting rice geographical distribution growth, development, and productivity. Cold stress mediates a series of physiological and metabolite changes, such as alterations in chlorophyll fluorescence, electrolyte leakage, reactive oxygen species (ROS), malondialdehyde (MAD), sucrose, lipid peroxides, proline, and other metabolites, plant endogenous hormones abscisic acid (ABA) and gibberellin (GA) also changes. In this review, we summarize the recent research progress on physiological and metabolic changes under low temperature, cold stress related loci and QTL reported by map-based cloning and genome-wide association analysis (GWAS), and some molecular mechanisms in response to low temperature in rice. We also discuss the future prospects on breeding cold tolerance varieties of rice.
Keywords：rice;cold stress;cold tolerance related genes;quantitative trait locus (QTL);signal transduction


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本文引用格式
刘次桃, 王威, 毛毕刚, 储成才. 水稻耐低温逆境研究：分子生理机制及育种展望. 遗传[J], 2018, 40(3): 171-185 doi:10.16288/j.yczz.18-007
Citao Liu, Wei Wang, Bigang Mao, Chengcai Chu. Cold stress tolerance in rice: physiological changes, molecular mechanism, and future prospects . Hereditas(Beijing)[J], 2018, 40(3): 171-185 doi:10.16288/j.yczz.18-007


水稻(Oryza sativa L.)是世界上最主要的粮食作物之一,世界上一半人口以水稻为主食[1, 2]。水稻起源于热带、亚热带,相较于小麦(Triticum aestivum L.)、大麦(Hordeum vulgare L.)等作物,水稻对低温更加敏感。随着对粮食需求的日益增加,水稻种植区域不断由热带、亚热带地区向高海拔、高纬度地区扩张,水稻遭受冷害的机率也逐渐增加,世界上约有24个国家如中国、日本、朝鲜等都遇到过严重冷害问题[3]。我国稻区南北分布相差约34度(最北端漠河53°27′N,最南端海南18°90′N),海拔分布相差2 700米(从东南沿海到云贵高原),均有不同程度低温冷害情况。低温对水稻的影响主要发生在幼苗期和生殖生长期。水稻最适宜的生长温度是25℃~ 30℃。苗期,当环境温度低于15℃,水稻的生理代谢就受到影响[2];孕穗期特别是花粉母细胞减数分裂期,当温度低于17℃时结实率受影响[4];开花授粉期,当环境温度持续3 d等于或者低于20℃时,将严重影响结实率[5, 6]。例如在华南和长江中下游双季稻区,早稻经常受寒潮侵袭,常发生温度急剧降低或持续低温阴雨天气造成的不同程度春寒,引起烂秧、死苗,最终影响水稻产量[7,8,9]。生殖生长期,低温影响水稻的结实和灌浆[10]。在高纬度(如东北地区)和高海拔(如云贵高原)地区,水稻在生殖生长期受低温影响最为严重,平均每3~4年就会遭遇一次较大规模的冷害[11,12,13]。除此之外,在华南及长江中下游地区的晚稻也易遭受“寒露风”的危害。每年秋季“寒露”节气前后(9月中下旬)是华南及长江中下游一带水稻抽穗扬花的关键时期,这时遇低温就会造成授粉和受精无法正常进行,水稻的瘪谷率大大增加,从而造成晚稻大幅减产[14]。尽管在全球气候变暖背景下,寒露风呈现减弱趋势,但近几年极端天气增多,寒露风等低温灾害又呈现增多趋势。低温灾害导致我国每年粮食减产3~5亿吨[3, 15],因此培育耐低温水稻品种是水稻育种工作的重要方向。

植物耐低温胁迫是一个复杂的遗传性状,受多个基因/数量性状基因座控制。与其他农艺性状相比,水稻低温耐受性的遗传研究进展缓慢,目前只鉴定出少数耐低温基因[2,16,17]。本文系统总结了低温胁迫下水稻的生理代谢变化、已克隆的耐低温基因/ QTL以及水稻应答低温胁迫的信号转导途径,以期为水稻耐低温育种研究提供帮助。

1 水稻低温胁迫下生理代谢变化指标

低温不仅对水稻造成明显的外部损伤,如发芽率低、幼苗生长发育迟缓甚至死亡、结实率低等,还会引起一系列生理及代谢物的变化,如叶绿素荧光的改变,电解质渗漏增加,活性氧、丙二醛、蔗糖、脂质过氧化物、脯氨酸其他代谢物的含量升高[2],植物激素脱落酸(abscisic acid, ABA)和赤霉素(gibberellins, GA)的改变等,这些生理水平的变化是衡量植物耐低温的生理指标。

1.1 叶绿素含量和叶绿素荧光

叶绿素含量高低和叶绿素荧光参数(最大荧光/可变荧光,Fv/Fm,反映光系统Ⅱ的最大光能转化效率)大小通常被作为判断植物耐受逆境胁迫的重要指标[18]。水稻在低温下会减少叶绿素合成以及叶绿体形成。因此,叶绿素含量以及Fv/Fm值变化是检验水稻对低温胁迫耐受能力的重要指标[18, 19]。研究表明,在水稻和烟草(Nicotiana tabacum L.)中过表达OsiSAP8可以在低温条件下显著提高植物叶绿素含量和对低温的耐受能力[20]。此外,研究还发现OsAsr1过表达的转基因水稻Fv/Fm值显著升高,并在低温条件下表现出明显的生长优势(表1)[21]。

1.2 电解质渗漏(electrolyte leakages, EL)

细胞膜是水稻细胞最先感知低温冷害的部位,其在低温下的理化性质容易发生变化,从而导致细胞内电解质渗漏,因此电介质渗漏率常作为植物耐受低温的重要指标[22]。研究发现,在水稻中分别过表达OsOVP1和OsNAC5等基因导致植物的电解质渗漏率降低,从而表现出对低温的耐受性(表1)[23, 24]。

1.3 活性氧(reactive oxygen species, ROS)与丙二醛(malondialdehyde, MDA)

水稻受低温等逆境胁迫时,细胞内氧代谢平衡失调,产生ROS(包括超氧阴离子O2-、过氧化氢H2O2和氢氧根离子OH-),引发膜脂过氧化,从而造成细胞膜系统损伤[25]。ROS还会促进多聚不饱和脂肪酸降解并产生MDA,进而对植物组织和细胞进一步损害[26]。水稻对氧化胁迫的保护机制有两大系统,即酶系统和非酶系统。酶系统包括各种抗氧化酶,催化清除ROS反应的酶。其中超氧化物歧化酶(SOD)和过氧化氢酶(CAT)是两种最有效的抗氧化酶,这两种酶可以使超氧阴离子和过氧化氢变成水和氧分 子,从而减轻ROS对细胞的伤害[27]。非酶系统包括各种抗氧化剂,其中还原型谷胱甘肽(GSH)和抗坏血酸(AsA)最为重要[28, 29]。如过量表达抗坏血酸过氧化物酶基因OsAPXa可以提高低温下抗坏血酸过氧化物酶的活性,减少细胞内脂类物质的过氧化反应 和MDA含量,从而提高水稻在低温下的结实率 (表1)[30]。

Table 1
表1
表1 水稻在低温下代谢和生理发生改变所涉及的基因
Table 1 Altered cold stress-related gene expression levels lead to metabolic and physiological changes in rice
生理代谢类型 基因 参考文献
光合作用
叶绿素含量 OsiSAP8 [20]
Fv/Fm CBF1/OsDREB1b、OsAsr1、OsASR3、OsCDPK7 [56~58]
电介质渗漏 CBF1/OsDREB1b、OsLti6、OsZFP245、OsOVP1、OsNAC5 [23, 24, 47, 56, 59]
活性氧与丙二醛
过氧化氢 OsAPXa、OsMKK6、OsMPK3、OsNAC6、OsTrx23 [29, 30, 60, 61]
超氧自由基 OsAPXa [30]
羟基自由基 OsAPXa [30]
丙二醛 OsAPXa、OsMKK6、OsMPK3、OsNAC6 [30, 60, 61]
可溶性糖
蔗糖 OsINV4、OsDREB1A [10, 62]
已糖 OsINV4 [10]
棉子糖 OsDREB1A [62]
葡萄糖 OsDREB1A [62]
果糖 OsDREB1A [62]
海藻糖 OsTPP1、OsTPP2、OsTPS1、OsNAC5 [24, 35, 63]
低温相关氨基酸
脯氨酸 OsCOIN、OsDREB1A、OsMYB2、OsOVP1、OsNAC5、OsMYB4、OsPRP3、OsZFP245、OsMYB3R-2 [23, 24, 42~47, 62, 64]
抗氧化剂
抗坏血酸 OsAPXa [30]
谷胱甘肽 OsTrx23、OsCPK24 [29, 65]
植物激素
ABA OsPHS1、OsPHS 2、OsPHS 3-1、OsPHS 4、OsPDS、OsABA8ox1 [51, 52]
GA SD1、D35 [37]

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1.4 可溶性糖与脯氨酸

水稻在低温逆境下会积累大量的可溶性糖(包括蔗糖、已糖、棉子糖、葡萄糖、果糖和海藻糖)。可溶性糖在低温逆境中可以作为细胞的渗透调节物质,稳定细胞膜和原生质胶体;同时还为其他有机物的合成提供碳骨架和能量[31,32,33]。过量表达海藻糖合成的关键基因OsTPP1、OsTPP2和OsTPS1均能显著提高水稻对低温的耐受性(表1)[34,35]。

水稻在孕穗期,特别是花粉母细胞减数分裂期遇低温时,糖(包括蔗糖、葡萄糖、果糖)在花药中积累,同时伴随着蔗糖分解酶活性降低,单糖转运蛋白表达量下调,从而导致供应到绒毡层和花粉粒的蔗糖不足。而蔗糖是合成淀粉的主要成分,这就造成了花粉不育[10, 36]。Sakata等[37]研究发现,外施蔗糖能极大地提高低温下水稻花粉的育性并提高结实率。这可能的解释是：在低温下,虽然流(从花药到花粉)受阻,但是增加外源的蔗糖,最终仍能增加库(花粉)的蔗糖含量,进而合成淀粉增多,可育花粉增多,结实率提高。

在低温下水稻还会积累大量的脯氨酸。脯氨酸广泛参与渗透调节、碳氮代谢、保护多数酶类物质,避免其变性失活[38]。同时,脯氨酸还具有稳定多聚核糖体、维持蛋白合成的作用[39]。在逆境条件下,脯氨酸还能够清除逆境反应产生的过量氢离子 (H+),维持细胞质中有氧呼吸的最佳pH[40]。另外,脯氨酸通过其疏水基团与蛋白质结合来提高蛋白的亲水性[41]。研究发现,OsCOIN、OsMYB2、OsMYB4、OsMYB3R-2和OsZFP245等基因的过表达植株均表现出脯氨酸含量显著提高和对低温耐受力的增强 (表1)[42,43,44,45,46,47]。

1.5 ABA与GA含量的改变

ABA在低温逆境中扮演着重要角色[48]。一般而言,低温下增加ABA含量可能放大ABA信号,启动下游ABA应答耐低温相关基因表达量改变,从而提高植物对低温的耐受性。然而植物适应逆境与生

长发育是两个相互矛盾的过程：适应逆境就会牺牲生长发育,反之亦然[49]。在持续性低温逆境下,植物需要保持正常生长与适应逆境之间的平衡,以避免逆境信号过度放大[49, 50]。因此在低温逆境下,维持相对低的ABA水平有利于提高水稻对低温的耐受力。Du等[51]研究发现,类胡卜素(ABA合成的前体)的突变体phs1、phs2、phs3-1、phs4与PDS-RNAi 干涉表达的转基因水稻在苗期和生殖生长期水稻内源ABA水平降低,但对低温的耐受性有所提高。过量表达水稻ABA代谢基因OsABA8ox1降低水稻幼苗内源ABA水平,在15℃低温处理下,提高了转基因株系对低温的耐受力;但在4℃条件下,转基因株系27-5提高了对低温的耐受力;株系27-3的OsABA8ox1表达量远比株系27-5的OsABA8ox1表达量高,相应的ABA含量27-3比27-5低很多,但是却表现出对低温敏感[52]。用水稻绒毡层特异性表达启动子OsG6B融合小麦ABA代谢基因TaABA8OH1的转基因水稻减少了花药中ABA含量,提高了花粉的育性和结实率[53]。过量表达OsNAC095提高水稻幼苗内源ABA水平,却使水稻对低温敏感[54]。Oliver等[36]报道水稻敏感品种Doongara与耐低温品种R31在孕穗期经低温处理,内源ABA含量均升高,但是耐低温品种R31内源ABA升高幅度要低得多,加之本身ABA含量低,表现出R31更耐低温。这些结果表明,无论是苗期还是抽穗期,在一定的阈值范围内,维持相对低的ABA水平,有利于提高水稻对低温的耐受性;但是极低内源ABA水平对植物适应低温逆境是极不利的。

外施ABA能提高苗期水稻低温耐受力[55]。这可能是在苗期外施ABA能促使ABA信号放大,促进低温相关基因表达,进而提高水稻对低温的耐受性。但是在孕穗期,外施ABA,导致ABA积累,而大量ABA抑制蔗糖转化酶和单糖转运蛋白基因表达,这就导致大量糖包括蔗糖在花药中积累,不能运输到花粉中,从而导致水稻花粉不育,结实率降低[10, 36, 53]。

孕穗期遇低温,水稻内源的赤霉素(gibberellins, GA)含量降低,GA合成突变体sd1和d35对低温敏感;外施GA处理,能使GA合成突变体在低温下花粉可育性得到恢复,进而结实率得到恢复[37]。

2 耐低温相关基因/QTL鉴定和克隆

2.1 图位克隆且精细定位的QTL

近20年来,以Silewah、Koshihikari、M202、Norin-PL8、东乡野生稻、昆明小白谷、丽江新团黑谷等高海拔或高纬度的耐低温材料作为供体亲本,构建了大量重组自交系(recombinant inbred lines, RIL)、双单倍体系(double haploid, DH)、回交自交系(backcross inbred lines, BIL)和近等基因系(nearly isogenic lines, NIL)等群体,在水稻的12条染色体上定位到250多个与低温相关的QTL[66,67,68,69,70,71,72],但大多数只局限在QTL的初步定位。目前,水稻中精细定位的仅有5个孕穗期耐低温QTL(qLTB3[73]、qCTB7[11]、qCTB8[12]、qCT-3-2[15]和qCTB10-2[74],表3),5个苗期耐低温QTL(qCTS4[75]、qCtss11[76]、qSCT1[77]、qSCT11[77]和qLOP2/qPSR2-1[78],表3),1个苗期和成熟期耐低温QTL(qRC10-2[79])以及1个萌发期耐低温QTL(qLTG-9[80],表3)。克隆且有功能鉴定的耐冷基因仅有7个,包括qLTG3-1、COLD1、qCTS-9、GSTZ2、LTG1、Ctb1和CTB4a(表2)。qLTG3-1编码一个富含甘氨酸(GRP)的保守结构域,第17位氨基酸的差异(L17H)决定在水稻在低温条件下种子萌发的强与弱[81]。COLD1编码一个G蛋白信号调节因子,它能与G蛋白α亚基RGA1互作,促进Ca2+内流并增强G蛋白的GTP酶活性,进而增强水稻的耐寒性[82]。qCTS-9在亲本丽江新团黑谷、黄山占2号及其重组自交系RIL群体的启动子区有一个InDel标记与低温的耐受性显著相关,过表达qCTS-9提高水稻对低温的耐受性[83]。OsGSTZ2第99位的苯丙氨酸(Ile)对提高水稻苗期耐低温起重要作用[84]。LTG1编码酪蛋白激酶Ⅰ,其编码区357位的氨基酸替换(I357K)对于低温环境下水稻的生长速率、抽穗期和产量都有重要的影响[85]。Ctb1编码一个含有F-box结构域的蛋白,它与一个E3泛素连接酶亚基Skp1互作,参与泛素-蛋白酶体途径的低温信号传导[86]。CTB4a编码一个保守的富亮氨酸受体样激酶LRR-RLK(leucine-rich repeat receptor-like kinase),其与ATP合成酶的β亚基AtpB互作,影响ATP合成酶的活性,为低温下水稻灌浆提供能量供应。Zhang等[3]分析了119个水稻品种的单倍型,结果显示CTB4a启动子区域的3个SNP(分别是ATG上游-2536、-2511与-1930)的差异决定不同水稻品种对低温耐受能力的强弱。

