Barkan Lab

Photosynthetic Mutant Library

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Maize Gene NameGene FunctionMaize Locus IDRice OrthologArabidopsis OrthologAlleleSeedling ColorPublications
At-ChlH SALK_062726 
At-THF1GRMZM2G033493NYC4 , Os07g0558500At2g20890 THF1-2WT 
THF1-1pg-cot
RBD1At1g54500  
TESTtesttesttest
Zm-rh3aGRMZM2G415491Os03g61220AT5G26742 Zm-rh3a-1
Zm-rh3a-2
Zm-rh3a-3
Zm-rh3a-4
Zm-rh3a-5
Zm-rh3a
Zm-rh3a
cp33aGRMZM2G026614At3g52380
ppr13GRMZM2G099604Os03g58100At2g02980 ppr13-1ivm 
ppr13-2iv-stripey
uvr8GRMZM2G003565Os04g35570At5g63860
ppr10 PPR protein; activates atpH translation, stabilizes atpH and psaJ mRNAs, binds atpI-atpH and psaJ-rpl33 intergenic regions. Classical name l15 (luteus 15)GRMZM2G177169Os05g19380AT2G18940 ppr10-1vpygm
ppr10-2vpyg
ppr10-3vpyg
ppr10-4
hcf108 assembly of the ATPase complex, BFA1 orthologGRMZM2G089696Os03g55164AT3G29185 hcf108-1spgm
Zm-mterf4 cp mTERF domain protein. Binds in vivo to multiple group II introns and promotes their splicing.GRMZM2G029933Os05g33500At4g02990 Zm-mterf4-2pyg
Zm-mterf4-1iv / iv runty
zm-mterf4
ak2Ankyrin repeat protein (Arab - EMB506)GRMZM2G019838Os06g13000At5g40160 ak2-1iv 
ak2-2iv
zb7AtCLB6, zb7 in maize, 4-hydroxy-3-methylbut-2-enyl diphosphate reductase. ISPH enzyme involved in isoprenoid biosynthesis GRMZM2G027059Os03g52170AT4G34350 zb7-1pyg
zb7
zb7-3iv-vpyg
zb7-mum2vpy-vir
cfm4CRM domain containing proteinGRMZM2G142740Os09g19850At4g29750 cfm4-1iv-runty 
cfm4-2iv-runty
caf1CRS2-associated factor 1; CRM domain protein,chloroplast group II intron splicingGRMZM2G173923Os01g31110AT2G20020 caf1-1iv
caf1-2iv
caf2CRS2-associated factor 2; CRM domain protein. chloroplast group II intron splicingAC199526.5_FGT003 Os01g21990AT1G23400 caf2-1iv
Oil yellow (OY)Chlorophyll synthesis, Magnesium chelatase IGRMZM2G419806Os03g36540AT5G45930, AT4G18480 CHLI-1yel or pyg
OY
OY
chlI-2yel
OY
Zm-chlhChlorophyll synthesis: magnesium-chelatase subunit H. GRMZM2G323024Os03g20700 AT5G13630 gun5-1yel-m
gun5-2vpyg
gun5-3yellow
gun5-4
crs1Chloroplast RNA Splicing 1;founding member of CRM domain family. Binds atpF intron, promotes atpF splicingGRMZM2G078412Os08g27150AT5G16180 crs1-1spg
crs1-2vpyg
crs1-3vpyg
crs1-4pg
crs1-5spg
crs1-6pg
crs2Chloroplast RNA Splicing 2; required for splicing of chloroplast subgroup IIB introns. peptidyl tRNA hydrolase homolog.GRMZM2G132021Os01g04130AT5G16140, AT5G38290 crs2-1iv, vir
crs2-2iv
wtf2Chloroplast group II intron splicing, PORR domain. manuscript in preparationZm00001d013029Os04g24710At5g62990 wtf2-1iv-vir 
wtf2-2iv-vir
wtf2-3vpyg
wtf2-4vpyg
wtf2-5iv-vpyg
wtf2-6ivm
wtf2-7ivm
wtf2-8vpygm
