Advances in the Mechanism of Leaf Width Regulation and Related Genes in Rice

doi:10.13560/j.cnki.biotech.bull.1985.2024-0311

Abstract

Abstract:

Rice leaf width is a crucial component of leaf morphology, which has biological significance to leaf construction and photosynthesis. Leaf width morphogenesis is accomplished in rice by cell division and cell expansion in the medio-lateral axis direction of the leaf primordium, a process that is influenced by phytohormones, cellular metabolism, and the expression levels of related genes. TDD1, NAL7 and FIB are involved in the tryptophan-dependent auxin biosynthesis pathway in rice, and GID1, GID2, and SLR1 are involved in the negative regulation of leaf width in rice by GA. Genes such as NAL21, NLG1, and NAL9 are essential for maintaining organelle homeostasis and normal cellular metabolism. While most rice leaf width genes act on cell division through the auxin signaling, CCC1 is involved in cell expansion by regulating cellular osmotic potential. The transcription factor WL1 negatively regulates the expression of the serine protease gene NAL1 by recruiting the co-repressor TOPLESS-RELATED PROTEIN, and consequently the auxin signaling pathway is affected. Exploring the genes related to leaf width development and applying them to breeding practice is of great significance for improving rice yield. Utilizing germplasm resources to explore superior haplotypes of rice leaf width gene and gene editing technology represented by CRISPR/Cas9 provide abundant means and ways to improve rice leaf width. When utilizing the rice leaf width gene for trait improvement, comprehensive consideration should be given to pleiotropy and genetic interactions, which should be combined with production practice to avoid adverse effects on other traits. In this review we summarized the genetic regulatory mechanism of rice leaf width from the aspects of histological characteristics, plant hormones, molecular mechanisms and leaf width-related genes, discussed the significance and strategies of rice leaf width improvement in breeding, and provided ideas for research on the molecular mechanism of rice leaf shape and the breeding of “ideal plant architecture”.

Key words: rice, leaf width, regulatory mechanism, improvement of trait

QIAO Cheng-bin, SONG Jia-wei, YANG Hui, DUAN Kai-rong, RAN Jie, KONG Wei-ru, FENG Pei-yuan, LUO Cheng-ke, LI Pei-fu, TIAN Lei. Advances in the Mechanism of Leaf Width Regulation and Related Genes in Rice[J]. Biotechnology Bulletin, 2024, 40(11): 88-102.