Table 2
表2
表2 基于图位克隆的耐低温基因
Table 2 Cold tolerance genes based on map-based cloning technique
基因 LOC号 是否有遗传功能验证 时期 表型鉴定 参考文献
qLTG3-1 LOC_Os03g01320 是 萌发期 萌发率 [81]
COLD1 LOC_Os04g51180 是 苗期 成活率 [82]
qCTS-9 LOC_Os09g24440 是 苗期 成活率 [83]
GSTZ2 LOC_Os12g10730 是 苗期 成活率 [84]
LTG1 LOC_Os02g40860 是 营养生长期 株高、分蘖数、产量 [85]
Ctb1 LOC_Os04g52830 是 孕穗期 结实率 [86]
CTB4a LOC_Os04g04330 是 孕穗期 结实率 [3]

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Table 3
表3
表3 精细定位的耐低温QTL
Table 3 Fine mapped cold tolerance QTL
QTL 候选基因LOC号或区间 是否有遗传功能验证 时期 表型鉴定 参考文献
qLTB3 LOC_Os03g57680
LOC_Os03g59200		 否 孕穗期 结实率 [73]
qCTB7 RI02905-RM21862 否 孕穗期 结实率 [11]
qCTB8 RM5647-PLA61 否 孕穗期 结实率 [12]
qCT-3-2 Chr.3:1770855-2199078 否 孕穗期 结实率 [15]
qCTB10-2 LOC_Os10g11820 LOC_Os10g11730 LOC_Os10g11770 LOC_Os10g11810 否 孕穗期 结实率 [74]
qCTS4 CT245-CT236 否 苗期 成活率 [75]
qCtss11 LOC_Os11g40130 LOC_Os11g40160 否 苗期 成活率 [76]
qSCT1 LOC_Os01g69910 否 苗期 成活率 [77]
qSCT11 LOC_Os11g37720 否 苗期 成活率 [77]
qLOP2/ qPSR2-1 LOC_Os02g45450 否 苗期 成活率 [78]
qRC10-2 LOC_Os10g34790
LOC_Os10g34840		 否 苗期和成熟期 成活率 [79]
qLTG-9 L9-25D-ID-1 否 萌发期 萌发率 [80]

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2.2 全基因组关联分析(genome-wide association study, GWAS)

虽然图位克隆是挖掘新基因的重要手段,但是图位克隆所得基因的遗传背景仅局限于构建遗传群体的双亲及它们的重组后代。利用图位克隆所进行的研究没有考虑到除亲本之外水稻种质资源的遗传多样性,并且局限于东亚的几个代表性的籼稻×粳稻亚种定位群体。GWAS是以自然群体的同一染色体或不同染色体上不同座位的等位基因之间的遗传连锁不平衡为基础,从而对目标性状与遗传标记(或候选基因)的相关性进行分析的方法。利用全基因组范围单核苷酸多态性(single nucleotide polymorphism, SNP)为遗传标记,进行全基因组水平的相关性分析,能够有效发掘影响复杂性状的基因变异和优异等位变异[87,88,89],从而为分子设计育种提供更为广泛的基因资源。

Fujino等[90]收集了日本北海道63个水稻品种,并利用GWAS分析鉴定了6个抽穗期相关的QTL和13个低温萌发相关QTL。Pan等[14]利用174个来源于中国的水稻核心种质进行低温下萌发实验,获得22个低温萌发相关的QTL(图1,表5)。Shakiba等[91]利用水稻种质资源库RDP1中400多个水稻核心种质进行萌发期和生殖生长期耐低温筛选,分析并鉴定到42个苗期耐低温QTL和29个生殖生长期耐低温QTL(图1,表5)。Wang等[7]采用RDP1中295份水稻品种,对三叶期的水稻进行低温处理3天,以成活率为耐低温指标,利用GWAS分析获得了67个分布在水稻11条染色体上的苗期耐低温QTL (图1,表4,表5)。此外,该研究还发现一个候选基因Osryh1在耐低温品种与敏感品种间有一个InDel标记跟低温耐受性相关(图1,表3)。Lv[92]等对529份水稻核心种质在四叶期时进行自然条件下的耐低温处理(5℃~12℃),以电解质渗漏为耐低温指标,获得了132个苗期低温关联位点。此外,该研究中的单倍型关联分析显示,OsMYB2具有明显的籼粳基因型分化,粳稻型OsMYB2的水稻品种具有更强的耐低温能力(图1,表4)。Schl?ppi等[8]利用来源于美国农业部(USDA)的202份水稻微核心种质进行苗期低温处理,通过5种苗期耐低温指标进行GWAS分析,获得48个耐低温相关QTL(图1,表5)。但是目前通过GWAS获得的多数耐低温的QTL区间仍然很大,侯选基因过多,耐低温主效基因仍不明确,还需要结合遗传学及转录组、蛋白组和代谢组等其他手段以获得更精确的定位和候选基因,并验证其遗传功能,为水稻分子育种提供优异的耐低温等位基因。

图1

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图1经图位克隆和GWAS定位的耐低温QTL

绿色代表苗期耐低温位点,红色代表孕穗期耐低温位点,蓝色代表萌发期耐低温位点。
Fig. 1Fine-mapped and GWAS-mapped cold tolerance QTL



Table 4
表4
表4 GWAS预测的侯选耐低温基因
Table 4 Cold tolerance candidate genes predicted by GWAS
QTL 候选基因 LOC号 是否有遗传功能验证 时期 表型鉴定 参考文献
qCTS1-6 OsDREB1F LOC_Os01g73770 是 苗期 成活率 [7, 93]
qCTS3-1 qLTG3-1 LOC_Os03g01320 是 苗期 成活率 [7, 81]
qCTS3-9 Osryh1 LOC_Os03g09140 否 苗期 成活率 [7]
qCTS6-5 OsSPX1 LOC_Os06g40120 是 苗期 成活率 [7, 94]
qCTS7-5 OsFAD8 LOC_Os07g49310 是 苗期 成活率 [7, 95]
qCTS9-1 OsCYL4a LOC_Os09g02270 是 苗期 成活率 [7, 96]
L33 OsMyb2 LOC_Os03g20090 是 苗期 电解质渗漏 [43, 92]
L48 Ctb1 LOC_Os04g52830 是 苗期 电解质渗漏 [6, 86, 92]
L56 OsRAN2 LOC_Os05g49890 是 苗期 电解质渗漏 [92, 97]
L66 OsiSAP8 LOC_Os06g41010 是 苗期 电解质渗漏 [20, 92]
L77 OsLti6a LOC_Os07g44180 是 苗期 电解质渗漏 [59, 92]
L：locus(位点);qCTS：QTL for cold tolerance at seedling stage(苗期耐低温QTL)。
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Table 5
表5
表5 通过GWAS获得的耐低温可重复定位的QTL区段
Table 5 Cold tolerance QTL co-localized by at least two independent GWAS
QTL 染色体 物理位置(Mb) SNP (Mb) 是否有遗传功能验证 时期 参考文献
L13/qLTG1 1 28.260~28.458 28.359 否 苗期、萌发期 [8, 92]
qCTS1-5/qCTGERM1-8 1 41.518~41.786 41.676 否 苗期、萌发期 [6, 7]
L21/qCTGERM2-2 2 6.557~6.624 6.557 否 苗期、萌发期 [6, 7]
L27/qCTS3-6 3 2.851~3.173 2.680 否 苗期 [7, 92]
L31/qCTS3-10 3 8.060~8.248 8.192 否 苗期 [7, 92]
L48/qCTGERM4-5 4 31.247~31.428 31.436 否 苗期 [6, 92]
L52/qCTS5-1 5 1.865~2.058 1.970 否 苗期 [7, 92]
qCTS6-1/qSWTPNCT6-1 6 5.927~6.229 6.194 否 苗期、孕穗期 [6, 7]
L63/qFERCT6-4 6 16.588~16.770 16.659 否 苗期、孕穗期 [6, 92]
L66/qSWTCT6 6 24.708~24.901 24.812 否 苗期、孕穗期 [6, 92]
L79/qCTGERM7-5 7 27.651~27.844 27.749 否 苗期、萌发期 [6, 92]
L72/qCTGERM7-1 7 10.448~10.627 10.525 否 苗期、萌发期 [6, 92]
L74/qFERCT7 7 13.981~14.471 14.316 否 苗期、孕穗期 [6, 92]
qCTS8-3/qCTGERM8-1 8 10.438~10.903 10.506 否 苗期、萌发期 [6, 7]
qCTS9-7/qPGCG9-1 9 6.303~6.503 6.403 否 苗期、萌发期 [7, 8]
L100/qLTS9 9 22.701~22.887 22.701 否 苗期 [8, 92]
L92/qCTS9-5 9 4.304~4.488 4.350 否 苗期 [7, 92]
L102/qCTGERM10-3 10 10.165~10.335 10.257 否 苗期、萌发期 [6, 92]
L107/qCTGERM10-4 10 22.258~22.298 22.298 否 苗期、萌发期 [6, 92]
qCTGERM11-2/qLTSS11-1 11 5.500~5.710 5.590 否 苗期、萌发期 [6, 8]
L112/qCTS11-6 11 19.811~20.004 19.898 否 苗期 [7, 92]
L125/qFERCT12 12 7.118~7.312 7.243 否 苗期、孕穗期 [6, 92]
L131/qLTSS12-1 12 23.082~23.272 23.173 否 苗期 [8, 92]
L：locus (位点);qCTS：QTL for cold tolerance at seedling stage (苗期耐低温QTL);LTG：low temperature tolerance at germination stage (萌发期耐低温);CTGERM：cold tolerance at germination stage (萌发期耐低温);FERCT：cold tolerance associated with spikelet fertility(以结实率为评价标准的耐低温);SWTPNCT：cold tolerance associated with seed weight per panicle(以单穗重量为评价标准的耐低温);SWTCT：cold tolerance associated with seed weight per plant(以单株产量为评价标准的耐低温);PGCG：plumule growth rate after cold germination(低温萌发后胚芽生长速率);LTS：low temperature survival (低温处理后存活率);LTSS：low temperature seedling survivability(苗期低温处理后存活率)。
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3 耐低温胁迫信号传导途径

植物作为不可移动的生物,在长期进化过程中产生了一系列适应逆境（如低温）的机制。植物适应低温的信号转导途径主要包括依赖ABA的信号转导途径和不依赖ABA的信号转导途径。低温信号转导途径主要组分包括核心转录因子和蛋白激酶。核心转录因子如CBF(C-repeat-binding factor),蛋白激酶如SNF1(sucrose nonfermenting 1 protein kinase)和MAPK (mitogen-activated protein kinase)等。低温信号促进植物体离子平衡、水分运输以及代谢和基因表达的重调,进而维持低温环境下离子平衡和细胞稳定[98]。

3.1 ABA依赖途径

植物激素ABA不仅在种子萌发、休眠、气孔开闭等生理过程中起重要作用,而且还对植物应对逆境条件起重要的调节作用,主要表现为ABA能在逆境下促进多数逆境应答基因的表达[99]。ABA信号传导通路由ABA受体PYR/PYL/RCAR (pyrabactin resistance/pyrabactin resistance-like/regulatory component of abscisic acid receptor)、负调控因子2C类蛋白磷酸酶(type 2C protein phosphatase, PP2C)、正调控因子SNF1相关的蛋白激酶2(SNFl related protein kinase 2,SnRK2)和转录因子AREB/ABF等4个核心组分共同组成一个双重负调控系统(PYR/PYL/RCAR┤PP2C┤SnRK2-ABF/AREB),调控ABA应答反应[100,101,102,103]。低温条件下,水稻内源的ABA升高,PYR/PYL/RCAR与ABA结合,并与PP2C互作,抑制PP2C与SnRK2结合,SnRK2便能磷酸化转录因子如AREB、ABF等,磷酸化的转录因子激活ABA响应基因表达,从而提高植物对低温的耐受性[104]。然而在正常条件下,内源ABA含量不变,PP2C与SnRK2互作阻止SnRK2磷酸化下游的底物,从而阻遏ABA应答基因的表达(图1)[104]。水稻中ABA受体OsPYL3和OsPYL9正向调节ABA信号,过表达OsPYL3和OsPYL9均能显著提高水稻耐冷性[105]。除了核心组分PYL-PP2C-SnRK2-ABF之外,ABA信号通路的组分还包括Ca2+、ROS、一氧化氮(NO)、磷脂分子以及其他的激酶如MAPK等[98]。

图2

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图2水稻耐低温胁迫的信号传导途径

水稻应答低温逆境信号分为ABA依赖途径和不依赖ABA途径。低温下水稻内源的ABA升高,与ABA受体PYR/PYL/RCAR和2C类蛋白磷酸酶(PP2C)相结合,SNF1相关蛋白激酶2(SnRK2)磷酸化下游转录因子如AREB/ABF等,进一步激活下游ABA响应低温应答基因表达,从而提高水稻对低温的耐受性[104];然而在常温下,由于PP2C与SnRK2相结合,SnRK2就丧失了磷酸激酶活性,下游底物也就不能被磷酸化,也就没有低温应答反应[104]。OsbHLH002/OsICE1受丝裂原活化蛋白激酶OsMAPK3磷酸化,磷酸化的OsbHLH002/OsICE1激活下游海藻糖磷酸酶基因OsTPP1的表达,水稻中海藻糖含量升高,进而提高水稻对低温的耐受性[110]。转录因子OsDREB1F可能被上游OsbHLH002/OsICE1激活,再激活下游含DRE/CRT低温响应基因 (COR)的表达,参与不依赖ABA的耐低温调控途径;也可能直接参与不依赖ABA的耐低温调控途径;还可能激活下游含AREB/ABF这类转录因子,参与ABA依赖的耐低温信号传导途径[93]。
Fig. 2Rice cold signal transduction pathway