wtf2-9pyg
wtf2-10vpygm
wtf2-11iv
Zm-rps9Chloroplast ribosomal protein S9AC234157.1_FGT002Os03g55930At1g74970 Zm-rps9-1vpyg-vir 
Zm-rps9
ppr53Chloroplast-localized PPR-SMR protein that binds to and stabilizes the 5’ end of a pre-23S rRNA isoform and also promotes the stabilization and translation of a specific processed ndhA mRNA. The absence of PPR53 results in severely chlorotic seedlings, a strong reduction in chloroplast ribosomes and a global loss of chloroplast translation products.GRMZM2G438524Os03g63910AT5G46580 ppr53
ppr53-2iv-vir
ppr53-3iv-vir
ppr53
ppr53-1vpyg-vir
ppr53-4iv-vir
psa2DnaJ-type zinc finger domain protein required for PSI assembly. PSI mutant. (Ortholog of EDA3)GRMZM2G021687Os08g36140AT2G34860 psa2-1spgm
Zm-hcf107HAT domain protein required to stabilize psbH 5? end and promote psbH translation. Binds psbH 5?UTRGRMZM2G121960Os03g19560AT3G17040 Zm-hcf107-1hcf
Zm-hcf107-2hcf
Zm-hcf107-3hcf
Zm-hcf107-4hcf
psb1HCF136 (High chlorophyll fluorescence 136) . Photosystem II assembly factorGRMZM2G102838AT5G23120 psb1-1spg-hcf
psb1-2unknown
psb1-3unknown
Zm-HEMCHEMC ortholog. porphobilinogen deaminase. necrotic seedling phenotype.GRMZM2G026117Os02g07230AT5G08280 Zm-HEMC-1pyg-nec
Zm-HEMC
iv186bLEUCYL-TRNA SYNTHETASE, orthologous to Arabidopsis EMBRYO DEFECTIVE 2369GRMZM5G858471Os01g03020AT4G04350 cps5-1
cps5-2iv
cps5
cps5-3iv
cps5-4vpygm
cps5-5iv-runty
Zm-apg1MPBQ/MSBQ methyltransferase involved in plastoquinone synthesisGRMZM2G082998Os12g42090AT3G63410 Zm-apg1-1pg-grny
Zm-apg1-2pg
Zm-apg1-3pyg
Zm-apg1-4pgm
ppp1Maize Classical Gene: ppp1 (pyrophosphate-energized proton pump1), AtPPa6 (Arabidopsis thaliana pyrophosphorylase 6); inorganic diphosphatase/ pyrophosphataseGRMZM2G090718Os02g52940AT5G09650 ppp1-1pyg 
hcf472Maize ortholog of Arabidopsis Pyg7. Required for PSI assembly. Shen et al, Plant Physiology 2017.GRMZM5G809292AT1G22700 hcf472spg 
Zm-murEMaize ortholog of MurE; PEP associated protein required for PEP-mediated transcription. GRMZM2G009070Os10g40130At1g63680 Zm-mure-1pygm
Zm-mure-2iv
Zm-murE-3
Zm-murE-4pyg-vir
Zm-prin2Maize ortholog of PRIN2; PEP associated protein required for PEP-mediated transcriptionGRMZM2G119906Os09g21460AT1G10522 Zm-prin2-1vpyg
Zm-prin2-2iv
Zm-ptac12Maize ortholog of PTAC12; PEP associated protein required for PEP-mediated transcriptionGRMZM5G897926Os01g56350AT2G34640 Zm-ptac12-1vpygm
Zm-ptac12-2vpyg
Zm-ptac12
undefinedundefined
undefinedundefined
Zm-ptac2Maize ortholog of PTAC2; PEP associated protein required for PEP-mediated transcriptionGRMZM2G122116Os03g60910At1g74850 Zm-ptac2-1vpyg
Zm-ptac2-2iv
Zm-ptac2-3iv
