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Rice Science ›› 2023, Vol. 30 ›› Issue (5): 379-404.DOI: 10.1016/j.rsci.2023.03.014

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  • 收稿日期:2022-07-05 接受日期:2023-03-08 出版日期:2023-09-28 发布日期:2023-08-14

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. [J]. Rice Science, 2023, 30(5): 379-404.

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链接本文: http://www.ricesci.org/CN/10.1016/j.rsci.2023.03.014

               http://www.ricesci.org/CN/Y2023/V30/I5/379

图/表 13

Table 1. Main regulators in G protein signaling for rice grain shape.
Gene ID Protein category Regulatory role Effect on cell Reference
Length Width Thickness Weight
RGA1 Os05g0333200 Gα subunit + None / + Proliferation Oki et al, 2005, 2009
GS3 Os03g0407400 Atypical Gγ subunit − + − − Proliferation
and elongation
Fan et al, 2006, 2009; Takano-Kai et al, 2009; Mao et al, 2010
DEP1 Os09g0441900 Atypical Gγ subunit + / / / / Huang et al, 2009; Zhou et al, 2009; Sun et al, 2014
RGB1 Os03g0669100 Gβ subunit + / / / Proliferation Utsunomiya et al, 2011
GGC2 Os08g0456600 Atypical Gγ subunit + / / / / Sun et al, 2014
RGG1 Os03g0635100 Typical Gγ subunit − − / − Proliferation Sun et al, 2014; Tao et al, 2020
RGG2 Os02g0137900 Typical Gγ subunit − − − − Expansion Sun et al, 2014; Miao et al, 2019
OsMADS1/OsLG3b Os03g0215400 Transcription factor downstream of GS3 and DEP1 + / + + Proliferation Liu et al, 2018; Yu et al, 2018

Table 1. Main regulators in G protein signaling for rice grain shape.

Gene ID Protein category Regulatory role Effect on cell Reference
Length Width Thickness Weight
RGA1 Os05g0333200 Gα subunit + None / + Proliferation Oki et al, 2005, 2009
GS3 Os03g0407400 Atypical Gγ subunit − + − − Proliferation
and elongation
Fan et al, 2006, 2009; Takano-Kai et al, 2009; Mao et al, 2010
DEP1 Os09g0441900 Atypical Gγ subunit + / / / / Huang et al, 2009; Zhou et al, 2009; Sun et al, 2014
RGB1 Os03g0669100 Gβ subunit + / / / Proliferation Utsunomiya et al, 2011
GGC2 Os08g0456600 Atypical Gγ subunit + / / / / Sun et al, 2014
RGG1 Os03g0635100 Typical Gγ subunit − − / − Proliferation Sun et al, 2014; Tao et al, 2020
RGG2 Os02g0137900 Typical Gγ subunit − − − − Expansion Sun et al, 2014; Miao et al, 2019
OsMADS1/OsLG3b Os03g0215400 Transcription factor downstream of GS3 and DEP1 + / + + Proliferation Liu et al, 2018; Yu et al, 2018
Table 2. Main regulators in ubiquitination-related pathway for rice grain shape.
Gene ID Protein category Regulatory role Effect on cell Reference
Length Width Thickness Weight
GW2 Os02g0244100 RING-type E3 ubiquitin ligase − − − − Proliferation and expansion Song et al, 2007; Choi et al, 2018; Verma et al, 2021
HGW Os06g0160400 Ubiquitin-associated domain protein None + + / Division and elongation Li et al, 2012
WTG1/
OsOTUB1
Os08g0537800 Deubiquitinating enzyme + − − − Proliferation and expansion Huang et al, 2017; Wang
et al, 2017
OsUBP15/LG1 Os02g0244300 Deubiquitinating enzyme + + + + Proliferation Shi et al, 2019
bZIP47 Os06g0265400 bZIP transcription factor + − / − Proliferation Hao et al, 2021
OsUBC13 Os01g0673600 E2 ubiquitin-conjugating protein / + / + / Hao et al, 2021
WG1/OsGRX8 Os02g0512400 Glutaredoxin protein/target of GW2 − + / + Proliferation Hao et al, 2021

Table 2. Main regulators in ubiquitination-related pathway for rice grain shape.