3.2 不依赖ABA途径

CBF/DREB1(C-repeat-binding factor/dehydration- responsive element binding factor)是一类非ABA依赖的低温应答信号途径中重要的转录因子。CBF属于AP2/ERF (APETALA 2/ethylene responsive transcription factors)转录因子家族的一个亚家族,AP2/ ERF家族主要分为AP2、ERF、DREB和RAV(related to VP1/ABI3)4个亚家族[106, 107]。CBF含有AP2保守结构域,在低温条件下,其能结合含有核心元件为CCGAC(又称CRT,C-repeat)的低温响应基因(CORs)启动区,并在转录水平上激活CORs[108]。CBF基因通常受bHLH类转录因子ICE1(inducer of CBF expression 1)调控[108]。因此,该低温调控途径又称为ICE-CBF-COR途径。现有研究证明,在水稻中过量表达DREB1亚家族的OsDREB1D、OsDREB1F均能提高水稻对低温的耐受性[93,109]。丝裂原活化蛋白激酶OsMAPK3磷酸化OsbHLH002/OsICE1,降低OsbHLH002的泛素化水平,进而促进活性OsbHLH002的积累;OsbHLH002能激活下游基因OsTPP1的表达,提高水稻海藻糖含量,进而提高水稻对低温的耐受性(图2)[110]。此外,还有一些CBF基因受ABA诱导,如OsDREB1F等。OsDREB1F既参与不依赖ABA信号传导途径,又参与ABA依赖信号传导途径[93](图1),这就暗示着ABA依赖的信号传导途径与ICE-CBF-COR途径有重叠[111]。

4 结语与展望

未来水稻耐低温育种可以主要集中在以下几个方向：(1)利用基因工程技术过表达耐低温基因或是敲除低温敏感基因。如过量表达OsMyb4[44]、OsNAC5[24]等;敲除低温敏感基因,如OsbZIP52[112];(2)基于分子标记和连锁图谱获得了大量的耐低温相关QTL或基因,而这些QTL遗传标记或基因是基于籼粳之间或栽培稻与野生稻之间差异而获得的,因此,可以利用分子标记辅助选择,选择携带耐低温相关QTL或等位基因的供体亲本,农艺性状优良主栽品种为轮回亲本,经多代回交后将供体耐低温QTL或等位基因转育到受体材料中,在育种群体世代繁殖过程中逐步开展表现型和基因型鉴定,选择耐低温且农艺性状优良的品系;(3)在改良多基因控制的复杂性状时,分子标记辅助选择和分子标记辅助选择育种存在两个方面的缺陷：一是基于双亲的QTL定位结果有时不具有普遍性,遗传群体中的QTL定位结果不能很好地应用于育种群体中;二是耐低温多由多个微效基因控制,不能将这些数量基因位点有效地应用于数量性状改良。全基因组选择(genomic or genome-wide selection,GS)是在高密度分子标记基因型鉴定的情况下,利用覆盖全基因组的SNP标记或等位基因型,以及群体中个体的表型数据,估计每个标记的遗传效应,从而得到基因组估计的育种值(genomic estimated breeding value,GEBV)。在后续的育种群体中,利用每一个标记的估计效应和个体的基因型鉴定数据,预测个体的表型或育种值,然后根据预测表型选择优良后代[113,114,115,116];(4)利用基因组编辑技术改良水稻品种,如将籼稻品种中的COLD1Ind基因型替换为粳稻型的COLD1Jap。

总之,随着对水稻低温胁迫应答机制的深入理解,水稻耐低温育种改良也逐渐成为可能。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。


参考文献 原文顺序
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Abstract Rice (Oryza sativa L.) has supported a greater number of people for a longer period of time than any other crop since it was domesti- cated between 8,000 to 10,000 years ago (Greenland, 1997). At present, rice is the staple food for more people than wheat, and 90 percent of total rice production is grown and consumed in Asia (Evans, 1998). Unlike maize or wheat, less than five percent of total rice production is traded on world markets, mainly within Asia and from Asia to Africa and Europe. Thus the emphasis in all rice economies is on self-sufficiency. In many Asian countries, rice self- sufficiency and political stability are interdependent issues. Wetland or paddy rice production has been sustained over millen- nia and can be considered one of the world's most sustainable and productive farming systems. On an annual basis, irrigated rice is often 100 times more productive than upland rice, over 12 times more pro- ductive than deep-water rice, and five times more productive than rainfed rice (Table 1). Irrigated rice accounts for 55 percent of the global har- vested area and contributes 75 percent of global rice production, which is about 410 million tonnes (M t) of rice per year (Dobermann and Fairhurst, 2000). Rice is now the staple food of 2.7 billion people, almost half the world's population, and is grown by more than half the world's farmers. The enormous productivity of intensified paddy rice systems accounts for the very high population densities and rich cul- tures that have developed alongside the major river systems of Asia. Rice culture is thus the cornerstone of cultural, social, and economic development in Asia. Until the middle part of the last century, yields increased slowly but steadily and crop failure became less frequent as improved methods to control water supply were developed and farmers selected varieties adapted to spe- cific agroecological conditions. Rice was adapted to fit a wide range of growing con- ditions, from the equatorial tropics to the high altitudes of Japan, from the tropical lowlands to the mountain terraces of the Himalayas, and from deep-water swamps to the uplands. This explains why over the past 35 years it has been possible to collect more than 80,000 local varieties now T

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Cold stress adversely affects rice ( Oryza sativa L.) growth and productivity, and has so far determined its geographical distribution. Dissecting cold stress-mediated physiological changes and understanding their genetic causes will facilitate the breeding of rice for cold tolerance. Here, we review recent progress in research on cold stress-mediated physiological traits and metabolites, and indicate their roles in the cold-response network and cold-tolerance evaluation. We also discuss criteria for evaluating cold tolerance and evaluate the scope and shortcomings of each application. Moreover, we summarize research on quantitative trait loci (QTL) related to cold stress at the germination, seedling, and reproductive stages that should provide useful information to accelerate progress in breeding cold-tolerant rice.

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Pan YH, Zhang HL, Zhang DL, Li JJ, Xiong HY, Yu JP, Li JL, Rashid MAR, Li GL, Ma XD, Cao GL, Han LZ, Li ZC . Genetic analysis of cold tolerance at the germination and booting stages in rice by association mapping
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PLoS One, 2012,7(7):e40899.
URLPMID:3397947 [本文引用: 1]
Genome wide transcriptional changes by cold stress, heat stress and oxidative stress in rice seedlings were analyzed. Heat stress resulted in predominant changes in transcripts of heat shock protein and heat shock transcription factor genes, as well as genes associated with synthesis of scavengers of reactive oxygen species and genes that control the level of sugars, metabolites and auxins. Cold stress treatment caused differential expression of transcripts of various transcription factors including desiccation response element binding proteins and different kinases. Transcripts of genes that are part of calcium signaling, reactive oxygen scavenging and diverse metabolic reactions were differentially expressed during cold stress. Oxidative stress induced by hydrogen peroxide treatment, resulted in significant up-regulation in transcript levels of genes related to redox homeostasis and down-regulation of transporter proteins. ROS homeostasis appeared to play central role in response to temperature extremes. The key transcription factors that may underlie the concerted transcriptional changes of specific components in various signal transduction networks involved are highlighted. Co-ordinated expression pattern and promoter architectures based analysis (promoter models and overrepresented transcription factor binding sites) suggested potential regulons involved in stress responses. A considerable overlap was noted at the level of transcription as well as in regulatory modules of differentially expressed genes.

Pamplona R . Advanced lipoxidation end-products
Chem Biol Interact, 2011,192(1-2):14-20.
URLPMID:21238437 [本文引用: 1]
Chemical and nonenzymatic molecular modifications induced by reactive carbonyl species (RCS) generated by peroxidation of membrane phospholipids acyl chains play a dual role as signaling molecules and as mediators of the aging process. Cytotoxic effects of RCS are due to their capacity to react with cellular constituents, forming advanced lipoxidation end-products (ALEs). Reactive carbonyl compounds are stable and can diffuse within or even escape from the cell and attack targets far from the site of formation. The consequent loss of function and structural integrity of modified biomolecules can have a wide range of downstream functional consequences and may be the cause of subsequent cellular dysfunctions and tissue damage.

Bonnecarrère V, Borsani O, Díaz P, Capdevielle F, Blanco P, Monza J . Response to photoxidative stress induced by cold in japonica rice is genotype dependent
Plant Sci, 2011,180(5):726-732.
[本文引用: 1]

Kim SI, Tai TH . Evaluation of seedling cold tolerance in rice cultivars: a comparison of visual ratings and quantitative indicators of physiological changes
Euphytica, 2011,178(3):437-447.
URL [本文引用: 1]
Rice ( Oryza sativa L.) is sensitive to prolonged exposure to low temperature, which at the seedling stage can result in significant chilling injury and mortality. The objective of this study was to quantify physiological and biochemical changes in rice seedlings undergoing chilling stress and compare those changes with visual evaluation of tolerance. Seedlings from the cultivars M-202 (tolerant) and IR50 (sensitive) were subjected to 9°C for 1402days in a controlled environment chamber. Leaf tissues were harvested at various time points for determination of electrolyte leakage, proline, malondialdehyde, ascorbic acid and reduced glutathione. Significant differences between M-202 and IR50 were detected in electrolyte leakage, proline, and ascorbic acid starting at 702days with IR50 exhibiting higher levels of these indicators. Most IR50 seedlings were dead at 1402days. A set of fifty rice accessions including M-202 and IR50 was evaluated at 1002days of cold treatment to examine the correlation of visual ratings with the physiological indices. Visual ratings were most highly correlated with electrolyte leakage and least correlated with proline content. Based on visual ratings and the physiological indices, we identified several cultivars that outperformed M-202 in cold tolerance while IR50 had the lowest tolerance of the cultivars tested.

Xie GS, Kato H, Sasaki K, Imai R . A cold-induced thioredoxin h of rice, OsTrx23, negatively regulates kinase activities of OsMPK3 and OsMPK6 in vitro
FEBS Lett, 2009,583(17):2734-2738.
URLPMID:19665023 [本文引用: 1]
Cytosolic thioredoxins are small conserved proteins that are involved in cellular redox regulation. Here, we report that a major and cold-induced thioredoxin h of rice, OsTrx23, has an inhibitory activity on stress-activated mitogen-activated protein kinases (MAPKs), OsMPK3 and OsMPK6 in vitro. This inhibition effects were redox-dependent and did not involve stable physical interaction. The data suggested a novel mechanism for redox regulation of MAPKs in plants.

Sato Y, Masuta Y, Saito K, Murayama S, Ozawa K . Enhanced chilling tolerance at the booting stage in rice by transgenic overexpression of the ascorbate peroxidase gene,OsAPXa
Plant Cell Rep, 2011,30(3):399-406.
URLPMID:21203887 [本文引用: 1]
Low temperatures during the booting stage reduce rice yields by causing cold-induced male sterility. To determine whether antioxidant capacity affects the ability of rice plants to withstand chilling at the booting stage, we produced transgenic rice plants that overexpress OsAPXa and have increased APX activity. The effect of increased APX activity on the levels of H 2 O 2 and lipid peroxidation were determined by measuring H 2 O 2 levels and malondialdehyde (MDA) contents in spikelets during cold treatments at the booting stage. The levels of H 2 O 2 and the MDA content increased by 1.5-fold and twofold, respectively in WT plants subjected to a 12 C treatment for 6/days. In contrast, transgenic lines showed small changes in H 2 O 2 levels and MDA content under cold stress, and H 2 O 2 levels and MDA content were significantly lower than in WT plants. APX activity showed negative correlations with levels of H 2 O 2 and MDA content, which increased during cold treatment. Cold tolerance at the booting stage in transgenic lines and WT plants was evaluated. Spikelet fertility was significantly higher in transgenic lines than in WT plants after a 12 C treatment for 6/days. These results indicate that higher APX activity enhances H 2 O 2 -scavenging capacity and protects spikelets from lipid peroxidation, thereby increasing spikelet fertility under cold stress.

Nagao M, Minami A, Arakawa K, Fujikawa S, Takezawa D . Rapid degradation of starch in chloroplasts and concomitant accumulation of soluble sugars associated with ABA-induced freezing tolerance in the moss Physcomitrella patens
J Plant Physiol, 2005,162(2):169-180.
URLPMID:15779827 [本文引用: 1]
Abscisic acid (ABA) has been postulated to play a role in the development of freezing tolerance during the cold acclimation process in higher plants, but its role in cold tolerance in tower land plants has not been elucidated. The moss Physcomitrella patens rapidly developed freezing tolerance when its protonemata were grown in a medium containing ABA, with dramatic changes in the LT50 value from -2 degrees C to over -10 degrees C. We examined physiological and morphological alterations in protonema cells caused by ABA treatment to elucidate early cellular events responsible for rapid enhancement of freezing tolerance. Microscopic observations revealed that ABA treatment for 1 day resulted in a dramatic alteration in the appearance of intracellular organelles. ABA-treated cells had slender chloroplasts, with a reduced amount of starch grains, in comparison with those of non-treated cells. The ABA-treated cells also had several segmented vacuoles while many of non-treated cells had one central vacuole. When frozen to -4 degrees C, freezing injury-associated ultrastructural changes such as formation of aparticulate domains and fracture-jump lesions were frequently observed in the plasma membrane of non-treated protonema cells but not in that of ABA-treated cells. The ABA treatment increased the osmotic concentration of the protonema cells, in correlation with accumulation of free soluble sugars. These results suggest that ABA-induced accumulation of soluble sugars, associated with morphological changes in organelles, mitigated freezing-induced structural damage in the plasma membrane, eventually leading to enhancement of freezing tolerance in the protonema cells.

Shao HB, Guo QJ, Chu LY, Zhao XN, Su ZL, Hu YC, Cheng JF . Understanding molecular mechanism of higher plant plasticity under abiotic stress
Colloids Surf B Biointerfaces, 2007,54(1):37-45.
URLPMID:16914294 [本文引用: 1]
Higher plants play the most important role in keeping a stable environment on the earth, which regulate global circumstances in many ways in terms of different levels (molecular, individual, community, and so on), but the nature of the mechanism is gene expression and control temporally and spatially at the molecular level. In persistently changing environment, there are many adverse stress conditions such as cold, drought, salinity and UV-B (280-320/mm), which influence plant growth and crop production greatly. Plants differ from animals in many aspects, but the important may be that plants are more easily influenced by environment than animals. Plants have a series of fine mechanisms for responding to environmental changes, which has been established during their long-period evolution and artificial domestication. These mechanisms are involved in many aspects of anatomy, physiology, biochemistry, genetics, development, evolution and molecular biology, in which the adaptive machinery related to molecular biology is the most important. The elucidation of it will extremely and purposefully promote the sustainable utilization of plant resources and make the best use of its current potential under different scales. This molecular mechanism at least include environmental signal recognition (input), signal transduction (many cascade biochemical reactions are involved in this process), signal output, signal responses and phenotype realization, which is a multi-dimensional network system and contain many levels of gene expression and regulation. We will focus on the molecular adaptive machinery of higher plant plasticity under abiotic stresses.

Ma YY, Zhang YL, Lu J, Shao HB . Roles of plant soluble sugars and their responses to plant cold stress
Afr J Biotechnol, 2009,8(10):2004-2010.
URL [本文引用: 1]
Plants can enhance freezing tolerance by gradual exposure to low temperature known as cold acclimation. Soluble sugars have been confirmed to play an important role during the process. This is based on the fact they are most commonly detected in various species of terrestrial plants that have undergone seasonal cold acclimation. Soluble sugars exert their positive effects to protect plant cells...