pet2Mediates heme attachment to cytochrome b6 (ortholog CCB2)GRMZM2G087063Os10g37840AT5G52110 pet2-1pyg
pet2-2pyg
pet2-3pyg
pet2-4pg
pet2-5spg
pet2-6pyg
wtf4No phenotype.GRMZM2G350157Os06g15550At2g31290 wtf4-1no phenotype 
wtf4-2no phenotype
wtf4-3no phenotype
wtf4-4no phenotype
wtf4
Zm-ptac17No phenotype. (Ortholog of PTAC17). RIP-chip was done together with mTERF4 long ago. Came down in HCF173 coIP from thylakoid. GRMZM2G180418Os02g55630At1g80480 , At1g15730 Zm-ptac17-1no phenotype 
orrm1ORRM-domain protein, involved in cp RNA editingGRMZM5G899787Os02g49820AT3G20930 orrm1-1spg
orrm1-2spg
orrm1-3spg
w2Organellar DNA polymerase gamma; Required for cpDNA replication.GRMZM2G480171Os08g07840, Os08g07850AT1G50840, AT3G20540 w2-mum1vpyg
w2-mum2ivory-runty
w2-Burnhamivory
Zm-tab2Ortholog of ATAB2 in Arabidopsis and TAB2 in Chlamydomonas. Required for PSI accumulation. Activates translation of cp psaB RNA in Chlamydomonas but not in maize.GRMZM2G081955Os02g39740AT3G08010 Zm-tab2-1pyg
Zm-tab2-2pyg
tha12-3pyg
Zm-tab2-4pygm
Zm-abcg11Ortholog of Arabidopsis ABCG11/COF1/WBC11. Plasma membrane- localized ATP-binding cassette transporter; cutin transport to extracellular matrixGRMZM2G177812Os04g44610AT1G17840 Zm-abcg11-1leaves adhered toget
Zm-abcg11
Zm-abcg11
Zm-s1Ortholog of Arabidopsis BSF. Stabilizes processed RNA termini in the chloroplast petB-petD intergenic regionGRMZM2G087628Os01g40640At1g71720 Zm-s1-1iv-vir
Zm-s1-2iv-vir
Zm-era1Ortholog of Arabidopsis ERA1. Required for cp ribosome accumulation. Putative cp ribosome assembly factor. ERA domain (IPR005662).GRMZM2G158024Os05g49220At5g66470 iv-vir18-1vpyg-vir
iv-vir18-2
iv-vir18-3
iv-vir18-4
undefinedundefined
Zm-era1-5vpyg-virm
Zm-gun4Ortholog of Arabidopsis GUN4; Mg-chelatase cofactor and porphyrin binding proteinGRMZM2G464328Os11g16550AT3G59400 gun4-1pyg
gun4
gun4
gun4-2yel
Zm-gun4-3yel-m
atp5Ortholog of Arabidopsis PAB, involved in ATP synthase assembly. Mutant specifically lacks the ATP synthase.GRMZM2G110258Os03g01920At4g34090 nta1-1spg
nta1
nta1-3unknown
nta1-4unknown
nta1-2unknown
crp17Ortholog of Arabidopsis PGR3. PPR protein that stabilizes 5'-end of cp petL RNA and is required for NDH accumulation. Additional functions in maize. PGR3 ortholog. Stabilizes 5'end of cp petL RNA and 3' end of rpl14 mRNA. GRMZM2G372632Os10g28600AT4G31850 crp17-1vpygm
Zm-pgr3
crp19Ortholog of Arabidopsis PRDA1. cp nucleoid protein involved in PEP-mediated transcription; cpRNA defects in maize similar to other PEP-deficient mutantsGRMZM2G079452Os11g23790AT5G48470 Zm-prda1-1vpyg/iv-vir
Zm-prda1-2
Zm-prda1-3
