Figures (3)  Tables (2)
    • Figure 1. 

      Pleiotropy of Wx gene in rice quality regulation. This pattern diagram presents the regulatory factors of the Wx gene at both the transcriptional and post-transcriptional levels that have been reported thus far. It also highlights the encoding product GBSSI of the Wx gene, which plays a role in regulating the three major components of rice endosperm (starch, protein, and lipids) and subsequently affects various physicochemical properties of rice grains. The pleiotropic effect of the Wx gene ultimately determines the overall quality of rice from multiple perspectives. GC, gel consistency; GT, gelatinization temperature. Yellow spheres with the letter P indicate phosphorylation.

    • Figure 2. 

      Determinants of cooking and eating quality of rice and external factors affecting protein content of rice grains. The six parts in the inner circle represent the physical and chemical characteristics or indicators that determine the cooking and eating quality of rice; the three parts in the outer circle represent the material composition that affects the physical and chemical characteristics of rice; the cartoon diagram in the outermost circle represents the ecological factors and environmental factors that affect the grain protein content, from clockwise from 1 o'clock to indicate light, moisture, carbon dioxide, salinity, cultivation, fertilizer and temperature.

    • Figure 3. 

      The nitrogen utilization ability of rice may determine the protein content of grains. Each step of nitrogen utilization in rice, which includes absorption, transportation, assimilation, distribution, and reuse, can potentially impact the final grain protein content.