Shima S, Matsui H, Tahara S, Imai R . Biochemical characterization of rice trehalose-6-phosphate phosphatases supports distinctive functions of these plant enzymes
FEBS J, 2007,274(5):1192-1201.
URLPMID:17257172 [本文引用: 1]
Substantial levels of trehalose accumulate in bacteria, fungi, and invertebrates, where it serves as a storage carbohydrate or as a protectant against environmental stresses. In higher plants, trehalose is detected at fairly low levels; therefore, a regulatory or signaling function has been proposed for this molecule. In many organisms, trehalose-6-phosphate phosphatase is the enzyme governing the final step of trehalose biosynthesis. Here we report that OsTPP1 and OsTPP2 are the two major trehalose-6-phosphate phosphatase genes expressed in vegetative tissues of rice. Similar to results obtained from our previous OsTPP1 study, complementation analysis of a yeast trehalose-6-phosphate phosphatase mutant and activity measurement of the recombinant protein demonstrated that OsTPP2 encodes a functional trehalose-6-phosphate phosphatase enzyme. OsTPP2 expression is transiently induced in response to chilling and other abiotic stresses. Enzymatic characterization of recombinant OsTPP1 and OsTPP2 revealed stringent substrate specificity for trehalose 6-phosphate and about 10 times lower K m values for trehalose 6-phosphate as compared with trehalose-6-phosphate phosphatase enzymes from microorganisms. OsTPP1 and OsTPP2 also clearly contrasted with microbial enzymes, in that they are generally unstable, almost completely losing activity when subjected to heat treatment at 50 C for 4 min. These characteristics of rice trehalose-6-phosphate phosphatase enzymes are consistent with very low cellular substrate concentration and tightly regulated gene expression. These data also support a plant-specific function of trehalose biosynthesis in response to environmental stresses.

Li HW, Zang BS, Deng XW, Wang XP . Overexpression of the trehalose-6-phosphate synthase gene OsTPS1 enhances abiotic stress tolerance in rice
Planta, 2011,234(5):1007-1018.
[本文引用: 1]

Oliver SN, Dennis ES, Dolferus R . ABA regulates apoplastic sugar transport and is a potential signal for cold-induced pollen sterility in rice
Plant Cell Physiol, 2007,48(9):1319-1330.
URLPMID:17693452 [本文引用: 3]
Abstract Cold temperatures cause pollen sterility and large reductions in grain yield in temperate rice growing regions of the world. Induction of pollen sterility by cold involves a disruption of sugar transport in anthers, caused by the cold-induced repression of the apoplastic sugar transport pathway in the tapetum. Here we demonstrate that the phytohormone ABA is a potential signal for cold-induced pollen sterility (CIPS). Cold treatment of the cold-sensitive cultivar Doongara resulted in increased anther ABA levels. Exogenous ABA treatment at the young microspore stage induced pollen sterility and affected cell wall invertase and monosaccharide transporter gene expression in a way similar to cold treatment. In the cold-tolerant cultivar R31, ABA levels were significantly lower under normal circumstances and remained low after cold treatment. The differences in endogenous ABA levels in Doongara and R31 correlated with differences in expression of the ABA biosynthetic genes encoding zeaxanthin epoxidase (OSZEP1) and 9-cis-epoxycarotenoid dioxygenase (OSNCED2, OSNCED3) in anthers. The expression of three ABA-8-hydroxylase genes (ABA8OX1, 2 and 3) in R31 anthers was higher under control conditions and was regulated differently by cold compared with Doongara. Our results indicate that the cold tolerance phenotype of R31 is correlated with lower endogenous ABA levels and a different regulation of ABA metabolism.

Sakata T, Oda S, Tsunaga Y, Shomura H, Kawagishi-Kobayashi M, Aya K, Saeki K, Endo T, Nagano K, Kojima M, Sakakibara H, Watanabe M, Matsuoka M, Higashitani A . Reduction of gibberellin by low temperature disrupts pollen development in rice
Plant Physiol, 2014,164(4):2011-2019.
URL [本文引用: 2]

Kishor PBK, Sangam S, Amrutha RN, Laxmi PS, Naidu KR, Rao KRSS, Rao S, Reddy KJ, Theriappan P, Sreenivasulu N . Regulation of proline biosynthesis, degradation, uptake and transport in higher plants: Its implications in plant growth and abiotic stress tolerance
Current Sci, 2005,88(3):424-438.
URL [本文引用: 1]
Dramatic accumulation of proline due to increased synthesis and decreased degradation under a variety of stress conditions such as salt, drought and metal has been documented in many plants. Similarly, a decrease in the level of accumulated proline in the rehydrated plants is due to both down regulation of proline biosynthetic pathway enzymes and upregulation of proline degrading enzymes. But, the role of proline during plant development and the molecular basis of the effect of proline accumulation during stress and upon relief of stress are still largely obscure. Here, we summarize the genes governing the proline biosynthetic pathway, its degradation and regulation. Sequentially, we provide an account on transgenics raised so far to engineer the overproduction of osmolyte proline. Also, the identification of specific cellular pathways involved in proline biosynthesis and metabolic changes occurring in transgenic plants developed for proline enhancements are discussed. Further, emphasis is also made on an untouched area of signal transduction of proline biosynthetic pathway.

Kandpal RP, Rao NA . Alterations in the biosynthesis of proteins and nucleic acids in finger millet (Eleucine coracana) seedlings during water stress and the effect of proline on protein biosynthesis
Plant Sci, 1985,40(2):73-79.
[本文引用: 1]

Venekamp JH . Regulation of cytosol acidity in plants under conditions of drought
Physiol Plant, 1989,76(1):112-117.
URL [本文引用: 1]
Abstract In plants under water stress, the activity of photosynthesis declines most. Stimulation of the oxidative respiration and fermentation results in an increase in the amount of related organic acids: citrate, malate and lactate. In spite of some decline in photo-respiratory activity, dehydration may enhance the concentration of related organic acids, glycerate and glycolate. The resulting amount of H + should stimulate NAD(P)H reduction of organic acids by dehydrogenases. Accumulation of proline could be the consequence of such reactions. In the oxidation of glycine, regeneration of NAD(P)H does not liberate H + but NH 4 + . Assimilation of NH 4 + by cytosolic glutamine synthetase (EC 6.3.1.2) results in positively charged glutamine. It is also conceivable that the charge is essential in the final asparagine synthesis by cytosolic asparagine synthetase (EC 6.3.1.1). At low pH the activity of the oxidative respiration declines. In water-stressed plants, maintenance of oxidative respiration will depend on the availability of sufficient amounts of carbohydrates and on adequate removal of excess H + by accumulation of proline and asparagine.

Schobert B, Tschesche H . Unusual solution properties of proline and its interaction with proteins
Biochim Biophys Acta, 1978,541(2):270-277.
URLPMID:667127 [本文引用: 1]
The presence of proline in solutions of the well soluble protein bovine albumin reduces the precipitation of this protein by ethanol and (NH 4 ) 2 SO 4 , presumably by an increased water-binding capacity of the proline-protein solution.

Liu KM, Wang L, Xu YY, Chen N, Ma QB, Li F, Chong K . Overexpression of OsCOIN, a putative cold inducible zinc finger protein, increased tolerance to chilling, salt and drought, and enhanced proline level in rice
Planta, 2007,226(4):1007-1016.
[本文引用: 1]

Yang A, Dai XY, Zhang WH . A R2R3-type MYB gene,OsMYB2, is involved in salt, cold, and dehydration tolerance in rice
J Exp Bot, 2012,63(7):2541-2556.
[本文引用: 1]

Park MR, Yun KY, Mohanty B, Herath V, Xu FY, Wijaya E, Bajic VB, Yun SJ , De Los Reyes BG. Supra-optimal expression of the cold-regulated OsMyb4 transcription factor in transgenic rice changes the complexity of transcriptional network with major effects on stress tolerance and panicle development
Plant Cell Environ, 2010,33(12):2209-2230.
[本文引用: 2]

Vannini C, Locatelli F, Bracale M, Magnani E, Marsoni M, Osnato M, Mattana M, Baldoni E, Coraggio I . Overexpression of the rice Osmyb4 gene increases chilling and freezing tolerance of Arabidopsis thaliana plants
Plant J, 2004,37(1):115-127.
[本文引用: 1]

Ma QB, Dai XY, Xu YY, Guo J, Liu YJ, Chen N, Xiao J, Zhang DJ, Xu ZH, Zhang XS, Chong K . Enhanced tolerance to chilling stress in OsMYB3R-2 transgenic rice is mediated by alteration in cell cycle and ectopic expression of stress genes
Plant Physiol, 2009,150(1):244-256.
[本文引用: 1]

Huang J, Sun SJ, Xu DQ, Yang X, Bao YM, Wang ZF, Tang HJ, Zhang HS . Increased tolerance of rice to cold, drought and oxidative stresses mediated by the overexpression of a gene that encodes the zinc finger protein ZFP245
Biochem Biophys Res Commun, 2009,389(3):556-561.
URLPMID:19751706 [本文引用: 1]
ZFP245 is a cold- and drought-responsive gene that encodes a zinc finger protein in rice. The ZFP245 protein localizes in the nucleus and exhibits trans-activation activity. Transgenic rice plants overexpressing ZFP245 were generated and found to display high tolerance to cold and drought stresses. The transgenic plants did not exhibit growth retardation, but showed growth sensitivity against exogenous abscisic acid, increased free proline levels and elevated expression of rice pyrroline-5-carboxylatesynthetase and proline transporter genes under stress conditions. Overproduction of ZFP245 enhanced the activities of reactive oxygen species-scavenging enzymes under stress conditions and increased the tolerance of rice seedlings to oxidative stress. Our data suggest that ZFP245 may contribute to the tolerance of rice plants to cold and drought stresses by regulating proline levels and reactive oxygen species-scavenging activities, and therefore may be useful for developing transgenic crops with enhanced tolerance to abiotic stress.

Mittler R, Blumwald E . The roles of ROS and ABA in systemic acquired acclimation
Plant Cell, 2015,27(1):64-70.
URLPMID:25604442 [本文引用: 1]
Systemic responses to environmental stimuli are essential for the survival of multicellular organisms. In plants, they are initiated in response to many different signals including pathogens, wounding, and abiotic stresses. Recent studies highlighted the importance of systemic acquired acclimation to abiotic stresses in plants and identified several different signals involved in this response. These included reactive oxygen species (ROS) and calcium waves, hydraulic waves, electric signals, and abscisic acid (ABA). Here, we address the interactions between ROS and ABA at the local and systemic tissues of plants subjected to abiotic stress and attempt to propose a model for the involvement of ROS, ABA, and stomata in systemic signaling leading to systemic acquired acclimation.

Scheres B , Van Der Putten WH. The plant perceptron connects environment to development
Nature, 2017,543(7645):337-345.
URLPMID:28300110 [本文引用: 2]
Plants cope with the environment in a variety of ways, and ecological analyses attempt to capture this through life-history strategies or trait-based categorization. These approaches are limited because they treat the trade-off mechanisms that underlie plant responses as a black box. Approaches that involve the molecular or physiological analysis of plant responses to the environment have elucidated intricate connections between developmental and environmental signals, but in only a few well-studied model species. By considering diversity in the plant response to the environment as the adaptation of an information-processing network, new directions can be found for the study of life-history strategies, trade-offs and evolution in plants.

Zong W, Tang N, Yang J, Peng L, Ma SQ, Xu Y, Li GL, Xiong LZ . Feedback regulation of ABA Signaling and biosynthesis by a bZIP transcription factor targets drought-resistance-related genes
Plant Physiol, 2016,171(4):2810-2825.
URLPMID:27325665 [本文引用: 1]
The OsbZIP23 transcription factor has been characterized for its essential role in drought resistance in rice (Oryza sativa), but the mechanism is unknown. In this study, we first investigated the transcriptional activation of OsbZIP23. A homolog of SnRK2 protein kinase (SAPK2) was found to interact with and phosphorylate OsbZIP23 for its transcriptional activation. SAPK2 also interacted with OsPP2C49, an ABI1 homolog, which deactivated the SAPK2 to inhibit the transcriptional activation activity of OsbZIP23. Next, we performed genome-wide identification of OsbZIP23 targets by immunoprecipitation sequencing and RNA sequencing analyses in the OsbZIP23-overexpression, osbzip23 mutant, and wild-type rice under normal and drought stress conditions. OsbZIP23 directly regulates a large number of reported genes that function in stress response, hormone signaling, and developmental processes. Among these targets, we found that OsbZIP23 could positively regulate OsPP2C49, and overexpression of OsPP2C49 in rice resulted in significantly decreased sensitivity of the abscisic acid (ABA) response and rapid dehydration. Moreover, OsNCED4 (9-cis-epoxycarotenoid dioxygenase4), a key gene in ABA biosynthesis, was also positively regulated by OsbZIP23. Together, our results suggest that OsbZIP23 acts as a central regulator in ABA signaling and biosynthesis, and drought resistance in rice.

Du H, Wu N, Chang Y, Li XH, Xiao JH, Xiong LZ . Carotenoid deficiency impairs ABA and IAA biosynthesis and differentially affects drought and cold tolerance in rice
Plant Mol Biol, 2013,83(4-5):475-488.
URL [本文引用: 1]
Plant responses to abiotic stresses are coordinated by arrays of growth and developmental programs. Phytohormones such as abscisic acid (ABA) and indole-3-acetic acid (IAA) play critical roles in deve

Mega R, Meguro-Maoka A, Endo A, Shimosaka E, Murayama S, Nambara E, Seo M, Kanno Y, Abrams SR, Sato Y . Sustained low abscisic acid levels increase seedling vigor under cold stress in rice (Oryza sativa L.).
Sci Rep, 2015,5:13819.
URLPMID:26350634 [本文引用: 1]
Abstract Stress-induced abscisic acid (ABA) is mainly catabolized by ABA 8'-hydroxylase (ABA8ox), which also strictly regulates endogenous ABA levels. Although three members of the ABA8ox gene family are conserved in rice, it is not clear which stressors induce expression of these genes. Here, we found that OsABA8ox1 was induced by cold stress within 24090009h and that OsABA8ox2 and OsABA8ox3 were not. In contrast, OsABA8ox2 and OsABA8ox3 were ABA-inducible, but OsABA8ox1 was not. OsABA8ox1, OsABA8ox2, and OsABA8ox3 restored germination of a cyp707a1/a2/a3 triple mutant of Arabidopsis to rates comparable to those of the wild type, indicating that OsABA8ox1, OsABA8ox2, and OsABA8ox3 function as ABA-catabolic genes in vivo. Transgenic rice lines overexpressing OsABA8ox1 showed decreased levels of ABA and increased seedling vigor at 1509000900°C. These results indicate that sustained low levels of ABA lead to increased seedling vigor during cold stress. On the other hand, excessively low endogenous ABA levels caused reduced drought and cold tolerance, although some of the transgenic rice lines expressing OsABA8ox1 at moderate levels did not show these harmful effects. Adequate regulation of endogenous ABA levels is thought to be crucial for maintaining seedling vigor under cold stress and for cold and drought tolerance in rice.