Figures/Tables 2

References 127

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基因名和ID Gene and ID	染色体 Chromosome	基因功能 Gene function	生物学功能 Biological function	亚细胞定位 Subcellular localization	突变体表型 Mutant phenotypes	参考文献 Reference
GID2 （LOC_Os02g36974）	2	编码SCF E3泛素连接酶复合体的F-box 亚基 Encodes an F-box subunit of the SCF E3 complex	GA信号传导的正调节因子 Positive regulators of GA signaling	细胞核 Nucleus	宽叶、矮化 Wide leaves, dwarf	[19]
NAL1 （LOC_Os04g52479）	4	编码丝氨酸蛋白酶 Encodes serine protease	通过降解TPR2调节生长素相关基因表达 Modulate auxin-related gene expression by degradation of TPR2	细胞核、细胞质 Nucleus, cytoplasm	窄叶、矮化 Narrow leaves, dwarf	[20-21]
NAL2 （LOC_Os11g01130）	11	编码WOX3A转录激活因子 Encodes WOX3A transcriptional activator	促进叶片横向生长和维管束发育 Promote leaf lateral growth and veins development	细胞核 Nucleus	窄卷叶 Narrow and rolled leaves	[17]
NAL3 （LOC_Os12g01120）	12	编码WOX3A转录激活因子 Encodes WOX3A transcriptional activator	促进叶片横向生长和维管束发育 Promote leaf lateral growth and veins development	细胞核 Nucleus	窄卷叶 Narrow and rolled leaves	[17]
NAL9 （LOC_Os03g29810）	3	编码酪蛋白酶水解亚基 Encodes caseinolytic protease proteolytic subunit	促进水稻叶绿体生物合成 Promote chloroplast biosynthesis	叶绿体 Chloroplast	窄叶、矮化 Narrow leaves, dwarf	[22-23]
NAL7 （LOC_Os03g06654）	3	编码黄素单加氧酶 Encodes flavin-containing monooxygenase	参与生长素生物合成 Involved in auxin biosynthesis	细胞质 Cytoplasm	窄卷叶 Narrow and rolled leaves	[24-25]
SNFL1 （LOC_Os05g50270）	5	编码GATA锌指蛋白 Encodes GATA zinc finger protein	参与叶片表皮细胞及维管束发育 Involved in the development of leaf epidermal cells and veins	细胞核 Nucleus	短窄剑叶 Short and narrow flag leaves	[26]
NAAL1 （LOC_Os07g31450）	7	染色质重塑因子 Chromatin-remodeling factor	参与生长素相关基因表达 Participate in the expression of auxin-related gene	细胞核 Nucleus	短窄叶、矮化 Short and narrow leaves, dwarf	[27-28]
WL1 （LOC_Os03g57240）	3	编码Cys-2/His-2型锌指蛋白 Encodes Cys-2/His-2-type zinc finger protein	负调控NAL1 Negatively regulate NAL1	细胞核 Nucleus	宽叶 Wide leaves	[29]
TAD1 （LOC_Os03g03150）	3	APC/C的共激活因子 Co-activator of the APC/C	激活APC/C泛素连接酶活性 Activate APC/C E3 ubiquitin ligase activity	细胞核 Nucleus	窄叶 Narrow leaves	[29]
ARF19 （LOC_Os06g48950）	6	生长素响应因子 Auxin response factor	参与生长素信号传导并正调控GH3-5 Involved in auxin signaling and positively regulates GH3-5	细胞核 Nucleus	窄叶 Narrow leaves	[30]
ARF11 （LOC_Os04g56850）	4	生长素响应因子 Auxin response factor	参与生长素信号传导 Involved in auxin signaling	细胞核 Nucleus	窄叶、矮化 Narrow leaves, dwarf	[31]
WOX3B （LOC_Os05g02730）	5	编码WOX3B转录因子 Encodes WOX3B transcription factor	参与叶片横向生长 Involved in leaf lateral growth	细胞核 Nucleus	窄叶或分叉叶 Narrow or forked leaf	[32]
SLL1 （LOC_Os09g23200）	9	编码MYB转录因子 Encodes MYB transcription factor	通过参与远轴端细胞发育调控水稻叶形 Regulate leaf shape by involving in leaf abaxial cell development	细胞核 Nucleus	窄卷叶 Narrow and rolled leaves	[33]
DCL1 （LOC_Os03g02970）	3	编码类Dicer蛋白 Encodes Dicer-like protein	负责miRNAs的生成 Responsible for the generation of miRNAs	细胞质 Cytoplasm	窄叶、矮化 Narrow leaves, dwarf	[34]
AGO1a/b/c/d （LOC_Os02g45070 LOC_Os04g47870 LOC_Os02g58490 LOC_Os06g51310）	2/4/2/6	编码Argonaute（AGO）蛋白 Encodes Argonaute（AGO）protein	抑制基因表达 Inhibit gene expression	细胞质 Cytoplasm	窄卷叶、矮化 Narrow and rolled leaves, dwarf	[35]