Gene ID Protein category Regulatory role Effect on cell Reference
Length Width Thickness Weight
GW2 Os02g0244100 RING-type E3 ubiquitin ligase − − − − Proliferation and expansion Song et al, 2007; Choi et al, 2018; Verma et al, 2021
HGW Os06g0160400 Ubiquitin-associated domain protein None + + / Division and elongation Li et al, 2012
WTG1/
OsOTUB1
Os08g0537800 Deubiquitinating enzyme + − − − Proliferation and expansion Huang et al, 2017; Wang
et al, 2017
OsUBP15/LG1 Os02g0244300 Deubiquitinating enzyme + + + + Proliferation Shi et al, 2019
bZIP47 Os06g0265400 bZIP transcription factor + − / − Proliferation Hao et al, 2021
OsUBC13 Os01g0673600 E2 ubiquitin-conjugating protein / + / + / Hao et al, 2021
WG1/OsGRX8 Os02g0512400 Glutaredoxin protein/target of GW2 − + / + Proliferation Hao et al, 2021
Fig. 1. Control of grain shape by ubiquitination-related pathway. GW2, a RING protein with E3 ubiquitin (Ub) ligase activity, interacts and catalyzes the ubiquitination of EXPLA1 and WG1/OsGRX8. EXPLA1 is a cell wall-loosening protein that increases cell expansion, while WG1/OsGRX8 acts as an adaptor protein to recruit the transcriptional co-repressor ASP1 to repress the transcription activity of OsbZIP47. OsbZIP47 limits grain growth by inhibiting cell proliferation. The deubiquitinase WTG1/OsTUB1 forms complex with E2 conjugating enzyme OsUBC13 to regulate grain width, maybe through promoting the ubiquitination-dependent proteasomal degradation of OsSPL14, and thus longitudinally increases cell length and transversely decreases cell width. Another deubiquitinase OsUBP15 positively regulates cell proliferation to determine grain shape. Moreover, a novel plant-specific ubiquitin-associated domain protein HGW (heading and grain weight) promotes latitudinal cell division to play a role in grain shape determination. Regular arrows indicate positive regulation, while block arrows indicate negative regulation.

Fig. 1. Control of grain shape by ubiquitination-related pathway. GW2, a RING protein with E3 ubiquitin (Ub) ligase activity, interacts and catalyzes the ubiquitination of EXPLA1 and WG1/OsGRX8. EXPLA1 is a cell wall-loosening protein that increases cell expansion, while WG1/OsGRX8 acts as an adaptor protein to recruit the transcriptional co-repressor ASP1 to repress the transcription activity of OsbZIP47. OsbZIP47 limits grain growth by inhibiting cell proliferation. The deubiquitinase WTG1/OsTUB1 forms complex with E2 conjugating enzyme OsUBC13 to regulate grain width, maybe through promoting the ubiquitination-dependent proteasomal degradation of OsSPL14, and thus longitudinally increases cell length and transversely decreases cell width. Another deubiquitinase OsUBP15 positively regulates cell proliferation to determine grain shape. Moreover, a novel plant-specific ubiquitin-associated domain protein HGW (heading and grain weight) promotes latitudinal cell division to play a role in grain shape determination. Regular arrows indicate positive regulation, while block arrows indicate negative regulation.

Table 3. Main regulators in mitogen-activated protein kinase (MAPK) signaling pathway for rice grain shape.
Gene ID Protein category Regulatory role Effect on cell Reference
Length Width Thickness Weight
OsMKK4 Os02g0787300 MAPK kinase + + / + Proliferation Duan et al, 2014; Xu et al, 2018
OsMAPK6 Os06g0154500 MAPK + + / + Proliferation Liu S Y et al, 2015; Xu et al, 2018
OsWRKY53 Os05g0343400 Substrate of OsMAPK6 + + / / Expansion Tian et al, 2017, 2021
GSN1/OsMKP1 Os05g0115800 MAPK phosphatase − − / − Proliferation Guo et al, 2018
OsMKKK10 Os04g0559800 MAPK kinase kinase + + / + Proliferation Xu et al, 2018

Table 3. Main regulators in mitogen-activated protein kinase (MAPK) signaling pathway for rice grain shape.