Ji XM, Dong BD, Shiran B, Talbot MJ, Edlington JE, Hughes T, White RG, Gubler F, Dolferus R . Control of abscisic acid catabolism and abscisic acid homeostasis is important for reproductive stage stress tolerance in cereals
Plant Physiol, 2011,156(2):647-662.
URL [本文引用: 2]

Huang L, Hong YB, Zhang HJ, Li DY, Song FM . Rice NAC transcription factor ONAC095 plays opposite roles in drought and cold stress tolerance
BMC Plant Biol, 2016,16:203.
URLPMID:5029094 [本文引用: 1]
The NAC (NAM, ATAF and CUC) transcriptional factors constitute a large family with more than 150 members in rice and some of them have been demonstrated to play crucial roles in plant abiotic stress response. Here, we report the characterization of a rice stress-responsive NAC gene,ONAC095, and the exploration of its function in drought and cold stress tolerance. Expression ofONAC095was up-regulated by drought stress and abscisic acid (ABA) but down-regulated by cold stress. ONAC095 protein had transactivation activity and the C2 domain in C-terminal was found to be critical for transactivation activity. Transgenic rice lines with overexpression ofONAC095(ONAC095-OE) and dominant chimeric repressor-mediated suppression ofONAC095(ONAC095-SRDX) were generated. The ONAC095-OE plants showed comparable phenotype to wild type under drought and cold stress conditions. However, the ONAC095-SRDX plants displayed an improved drought tolerance but exhibited an attenuated cold tolerance. The ONAC095-SRDX plants had decreased water loss rate, increased proline and soluble sugar contents, and up-regulated expression of drought-responsive genes under drought condition, whereas the ONAC095-SRDX plants accumulated excess reactive oxygen species, increased malondialdehyde content and down-regulated expression of cold-responsive genes under cold condition. Furthermore, ONAC095-SRDX plants showed an increased ABA sensitivity, contained an elevated ABA level, and displayed altered expression of ABA biosynthetic and metabolic genes as well as some ABA signaling-related genes. Functional analyses through dominant chimeric repressor-mediated suppression ofONAC095demonstrate that ONAC095 plays opposite roles in drought and cold stress tolerance, acting as a negative regulator of drought response but as a positive regulator of cold response in rice. The online version of this article (doi:10.1186/s12870-016-0897-y) contains supplementary material, which is available to authorized users.

Qi G . Identification of cold tolerance at seedling stage among main rice cultivars in Heilongjiang and regulation of abscisic acid[Dissertation]
Harbin: Northeast Agricultural University, 2008.
URL [本文引用: 1]
水稻为喜温植物,其生长发育各个时期都可能受到低温伤害,而苗期耐冷性与其它时期耐冷性密切相关,是种质资源耐冷性鉴定的主要依据,同时也是研究水稻耐冷性机理的重要内容,因此受到人们越来越多的重视。黑龙江省为高纬度稻作区,水稻资源丰富,但冷害较为严重,对黑龙江省主栽水稻品种苗期耐冷性的研究将为水稻遗传育种和耐冷机制的阐明提供理论和试验依据。 本研究以116份黑龙江省主栽水稻品种为材料,根据死苗率、叶片卷曲度、叶片萎蔫度、叶片失绿度4种苗期耐冷性鉴定指标,对供试116份品种进行苗期耐冷性级别鉴定,从中筛选出耐冷品种和冷敏感品种进行外源ABA和低温胁迫,在不同处理时期测定了叶片SOD酶活性、POD酶活性、MDA含量等生理指标,最后对外源ABA和低温诱导两品种叶片细胞H2O2积累进行亚细胞定位,主要结果如下: 1.以日本晴(强耐冷型)和kasalath(冷敏感型)为参照品种,以死苗率、叶片卷曲度、叶片萎蔫度、叶片失绿度为评价指标将116份黑龙江省主栽水稻品种聚为耐冷、中间型和冷敏感三大类群。 2.以苗期强耐冷品种东农418和苗期冷敏感品种松粳9号为材料,喷施ABA合成抑制剂钨酸钠(Tungstate)及ABA后进行低温胁迫,连续测定了处理后不同时期的生理指标。外施ABA减缓了叶片SOD活性的降低,并可减少POD活性和MDA含量升高的幅度,而钨酸钠处理则加剧了SOD活性降低及POD活性和MDA含量的升高,说明ABA处理可增强水稻幼苗耐冷性。ABA对耐冷性的调节作用因品种而异,对耐冷性品种东农418更明显。 3. DAB染色的组织化学定位初步揭示低温和外源ABA诱导叶片H2O2主要积累在叶脉附近,其中ABA引起H2O2迅速积累,而冷胁迫诱导的H2O2积累速度稍慢。电镜技术结合CeC13染色的细胞化学定位表明低温和外源ABA诱导的H2O2在水稻叶肉细胞中主要积累于朝向细胞间隙的细胞壁上,而细胞器如叶绿体、线粒体、细胞核、内质网和液泡中并未观察到H2O2积累。
齐光 . 黑龙江省主栽水稻品种苗期耐冷鉴定及ABA对苗期耐冷的调节作用[学位论文]
哈尔滨: 东北农业大学, 2008.
URL [本文引用: 1]
水稻为喜温植物,其生长发育各个时期都可能受到低温伤害,而苗期耐冷性与其它时期耐冷性密切相关,是种质资源耐冷性鉴定的主要依据,同时也是研究水稻耐冷性机理的重要内容,因此受到人们越来越多的重视。黑龙江省为高纬度稻作区,水稻资源丰富,但冷害较为严重,对黑龙江省主栽水稻品种苗期耐冷性的研究将为水稻遗传育种和耐冷机制的阐明提供理论和试验依据。 本研究以116份黑龙江省主栽水稻品种为材料,根据死苗率、叶片卷曲度、叶片萎蔫度、叶片失绿度4种苗期耐冷性鉴定指标,对供试116份品种进行苗期耐冷性级别鉴定,从中筛选出耐冷品种和冷敏感品种进行外源ABA和低温胁迫,在不同处理时期测定了叶片SOD酶活性、POD酶活性、MDA含量等生理指标,最后对外源ABA和低温诱导两品种叶片细胞H2O2积累进行亚细胞定位,主要结果如下: 1.以日本晴(强耐冷型)和kasalath(冷敏感型)为参照品种,以死苗率、叶片卷曲度、叶片萎蔫度、叶片失绿度为评价指标将116份黑龙江省主栽水稻品种聚为耐冷、中间型和冷敏感三大类群。 2.以苗期强耐冷品种东农418和苗期冷敏感品种松粳9号为材料,喷施ABA合成抑制剂钨酸钠(Tungstate)及ABA后进行低温胁迫,连续测定了处理后不同时期的生理指标。外施ABA减缓了叶片SOD活性的降低,并可减少POD活性和MDA含量升高的幅度,而钨酸钠处理则加剧了SOD活性降低及POD活性和MDA含量的升高,说明ABA处理可增强水稻幼苗耐冷性。ABA对耐冷性的调节作用因品种而异,对耐冷性品种东农418更明显。 3. DAB染色的组织化学定位初步揭示低温和外源ABA诱导叶片H2O2主要积累在叶脉附近,其中ABA引起H2O2迅速积累,而冷胁迫诱导的H2O2积累速度稍慢。电镜技术结合CeC13染色的细胞化学定位表明低温和外源ABA诱导的H2O2在水稻叶肉细胞中主要积累于朝向细胞间隙的细胞壁上,而细胞器如叶绿体、线粒体、细胞核、内质网和液泡中并未观察到H2O2积累。

Lee SC, Huh KW, An K, An G, Kim SR . Ectopic expression of a cold-inducible transcription factor, CBF1/DREB1b, in transgenic rice (
Oryza sativa L.). Mol Cells, 2004,18(1):107-114.
URLPMID:15359131
Abstract The gene encoding C-repeat/dehydration-responsive element binding factor 1 (CBF1/DREB1b) of Arabidopsis was introduced into rice (Oryza sativa L.) under the control of the maize ubiquitin promoter. Its incorporation and expression in transgenic rice plants were confirmed by DNA and RNA gel-blot analyses. Cold tolerance in the transgenics was not significantly different from that of the wild-type plants, as determined by ion leakage, chlorophyll fluorescence, and survival rates. However, the cold-responsive genes lip5, lip9, and OsDhn1 were up-regulated in the transgenic plants, suggesting that the cold signal transduction pathway involving CBF1 is partially conserved in this cold-labile plant.

Joo J, Lee YH, Kim YK, Nahm BH, Song SI . Abiotic stress responsive rice ASR1 and ASR3 exhibit different tissue-dependent sugar and hormone-sensitivities.
Mol Cells, 2013,35(5):421-435.


Saijo Y, Hata S, Kyozuka J, Shimamoto K, Izui K . Over-expression of a single Ca 2+-dependent protein kinase confers both cold and salt/drought tolerance on rice plants
Plant J, 2000,23(3):319-327.
URL

Morsy MR, Almutairi AM, Gibbons J, Yun SJ , De Los Reyes BG. The OsLti6 genes encoding low-molecular- weight membrane proteins are differentially expressed in rice cultivars with contrasting sensitivity to low temperature.
Gene, 2005,344:171-180.


Xie GS, Kato H, Imai R . Biochemical identification of the OsMKK6-OsMPK3 signalling pathway for chilling stress tolerance in rice
Biochem J, 2012,443(1):95-102.
URLPMID:22248149
Abstract MAPK (mitogen-activated protein kinase) pathways have been implicated in stress signalling in plants. In the present study, we performed yeast two-hybrid screening to identify partner MAPKs for OsMKK (Oryza sativa MAPK kinase) 6, a rice MAPK kinase, and revealed specific interactions of OsMKK6 with OsMPK3 and OsMPK6. OsMPK3 and OsMPK6 each co-immunoprecipitated OsMKK6, and both were directly phosphorylated by OsMKK6 in vitro. An MBP (myelin basic protein) kinase assay of the immunoprecipitation complex indicated that OsMPK3 and OsMPK6 were activated in response to a moderately low temperature (12 C), but not a severely low temperature (4 C) in rice seedlings. A constitutively active form of OsMKK6, OsMKK6DD, showed elevated phosphorylation activity against OsMPK3 and OsMPK6 in vitro. OsMPK3, but not OsMPK6, was constitutively activated in transgenic plants overexpressing OsMKK6DD, indicating that OsMPK3 is an in vivo target of OsMKK6. Enhanced chilling tolerance was observed in the transgenic plants overexpressing OsMKK6DD. Taken together, our data suggest that OsMKK6 and OsMPK3 constitute a moderately low-temperature signalling pathway and regulate cold stress tolerance in rice.

Nakashima K, Tran LS, Van Nguyen D, Fujita M, Maruyama K, Todaka D, Ito Y, Hayashi N, Shinozaki K, Yamaguchi-Shinozaki K . Functional analysis of a NAC-type transcription factor OsNAC6 involved in abiotic and biotic stress-responsive gene expression in rice
Plant J, 2007,51(4):617-630.
URLPMID:17587305
The OsNAC6 gene is a member of the NAC transcription factor gene family in rice. Expression of OsNAC6 is induced by abiotic stresses, including cold, drought and high salinity. OsNAC6 gene expression is also induced by wounding and blast disease. A transactivation assay using a yeast system demonstrated that OsNAC6 functions as a transcriptional activator, and transient localization studies with OsNAC6-sGFP fusion protein revealed its nuclear localization. Transgenic rice plants over-expressing OsNAC6 constitutively exhibited growth retardation and low reproductive yields. These transgenic rice plants showed an improved tolerance to dehydration and high-salt stresses, and also exhibited increased tolerance to blast disease. By utilizing stress-inducible promoters, such as the OsNAC6 promoter, it is hoped that stress-inducible over-expression of OsNAC6 in rice can improve stress tolerance by suppressing the negative effects of OsNAC6 on growth under normal growth conditions. The results of microarray analysis revealed that many genes that are inducible by abiotic and biotic stresses were upregulated in rice plants over-expressing OsNAC6. A transient transactivation assay showed that OsNAC6 activates the expression of at least two genes, including a gene encoding peroxidase. Collectively, these results indicate that OsNAC6 functions as a transcriptional activator in response to abiotic and biotic stresses in plants. We conclude that OsNAC6 may serve as a useful biotechnological tool for the improvement of stress tolerance in various kinds of plants.

Ito Y, Katsura K, Maruyama K, Taji T, Kobayashi M, Seki M, Shinozaki K, Yamaguchi-Shinozaki K . Functional analysis of rice DREB1/CBF-type transcription factors involved in cold-responsive gene expression in transgenic rice
Plant Cell Physiol, 2006,47(1):141-153.
URLPMID:16284406
Abstract The transcription factors dehydration-responsive element-binding protein 1s (DREB1s)/C-repeat-binding factors (CBFs) specifically interact with the DRE/CRT cis-acting element and control the expression of many stress-inducible genes in Arabidopsis. The genes for DREB1 orthologs, OsDREB1A and OsDREB1B from rice, are induced by cold stress, and overexpression of DREB1 or OsDREB1 induced strong expression of stress-responsive genes in transgenic Arabidopsis plants, resulting in increased tolerance to high-salt and freezing stresses. In this study, we generated transgenic rice plants overexpressing the OsDREB1 or DREB1 genes. These transgenic rice plants showed not only growth retardation under normal growth conditions but also improved tolerance to drought, high-salt and low-temperature stresses like the transgenic Arabidopsis plants overexpressing OsDREB1 or DREB1. We also detected elevated contents of osmoprotectants such as free proline and various soluble sugars in the transgenic rice as in the transgenic Arabidopsis plants. We identified target stress-inducible genes of OsDREB1A in the transgenic rice using microarray and RNA gel blot analyses. These genes encode proteins that are thought to function in stress tolerance in the plants. These results indicate that the DREB1/CBF cold-responsive pathway is conserved in rice and the DREB1-type genes are quite useful for improvement of stress tolerance to environmental stresses in various kinds of transgenic plants including rice.

Ge LF, Chao DY, Shi M, Zhu MZ, Gao JP, Lin HX . Overexpression of the trehalose-6-phosphate phosphatase gene OsTPP1 confers stress tolerance in rice and results in the activation of stress responsive genes.
Planta, 2008,228(1):191-201.


Gothandam KM, Nalini E, Karthikeyan S, Shin JS . OsPRP3, a flower specific proline-rich protein of rice, determines extracellular matrix structure of floral organs and its overexpression confers cold-tolerance
Plant Mol Biol, 2010,72(1-2):125-135.


Liu Y, Xu CJ, Zhu YF, Zhang LN, Chen TY, Zhou F, Chen H, Lin YJ . The calcium-dependent kinase OsCPK24 functions in cold stress responses in rice
J Integr Plant Biol, 2018,60(2):173-188, doi: 10.1111/ jipb.12614.
URLPMID:29193704
Summary Calcium-dependent protein kinases (CPKs) are serine/threonine protein kinases that function in plant stress responses. Although CPKs are recognized as key messengers in signal transduction, the specific roles of CPKs and the molecular mechanisms underlying their activity remain largely unknown. Here, we characterized the function of OsCPK24, a cytosol-localized calcium-dependent protein kinase in rice. OsCPK24 was universally and highly expressed in rice plants and was induced by cold treatment. While OsCPK24 knockdown plants exhibited increased sensitivity to cold compared to wild type, OsCPK24 -overexpressing plants exhibited increased cold tolerance. Plants overexpressing OsCPK24 exhibited increased accumulation of proline (an osmoprotectant) and glutathione (an antioxidant) and maintained a higher GSH/GSSG (reduced glutathione to oxidized glutathione) ratio during cold stress compared to wild type. In addition to these effects in responses to cold stress, we found that the kinase activity of OsCPK24 varied under different calcium concentrations. Further, OsCPK24 phosphorylated OsGrx10, a glutathione-dependent thioltransferase, at rates modulated by changes in calcium concentration. Together, our results suggest that OsCPK24 functions as a positive regulator of cold stress tolerance in rice, a process mediated by calcium signaling and involving phosphorylation and the inhibition of OsGrx10 to sustain higher glutathione levels.