ptac14Ortholog of Arabidopsis PTAC14. PEP-associated protein, required for PEP-mediated transcription; cpRNA defects in maize similar to other PEP-deficient mutants. ptac14-2 x ptac14-3 are iv, both exon insertions.GRMZM5G807767Os05g50980AT4G20130 ptac14-1iv
ptac14-2iv
pTac14-3iv
Zm-y3ip1Ortholog of Arabidopsis Y3IP1. PSI assembly factor, cooperates with Ycf3GRMZM2G002165Os01g58470AT5G44650 hcf148-1pyg
hcf148-2
hcf148-3
hcf148-4pyg
Zm-y3ip1-5
Zm-ycf54Ortholog of barley/cyanobacterial Ycf54; subunit of Mg-protoporphyrin IX monomethyl ester cyclaseGRMZM2G010196Os03g21370AT5G58250 Zm-ycf54-1pyg
Zm-ycf54-2pg
ppr102P-type PPR protein. Highest expression near leaf base and in sxn. 4. GRMZM2G012174Os05g22870At5g50280 ppr102-1 
ppr102-2
ppr102-3
ppr102-4
ppr102
ppr102-6iv-runty
ppr102
ppr102-5
ppr102-7
ppr102-8
ppr36P-type PPR. No phenotype. Highest expression in very leaf base. Coexpressed with mito editing factors in Arabidopsis. Probably mito. GRMZM2G333678Os11g14980AT4G19191 ppr36-1no pnenotype 
ppr61PPR proteinGRMZM2G341479Os02g49830AT1G09220 ppr61-1iv 
ppr61
ppr61
ppr2PPR protein required for plastid ribosome accumulation.GRMZM2G341621Os03g19650AT3G06430 ppr2-1iv
ppr2-4iv
ppr4PPR protein required for rps12 trans-splicing; binds rps12 intron 1.GRMZM2G404066Os04g58780AT5G04810 ppr4-1vpyg
ppr4-2spg
tpj1PPR protein required for translation of the chloroplast psbJ RNA in maize. Orthologous to Arabidopsis LPE1, which is required for psbJ and psbN translation.GRMZM2G056116Os12g18640AT3G46610 ppr66
tpj1-1
tpj1-2mild hcf
ppr66
ppr6PPR protein. No phenotype.GRMZM2G093291Os06g02120At5g27270 ppr6-1no phenotype 
ppr6-2no phenotype
ppr6-3no phenotype
crp1PPR protein; Activates petA and psaC translation; stabilizes processed petB, petD mRNAs; binds petA, psaC, petD 5'UTRsGRMZM2G083950Os07g36390AT5G42310 crp1-1pyg
crp1-2pyg
crp1-3pyg
crp1-4pyg
crp1-5pyg
crp1-6pyg
crp1-7pyg
tha8PPR protein; required for splicing ycf3-int2 and trnA introns and associates with those introns in vivo.AC217965.2_FGT012At3g27750 tha8-1pyg
tha8-2pyg
tha8-3pyg
tha8-4pyg
tha8-5pyg
tha8-6pyg
tha8-7pyg or pg
ppr5PPR protein; stabilizes unspliced trnG precursor and stimulates its splicing; binds the trnG intronGRMZM2G025409Os02g51480AT4G39620 ppr5-1iv
ppr5-2pg
Zm-otp51PPR-LAGLIDADG protein; required for splicing cp ycf3-int2 (Ortholog of otp51)GRMZM2G325019Os02g47360At2g15820 Zm-otp51-1pygm
Zm-otp51-2pygm
Zm-otp51-3pgm
Zm-otp51-4pyg
Zm-otp51-5pyg
atp4PPR-SMR protein; activates atpB translation, binds atpB 5?UTR GRMZM2G128665Os03g11670AT4G16390 atp4-1spg
atp4-2spg
atp4-3spg
ppr-smr4PPR/SMR protein. No phenotype.GRMZM2G164202Os02g02770At2g17033 pprsmr4-1no phenotype 