miR319a/b （LOC_Os01g46984 Os01g0222001）	1	微RNA microRNA	抑制TCP转录本翻译 Inhibite translatie of TCP transcripts	细胞核 Nucleus	过表达株系表现为宽叶 Overexpression lines show wider leaves	[36]
HDT702 （LOC_Os01g68104）	1	组蛋白去乙酰化酶 Histone deacetylase	参与组蛋白修饰 Involved in histone modification	细胞核 Nucleus	窄叶、矮化 Narrow leaves, dwarf	[37]
TDD1 （LOC_Os04g38950）	4	邻氨基苯甲酸合酶β亚基 Anthranilate synthase β-Subunit	催化色氨酸生物合成的第一步反应 Catalyze the first step of the Trp biosynthesis	叶绿体 Chloroplast	窄叶、矮化 Narrow leaves, dwarf	[38]
FIB （LOC_Os01g07500）	1	色氨酸氨基转移酶 Tryptophan aminotransferase	参与生长素生物合成 Involved in auxin biosynthesis	细胞质 Cytoplasm	窄叶 Narrow leaves	[39]
CKX1 （LOC_Os01g09260）	1	细胞分裂素氧化酶/脱氢酶 Cytokinin oxidase/dehydrogenase	负调控叶长、叶宽及籽粒大小 Negatively regulate leaf length, leaf width and grain size	细胞质 Cytoplasm	长宽叶 Long and wide leaves	[40]
CKX2 （LOC_Os01g10110）	1	细胞分裂素氧化酶/脱氢酶 Cytokinin oxidase/dehydrogenase	负调控叶长、叶宽及籽粒大小 Negatively regulate leaf length, leaf width and grain size	细胞质 Cytoplasm	长宽叶 Long and wide leaves	[40]
CKX4 （LOC_Os01g71310）	1	细胞分裂素氧化酶/脱氢酶 Cytokinin oxidase/dehydrogenase	正调控剑叶、籽粒大小 Positively regulate flag leaf and grain size	细胞质 Cytoplasm	短窄叶、矮化 Short and narrow leaves, dwarf	[40]
GID1 （LOC_Os05g33730）	5	赤霉素受体 GA receptor	参与GA信号传导 Involved in GA signaling	细胞核 Nucleus	宽叶、矮化 Wide leaves, dwarf	[41]
SLR1 （LOC_Os03g49990）	3	编码GRAS转录因子 Encodes GRAS transcription factor	抑制GA信号传导 Inhibit GA signaling	细胞核 Nucleus	窄叶 Narrow leaves	[42]
GH3-5 （LOC_Os05g50890）	5	茉莉酸-异亮氨酸（JA-Ile）合酶 Jasmonoyl-isoleucine（JA-Ile）synthase	催化茉莉酸-异亮氨酸复合物的合成 Catalyse JA-Ile synthesis	内质网 Endoplasmic reticulum	过表达株系表现为窄叶、矮化 Overexpression lines show narrow leaves, dwarf	[30]
CSLD4 （LOC_Os12g36890）	12	类纤维素合酶 Cellulose synthase-like	参与细胞壁形成 Involved in cell wall synthesis	高尔基体 Golgi	窄卷叶、矮化 Narrow and rolled leaves, dwarf	[43⇓⇓⇓⇓⇓-49]
NSL2 （LOC_Os06g14620）	6	编码核糖核苷酸还原酶小亚基 Encodes small subunits of ribonucleotide reductase	参与dNTPs合成 Involved in dNTPs synthesis	细胞核、细胞质 Nucleus, cytoplasm	窄叶 Narrow leaves	[50]
DLT （LOC_Os06g03710）	6	编码GRAS转录因子 Encodes GRAS transcription factor	通过抑制细胞周期蛋白相关基因的表达调控细胞分裂 Regulate cell division by suppressing the expression of cyclin related genes	细胞核 Nucleus	短宽叶 Short and wide leaves	[51]
AVB （LOC_Os03g19520）	3	未知功能蛋白 Unknown function protein	维持细胞分裂 Maintain cell division	细胞核、细胞质 Nucleus, cytoplasm	窄卷叶 Narrow and rolled leaves	[52]
CCC1 （LOC_Os08g23440）	8	阳离子-氯离子协同转运蛋白 Cation-chloride cotransporter	通过调控离子平衡维持细胞渗透势 Maintain cellular osmotic potential by regulating ion homeostasis	细胞膜 Cell membrane	窄叶 Narrow leaves	[53]
MKB3 （LOC_Os03g52320）	3	GRF互作因子 GRF-interacting factor	正调控细胞增殖和细胞大小 Positively regulate cell proliferation and cell size	细胞核 Nucleus	短窄叶 Short and narrow leaves	[54-55]
NLG1 （LOC_Os03g14890）	3	编码TIM23亚基 Encodes subunits of TIM23	维持线粒体代谢 Maintain mitochondrion metabolism	线粒体 Mitochondrion	窄叶、矮化 Narrow leaves, dwarf	[56]
NAL8 （LOC_Os07g15880）	7	编码PHB亚基 Encodes subunits of PHB	维持线粒体和叶绿体稳定性 Maintain mitochondrion and chloroplast stability	线粒体、叶绿体 Mitochondrion, chloroplast	窄叶、矮化 Narrow leaves, dwarf	[57]