Liu FX, Sun QC, Tan LB, Fu YC, Li DJ, Wang XK . Identification and mapping of quantitative trait loci controlling cold-tolerance of Chinese common wild rice (O. rufipogon Griff.) at booting to flowering stages.
Chin Sci Bull, 2003,48(19):2068-2071.
[本文引用: 1]

Andaya V, Mackill D . QTLs conferring cold tolerance at the booting stage of rice using recombinant inbred lines from a
japonica × indica cross. Theor Appl Genet, 2003,106(6):1084-1090.
URLPMID:12671757 [本文引用: 1]
Low temperature stress is common for rice grown in temperate regions and at high elevations in the tropics. The most senstive stage to this stress is booting, about 11 days before heading. Japonica cultivars are known to be more tolerant than indicas. We constructed a genetic map using 191 recombinant inbred lines derived from a cross between a temperate japonica , M-202, and a tropical indica , IR50, in order to locate quantitative trait loci (QTLs) conferring cold tolerance. The map with a total length of 1,276.8/cM and an average density of one marker every 7.1/cM was developed from 181 loci produced by 175 microsatellite markers. Cold tolerance was measured as the degree of spikelet sterility of treated plants at a 12 C temperature for 5 days in the growth chamber. QTLs on chromosomes 1, 2, 3, 5, 6, 7, 9 and 12 were identified to confer cold tolerance at the booting stage. The QTL contribution to the phenotypic variation ranged from 11 to 17%. The two QTLs with the highest contribution to variation, designated qCTB2a and qCTB3 , were derived from the tolerant parent, M-202, each explaining approximately 17% of the phenotypic variance. Two of the eight QTLs for cold tolerance were contributed by IR50.

Xu LM, Zhou L, Zeng YW, Wang FM, Zhang HL, Shen SQ, Li ZC . Identification and mapping of quantitative trait loci for cold tolerance at the booting stage in a
japonica rice near-isogenic line. Plant Sci, 2008,174(3):340-347.
[本文引用: 1]

Kuroki M, Saito K, Matsuba S, Yokogami N, Shimizu H, Ando I, Sato Y . Quantitative trait locus analysis for cold tolerance at the booting stage in a rice cultivar, Hatsushizuku
Jpn Agr Res Q, 2009,43(2):115-121.
URL [本文引用: 1]

Mori M, Onishi K, Tokizono Y, Shinada H, Yoshimura T, Numao Y, Miura H, Sato T . Detection of a novel quantitative trait locus for cold tolerance at the booting stage derived from a tropical japonica rice variety Silewah.
Breed Sci, 2011,61(1):61-68.
[本文引用: 1]

Shirasawa S, Endo T, Nakagomi K, Yamaguchi M, Nishio T . Delimitation of a QTL region controlling cold tolerance at booting stage of a cultivar, ‘Lijiangxintuanheigu’, in rice,
Oryza sativa L. Theor Appl Genet, 2012,124(5):937-946.
URLPMID:22113591 [本文引用: 1]
Low temperature at the booting stage of rice causes male sterility resulting in severe yield loss. Cold tolerance has long been an important objective in rice breeding. We identified a quantitative trait locus (QTL) for cold tolerance on the long arm of chromosome 3 from the cold-tolerant breeding line ‘Ukei 840’ by using F 2 and BC 1 F 2 populations from crosses between ‘Ukei 840’ and ‘Hitomebore’. The cold tolerance of ‘Ukei 840’ is derived from the Chinese cultivar ‘Lijiangxintuanheigu’. The effect of this QTL on cold tolerance was confirmed by developing ‘Hitomebore’ chromosome segment substitution lines having ‘Lijiangxintuanheigu’ alleles on chromosome 3. By producing recombinants in chromosome 3, the QTL region for cold tolerance was delimited to the region of about 1.2-Mb region between RM3719 and RM7000. All lines heterozygous for the QTL showed seed fertilities as low as that of ‘Hitomebore’, suggesting that the ‘Lijiangxintuanheigu’ allele for cold tolerance in the QTL region is recessive. Determination of a 1.2-Mb nucleotide sequence of ‘Ukei 840’ and comparison with the published genomic sequence of ‘Nipponbare’ showed 254 SNPs, of which 11 were in coding regions of genes, seven in five genes being non-synonymous. SNPs were detected in the 5-kb upstream regions of 89 genes, but no differences of gene expression levels were detected between alleles of these genes. Although further delimitation is required to identify the gene responsible for cold tolerance of ‘Lijiangxintuanheigu’, SNP markers developed here will be useful for marker-assisted selection in a breeding program using ‘Lijiangxintuanheigu’ as a donor of cold tolerance.

Biswas PS, Khatun H, Das N, Sarker MM, Anisuzzaman M . Mapping and validation of QTLs for cold tolerance at seedling stage in rice from an indica cultivar Habiganj Boro VI (Hbj.BVI).
3 Biotech, 2017,7(6):359.
[本文引用: 1]

Endo T, Chiba B, Wagatsuma K, Saeki K, Ando T, Shomura A, Mizubayashi T, Ueda T, Yamamoto T, Nishio T . Detection of QTLs for cold tolerance of rice cultivar 'Kuchum' and effect of QTL pyramiding
Theor Appl Genet, 2016,129(3):631-640.
URLPMID:26747044 [本文引用: 1]
Abstract KEY MESSAGE: A QTL for cold tolerance at the booting stage of rice cultivar 'Kuchum' was detected and delimited into a 1.36 Mb region, and a cold-tolerant line was developed by QTL pyramiding. Low temperature in summer causes pollen sterility in rice, resulting in a serious loss of yield. The second most widely grown rice cultivar in Japan, 'Hitomebore', has been developed as a cultivar highly tolerant to low temperature at the booting stage. However, even 'Hitomebore' exhibits sterility at a temperature lower than 18.5 C. Further improvement of cold tolerance of rice is required. In the present study, QTLs for cold tolerance in a Bhutanese rice variety, 'Kuchum', were analyzed using backcrossed progenies and a major QTL, named qCT-4, was detected on chromosome 4. Evaluating cold tolerance of seven types of near isogenic lines having 'Kuchum' alleles around qCT-4 with a 'Hitomebore' genetic background, qCT-4 was delimited to a region of ca. 1.36 Mb between DNA markers 9_1 and 10_13. Homozygous 'Kuchum' alleles at qCT-4 showed an effect of increasing seed fertility by ca. 10 % under cold-water treatment. Near isogenic lines of 'Hitomebore' having 'Silewah' alleles of Ctb1 and Ctb2 and a 'Hokkai PL9' allele of qCTB8 did not exhibit higher cold tolerance than that of 'Hitomebore'. On the other hand, a qLTB3 allele derived from a Chinese cultivar 'Lijiangxintuanheigu' increased cold tolerance of 'Hitomebore', and pyramiding of the qCT-4 allele and the qLTB3 allele further increased seed fertility under cold-water treatment. Since NILs of 'Hitomebore' with the 'Kuchum' allele of qCT-4 were highly similar to 'Hitomebore' in other agronomic traits, the qCT-4 allele is considered to be useful for developing a cold-tolerant cultivar.

Li JL, Pan YH, Guo HF, Zhou L, Yang SM, Zhang YZ, Yang JZ, Zhang HL, Li JJ, Zeng YW, Li ZC . Fine mapping of QTL qCTB10-2 that confers cold tolerance at the booting stage in rice.
Theor Appl Genet, 2018,131(1):157-166 .
[本文引用: 1]

Andaya VC, Tai TH . Fine mapping of the qCTS4 locus associated with seedling cold tolerance in rice(Oryza sativa L.).
Mol Breed, 2007,20(4):349-358.
[本文引用: 1]

Koseki M, Kitazawa N, Yonebayashi S, Maehara Y, Wang ZX, Minobe Y . Identification and fine mapping of a major quantitative trait locus originating from wild rice, controlling cold tolerance at the seedling stage
Mol Genet Genomics, 2010,284(1):45-54.
URLPMID:20526617 [本文引用: 1]
Cold tolerance at the seedling stage (CTSS) is an important trait affecting stable rice production in temperate climates and areas of high elevation. In this study, 331 single nucleotide polymorphism (SNP) markers were developed and used along with phenotypic evaluation to identify quantitative trait loci (QTLs) associated with CTSS from a mapping population of 184 F 2 plants derived from a cold tolerant wild rice, W1943 ( Oryza rufipogon ), and a sensitive indica cultivar, Guang-lu-ai 4 (GLA4). Three QTLs were detected on chromosomes 3, 10 and 11. A major locus, qCtss11 ( Q TL for c old t olerance at s eedling s tage), was located on the long arm of chromosome 11 explaining about 40% of the phenotypic variation. Introduction of the W1943 allele of qCtss11 to the GLA4 genetic background increased CTSS. Based on the phenotypic and genotypic assessment of advanced backcross progenies, qCtss11 was dissected as a single Mendelian factor. A high-resolution genetic map was constructed using 23 markers across the qCtss11 locus. As a result, qCtss11 was fine mapped to a 60-kb candidate region defined by marker AK24 and GP0030 on chromosome 11, in which six genes were annotated. Expression and resequence analyses of the six candidates supported the hypothesis that Os11g0615600 and/or Os11g0615900 are causal gene(s) of the CTSS.

Kim SM, Suh JP, Lee CK, Lee JH, Kim YG, Jena KK . QTL mapping and development of candidate gene- derived DNA markers associated with seedling cold tolerance in rice (
Oryza sativa L.). Mol Genet Genomics, 2014,289(3):333-343.
URLPMID:24464311 [本文引用: 2]
Cold stress at the seedling stage is a major threat to rice production. Cold tolerance is controlled by complex genetic factors. We used an F 7 recombinant inbred line (RIL) population of 123 individuals derived from a cross of the cold-tolerant japonica cultivar Jinbu and the cold-susceptible indica cultivar BR29 for QTL mapping. Phenotypic evaluation of the parents and RILs in an 18/8 C (day/night) cold stress regime revealed continuous variation for cold tolerance. Six QTLs including two on chromosome 1 and one each on chromosomes 2, 4, 10, and 11 for seedling cold tolerance were identified with phenotypic variation ( R 2 ) ranging from 6.1 to 16.5/%. The QTL combinations ( qSCT1 and qSCT11 ) were detected in all stable cold-tolerant RIL groups, which explained the critical threshold of 27.1/% for the R 2 value determining cold tolerance at the seedling stage. Two QTLs ( qSCT1 and qSCT11 ) on chromosomes 1 and 11, respectively, were fine mapped. The markers In1-c3, derived from the open reading frame (ORF) LOC_Os01g69910 encoding calmodulin-binding transcription activator ( CAMTA ), and In11-d1, derived from ORF LOC_Os11g37720 ( Duf6 gene), co-segregated with seedling cold tolerance. The result may provide useful information on seedling cold tolerance mechanism and provide DNA markers for a marker-assisted breeding program to improve seedling cold tolerance in indica rice varieties.

Xiao N, Huang WN, Li AH, Gao Y, Li YH, Pan CH, Ji HJ, Zhang XX, Dai Y, Dai ZY, Chen JM . Fine mapping of the qLOP2 and qPSR2-1 loci associated with chilling stress tolerance of wild rice seedlings.
Theor Appl Genet, 2015,128(1):173-185.
[本文引用: 1]

Xiao N, Huang WN, Zhang XX, Gao Y, Li AH, Dai Y, Yu L, Liu GQ, Pan CH, Li YH, Dai ZY, Chen JM . Fine mapping of qRC10-2, a quantitative trait locus for cold tolerance of rice roots at seedling and mature stages.
PLoS One, 2014,9(5):e96046.
[本文引用: 1]

Li LF, Liu X, Xie K, Wang YH, Liu F, Lin QY, Wang WY, Yang CY, Lu BY, Liu SJ, Chen LM, Jiang L, Wan JM . qLTG-9, a stable quantitative trait locus for low- temperature germination in rice (Oryza sativa L.)
Theor Appl Genet, 2013,126(9):2313-2322.
[本文引用: 1]

Fujino K, Sekiguchi H, Matsuda Y, Sugimoto K, Ono K, Yano M . Molecular identification of a major quantitative trait locus,qLTG3-1, controlling low-temperature germinability in rice.
Proc Natl Acad Sci USA, 2008,105(34):12623-12628.
[本文引用: 1]

Ma Y, Dai XY, Xu YY, Luo W, Zheng XM, Zeng DL, Pan YJ, Lin XL, Liu HH, Zhang DJ, Xiao J, Guo XY, Xu SJ, Niu YD, Jin JB, Zhang H, Xu X, Li LG, Wang W, Qian Q, Ge S, Chong K . COLD1 confers chilling tolerance in rice
Cell, 2015,160(6):1209-1221.
[本文引用: 1]

Zhao JL, Zhang SH, Dong JF, Yang TF, Mao XX, Liu Q, Wang XF, Liu B . A novel functional gene associated with cold tolerance at the seedling stage in rice
Plant Biotechnol J, 2017,15(9):1141-1148.
URLPMID:28173633 [本文引用: 1]
Summary Identification and cloning of cold???tolerant genes that can stably express under different cold environments are crucial for molecular rice breeding for cold tolerance. In the previous study, we identified a cold???tolerant QTL at the seedling stage, qCTS???9 which could be detected under different cold environments using a recombinant inbred line (RIL) population derived from a cold???tolerant variety Lijiangxintuanheigu (LTH) and a cold???sensitive variety Shanhuangzhan 2 (SHZ???2). In this study, eight candidate genes within the qCTS???9 interval were identified through integrated analysis of QTL mapping with genomewide differential expression profiling of LTH. The qRT???PCR assay showed that only Os09g0410300 exhibited different expression patterns between LTH and SHZ???2 during cold stress, and significantly positive correlation was found between cold induction of Os09g0410300 and seedling cold tolerance in the RI lines. Five SNPs and one InDel in the promoters of Os09g0410300 were detected between LTH and SHZ???2, and the InDel marker ID410300 designed based on the insertion???deletion polymorphism in the promoter was significantly associated with seedling cold tolerance in RIL population. Further, Os09g0410300 over???expression plants exhibited enhanced cold tolerance at the seedling stage compared with the wild???type plants. Thus, our results suggest that Os09g0410300 is the functional gene underlying qCTS???9. To our knowledge, it is a novel gene contributed to enhance cold tolerance at the seedling stage in rice. Identification of the functional gene underlying qCTS???9 and development of the gene???specific marker will facilitate molecular breeding for cold tolerance at the seedling stage in rice through transgenic approach and marker???assisted selection (MAS).