    • ClassificationNameGene IDEffect of gene knockout
      or knockdown on grain
      protein content
      Annotation
      Biochemical metabolismflo4/OsPPDKB/OsC4PPDKLOC_Os05g33570Increased[130]Pyruvate, phosphate dikinase
      OsSSIIIa/Flo5/SS3aLOC_Os08g09230Increased[103]Starch synthase III
      FLO8/OsUgp1LOC_Os09g38030Increased[131]UTP--glucose-1-phosphate uridylyltransferase
      FLO12/OsAlaAT1LOC_Os10g25130Increased[132,133]Aminotransferase
      FLO15/OsGLYI7LOC_Os05g14194Increased[134]Glyoxalase family protein
      FLO16LOC_Os10g33800Increased[135]lactate/malate dehydrogenase
      FSE1LOC_Os08g01920Increased[136]Phospholipase-like protein
      OsAGPL2/OsAPL2/shr1/GIF2/osagpl2-3LOC_Os01g44220Decreased[137]ADP-glucose pyrophosphorylase large subunit 2
      OsBEIIb/be2bLOC_Os02g32660Unknown[138]Starch branching enzyme IIb
      Pho1LOC_Os03g55090Unknown[139]Plastidial phosphorylase
      OsGINT1/FSE6LOC_Os05g46260Increased[140]Glycosyltransferase
      OsPK2/OsPKpα1LOC_Os07g08340Increased[141]Plastidic pyruvate kinase
      GIF1/OsCIN2LOC_Os04g33740Unknown[142]Glycosyl hydrolases
      PDIL1-1LOC_Os11g09280Increased[143]Protein disulphide isomerase-like enzyme
      OsACS6/SSG6LOC_Os06g03990Unknown[144]Aminotransferase
      PFPβ/PFP1LOC_Os06g13810Unknown[145]Pyrophosphate-fructose 6-phosphate
      1-phosphotransferase subunit beta
      FLO19LOC_Os03g48060Increased[28]Class I glutamine amidotransferase
      FLO19LOC_Os04g02900Decreased[146]Plastid-localized pyruvate dehydrogenase complex E1 component subunit α1
      FLO23/OsF2KP2LOC_Os03g18310Decreased[147]Fructose-6-phosphate-2-kinase/fructose-2, 6-bisphosphatase
      OsDPE1LOC_Os07g43390Unknown[148]Disproportionating enzyme
      Transcriptional regulation and protein interactionOsNF-YC12LOC_Os10g11580Increased[19,149]CCAAT-box-binding transcription factor
      OsNF-YB1/OsHAP3K/OsEnS-41LOC_Os02g49410Increased[19]Nuclear transcription factor Y subunit B
      bHLH144LOC_Os04g35010Increased[19]Helix-loop-helix DNA-binding domain containing protein
      RISBZ1/OsbZIP58LOC_Os07g08420Decreased[150]bZIP transcription factor
      REB/OsbZIP33/RISBZ2LOC_Os03g58250Unknown[151]bZIP transcription factor
      RPBF/OsDof3/OsDof-10/OsDof7LOC_Os02g15350Decreased[150,152]Dof transcription factor
      FLO2LOC_Os04g55230Unknown[153]Tetratricopeptide repeat domain containing protein
      FLO6LOC_Os03g48170Increased[154]CBM48 domain-containing protein
      OsGBPLOC_Os02g04330No change[29]GBSS-binding protein
      FLO7LOC_Os10g32680Unknown[155]DUF1388 domain protein
      OsHsp70cp-2/cpHSP70-2/ flo11-2/ FLO11LOC_Os12g14070Unknown[156]Plastid heat shock protein 70
      RSR1LOC_Os05g03040Unknown[157]Transcription factor of the AP2/EREBP family
      OsNAC20; OsNAC26LOC_Os01g01470; LOC_Os01g29840Decreased when knock out together[18]NAC transcription factor
      OsNAC23/ONAC023LOC_Os02g12310Decreased when gene knock out and increased when gene overexpression[158]NAC transcription factor
      OsNAC24/OsNAC024LOC_Os05g34310Unknown[17]NAC transcription factor
      OsNAC127LOC_Os11g31340Unknown[159]NAC transcription factor
      OsNAC129LOC_Os11g31380Unknown[159,160]NAC transcription factor
      Du13/TL1LOC_Os06g48530No change[12]C2H2 zinc finger protein
      OsbZIP60/O3/OPAQUE3LOC_Os07g44950Decreased[24]Basic leucine zipper transcription factor
      OsMADS6/MFO1LOC_Os02g45770Increased[161]MADS-box transcription factor
      OsMADS29LOC_Os02g07430Unknown[162]MADS-box transcription factor
      OsMADS14LOC_Os03g54160Unknown[21]MADS-box transcription factor
      EpigeneticsOsROS1/ROS1a/DNG702LOC_Os01g11900Increased[163]DNA demethylase
      OsCADT1/FLO20/SHMT4LOC_Os01g65410Decreased[164]Serine hydroxymethyltransferase
      Energy supplyFLO10LOC_Os03g07220Increased[165]Pentatricopeptide repeat protein
      FLO14/OsNPPR3LOC_Os03g51840Unknown[166]Pentatricopeptide repeat protein
      FLO18LOC_Os07g48850Decreased[167]Pentatricopeptide repeat protein
      OGR1LOC_Os12g17080Unknown[168]Pentatricopeptide repeat–DYW protein
      FGR1/OsNPPR1LOC_Os08g19310Unknown[169]Pentatricopeptide repeat protein
      FLO13/OsNDUFA9LOC_Os02g57180Unknown[169]Mitochondrial complex I subunit
      FLO22LOC_Os07g08180Unknown[170]P-type pentatricopeptide repeat (PPR) protein
      Material transportESG1LOC_Os04g46700Decreased[171]Bcterial-type ABC (ATP-binding cassette) lipid transporter
      OsBT1LOC_Os02g10800Unknown[172]ADP-Glucose Transporter
      OsBip1/BiP3LOC_Os02g02410Decreased[173]Endoplasmic riculum caperone
      OsLTPL36LOC_Os03g25350Decreased[174]Lipid transfer protein
      OsRab5a/gpa1/glup4LOC_Os12g43550No change, but pro-glutelin accumulation[175]Small GTPase
      Sar1a; Sar1b; Sar1cLOC_Os01g23620; LOC_Os12g37360; LOC_Os01g15010Unknown but pro-glutelin accumulation when three genes knockdown together[176]Small GTPase
      GPA3LOC_Os03g61950Increased and pro-glutelin accumulation[177]Regulator of post-Golgi vesicular Traffic
      GPA4/GLUP2/GOT1BLOC_Os03g11100Decreased but pro-glutelin accumulation[178]Golgi Transport 1
      GPA5LOC_Os06g43560Unknown but pro-glutelin accumulation[179]Rab5a Effector
      OsVPS9A/GPA2/GLUP6/GEFLOC_Os03g15650Decreased but pro-glutelin accumulation[180]Guanine nucleotide exchange factor
      GPA6/OsNHX5LOC_Os09g11450No change, but pro-glutelin accumulation[181]Vacuolar Na+/H+ antiporter
      Function unknownSSG4LOC_Os01g08420Unknown[182]Unknown function protein

      Table 1. 

      Reported floury endosperm genes that may affect grain protein content.

