生长素代谢、运输及信号转导调控水稻粒型研究进展

doi:10.11983/CBB21227

植物学报 ›› 2022, Vol. 57 ›› Issue (3): 263-275.DOI: 10.11983/CBB21227

• 特邀综述 • 下一篇

生长素代谢、运输及信号转导调控水稻粒型研究进展

贾利霞¹, 齐艳华¹^,²^,^*()

¹浙江大学生命科学学院, 植物生理与生物化学国家重点实验室, 杭州 310058
²内蒙古大学生命科学学院, 牧草与特色作物生物学教育部重点实验室, 内蒙古自治区牧草与特色作物生物技术重点实验室, 呼和浩特 010000

收稿日期:2021-12-24 接受日期:2022-03-18 出版日期:2022-05-01 发布日期:2022-05-18
通讯作者: 齐艳华
作者简介:* E-mail: qyhjp@zju.edu.cn
基金资助:
国家自然科学基金(32060451);浙江省自然科学基金(LZ19C020001)

Advances in the Regulation of Rice (Oryza sativa) Grain Shape by Auxin Metabolism, Transport and Signal Transduction

Lixia Jia¹, Yanhua Qi¹^,²^,^*()

¹State Key Laboratory of Plant Physiol and Biochemistry, College of Life Sciences, Zhejiang University, Hangzhou 310058, China
²Inner Mongolia Key Laboratory of Herbage & Endemic Crop Biotechnology, Key Laboratory of Herbage & Endemic Crop Biology of Ministry of Education, School of Life Sciences, Inner Mongolia University, Hohhot 010000, China

Received:2021-12-24 Accepted:2022-03-18 Online:2022-05-01 Published:2022-05-18
Contact: Yanhua Qi

摘要/Abstract

摘要： 水稻(Oryza sativa)是世界主要粮食作物。随着我国经济飞速发展, 耕地面积逐年减少, 提高水稻总产量唯有依靠单产的增加。粒重是决定水稻产量的重要因素之一, 其遗传稳定, 受外界环境因素影响较小。粒重由粒型和灌浆程度决定, 而粒型性状包括粒长、粒宽、粒厚和长宽比。水稻种子颖壳和胚乳发育决定了粒型和粒重, 颖壳细胞的增殖和扩张限制籽粒发育, 胚乳占据成熟种子的大部分体积。而生长素调控受精后颖壳和胚乳的发育, 是调控种子发育和影响水稻产量的重要植物激素。生长素的时空分布受生长素代谢、运输和信号转导的动态调节, 以维持生长素在种子发育中的最适水平。该文综述了生长素代谢、运输和信号转导调控水稻粒型的研究进展, 以期为深入探究生长素调控水稻粒型发育机制和提高水稻产量提供线索。

关键词: 水稻, 生长素代谢, 生长素运输, 生长素信号转导, 粒型

Abstract: Rice (Oryza sativa) is a major food crop in the world. The optimization and utilization of the major yield-determining factors are important for increasing yield potential. Among these factors, seed weight is one of the most important factors determining rice production. The heritability of rice grain weight is stable, which is largely unaffected by environmental factors. Grain weight depends on grain shape, which is determined by grain length, grain width, and grain thickness, and the degree of grain filling. The growth of rice glumes and seed endosperm determines the grain shape and weight. The proliferation and expansion of glume cells affect grain development, and endosperm occupies most of the volume of mature seeds. Auxin is an important plant hormone that affects rice yield, which regulates the development of glume and endosperm after fertilization. The spatial-temporal distribution of active auxin is dynamically modulated by auxin metabolism, auxin transport and signal transduction, all of which maintain auxin at the optimal level for seed development. Here we reviewed the research progress of auxin pathways regulating rice grain shape from three aspects, auxin metabolism, auxin transport and auxin signal transduction, to provide clues for exploring the auxin regulation mechanism of grain shape and improve yield in rice.

Key words: rice, auxin metabolism, auxin transport, auxin signal transduction, grain shape

贾利霞, 齐艳华. 生长素代谢、运输及信号转导调控水稻粒型研究进展. 植物学报, 2022, 57(3): 263-275.

Lixia Jia, Yanhua Qi. Advances in the Regulation of Rice (Oryza sativa) Grain Shape by Auxin Metabolism, Transport and Signal Transduction. Chinese Bulletin of Botany, 2022, 57(3): 263-275.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB21227

https://www.chinbullbotany.com/CN/Y2022/V57/I3/263

图/表 5

图1 生长素代谢(参考Staswick et al., 2005; Dai et al., 2013; Kakei et al., 2017; Wang et al., 2018; Hayashi et al., 2021) TAA转氨酶家族催化色氨酸(Trp)产生吲哚-3-丙酮酸(IPA), 而后YUC (YUCCA)家族催化IPA形成吲哚-3-乙酸(IAA)。TGW6水解IAA-葡萄糖释放IAA, IAA可以在GH3 (IAA-酰胺合成酶)作用下与氨基酸结合, 或通过DAO分解代谢为2-氧代吲哚-3-乙酸(oxIAA), 在体外DAO12可将IAA-天冬氨酸(IAA-Asp)和IAA-谷氨酸(IAA-Glu)氧化为2-氧代吲哚-3-乙酸-天冬氨酸(oxIAA-Asp)和2-氧代吲哚-3-乙酸-谷氨酸(oxIAA-Glu)。