Gene ID Protein category Regulatory role Effect on cell Reference
Length Width Thickness Weight
OsMKK4 Os02g0787300 MAPK kinase + + / + Proliferation Duan et al, 2014; Xu et al, 2018
OsMAPK6 Os06g0154500 MAPK + + / + Proliferation Liu S Y et al, 2015; Xu et al, 2018
OsWRKY53 Os05g0343400 Substrate of OsMAPK6 + + / / Expansion Tian et al, 2017, 2021
GSN1/OsMKP1 Os05g0115800 MAPK phosphatase − − / − Proliferation Guo et al, 2018
OsMKKK10 Os04g0559800 MAPK kinase kinase + + / + Proliferation Xu et al, 2018
Fig. 2. Control of grain shape by brassinosteroid (BR) and mitogen-activated protein kinase (MAPK) signaling pathway. The rate limiting enzymes in BR synthesis, including SLG, D2/CYP90D2, DWARF11, DWARF4 and brd1/OsBR6ox/OsDWARF, are involved in grain shape determination through regulation of BR homeostasis. The BR signal is perceived by OsBRI1 and its co-receptor OsBAK1 and transduced to OsBSK3, which phosphorylates an unidentified phosphatase and further represses OsGSK2/OsSK22 and OsGSK3. OsGSK2 and OsGSK3 repress BR signaling by negatively regulating the transcription factors including OsBZR1, DLT, GL2/GS2/OsGRF4, OsWRKY53 and OFPs. OFPs repress the transcription activity of GS9. A protein phosphatase GL3.1/OsPPKL1 dephosphorylates but stabilizes OsGSK3, leading to accumulation of phosphorylated OsBZR1, which cannot activate BR-induced genes and thus suppressing BR signaling. GL3.1/OsPPKL1 also induces the phosphorylation of OsGF14b, which inhibits OsBZR1 to translocate to nucleus. OsGSK2 phosphorylates OsMKK4 to suppress OsMAPK6 activity, while the phosphorylation of OsWRKY53 by GSK2 lowers OsWRKY53 protein stability to regulate grain shape. GSN1/OsMKP1 inactivates OsMAPK6 via dephosphorylation and thus negatively regulates grain length and width formation. Regular arrows indicate positive regulation, while block arrows indicate negative regulation.

Fig. 2. Control of grain shape by brassinosteroid (BR) and mitogen-activated protein kinase (MAPK) signaling pathway. The rate limiting enzymes in BR synthesis, including SLG, D2/CYP90D2, DWARF11, DWARF4 and brd1/OsBR6ox/OsDWARF, are involved in grain shape determination through regulation of BR homeostasis. The BR signal is perceived by OsBRI1 and its co-receptor OsBAK1 and transduced to OsBSK3, which phosphorylates an unidentified phosphatase and further represses OsGSK2/OsSK22 and OsGSK3. OsGSK2 and OsGSK3 repress BR signaling by negatively regulating the transcription factors including OsBZR1, DLT, GL2/GS2/OsGRF4, OsWRKY53 and OFPs. OFPs repress the transcription activity of GS9. A protein phosphatase GL3.1/OsPPKL1 dephosphorylates but stabilizes OsGSK3, leading to accumulation of phosphorylated OsBZR1, which cannot activate BR-induced genes and thus suppressing BR signaling. GL3.1/OsPPKL1 also induces the phosphorylation of OsGF14b, which inhibits OsBZR1 to translocate to nucleus. OsGSK2 phosphorylates OsMKK4 to suppress OsMAPK6 activity, while the phosphorylation of OsWRKY53 by GSK2 lowers OsWRKY53 protein stability to regulate grain shape. GSN1/OsMKP1 inactivates OsMAPK6 via dephosphorylation and thus negatively regulates grain length and width formation. Regular arrows indicate positive regulation, while block arrows indicate negative regulation.