psa3PSI assembly factor. Shen et al, Plant Physiology 2017. Our PSA3 antibody does not work in Arabidopsis. Chlamy ortholog is Cre03.g183400.t1.1GRMZM2G051403Os05g45030AT3G55250 spg184-1spg
spg184-3wt
spg184
spg184
spg184-4
spg184-2spg
psa1Required for PSI accumulation. Harbors NifU domain implicated in Fe-S cluster assembly. Ortholog of Arabidopsis Nfu-IVa.GRMZM2G100976Os06g47940AT4G25910 psa1-1spg
psa1-2spg
raf1Rubisco Accumulation Factor1; promotes Rubisco assembly.GRMZM2G457621 LOC_Os10g30870AT3G04550, AT5G28500 raf1-1pg
raf1-2pg
raf1-3pg
raf2Rubisco Accumulation Factor2; promotes Rubisco assembly.GRMZM2G139123At5g51110 raf2-1
Zm-rbd1Rubredoxin, involved in PSII accumulation RBD1 is At1g54500 see Calderon JBC 2013GRMZM2G093815Os07g08770AT4G27990 Zm-rbd1-1pyg-vir 
Zm-rbd1-2pyg-vir
Zm-paacTAAC, thylakoid ATP/ADP carrierGRMZM2G178460Os01g16040AT5G01500, AT3G51870 Zm-paac-1no phenotype 
Zm-paac-2vpyg
Zm-paac-3vpyg
Zm-pyg7TPR protein, PSI biogenesis, ortholog of pyg7 in Arabidopsis.GRMZM5G809292Os10g32300AT1G22700 pg26-1pg
pg26-2pgm
pg26-3pg
tha4Thylakoid protein targeting: tat pathwayGRMZM2G472651Os03g43430 AT5G28750 tha4-1spg
tha5Thylakoid targeting. SecE subunit of protein translocation complex.GRMZM2G300408Os08g30830AT4G14870 tha5-1pg
tha5-2pg
tha5-3pg
tha5-4
not pyg18Unknown functionGRMZM2G168375Os01g74152AT5G59480 pyg18-1pyg 
pyg18-2
csy1Zm00001d000350, thylakoid protein translocase (ortholog CpSecY)GRMZM5G809546Os08g15460AT2G18710 csy1-1vpyg
csy1-2vpyg
csy1-3
crp11cp 3'-tRNA processing endoribonuclease (ortholog of CPZ)GRMZM5G802566Os09g30466AT2G04530 crp11-1iv-vir grny 
crp11-2
crp11
crp11
crp11
ZmClpP5cp ATP-dependent Clp protease subunit 5. pale yellow phenotype confirmed by complementation crosses by Bailin Li groupGRMZM2G121456Os03g19510AT1G02560 clpP5-1iv
clpP5
vylvpyg
cplP5-4pyg-vir
vdl1cp DEAD box protein (ortholog of tobacco VDL helicase). Required for biogenesis of plastid translation machinery.GRMZM2G415538Os01g73900AT5G19210 vdl1-1iv & stripey 
vdl1-2iv
ppdkcp PPDK, pyruvate orthophosphate dikinaseGRMZM2G306345Os03g31750AT4G15530 ppdk-1pgm
ppdk-2pg
ppdk-3pg
ppr100cp PPR protein: required for biogenesis of plastid translation machinery. Highest expression near leaf base.GRMZM2G428579Os02g02590At2g01860 ppr100-1iv  
ppr100-2iv
cfm1cp RNA splicing factor, CRM domain containing proteinGRMZM5G813259Os05g47850At3g18390 cfm1-1possibly embryo leth 
wtf1cp RNA splicing factor. Founding member of PORR domain family. Interacts with RNC1GRMZM2G403797Os05g49610AT4G01037 wtf1-1pg
wtf1-2ivm
wtf1-3iv-runty
wtf1-4iv-runty
wtf1-5iv-vir