基因名和ID Gene and ID	染色体 Chromosome	基因功能 Gene function	生物学功能 Biological function	亚细胞定位 Subcellular localization	突变体表型 Mutant phenotypes	参考文献 Reference
GID2 （LOC_Os02g36974）	2	编码SCF E3泛素连接酶复合体的F-box 亚基 Encodes an F-box subunit of the SCF E3 complex	GA信号传导的正调节因子 Positive regulators of GA signaling	细胞核 Nucleus	宽叶、矮化 Wide leaves, dwarf	[19]
NAL1 （LOC_Os04g52479）	4	编码丝氨酸蛋白酶 Encodes serine protease	通过降解TPR2调节生长素相关基因表达 Modulate auxin-related gene expression by degradation of TPR2	细胞核、细胞质 Nucleus, cytoplasm	窄叶、矮化 Narrow leaves, dwarf	[20-21]
NAL2 （LOC_Os11g01130）	11	编码WOX3A转录激活因子 Encodes WOX3A transcriptional activator	促进叶片横向生长和维管束发育 Promote leaf lateral growth and veins development	细胞核 Nucleus	窄卷叶 Narrow and rolled leaves	[17]
NAL3 （LOC_Os12g01120）	12	编码WOX3A转录激活因子 Encodes WOX3A transcriptional activator	促进叶片横向生长和维管束发育 Promote leaf lateral growth and veins development	细胞核 Nucleus	窄卷叶 Narrow and rolled leaves	[17]
NAL9 （LOC_Os03g29810）	3	编码酪蛋白酶水解亚基 Encodes caseinolytic protease proteolytic subunit	促进水稻叶绿体生物合成 Promote chloroplast biosynthesis	叶绿体 Chloroplast	窄叶、矮化 Narrow leaves, dwarf	[22-23]
NAL7 （LOC_Os03g06654）	3	编码黄素单加氧酶 Encodes flavin-containing monooxygenase	参与生长素生物合成 Involved in auxin biosynthesis	细胞质 Cytoplasm	窄卷叶 Narrow and rolled leaves	[24-25]
SNFL1 （LOC_Os05g50270）	5	编码GATA锌指蛋白 Encodes GATA zinc finger protein	参与叶片表皮细胞及维管束发育 Involved in the development of leaf epidermal cells and veins	细胞核 Nucleus	短窄剑叶 Short and narrow flag leaves	[26]
NAAL1 （LOC_Os07g31450）	7	染色质重塑因子 Chromatin-remodeling factor	参与生长素相关基因表达 Participate in the expression of auxin-related gene	细胞核 Nucleus	短窄叶、矮化 Short and narrow leaves, dwarf	[27-28]
WL1 （LOC_Os03g57240）	3	编码Cys-2/His-2型锌指蛋白 Encodes Cys-2/His-2-type zinc finger protein	负调控NAL1 Negatively regulate NAL1	细胞核 Nucleus	宽叶 Wide leaves	[29]
TAD1 （LOC_Os03g03150）	3	APC/C的共激活因子 Co-activator of the APC/C	激活APC/C泛素连接酶活性 Activate APC/C E3 ubiquitin ligase activity	细胞核 Nucleus	窄叶 Narrow leaves	[29]
ARF19 （LOC_Os06g48950）	6	生长素响应因子 Auxin response factor	参与生长素信号传导并正调控GH3-5 Involved in auxin signaling and positively regulates GH3-5	细胞核 Nucleus	窄叶 Narrow leaves	[30]
ARF11 （LOC_Os04g56850）	4	生长素响应因子 Auxin response factor	参与生长素信号传导 Involved in auxin signaling	细胞核 Nucleus	窄叶、矮化 Narrow leaves, dwarf	[31]
WOX3B （LOC_Os05g02730）	5	编码WOX3B转录因子 Encodes WOX3B transcription factor	参与叶片横向生长 Involved in leaf lateral growth	细胞核 Nucleus	窄叶或分叉叶 Narrow or forked leaf	[32]
SLL1 （LOC_Os09g23200）	9	编码MYB转录因子 Encodes MYB transcription factor	通过参与远轴端细胞发育调控水稻叶形 Regulate leaf shape by involving in leaf abaxial cell development	细胞核 Nucleus	窄卷叶 Narrow and rolled leaves	[33]
DCL1 （LOC_Os03g02970）	3	编码类Dicer蛋白 Encodes Dicer-like protein	负责miRNAs的生成 Responsible for the generation of miRNAs	细胞质 Cytoplasm	窄叶、矮化 Narrow leaves, dwarf	[34]
AGO1a/b/c/d （LOC_Os02g45070 LOC_Os04g47870 LOC_Os02g58490 LOC_Os06g51310）	2/4/2/6	编码Argonaute（AGO）蛋白 Encodes Argonaute（AGO）protein	抑制基因表达 Inhibit gene expression	细胞质 Cytoplasm	窄卷叶、矮化 Narrow and rolled leaves, dwarf	[35]