Kim SI, Andaya VC, Tai TH . Cold sensitivity in rice (Oryza sativa L.) is strongly correlated with a naturally occurring I99V mutation in the multifunctional glutathione transferase isoenzyme GSTZ2.
Biochem J, 2011,435(2):373-380.
[本文引用: 1]

Lu GW, Wu FQ, Wu WX, Wang HJ, Zheng XM, Zhang YH, Chen XL, Zhou KN, Jin MN, Cheng ZJ, Li XY, Jiang L, Wang HY, Wan JM . Rice LTG1 is involved in adaptive growth and fitness under low ambient temperature.
Plant J, 2014,78(3):468-480.
[本文引用: 1]

Saito K, Hayano-Saito Y, Kuroki M, Sato Y . Map-based cloning of the rice cold tolerance gene
Ctb1. Plant Sci, 2010,179(1-2):97-102.
URL [本文引用: 1]
Low temperatures during the booting stage reduce rice yields by causing cold-induced male sterility. We previously mapped a quantitative trait locus for cold tolerance, Ctb1, to a 56-kb region containing 7 putative genes. In this study we further mapped Ctb1 to a 17-kb region containing two genes that encode an F-box protein and a ser/thr protein kinase. The F-box protein gene was preferentially expressed in young panicles, while the ser/thr protein kinase gene was expressed in leaves and young panicles. Both genes were cloned from a cold-tolerant variety, Norin-PL8, and introduced into a cold-sensitive variety, Hokkai241, and a cold-sensitive line, BT4-74-8. The cold tolerance of T 2 and T 3 progenies was assessed by measuring the degree of spikelet fertility in plants treated with cool water irrigation (19 C, 25 cm) or cool air (12 C, 4 days). The results indicated that the F-box protein gene confers cold tolerance. Cold tolerance is associated with greater anther length, and the transgenic plants had longer anthers than non-transformed controls. The F-box protein interacts with a subunit of the E3 ubiquitin ligase, Skp1, suggesting that an ubiquitin roteasome pathway is involved in cold tolerance at the booting stage.

Huang XH, Zhao Y, Wei XH, Li CY, Wang AH, Zhao Q, Li WJ, Guo YL, Deng LW, Zhu CR, Fan DL, Lu YQ, Weng QJ, Liu KY, Zhou TY, Jing YF, Si LZ, Dong GJ, Huang T, Lu TT, Feng Q, Qian Q, Li JY, Han B . Genome-wide association study of flowering time and grain yield traits in a worldwide collection of rice germplasm
Nat Genet, 2012,44(1):32-39.
URLPMID:22138690 [本文引用: 1]
A high-density haplotype map recently enabled a genome-wide association study (GWAS) in a population of indica subspecies of Chinese landraces. Here we extend this methodology to a larger and more diverse sample of 950 worldwide varieties, including the and subspecies, to perform an additional GWAS. We identified a total of 32 new loci associated with flowering time and with ten grain-related traits, indicating that the larger sample increased the power to detect trait-associated variants using GWAS. To characterize various alleles and complex genetic variation, we developed an analytical framework for haplotype-based de novo assembly of the low-coverage sequencing data in . We identified candidate genes for 18 associated loci through detailed annotation. This study shows that the integrated approach of sequence-based GWAS and functional genome annotation has the potential to match complex traits to their causal polymorphisms in .

Meijón M, Satbhai SB, Tsuchimatsu T, Busch W . Genome-wide association study using cellular traits identifies a new regulator of root development in
Arabidopsis. Nat Genet, 2014,46(1):77-81.
URLPMID:24212884 [本文引用: 1]
Abstract With the increased availability of high-resolution sequence information, genome-wide association (GWA) studies have become feasible in a number of species. The vast majority of these studies are conducted in human populations, where it is difficult to provide strong evidence for the functional involvement of unknown genes that are identified using GWA. Here we used the model organism Arabidopsis thaliana to combine high-throughput confocal microscopy imaging of traits at the cellular level, GWA and expression analyses to identify genomic regions that are associated with developmental cell-type traits. We identify and characterize a new F-box gene, KUK, that regulates meristem and cell length. We further show that polymorphisms in the coding sequence are the major causes of KUK allele-dependent natural variation in root development. This work demonstrates the feasibility of GWA using cellular traits to identify causal genes for basic biological processes such as development.

Price AL, Zaitlen NA, Reich D, Patterson N . New approaches to population stratification in genome-wide association studies
Nat Rev Genet, 2010,11(7):459-463.
URLPMID:20548291 [本文引用: 1]
Abstract Genome-wide association (GWA) studies are an effective approach for identifying genetic variants associated with disease risk. GWA studies can be confounded by population stratification--systematic ancestry differences between cases and controls--which has previously been addressed by methods that infer genetic ancestry. Those methods perform well in data sets in which population structure is the only kind of structure present but are inadequate in data sets that also contain family structure or cryptic relatedness. Here, we review recent progress on methods that correct for stratification while accounting for these additional complexities.

Fujino K, Obara M, Shimizu T, Koyanagi KO, Ikegaya T . Genome-wide association mapping focusing on a rice population derived from rice breeding programs in a region
Breed Sci, 2015,65(5):403-410.
URLPMID:4671701 [本文引用: 1]
Plant breeding programs in local regions may generate genetic variations that are desirable to local populations and shape adaptability during the establishment of local populations. To elucidate genetic bases for this process, we proposed a new approach for identifying the genetic bases for the traits improved during rice breeding programs; association mapping focusing on a local population. In the present study, we performed association mapping focusing on a local rice population, consisting of 63 varieties, in Hokkaido, the northernmost region of Japan and one of the northern limits of rice cultivation worldwide. Six and seventeen QTLs were identified for heading date and low temperature germinability, respectively. Of these, 13 were novel QTLs in this population and 10 corresponded to the QTLs previously reported based on QTL mapping. The identification of QTLs for traits in local populations including elite varieties may lead to a better understanding of the genetic bases of elite traits. This is of direct relevance for plant breeding programs in local regions.

Shakiba E, Edwards JD, Jodari F, Duke SE, Baldo AM, Korniliev P , McCouch SR, Eizenga GC. Genetic architecture of cold tolerance in rice (Oryza sativa) determined through high resolution genome-wide analysis.
PLoS One, 2017,12(3):e0172133.
[本文引用: 1]

Lv Y, Guo ZL, Li XK, Ye HY, Li XH, Xiong LZ . New insights into the genetic basis of natural chilling and cold shock tolerance in rice by genome-wide association analysis
Plant Cell Environ, 2016,39(3):556-570.
URLPMID:26381647 [本文引用: 1]
Abstract In order to understand cold adaptability and explore additional genetic resources for the cold tolerance improvement of rice, we investigated the genetic variation of 529 rice accessions under natural chilling and cold shock stress conditions at the seedling stage using genome-wide association studies; a total of 132 loci were identified. Among them, 12 loci were common for both chilling and cold shock tolerance, suggesting that rice has a distinct and overlapping genetic response and adaptation to the two stresses. Haplotype analysis of a known gene OsMYB2 , which is involved in cold tolerance, revealed indica aponica differentiation and latitude tendency for the haplotypes of this gene. By checking the subpopulation and geographical distribution of accessions with tolerance or sensitivity under these two stress conditions, we found that the chilling tolerance group, which mainly consisted of japonica accessions, has a wider latitudinal distribution than the chilling sensitivity group. We conclude that the genetic basis of natural chilling stress tolerance in rice is distinct from that of cold shock stress frequently used for low-temperature treatment in the laboratory and the cold adaptability of rice is associated with the subpopulation and latitudinal distribution.

Wang QY, Guan YC, Wu YR, Chen HL, Chen F, Chu CC . Overexpression of a rice OsDREB1F gene increases salt, drought, and low temperature tolerance in both Arabidopsis and rice.
Plant Mol Biol, 2008,67(6):589-602.
[本文引用: 3]

Wang CC, Wei Q, Zhang K, Wang L, Liu FX, Zhao NL, Tan YJ, Di C, Yan H, Yu JJ, Sun CQ, Chen WJ, Xu WY, Su Z . Down-regulation of
OsSPX1 causes high sensitivity to cold and oxidative stresses in rice seedlings. PLoS One, 2013,8(12):e81849.


Tovuu A, Zulfugarov IS, Wu GX, Kang IS, Kim C, Moon BY, An G, Lee CH . Rice mutants deficient in ω-3 fatty acid desaturase (FAD8) fail to acclimate to cold temperatures.
Plant Physiol Biochem, 2016,109:525-535.


Qin YH, Shen X, Wang NL, Ding XP . Characterization of a novel cyclase-like gene family involved in controlling stress tolerance in rice
J Plant Physiol, 2015,181:30-41.
URLPMID:25974367
A novel cyclase-like gene family ( CYL ) encodes proteins containing cyclase domain, but their functions are largely unknown. We report the systematic identification and characterization of CYL genes in the rice genome. Five putative CYL protein sequences (OsCYL1 to 4b) were identified. These sequences and other CYL homologs were classified into four subgroups based on phylogenetic analysis. Distinct diversification of these CYL proteins exists between plants and non-plants. The CYL family has conserved exon-intron structures, and the organizations of putative motifs in plants are specifically diverse. All OsCYL genes were expressed in a wide range of tissues or organs and were responsive to at least one of the abiotic stresses and hormone treatments applied. Protein OsCYL4a is targeted to the cell membrane. The overexpression of one stress-responsive gene OsCYL4a in rice resulted in decreased tolerance to salt, drought, cold, and oxidative stress. The expression levels of some abiotic stress-responsive factors, including H 2 O 2 -accumulating negative factors DST and OsSKIPa in OsCYL4a -overexpressing plants, were reduced compared with the wild type under normal condition and drought stress. These results suggest that rice CYL family may be functionally conserved polyketide cyclase, resulting in the rapid accumulation of reactive oxygen species to decrease tolerance to abiotic stresses.

Zang AP, Xu XJ, Neill S, Cai WM . Overexpression of OsRAN2 in rice and Arabidopsis renders transgenic plants hypersensitive to salinity and osmotic stress.
J Exp Bot, 2010,61(3):777-789.


Zhu JK . Abiotic stress signaling and responses in plants
Cell, 2016,167(2):313-324.
URLPMID:27716505 [本文引用: 2]
Abstrat: As sessile organisms, plants must cope with abiotic stress such as soil salinity, drought, and extreme temperatures. Core stress-signaling pathways involve protein kinases related to the yeast SNF1 and mammalian AMPK, suggesting that stress signaling in plants evolved from energy sensing. Stress signaling regulates proteins critical for ion and water transport and for metabolic and geneexpression reprogramming to bring about ionic and water homeostasis and cellular stability under stress conditions. Understanding stress signaling and responses will increase our ability to improve stress resistance in crops to achieve agricultural sustainability and food security for a growing world population.

Vishwakarma K, Upadhyay N, Kumar N, Yadav G, Singh J, Mishra RK, Kumar V, Verma R, Upadhyay RG, Pandey M, Sharma S . Abscisic acid signaling and abiotic stress tolerance in plants: a review on current knowledge and future prospects
Front Plant Sci, 2017,8:161.
URLPMID:5316533 [本文引用: 1]
Abiotic stress is one of the severe stresses of environment that lowers the growth and yield of any crop even on irrigated land throughout the world. A major phytohormone abscisic acid (ABA) plays an essential part in acting toward varied range of stresses like heavy metal stress, drought, thermal or heat stress, high level of salinity, low temperature, and radiation stress. Its role is also elaborated in various developmental processes including seed germination, seed dormancy, and closure of stomata. ABA acts by modifying the expression level of gene and subsequent analysis ofcis- andtrans-acting regulatory elements of responsive promoters. It also interacts with the signaling molecules of processes involved in stress response and development of seeds. On the whole, the stress to a plant can be susceptible or tolerant by taking into account the coordinated activities of various stress-responsive genes. Numbers of transcription factor are involved in regulating the expression of ABA responsive genes by acting together with their respectivecis-acting elements. Hence, for improvement in stress-tolerance capacity of plants, it is necessary to understand the mechanism behind it. On this ground, this article enlightens the importance and role of ABA signaling with regard to various stresses as well as regulation of ABA biosynthetic pathway along with the transcription factors for stress tolerance.

Ma Y, Szostkiewicz I, Korte A, Moes D, Yang Y, Christmann A, Grill E . Regulators of PP2C phosphatase activity function as abscisic acid sensors
Science, 2009,324(5930):1064-1068.
URLPMID:19407143 [本文引用: 1]
Abstract The plant hormone abscisic acid (ABA) acts as a developmental signal and as an integrator of environmental cues such as drought and cold. Key players in ABA signal transduction include the type 2C protein phosphatases (PP2Cs) ABI1 and ABI2, which act by negatively regulating ABA responses. In this study, we identify interactors of ABI1 and ABI2 which we have named regulatory components of ABA receptor (RCARs). In Arabidopsis, RCARs belong to a family with 14 members that share structural similarity with class 10 pathogen-related proteins. RCAR1 was shown to bind ABA, to mediate ABA-dependent inactivation of ABI1 or ABI2 in vitro, and to antagonize PP2C action in planta. Other RCARs also mediated ABA-dependent regulation of ABI1 and ABI2, consistent with a combinatorial assembly of receptor complexes.

Park SY, Fung P, Nishimura N, Jensen DR, Fujii H, Zhao Y, Lumba S, Santiago J, Rodrigues A, Chow TFF, Alfred SE, Bonetta D, Finkelstein R, Provart NJ, Desveaux D, Rodriguez PL , McCourt P, Zhu JK, Schroeder JI, Volkman BF, Cutler SR. Abscisic acid inhibits type 2C protein phosphatases via the PYR/PYL family of START proteins
Science, 2009,324(5930):1068-1071.
[本文引用: 1]

Fujii H, Chinnusamy V, Rodrigues A, Rubio S, Antoni R, Park SY, Cutler SR, Sheen J, Rodriguez PL, Zhu JK . In vitro reconstitution of an abscisic acid signalling pathway. Nature, 2009,462(7273):660-664.
[本文引用: 1]

Kim H, Hwang H, Hong JW, Lee YN, Ahn IP, Yoon IS, Yoo SD, Lee S, Lee SC, Kim BG . A rice orthologue of the ABA receptor, OsPYL/RCAR5, is a positive regulator of the ABA signal transduction pathway in seed germination and early seedling growth
J Exp Bot, 2012,63(2):1013-1024.
URLPMID:22071266 [本文引用: 1]
Abstract Abscisic acid (ABA) is a phytohormone that positively regulates seed dormancy and stress tolerance. PYL/RCARs were identified an intracellular ABA receptors regulating ABA-dependent gene expression in Arabidopsis thaliana. However, their function in monocot species has not been characterized yet. Herein, it is demonstrated that PYL/RCAR orthologues in Oryza sativa function as a positive regulator of the ABA signal transduction pathway. Transgenic rice plants expressing OsPYL/RCAR5, a PYL/RCAR orthologue of rice, were found to be hypersensitive to ABA during seed germination and early seedling growth. A rice ABA signalling unit composed of OsPYL/RCAR5, OsPP2C30, SAPK2, and OREB1 for ABA-dependent gene regulation was further identified, via interaction assays and a transient gene expression assay. Thus, a core signalling unit for ABA-responsive gene expression modulating seed germination and early seedling growth in rice has been unravelled. This study provides substantial contributions toward understanding the ABA signal transduction pathway in rice.