Table 4. Main regulators in microRNA process for rice grain shape.
Gene ID Protein category Regulatory role Effect on cell Reference
Length Width Thickness Weight
GW8/OsSPL16 Os08g0531600 Target of miR156 − + / − Proliferation and elongation Wang et al, 2012
GLW7/OsSPL13 Os07g0505200 Transcription factor, target of miR156 + None + + Expansion Si et al, 2016
OsmiR396e Os04g0674500 miRNA396 − − − − Elongation and expansion Miao et al, 2020; Yang et al, 2021
OsmiR396f Os02g0829000 miRNA396 − − − − Elongation and expansion Miao et al, 2020; Yang et al, 2021
SPL4 Os02g0829000 Target of miR156 − − + − Elongation and division Hu et al, 2021
OsmiR167a miRNA167 + / / + Elongation Qiao et al, 2021
OsmiR529a miRNA529 + − / − / Yan et al, 2021
OsmiR408 miRNA408 + + / + Elongation Yang et al, 2021

Table 4. Main regulators in microRNA process for rice grain shape.

Gene ID Protein category Regulatory role Effect on cell Reference
Length Width Thickness Weight
GW8/OsSPL16 Os08g0531600 Target of miR156 − + / − Proliferation and elongation Wang et al, 2012
GLW7/OsSPL13 Os07g0505200 Transcription factor, target of miR156 + None + + Expansion Si et al, 2016
OsmiR396e Os04g0674500 miRNA396 − − − − Elongation and expansion Miao et al, 2020; Yang et al, 2021
OsmiR396f Os02g0829000 miRNA396 − − − − Elongation and expansion Miao et al, 2020; Yang et al, 2021
SPL4 Os02g0829000 Target of miR156 − − + − Elongation and division Hu et al, 2021
OsmiR167a miRNA167 + / / + Elongation Qiao et al, 2021
OsmiR529a miRNA529 + − / − / Yan et al, 2021
OsmiR408 miRNA408 + + / + Elongation Yang et al, 2021
Fig. 3. MicroRNAs control rice grain shape by targeting key transcription factor-encoding genes. MicroRNA156 (miR156) targets and down-regulates the genes encoding transcription factors OsSPL4, GW8/OsSPL16 and GLW7/OsSPL13. GLW7/OsSPL13 promotes the gene expression of SRS5. miR167a targets and inhibits OsARF6, which encodes an upstream transcription factor promoting the expression of OsAUX3 in the auxin signaling pathway. miR396 directs OsGRF4/GL2/GS2 and OsGRF8 mRNA silencing. OsGRF4/GL2/GS2 is an important cross-connect node of brassinosteroid (BR) signaling and miRNA regulatory pathway to determine the grain shape. OsGRF8 directly binds to the OsmiR408 promoter and may promote the accumulation of OsmiR408. Moreover, miR529a is involved in the grain shape determination by directing other OsSPL family genes. Regular arrows indicate positive regulation, while block arrows indicate negative regulation.

Fig. 3. MicroRNAs control rice grain shape by targeting key transcription factor-encoding genes. MicroRNA156 (miR156) targets and down-regulates the genes encoding transcription factors OsSPL4, GW8/OsSPL16 and GLW7/OsSPL13. GLW7/OsSPL13 promotes the gene expression of SRS5. miR167a targets and inhibits OsARF6, which encodes an upstream transcription factor promoting the expression of OsAUX3 in the auxin signaling pathway. miR396 directs OsGRF4/GL2/GS2 and OsGRF8 mRNA silencing. OsGRF4/GL2/GS2 is an important cross-connect node of brassinosteroid (BR) signaling and miRNA regulatory pathway to determine the grain shape. OsGRF8 directly binds to the OsmiR408 promoter and may promote the accumulation of OsmiR408. Moreover, miR529a is involved in the grain shape determination by directing other OsSPL family genes. Regular arrows indicate positive regulation, while block arrows indicate negative regulation.