miR319a/b （LOC_Os01g46984 Os01g0222001）	1	微RNA microRNA	抑制TCP转录本翻译 Inhibite translatie of TCP transcripts	细胞核 Nucleus	过表达株系表现为宽叶 Overexpression lines show wider leaves	[36]
HDT702 （LOC_Os01g68104）	1	组蛋白去乙酰化酶 Histone deacetylase	参与组蛋白修饰 Involved in histone modification	细胞核 Nucleus	窄叶、矮化 Narrow leaves, dwarf	[37]
TDD1 （LOC_Os04g38950）	4	邻氨基苯甲酸合酶β亚基 Anthranilate synthase β-Subunit	催化色氨酸生物合成的第一步反应 Catalyze the first step of the Trp biosynthesis	叶绿体 Chloroplast	窄叶、矮化 Narrow leaves, dwarf	[38]
FIB （LOC_Os01g07500）	1	色氨酸氨基转移酶 Tryptophan aminotransferase	参与生长素生物合成 Involved in auxin biosynthesis	细胞质 Cytoplasm	窄叶 Narrow leaves	[39]
CKX1 （LOC_Os01g09260）	1	细胞分裂素氧化酶/脱氢酶 Cytokinin oxidase/dehydrogenase	负调控叶长、叶宽及籽粒大小 Negatively regulate leaf length, leaf width and grain size	细胞质 Cytoplasm	长宽叶 Long and wide leaves	[40]
CKX2 （LOC_Os01g10110）	1	细胞分裂素氧化酶/脱氢酶 Cytokinin oxidase/dehydrogenase	负调控叶长、叶宽及籽粒大小 Negatively regulate leaf length, leaf width and grain size	细胞质 Cytoplasm	长宽叶 Long and wide leaves	[40]
CKX4 （LOC_Os01g71310）	1	细胞分裂素氧化酶/脱氢酶 Cytokinin oxidase/dehydrogenase	正调控剑叶、籽粒大小 Positively regulate flag leaf and grain size	细胞质 Cytoplasm	短窄叶、矮化 Short and narrow leaves, dwarf	[40]
GID1 （LOC_Os05g33730）	5	赤霉素受体 GA receptor	参与GA信号传导 Involved in GA signaling	细胞核 Nucleus	宽叶、矮化 Wide leaves, dwarf	[41]
SLR1 （LOC_Os03g49990）	3	编码GRAS转录因子 Encodes GRAS transcription factor	抑制GA信号传导 Inhibit GA signaling	细胞核 Nucleus	窄叶 Narrow leaves	[42]
GH3-5 （LOC_Os05g50890）	5	茉莉酸-异亮氨酸（JA-Ile）合酶 Jasmonoyl-isoleucine（JA-Ile）synthase	催化茉莉酸-异亮氨酸复合物的合成 Catalyse JA-Ile synthesis	内质网 Endoplasmic reticulum	过表达株系表现为窄叶、矮化 Overexpression lines show narrow leaves, dwarf	[30]
CSLD4 （LOC_Os12g36890）	12	类纤维素合酶 Cellulose synthase-like	参与细胞壁形成 Involved in cell wall synthesis	高尔基体 Golgi	窄卷叶、矮化 Narrow and rolled leaves, dwarf	[43⇓⇓⇓⇓⇓-49]
NSL2 （LOC_Os06g14620）	6	编码核糖核苷酸还原酶小亚基 Encodes small subunits of ribonucleotide reductase	参与dNTPs合成 Involved in dNTPs synthesis	细胞核、细胞质 Nucleus, cytoplasm	窄叶 Narrow leaves	[50]
DLT （LOC_Os06g03710）	6	编码GRAS转录因子 Encodes GRAS transcription factor	通过抑制细胞周期蛋白相关基因的表达调控细胞分裂 Regulate cell division by suppressing the expression of cyclin related genes	细胞核 Nucleus	短宽叶 Short and wide leaves	[51]
AVB （LOC_Os03g19520）	3	未知功能蛋白 Unknown function protein	维持细胞分裂 Maintain cell division	细胞核、细胞质 Nucleus, cytoplasm	窄卷叶 Narrow and rolled leaves	[52]
CCC1 （LOC_Os08g23440）	8	阳离子-氯离子协同转运蛋白 Cation-chloride cotransporter	通过调控离子平衡维持细胞渗透势 Maintain cellular osmotic potential by regulating ion homeostasis	细胞膜 Cell membrane	窄叶 Narrow leaves	[53]
MKB3 （LOC_Os03g52320）	3	GRF互作因子 GRF-interacting factor	正调控细胞增殖和细胞大小 Positively regulate cell proliferation and cell size	细胞核 Nucleus	短窄叶 Short and narrow leaves	[54-55]
NLG1 （LOC_Os03g14890）	3	编码TIM23亚基 Encodes subunits of TIM23	维持线粒体代谢 Maintain mitochondrion metabolism	线粒体 Mitochondrion	窄叶、矮化 Narrow leaves, dwarf	[56]
NAL8 （LOC_Os07g15880）	7	编码PHB亚基 Encodes subunits of PHB	维持线粒体和叶绿体稳定性 Maintain mitochondrion and chloroplast stability	线粒体、叶绿体 Mitochondrion, chloroplast	窄叶、矮化 Narrow leaves, dwarf	[57]