    • NameGene IDEffect of altered gene function on grain protein contentAnnotation
      MYB61/qNLA1/qCel1LOC_Os01g18240Unknown[183]MYB family transcription factor
      OsGRF4/GS2/GL2/PT2/LGS1/GLW2LOC_Os02g47280Increased when gene overexpression[108]Growth-regulating factor
      TOND1LOC_Os12g43440Unknown[184]Unkonwn function protein
      OsDEP1/DN1/qPE9-1/qNGR9LOC_Os09g26999No change when gene overexpression[185]Gγ subunit
      OsTCP19LOC_Os06g12230Unknown[186]Class-I TCP transcription factor
      SMOS1/shb/RLA1/NGR5LOC_Os05g32270Unknown[187]GRAS protein
      OsNPF6.1LOC_Os01g01360Unknown[188]Nitrate transporter
      OsNAC42LOC_Os09g32040Unknown[188]No apical meristem protein
      OsNR2/qCR2LOC_Os02g53130Unknown[189]Nitrate reductase
      OsNRT1.1B/OsNPF6.5/qCHR-10LOC_Os10g40600Unknown[190]Peptide transporter PTR2
      Ghd7/E1/Hd4LOC_Os07g15770Unknown[191]CCT motif family protein
      ARE1LOC_Os08g12780Unknown[192]Chloroplast envelope membrane protein
      OsCCA1/OsLHY/Nhd1LOC_Os08g06110Decreased when gene expression enhanced[107]MYB transcription factor
      Ef-cdUnknown[193]Long noncoding RNA
      OsAtg8/OsATG8aLOC_Os07g32800Increased when gene overexpression[194]Autophagy-related protein
      OsPIN9LOC_Os01g58860Unknown[195]Auxin efflux transporter
      OsNLP4LOC_Os09g37710Unknown[196,197]NIN-like protein
      OsNLP3LOC_Os01g13540Unknown[198]NIN-like protein
      OsNLP1LOC_Os03g03900Unknown[199]NIN-like protein
      OsAAP6/qPC1LOC_Os01g65670Increased when gene overexpression[25]Amino acid permease
      OsAAP10LOC_Os02g49060Decreased when gene knock out[200]Amino acid permease
      OsAAP5LOC_Os01g65660Unknown[201]Amino acid permease
      OsAAP3LOC_Os06g36180Increased when gene overexpression[202]Amino acid permease
      OsAAP1LOC_Os07g04180Increased when gene overexpression and decreased when gene interference[203]Amino acid permease
      OsAMT1;1/OsAMT1-1LOC_Os04g43070Unknown[204]High-affinity ammonium transporter
      OsAMT2;1LOC_Os05g39240Unknown[205]Ammonium transporter
      OsGS1/OsGS1;1/OsGLN1;1/λGS28LOC_Os02g50240Decreased when OsGS1;1b overexpression[206]Glutamine synthetase
      OsGS2/OsGLN2/λGS31LOC_Os04g56400Increased when concurrent overexpression of OsGS1 and OsGS2[207]Glutamine synthetase
      OsAMT1;3/OsAMT1.3/OsAMT1;2LOC_Os02g40710Increased when gene overexpression[208]Ammonium transporter
      OsGOGAT1LOC_Os01g48960Increased when gene overexpression[208]Glutamate synthetase 1
      OsAS1/ OsASN1LOC_Os03g18130Increased when gene overexpression[209]Asparagine synthetase
      OsSHM1/OsSHMT1LOC_Os03g52840Unknown[210]Serine hydroxymethyltransferase 1
      OsENOD93-1LOC_Os06g05010Unknown[211]Early nodulin 93 ENOD93 protein
      OsNRT2.3/OsNRT2.3a/OsNRT2.3bLOC_Os01g50820Unknown[212]High-affinity nitrate transporter
      OsNAR2.1LOC_Os02g38230Unknown[213]Partner protein for high-affinity nitrate transport
      OsNRT2.1LOC_Os02g02170Unknown[214]High-affinity nitrate transporter
      OsNRT1.1A/OsNPF6.3LOC_Os08g05910Unknown[215]Nitrate transporter
      DNR1LOC_Os01g08270Decreased when gene knock out[216]Amino transferase
      qSBM1LOC_Os01g65120Unknown[217]Peptide transporter
      OsRBCS2LOC_Os12g17600Unknown[218]Small subunit of Rubisco
      OSA1LOC_Os03g48310Unknown[219]Plasma membrane H+-ATPase
      OsDREB1CLOC_Os06g03670Increased when gene overexpression[111]AP2/EREBP transcription factor
      OsPTR6/OsNPF7.3LOC_Os04g50950Unknown[220]Peptide transporter
      OsMADS1/LHS1/AFO/LGY3/GW3p6LOC_Os03g11614Unknown[221,222]MADS-domain transcription factor

      Table 2. 

      Reported nitrogen use efficiency genes that may also affect grain protein content.