Figure 1 Auxin metabolism (refer to Staswick et al., 2005; Dai et al., 2013; Kakei et al., 2017; Wang et al., 2018; Hayashi et al., 2021) The TAA family of aminotransferases catalyzes tryptophan (Trp) to produce indole-3-pyruvate (IPA), and the YUCCA (YUC) family catalyzes IPA to form indole-3-acetic acid (IAA). TGW6 hydrolyzes IAA-glucose to release IAA. Conjugation of IAA with amino acids is achieved through the GH3 IAA-amido synthetases or catabolized into 2-oxoindole-3-acetic acid (oxIAA) by DAO. In vitro, DAO12 oxidized IAA-aspartate (IAA-Asp) and IAA-glutamate (IAA-Glu) to yield 2-oxindole-3-acetic acid-aspartate (oxIAA-Asp) and 2-oxindole-3-acetic acid-glutamate (oxIAA-Glu).

图2 生长素运输载体PINs、AUX1/LAXs、ABCBs和PILs及BG1 (推定的)的亚细胞定位模式(参考Garcia et al., 2004; Zhang et al., 2012; Balzan et al., 2014; Liu et al., 2015; Wang et al., 2018; Hou et al., 2021) OsPIN1、OsPIN2、OsPIN3t (OsPIN3a/OsPIN10a)和OsPIN9定位于细胞膜, OsPIN5定位于内质网膜。箭头表示生长素流动的方向。

Figure 2 Model of subcellular localization of auxin transport vectors PINs, AUX1/LAXs, ABCBs and PILs or BG1 (putative) (refer to Garcia et al., 2004; Zhang et al., 2012; Balzan et al., 2014; Liu et al., 2015; Wang et al., 2018; Hou et al., 2021) OsPIN1, OsPIN2, OsPIN3t (OsPIN3a/OsPIN10a) and OsPIN9 are localized on the plasma membrane, OsPIN5 is localized on the endoplasmic reticulum (ER) membrane. The arrows indicate the direction of auxin flow.

图3 生长素信号转导的分子机制(参考Tan et al., 2007; Szemenyei et al., 2008; Lee et al., 2010; Shen et al., 2010b; Jing et al., 2015; Salehin et al., 2015) 在浓度低生长素下, AUX/IAAs与TPL结合, 抑制生长素响应因子ARFs基因的转录活性。高浓度生长素下, 肽基辅氨酸异构酶LRT2催化AUX/IAAs蛋白顺式和反式异构体之间相互转换, 生长素促进TIR1/AFBs和AUX/IAAs之间的相互作用, AUX/IAAs泛素化介导降解。OsARF家族包括9个推定的转录激活子, 即OsARF5、6、11、12、16、17、19、21和25, 其它OsARFs是推定的转录抑制子。

Figure 3 Molecular mechanisms of auxin signal transduction (refer to Tan et al., 2007; Szemenyei et al., 2008; Lee et al., 2010; Shen et al., 2010b; Jing et al., 2015; Salehin et al., 2015) Under low concentration auxin, AUX/IAAs combine with TPL to inhibit the transcriptional activity of auxin response factors (ARFs) genes. Under high concentration auxin, the peptidylprolyl isomerase LRT2 catalyzes the interconversion between the cis and trans isoforms of AUX/IAAs proteins. Auxin promotes the interaction between TIR1/AFBs and AUX/IAAs, leading to ubiquitin-mediated degradation of AUX/IAAs. OsARF family includes 9 putative transcriptional activators, OsARF5, 6, 11, 12, 16, 17, 19, 21, and 25, the other OsARFs are putative transcriptional repressors.

图4 生长素代谢、运输及信号转导调控水稻粒型(参考Zhang et al., 2010, 2018b, 2021; Chen et al., 2012; Huang et al., 2016; Guo et al., 2020; Li et al., 2020; Paul et al., 2020; Qiao et al., 2021; Sims et al., 2021) 蓝色和绿色框图分别显示生长素代谢、运输及信号转导相关基因。参与调控粒型的途径包括通过影响胚乳或颖壳发育及二者不明确的途径(虚线框)。

Figure 4 The regulation of grain shape by auxin metabolism, transport and signal transduction in rice (refer to Zhang et al., 2010, 2018b, 2021; Chen et al., 2012; Huang et al., 2016; Guo et al., 2020; Li et al., 2020; Paul et al., 2020; Qiao et al., 2021; Sims et al., 2021) Blue and green block diagrams show genes related to auxin metabolism, transport and signal transduction, respectively. The pathways involved in the regulation of grain shape include those affecting the development of either endosperm or glume and those undetermined between them (dotted box).