rnc1cp RNAse III domain protein: required for the splicing of many chloroplast introns. Interacts with WTF1.GRMZM2G035820Os01g59510 AT4G37510 rnc1-1spg
rnc1-2iv
rnc1-3iv-runty
rnc1-4vpyg
tha1cp SecA homolog: required for thylakoid protein targetingGRMZM5G880102?Os01g21820At4g01800 tha1-1pyg
cpx1cp coproporphyrinogen oxidase (ortholog of LIN2; LESION INITIATION 2)GRMZM5G870342Os04g52130AT1G03475 cpx1
cpx1-2necrotic
cpx1pg-nec
cps1cp cysteinyl-tRNA synthetaseGRMZM2G156565Os09g38420AT2G31170 cps1-1pygm
cps1-2pyg
cps1-3
crp4cp polynucleotide phosphorylase (PNPase), cpRNA processing and decayGRMZM2G377761Os07g07310AT3G03710 crp4-1pale green
crp4-2spgm
crp4-3pygm
crp4-4spgm
pet3cp protein involved in cytb6f assembly (ortholog is CCB4). Allelic to maize gene hcf6Zm00001d002054Os04g55690AT1G59840 pet3-1spg
pet3-2pyg
hcf6spg-lethal
Zm-ccs1cp protein required for the accumulation of the cyt b6f complex. (ortholog likely chlamydomonas CCS1). Involved in attaching heme to petAGRMZM2G038301Os03g55970At1g49380 Zm-ccs1-1pg
Zm-ccs1-2pg
Zm-ccs1-3pyg
cptk1cp thymidine kinase 1GRMZM2G048821Os03g02200AT3G07800 cptk1-1iv 
cptk1
cptk1-3WT
cptk1-2
cptk
Zm-ptac10cp transcription. (ortholog of PTAC10) Contains S1 RNA-binding domain.GRMZM2G091419Os01g67570At3g48500 Zm-ptac10-1vpyg-mutable
Zm-ptac10-2iv
Zm-ptac10-3vpyg
Zm-ptac18cp transcription. (ortholog of PTAC18)GRMZM2G306104Os02g54360At2g32180 , At2g32650 Zm-ptac18-1spg 
Zm-ptac18-2spg
znfn1cp transcription. zinc finger domain.GRMZM2G086277Os06g07020At5g52010 znfn1-1iv 
znfn1-2iv
znfn1-3ivm
znfn1-4
Zm-prep2cp transit peptide degrading enzyme, (ortholog of ATPREP2)GRMZM2G072861Os02g52390AT1G49630 Zm-prep2-1iv-green-tip-m 
Zm-prep2-2
csr1cpSRP receptor. Involved in thylakoid protein targeting (ortholog of CPFTSY)Zm00001d042044Os01g72800AT2G45770 csr1-1vpyg
csr1-2vpyg
csr1-3vpyg
csr1-4yelm
hcf106cpTAT transport. GRMZM5G898735=hcf106 GRMZM2G151529=hcf106CGRMZM5G898735Os11g37130AT5G52440
Zm-atpcgamma subunit of chloroplast ATP synthase (ATPC1)GRMZM2G048907Os07g32880AT4G04640 Zm-atpc-1spg
Zm-atpc-2unknown
Zm-atpc-3unknown
Zm-atpc-4spg
Zm-atpc-5vpygm
Zm-atpc-6
Zm-rh3bglobal cp Dead box RNA helicase involved in cp intron splicing and assembly of the 50S ribosomal subunitAC198418.3_FGT005Os03g61220AT5G26742 Zm-rh3b-1vpyg-vir
Zm-rh3b-2vpyg-vir
Zm-PPOXlesion mimic phenotype, cp protoporphyrinogen oxidase, Ortholog of Arabidopsis PPOX/ PPO1. GRMZM2G039396Os01g18320AT4G01690 Zm-ppox-1pg-with lesions
Zm-ppox-2necrotic
Zm-ppox-3unknown
why1single-stranded nucleic acid binding protein. Multifunctional: promotes atpF splicing, 23S rRNA maturation, stabilizes cp genome.GRMZM2G155662Os06g05350AT2G02740, AT1G14410 why1-2grainy
why1-1iv