Table 5. Main regulators in brassinosteroid (BR) biosynthesis and signaling pathways for rice grain shape.
Gene ID Protein category Regulatory role Effect on cell Reference
Length Width Thickness Weight
brd1/OsBR6ox/
OsDWARF
Os03g0602300 BR-6-oxidase + + + + / Mori et al, 2002
D2 Os01g0197100 Cytochrome P450 CYP90D2 + + + + / Hong et al, 2003
D11 Os04g0469800 Cytochrome P450 CYP724B1 + + + + / Tanabe et al, 2005; Zhou et al, 2017
OsBRI1/D61 Os01g0718300 BR receptor + + None + / Morinaka et al, 2006; Zhao et al, 2013
GW5/GSE5 Os05g0187500 Calmodulin binding protein − − − − Proliferation Wan et al, 2008; Weng et al, 2008; Duan et al, 2017; Liu et al, 2017
GS5 Os05g0158500 Putative serine carboxypeptidase None + / + Division and expansion Li et al, 2011; Xu et al, 2015
GL3.1/OsPPKL1 Os03g0646900 Phosphatase kelch (PPKL) family-Ser/Thr phosphatase + + + + Proliferation Hu et al, 2012; Qi et al, 2012; Zhang X J et al, 2012
GL2/GS2/
OsGRF4/GLW2
Os02g0701300 Transcription factor − − + − Division and expansion Che et al, 2015; Duan et al, 2015; Hu et al, 2015; Li S C et al, 2016
SLG Os08g0562500 BAHD aceltransferase-like protein + - / + Elongation Feng et al, 2016
GS9 Os09g0448500 Transcription factor − + None None Proliferation Zhao et al, 2018

Table 5. Main regulators in brassinosteroid (BR) biosynthesis and signaling pathways for rice grain shape.

Gene ID Protein category Regulatory role Effect on cell Reference
Length Width Thickness Weight
brd1/OsBR6ox/
OsDWARF
Os03g0602300 BR-6-oxidase + + + + / Mori et al, 2002
D2 Os01g0197100 Cytochrome P450 CYP90D2 + + + + / Hong et al, 2003
D11 Os04g0469800 Cytochrome P450 CYP724B1 + + + + / Tanabe et al, 2005; Zhou et al, 2017
OsBRI1/D61 Os01g0718300 BR receptor + + None + / Morinaka et al, 2006; Zhao et al, 2013
GW5/GSE5 Os05g0187500 Calmodulin binding protein − − − − Proliferation Wan et al, 2008; Weng et al, 2008; Duan et al, 2017; Liu et al, 2017
GS5 Os05g0158500 Putative serine carboxypeptidase None + / + Division and expansion Li et al, 2011; Xu et al, 2015
GL3.1/OsPPKL1 Os03g0646900 Phosphatase kelch (PPKL) family-Ser/Thr phosphatase + + + + Proliferation Hu et al, 2012; Qi et al, 2012; Zhang X J et al, 2012
GL2/GS2/
OsGRF4/GLW2
Os02g0701300 Transcription factor − − + − Division and expansion Che et al, 2015; Duan et al, 2015; Hu et al, 2015; Li S C et al, 2016
SLG Os08g0562500 BAHD aceltransferase-like protein + - / + Elongation Feng et al, 2016
GS9 Os09g0448500 Transcription factor − + None None Proliferation Zhao et al, 2018
Table 6. Main regulators in auxin biosynthesis and signaling pathway for rice grain shape.
Gene ID Protein category Regulatory role Effect on cell Reference
Length Width Thickness Weight
TGW6 Os06g0623700 Protein with IAA-glucose hydrolase activity − None None − Proliferation and elongation Ishimaru et al, 2013
OsARF4 Os01g0927600 Transcription factor in auxin signaling − − None − Expansion Hu et al, 2018
OsSK41/OsGSK5/GL3.3 Os03g0841800 GSK-like family member − − / − Proliferation and elongation Hu et al, 2018; Xia
et al, 2018
qGL5/OsAUX3 Os05g0447200 Target genes in auxin signaling − None / − Expansion Qiao et al, 2021

Table 6. Main regulators in auxin biosynthesis and signaling pathway for rice grain shape.