表1 调控水稻粒型的生长素代谢、运输及信号转导基因

Table 1 Genes of auxin metabolism, transport and signal transduction regulating grain shape in rice

基因	作用途径	粒型	参考文献
OsTSG1	参与IAA代谢	突变体粒长和粒宽减小	Guo et al., 2020
OsYUCs	参与IAA代谢	表达量过高导致不结实	Zhang et al., 2018a
OsRGB1	参与IAA代谢	RGB1-Ri株系粒长、粒宽和粒厚减小	Zhang et al., 2021
OsDEP1/qPE9-1	参与IAA代谢	DEP1/qPE9-1转基因株系粒长增加	Zhang et al., 2019
OsMADS1	参与IAA代谢	RNAi株系粒长增加, 粒宽减小, 长宽比增加	Li et al., 2020
OsMADS6	参与IAA代谢	突变体粒长减小, 粒宽增加	Zhang et al., 2010
OsGH3.13/TLD1	参与IAA代谢	突变体种子变薄	Zhang et al., 2009b
OsTGW6	参与IAA代谢	功能缺失株系粒长增加	Ishimaru et al., 2013
OsPIN2	参与IAA运输	过表达株系粒长和粒宽减小	Chen et al., 2012
OsAUX3	参与IAA运输	突变体粒长增加	Qiao et al., 2021
OsBG1	参与IAA运输	过表达株系粒长、粒宽和千粒重增加; BG1-Ri株系粒长、粒宽和千粒重减小	Liu et al., 2015
OsARF4	参与IAA信号转导	突变体粒长增加	Hu et al., 2018
OsARF6	参与IAA信号转导	突变体粒长增加	Qiao et al., 2021
OsARF11	参与IAA信号转导	T-DNA插入突变体种子变薄, 粒长和粒宽减小	Sims et al., 2021
OsARF18	参与IAA信号转导	过表达株系粒宽减小	Huang et al., 2016
OsARF25	参与IAA信号转导	T-DNA插入突变体粒长减小	Zhang et al., 2018b
OsERF142/smos1	参与IAA信号转导	突变体粒长减小, 粒宽增加	Aya et al., 2014
OsGnp4/LAX2	参与IAA信号转导	过表达株系粒长增加	Zhang et al., 2018b
OsIAA3	参与IAA信号转导	RNAi株系粒长增加	Zhang et al., 2018b
OsMADS78/79	参与IAA代谢、运输和信号转导	双敲除突变体株系完全不育, 单敲除突变体和过表达株系籽粒长宽比增加	Paul et al., 2020

表1 调控水稻粒型的生长素代谢、运输及信号转导基因

Table 1 Genes of auxin metabolism, transport and signal transduction regulating grain shape in rice

基因	作用途径	粒型	参考文献
OsTSG1	参与IAA代谢	突变体粒长和粒宽减小	Guo et al., 2020
OsYUCs	参与IAA代谢	表达量过高导致不结实	Zhang et al., 2018a
OsRGB1	参与IAA代谢	RGB1-Ri株系粒长、粒宽和粒厚减小	Zhang et al., 2021
OsDEP1/qPE9-1	参与IAA代谢	DEP1/qPE9-1转基因株系粒长增加	Zhang et al., 2019
OsMADS1	参与IAA代谢	RNAi株系粒长增加, 粒宽减小, 长宽比增加	Li et al., 2020
OsMADS6	参与IAA代谢	突变体粒长减小, 粒宽增加	Zhang et al., 2010
OsGH3.13/TLD1	参与IAA代谢	突变体种子变薄	Zhang et al., 2009b
OsTGW6	参与IAA代谢	功能缺失株系粒长增加	Ishimaru et al., 2013
OsPIN2	参与IAA运输	过表达株系粒长和粒宽减小	Chen et al., 2012
OsAUX3	参与IAA运输	突变体粒长增加	Qiao et al., 2021
OsBG1	参与IAA运输	过表达株系粒长、粒宽和千粒重增加; BG1-Ri株系粒长、粒宽和千粒重减小	Liu et al., 2015
OsARF4	参与IAA信号转导	突变体粒长增加	Hu et al., 2018
OsARF6	参与IAA信号转导	突变体粒长增加	Qiao et al., 2021
OsARF11	参与IAA信号转导	T-DNA插入突变体种子变薄, 粒长和粒宽减小	Sims et al., 2021
OsARF18	参与IAA信号转导	过表达株系粒宽减小	Huang et al., 2016
OsARF25	参与IAA信号转导	T-DNA插入突变体粒长减小	Zhang et al., 2018b
OsERF142/smos1	参与IAA信号转导	突变体粒长减小, 粒宽增加	Aya et al., 2014
OsGnp4/LAX2	参与IAA信号转导	过表达株系粒长增加	Zhang et al., 2018b
OsIAA3	参与IAA信号转导	RNAi株系粒长增加	Zhang et al., 2018b
OsMADS78/79	参与IAA代谢、运输和信号转导	双敲除突变体株系完全不育, 单敲除突变体和过表达株系籽粒长宽比增加	Paul et al., 2020

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生长素代谢、运输及信号转导调控水稻粒型研究进展

Advances in the Regulation of Rice (Oryza sativa) Grain Shape by Auxin Metabolism, Transport and Signal Transduction

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