Sah SK, Reddy KR, Li JX . Abscisic acid and abiotic stress tolerance in crop plants
Front Plant Sci, 2016,7:571.
URLPMID:4855980 [本文引用: 4]
Abiotic stress is a primary threat to fulfill the demand of agricultural production to feed the world in coming decades. Plants reduce growth and development process during stress conditions, which ultimately affect the yield. In stress conditions, plants develop various stress mechanism to face the magnitude of stress challenges, although that is not enough to protect them. Therefore, many strategies have been used to produce abiotic stress tolerance crop plants, among them, abscisic acid (ABA) phytohormone engineering could be one of the methods of choice. ABA is an isoprenoid phytohormone, which regulates various physiological processes ranging from stomatal opening to protein storage and provides adaptation to many stresses like drought, salt, and cold stresses. ABA is also called an important messenger that acts as the signaling mediator for regulating the adaptive response of plants to different environmental stress conditions. In this review, we will discuss the role of ABA in response to abiotic stress at the molecular level and ABA signaling. The review also deals with the effect of ABA in respect to gene expression.

Tian XJ, Wang ZY, Li XF, Lv TX, Liu HZ, Wang LZ, Niu HB, Bu QY . Characterization and functional analysis of pyrabactin resistance-like abscisic acid receptor family in rice
Rice, 2015,8:28.
URLPMID:4567572 [本文引用: 1]
Background Abscisic acid (ABA) plays crucial roles in regulating plant growth and development, especially in responding to abiotic stress. The pyrabactin resistance-like (PYL) abscisic acid receptor...

Muhammad R, He GY, Yang GY, Javeed H, Yan X . AP2/ERF transcription factor in rice: genome-wide canvas and syntenic relationships between monocots and eudicots
Evol Bioinform Online, 2012,8:321-355.
URLPMID:3396566 [本文引用: 1]
The transcription factor family intimately regulates gene expression in response to hormones, biotic and abiotic factors, symbiotic interactions, cell differentiation, and stress signalling pathways in plants. In this study, 170 AP2/ERF family genes are identified by phylogenetic analysis of the rice genome (Oryza sativa l. japonica) and they are divided into a total of 11 groups, including four major groups (AP2, ERF, DREB, and RAV), 10 subgroups, and two soloists. Gene structure analysis revealed that, at position-6, the amino acid threonine (Thr-6) is conserved in the double domain AP2 proteins compared to the amino acid arginine (Arg-6), which is preserved in the single domain of ERF proteins. In addition, the histidine (His) amino acid is found in both domains of the double domain AP2 protein, which is missing in single domain ERF proteins. Motif analysis indicates that most of the conserved motifs, apart from the AP2/ERF domain, are exclusively distributed among the specific clades in the phylogenetic tree and regulate plausible functions. Expression analysis reveals a widespread distribution of the rice AP2/ERF family genes within plant tissues. In the vegetative organs, the transcripts of these genes are found most abundant in the roots followed by the leaf and stem; whereas, in reproductive tissues, the gene expression of this family is observed high in the embryo and lemma. From chromosomal localization, it appears that repetition and tandem-duplication may contribute to the evolution of new genes in the rice genome. In this study, interspecies comparisons between rice and wheat reveal 34 rice loci and unveil the extent of collinearity between the two genomes. It was subsequently ascertained that chromosome-9 has more orthologous loci for CRT/DRE genes whereas chromosome-2 exhibits orthologs for ERF subfamily members. Maximum conserved synteny is found in chromosome-3 for AP2 double domain subfamily genes. Macrosynteny between rice and Arabidopsis, a distant, related genome, uncovered 11 homologs/orthologs loci in both genomes. The distribution of AP2/ERF family gene paralogs in Arabidopsis was most frequent in chromosome-1 followed by chromosome-5. In Arabidopsis, ERF subfamily gene orthologs are found on chromosome-1, chromosome-3, and chromosome-5, whereas DRE subfamily genes are found on chromosome-2 and chromosome-5. Orthologs for RAV and AP2 with double domains in Arabidopsis are located on chromosome-1 and chromosome-3, respectively. In conclusion, the data generated in this survey will be useful for conducting genomic research to determine the precise role of the AP2/ERF gene during stress responses with the ultimate goal of improving crops.

Liu JY, Shi YT, Yang SH . CBF: a key factor balancing plant cold stress responses and growth.Chin Bull Bot, 52(6):689-698.
URL [本文引用: 1]
低温是影响植物生长发育以及植被分布的重要环境因子。目前,低温信号研究中比较清楚的是CBF依赖的低温信号途径。该文总结了近年来有关CBF的研究成果,详细介绍了CBF家族成员在植物耐寒性中的重要作用,着重分析与讨论CBF介导的低温调控网络及一系列复杂调控机制。理解CBF的复杂作用机制有助于了解植物中CBF介导的冷信号如何平衡耐寒性与生长发育,进而有助于耐寒作物的培育。
刘静妍, 施怡婷, 杨淑华 . CBF: 平衡植物低温应答与生长发育的关键
植物学报, 2017,52(6):689-698.
URL [本文引用: 1]
低温是影响植物生长发育以及植被分布的重要环境因子。目前,低温信号研究中比较清楚的是CBF依赖的低温信号途径。该文总结了近年来有关CBF的研究成果,详细介绍了CBF家族成员在植物耐寒性中的重要作用,着重分析与讨论CBF介导的低温调控网络及一系列复杂调控机制。理解CBF的复杂作用机制有助于了解植物中CBF介导的冷信号如何平衡耐寒性与生长发育,进而有助于耐寒作物的培育。

Chinnusamy V, Zhu JH, Zhu JK . Cold stress regulation of gene expression in plants
Trends Plant Sci, 2007,12(10):444-451.
URLPMID:17855156 [本文引用: 2]
Cold stress adversely affects plant growth and development. Most temperate plants acquire freezing tolerance by a process called cold acclimation. Here, we focus on recent progress in transcriptional, post-transcriptional and post-translational regulation of gene expression that is critical for cold acclimation. Transcriptional regulation is mediated by the inducer of C-repeat binding factor (CBF) expression 1 (ICE1), the CBF transcriptional cascade and CBF-independent regulons during cold acclimation. ICE1 is negatively regulated by ubiquitination-mediated proteolysis and positively regulated by SUMO (small ubiquitin-related modifier) E3 ligase-catalyzed sumoylation. Post-transcriptional regulatory mechanisms, such as pre-mRNA splicing, mRNA export and small RNA-directed mRNA degradation, also play important roles in cold stress responses.

Zhang Y, Chen C, Jin XF, Xiong AS, Peng RH, Hong YH, Yao QH, Chen JM . Expression of a rice DREB1 gene,OsDREB1D, enhances cold and high-salt tolerance in transgenic Arabidopsis.
BMB Rep, 2009,42(8):486-492.
[本文引用: 1]

Zhang ZY, Li JH, Li F, Liu HH, Yang WS, Chong K, Xu YY . OsMAPK3 phosphorylates OsbHLH002/OsICE1 and inhibits its ubiquitination to activate OsTPP1 and enhances rice chilling tolerance
Dev Cell, 2017, 43(6): 731-743. e5.
[本文引用: 2]

Wang DZ, Jin YN, Ding XH, Wang WJ, Zhai SS, Bai LP, Guo ZF . Gene regulation and signal transduction in the ICE-CBF-COR signaling pathway during cold stress in plants
Biochemistry (Moscow), 2017,82(10):1103-1117.
URLPMID:29037131 [本文引用: 1]
Low temperature is an abiotic stress that adversely affects the growth and production of plants. Resistance and adaptation of plants to cold stress is dependent upon the activation of molecular networks and pathways involved in signal transduction and the regulation of cold-stress related genes. Because it has numerous and complex genes, regulation factors, and pathways, research on the ICE–CBF–COR signaling pathway is the most studied and detailed, which is thought to be rather important for cold resistance of plants. In this review, we focus on the function of each member, interrelation among members, and the influence of manipulators and repressors in the ICE–CBF–COR pathway. In addition, regulation and signal transduction concerning plant hormones, circadian clock, and light are discussed. The studies presented provide a detailed picture of the ICE–CBF–COR pathway.

Liu CT, Wu YB ,Wang XP . bZIP transcription factor OsbZIP52/RISBZ5: a potential negative regulator of cold and drought stress response in rice
Planta, 2012,235(6):1157-1169.
[本文引用: 1]

Meuwissen TH, Hayes BJ, Goddard ME . Prediction of total genetic value using genome-wide dense marker maps
Genetics, 2001,157(4):1819-1829.
PMID:11290733 [本文引用: 1]
Recent advances in molecular genetic techniques will make dense marker maps available and genotyping many individuals for these markers feasible. Here we attempted to estimate the effects of approximately 50,000 marker haplotypes simultaneously from a limited number of phenotypic records. A genome of 1000 cM was simulated with a marker spacing of 1 cM. The markers surrounding every 1-cM region were combined into marker haplotypes. Due to finite population size N(e) = 100, the marker haplotypes were in linkage disequilibrium with the QTL located between the markers. Using least squares, all haplotype effects could not be estimated simultaneously. When only the biggest effects were included, they were overestimated and the accuracy of predicting genetic values of the offspring of the recorded animals was only 0.32. Best linear unbiased prediction of haplotype effects assumed equal variances associated to each 1-cM chromosomal segment, which yielded an accuracy of 0.73, although this assumption was far from true. Bayesian methods that assumed a prior distribution of the variance associated with each chromosome segment increased this accuracy to 0.85, even when the prior was not correct. It was concluded that selection on genetic values predicted from markers could substantially increase the rate of genetic gain in animals and plants, especially if combined with reproductive techniques to shorten the generation interval.

Sonesson AK, Meuwissen THE . Testing strategies for genomic selection in aquaculture breeding programs
Genet Sel Evol, 2009,41:37.
URLPMID:2714299 [本文引用: 1]
Background Genomic selection is a selection method where effects of dense genetic markers are first estimated in a test population and later used to predict breeding values of selection candidates. The aim of this paper was to investigate genetic gains, inbreeding and the accuracy of selection in a general genomic selection scheme for aquaculture, where the test population consists of sibs of the candidates. Methods The selection scheme started after simulating 4000 generations in a Fisher-Wright population with a size of 1000 to create a founder population. The basic scheme had 3000 selection candidates, 3000 tested sibs of the candidates, 100 full-sib families, a trait heritability of 0.4 and a marker density of 0.5Ne/M. Variants of this scheme were also analysed. Results The accuracy of selection in generation 5 was 0.823 for the basic scheme when the sib-testing was performed every generation. The accuracy was hardly reduced by selection, probably because the increased frequency of favourable alleles compensated for the Bulmer effect. When sib-testing was performed only in the first generation, in order to reduce costs, accuracy of selection in generation 5 dropped to 0.304, the main reduction occurring in the first generation. The genetic level in generation 5 was 6.35??a when sib-testing was performed every generation, which was 72%, 12% and 9% higher than when sib-testing was performed only in the first generation, only in the first three generations or every second generation, respectively. A marker density above 0.5Ne/M hardly increased accuracy of selection further. For the basic scheme, rates of inbreeding were reduced by 81% in these schemes compared to traditional selection schemes, due to within-family selection. Increasing the number of sibs to 6000 hardly affected the accuracy of selection, and increasing the number of candidates to 6000 increased genetic gain by 10%, mainly because of increased selection intensity. Conclusion Various strategies were evaluated to reduce the amount of sib-testing and genotyping, but all resulted in loss of selection accuracy and thus of genetic gain. Rates of inbreeding were reduced by 81% in genomic selection schemes compared to traditional selection schemes for the parameters of the basic scheme, due to within-family selection.

Li HD, Bao ZM, Sun XW . Genomic selection and its application
Hereditas (Beijing), 2011,33(12):1308-1316.
URLMagsci [本文引用: 1]
品种选育在农业生产中占有十分重要的地位, 育种值估计是品种选育的核心。随着遗传标记的发展, 尤其是高通量的基因分型技术, 使得从基因组水平估计育种值成为可能, 即基因组选择。文章将基因组选择的方法分为两类：一是基于估计等位基因效应来预测基因组估计育种值(GEBV), 如最小二乘法, 随机回归-最佳线性无偏预测(RR-BLUP)、Bayes、主成分分析等方法; 二是基于遗传关系矩阵来预测GEBV, 通过采用高通量标记构建个体间的遗传关系矩阵, 然后用线性混合模型来预测育种值, 即GBLUP法, 并以这两种分类简要介绍了基因组选择各种方法的大致原理。影响基因组选择准确性的因素主要有标记类型和密度、单倍型长度、参考群体大小和标记-数量性状基因座(QTL)连锁不平衡(LD)大小等; 在基因组选择的各种方法中, 一般说来Bayes方法和GBLUP方法具有较高的准确性, 最小二乘法最差; GBLUP计算速度快, 能够将标记和系谱结合起来, 因而比其他方法更具优势。尽管基因组选择取得了很大进展, 但在理论方面还面临着一些挑战, 如联合育种、长期选择的遗传进展及如何解析与性状有关和无关的标记等。基因组选择在一些动植物育种上已经开始应用, 在人类遗传倾向预测和进化动力学研究中也有潜在的应用前景。基因组选择在个体间亲缘关系的量化上有了突破, 比传统方法更加精确, 因此, 基因组选择将会是动植物育种史上革命性的事件。
李恒德, 包振民, 孙效文 . 基因组选择及其应用
遗传, 2011,33(12):1308-1316.
URLMagsci [本文引用: 1]
品种选育在农业生产中占有十分重要的地位, 育种值估计是品种选育的核心。随着遗传标记的发展, 尤其是高通量的基因分型技术, 使得从基因组水平估计育种值成为可能, 即基因组选择。文章将基因组选择的方法分为两类：一是基于估计等位基因效应来预测基因组估计育种值(GEBV), 如最小二乘法, 随机回归-最佳线性无偏预测(RR-BLUP)、Bayes、主成分分析等方法; 二是基于遗传关系矩阵来预测GEBV, 通过采用高通量标记构建个体间的遗传关系矩阵, 然后用线性混合模型来预测育种值, 即GBLUP法, 并以这两种分类简要介绍了基因组选择各种方法的大致原理。影响基因组选择准确性的因素主要有标记类型和密度、单倍型长度、参考群体大小和标记-数量性状基因座(QTL)连锁不平衡(LD)大小等; 在基因组选择的各种方法中, 一般说来Bayes方法和GBLUP方法具有较高的准确性, 最小二乘法最差; GBLUP计算速度快, 能够将标记和系谱结合起来, 因而比其他方法更具优势。尽管基因组选择取得了很大进展, 但在理论方面还面临着一些挑战, 如联合育种、长期选择的遗传进展及如何解析与性状有关和无关的标记等。基因组选择在一些动植物育种上已经开始应用, 在人类遗传倾向预测和进化动力学研究中也有潜在的应用前景。基因组选择在个体间亲缘关系的量化上有了突破, 比传统方法更加精确, 因此, 基因组选择将会是动植物育种史上革命性的事件。

Wang JK, Li HH, Zhang LY. Genetic mapping and breeding design. Beijing: Science Press, 2014.
[本文引用: 1]

王建康, 李慧慧, 张鲁燕 . 基因定位与育种设计. 北京: 科学出版社, 2014.
[本文引用: 1]

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