Gene ID Protein category Regulatory role Effect on cell Reference
Length Width Thickness Weight
TGW6 Os06g0623700 Protein with IAA-glucose hydrolase activity − None None − Proliferation and elongation Ishimaru et al, 2013
OsARF4 Os01g0927600 Transcription factor in auxin signaling − − None − Expansion Hu et al, 2018
OsSK41/OsGSK5/GL3.3 Os03g0841800 GSK-like family member − − / − Proliferation and elongation Hu et al, 2018; Xia
et al, 2018
qGL5/OsAUX3 Os05g0447200 Target genes in auxin signaling − None / − Expansion Qiao et al, 2021
Table 7. Other plant hormones regulation for rice grain shape.
Gene
ID Protein category Regulatory role Effect on cell Reference
Length Width Thickness Weight
OsCKX4 Os01g0940000 Cytokinin oxidase/dehydrogenase − − / − Division Gao et al, 2014
OsVQ13 Os03g0100020 JA responsive protein + + / / / Uji et al, 2019
BG3/OsPUP4 Os01g0680200 Purine permease 4 + + + + Division Xiao et al, 2019
OsPUP7 Os05g0556800 Purine permease 4 + + / + Division Xiao et al, 2019
GW6 Os06g0266800 GA-regulated GAST family protein + + / + Expansion Shi et al, 2020
OsPUP1 Os03g0187800 Purine permease 1 None − / - Division Xiao et al, 2020a, b
AGO2 Os04g0615700 ARGONAUTE protein + None / + Division Yin et al, 2020

Table 7. Other plant hormones regulation for rice grain shape.

Gene
ID Protein category Regulatory role Effect on cell Reference
Length Width Thickness Weight
OsCKX4 Os01g0940000 Cytokinin oxidase/dehydrogenase − − / − Division Gao et al, 2014
OsVQ13 Os03g0100020 JA responsive protein + + / / / Uji et al, 2019
BG3/OsPUP4 Os01g0680200 Purine permease 4 + + + + Division Xiao et al, 2019
OsPUP7 Os05g0556800 Purine permease 4 + + / + Division Xiao et al, 2019
GW6 Os06g0266800 GA-regulated GAST family protein + + / + Expansion Shi et al, 2020
OsPUP1 Os03g0187800 Purine permease 1 None − / - Division Xiao et al, 2020a, b
AGO2 Os04g0615700 ARGONAUTE protein + None / + Division Yin et al, 2020
Table 8. Main regulators in other transcriptional regulatory pathway for rice grain shape.
Gene ID Protein category Regulatory role Effect on cell Reference
Length Width Thickness Weight
OsGIF1 Os11g0615200 Transcript co-activator + / / + Proliferation Duan et al, 2015; Li S C et al, 2016
OsGIF2 Os12g0496900 Transcript co-activator + / / + Proliferation Duan et al, 2015; Li S C et al, 2016
GW7/GL7/SLG7 Os07g0603300 TONNEAU1-recruiting motif protein + − + + Proliferation Wang S K et al, 2015; Wang Y X et al, 2015; Zhou et al, 2015
OsFD2 Os06g0720900 bZIP transcription factor − − − − Expansion and division He et al, 2022

Table 8. Main regulators in other transcriptional regulatory pathway for rice grain shape.

Gene ID Protein category Regulatory role Effect on cell Reference
Length Width Thickness Weight
OsGIF1 Os11g0615200 Transcript co-activator + / / + Proliferation Duan et al, 2015; Li S C et al, 2016
OsGIF2 Os12g0496900 Transcript co-activator + / / + Proliferation Duan et al, 2015; Li S C et al, 2016
GW7/GL7/SLG7 Os07g0603300 TONNEAU1-recruiting motif protein + − + + Proliferation Wang S K et al, 2015; Wang Y X et al, 2015; Zhou et al, 2015
OsFD2 Os06g0720900 bZIP transcription factor − − − − Expansion and division He et al, 2022
Table 9. Other regulators in rice grain shape determination.
Gene ID Protein category Regulatory role Effect on cell Reference
Length Width Thickness Weight
Cyclin-T1;3 Os11g0157100 Cell cycle regulator + / / / Proliferation Qi et al, 2012
FUWA Os02g0234200 NHL domain-containing protein + − + − Division Chen et al, 2015
GW6a/OsglHAT1 Os06g0650300 Histone H4 acetyltransferase / / / + / Song et al, 2015
GF14f Os03g0710800 14-3-3 protein − None None − Expansion and division Zhang et al, 2019
MIS2/OsCR4 Os03g0637800 Receptor-like kinase + + + + Expansion and division Chun et al, 2020
qTGW12a Os12g0552600 MATE transporter / + / + / Du et al, 2021
OsDDM1b Os03g0722400 Subunit of chromatin remoeling complexes + + / / Proliferation Guo et al, 2022
OsCEN2 Os11g0152500 CENTRORADIALIS family member − − − − Expansion He et al, 2022
SMG3 Os03g0410000 MYB-like protein − − / − Elongation and
proliferation
Li et al, 2022
qGSN5 Os05g054860 P-type PPR protein + + / + Proliferation Yuan et al, 2022a
SDR7-6 Os07g0663900 Short-chain alcohol dehydrogenase/reductase None + + None / Zheng et al, 2022

Table 9. Other regulators in rice grain shape determination.

Gene ID Protein category Regulatory role Effect on cell Reference
Length Width Thickness Weight
Cyclin-T1;3 Os11g0157100 Cell cycle regulator + / / / Proliferation Qi et al, 2012
FUWA Os02g0234200 NHL domain-containing protein + − + − Division Chen et al, 2015
GW6a/OsglHAT1 Os06g0650300 Histone H4 acetyltransferase / / / + / Song et al, 2015
GF14f Os03g0710800 14-3-3 protein − None None − Expansion and division Zhang et al, 2019
MIS2/OsCR4 Os03g0637800 Receptor-like kinase + + + + Expansion and division Chun et al, 2020
qTGW12a Os12g0552600 MATE transporter / + / + / Du et al, 2021
OsDDM1b Os03g0722400 Subunit of chromatin remoeling complexes + + / / Proliferation Guo et al, 2022
OsCEN2 Os11g0152500 CENTRORADIALIS family member − − − − Expansion He et al, 2022
SMG3 Os03g0410000 MYB-like protein − − / − Elongation and
proliferation
Li et al, 2022
qGSN5 Os05g054860 P-type PPR protein + + / + Proliferation Yuan et al, 2022a
SDR7-6 Os07g0663900 Short-chain alcohol dehydrogenase/reductase None + + None / Zheng et al, 2022
Fig. 4. Main genes regulating three-dimension of grain shape and their potential value in mechanized production of hybrid seeds. Genes with regulatory roles in the formation of grain length, grain width and grain thickness are highlighted in green, orange and purple, respectively. Red arrows indicate positive regulation, and blue block arrows indicate negative regulation. Red rectangles frame the genes with positive roles while blue rectangles frame genes with negative roles. The cylinder sorting, sieve sorting, gravity sorting systems can be used to separate hybrid rice seeds in dependence of grain length, grain thickness, and grain weight differences, respectively.

Fig. 4. Main genes regulating three-dimension of grain shape and their potential value in mechanized production of hybrid seeds. Genes with regulatory roles in the formation of grain length, grain width and grain thickness are highlighted in green, orange and purple, respectively. Red arrows indicate positive regulation, and blue block arrows indicate negative regulation. Red rectangles frame the genes with positive roles while blue rectangles frame genes with negative roles. The cylinder sorting, sieve sorting, gravity sorting systems can be used to separate hybrid rice seeds in dependence of grain length, grain thickness, and grain weight differences, respectively.

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