Gene Details:

• Gene ID: AT2G30950
• Gene Symbol: FTSH2, VAR2
• Gene Name: VARIEGATED 2
• Description: FtsH extracellular protease family;(source:Araport11)
• TAIR Accession: locus:2052806
• Genome: Araport11_genome_release
• Species: Arabidopsis thaliana

Transcripts:

• AT2G30950.1     AT2G30950.4     AT2G30950.2     AT2G30950.3

Plant Ontology Annotations:

• PO:0000037  — shoot axis apex — ápice del epiblasto (epiblastema) (Spanish, exact), シュート頂、茎頂 (Japanese, exact)
• PO:0008019  — leaf lamina base — base de la lámina de la hoja (Spanish, exact), 葉身基部 (Japanese, exact)
• PO:0009005  — root — raíz (Spanish, exact), radices (exact, plural), radix (exact), 根 (Japanese, exact), aerial root (narrow), climbing root (narrow)
• PO:0009006  — shoot system — sistema de epiblasto (epiblastema) (Spanish, exact), シュート系、苗条系 (Japanese, exact), Poaceae crown (related), shoot (related), thalli (related), thallus (related), tree crown (narrow)
• PO:0009009  — plant embryo — embrión (Spanish, exact), 植物胚 (Japanese, exact), germ (related), embryo (broad)
• PO:0009025  — vascular leaf — foliage leaf (exact), hoja vascular (Spanish, exact), leaf, vascular (exact), vascular leaves (exact, plural), 維管束のある葉, または維管束植物の葉 (Japanese, exact), crozier (related), macrophyll (related), megaphyll (related), ascidia (narrow), ascidium (narrow), fiddlehead (narrow), frond (narrow), needle-like leaf (narrow), pitcher (narrow), pitcher blade (narrow), pitcher-blade (narrow), scale-like leaf (narrow), sterile frond (narrow), trophophyll (narrow)
• PO:0009029  — stamen — estambre (Spanish, exact), 雄蕊 (Japanese, exact), Poaceae stamen (narrow), Zea stamen (narrow)
• PO:0009031  — sepal — sépalo (Spanish, exact), がく片 (Japanese, exact)
• PO:0009046  — flower — flor (Spanish, exact), 花 (Japanese, exact), floret (related), Asteraceae floret (narrow), basal flower (narrow), double flower (narrow), hermaphrodite flower (narrow), monoclinous flower (narrow), perfect flower (narrow)
• PO:0009052  — inflorescence flower pedicel — 小花柄 (Japanese, related), pedicelo (Spanish, broad)
• PO:0020030  — cotyledon — cotiledón (Spanish, exact), seed leaf (exact), 子葉 (Japanese, exact)
• PO:0020038  — petiole — pecíolo (Spanish, exact), 葉柄 (Japanese, exact)
• PO:0020100  — hypocotyl — hipocótile (Spanish, exact), 胚軸 (Japanese, exact)
• PO:0020137  — leaf apex — ápice de la hoja (Spanish, exact), 葉先 (Japanese, exact), leaf lamina apex (narrow), phyllid apex (narrow)
• PO:0025022  — collective leaf structure — estructura colectiva de hoja (Spanish, exact), leaf series (exact), 葉が集まった構造 (Japanese, exact), leaf whorl (narrow), rosette (narrow), cycle (broad), verticil (broad)
• PO:0025281  — pollen — polen (Spanish, exact), pollen grain (exact), 花粉 (Japanese, exact)
• PO:0000084  — plant sperm cell — célula espermática o esperma (Spanish, exact), male gamete (exact), microgamete (exact), 植物精子細胞 (Japanese, exact), sperm nucleus (related), sperm cell (broad)
• PO:0025195  — pollen tube cell — célula del tubo polínico (Spanish, exact), 花粉管細胞 (Japanese, exact)

Gene Ontology:

• GO:0009941  — located in — chloroplast envelope
• GO:0009579  — located in — thylakoid
• GO:0009535  — located in — chloroplast thylakoid membrane
• GO:0010205  — acts upstream of or within — photoinhibition
• GO:0004176  — enables — ATP-dependent peptidase activity
• GO:0009507  — located in — chloroplast
• GO:0009534  — located in — chloroplast thylakoid

Germplasm Phenotype:

• CS272  — variegated.
• CS73030  — variegated, leaves predominantly white; cotyledons in var2 appear normal, but first true leaves are bright yellow; yellow sectors fade to white as leaf expansion proceeds
• Ftsh2  — Lesser ability to deal with photoinhibition.
• Ftsh2  — variegated.
• evr1-1/var2-5  — Enhanced leaf varigation.
• evr1-2/var2-5  — Enhanced leaf varigation.
• ftsh1 ftsh2  — More variegated than ftsh2.
• ftsh2 ftsh5  — More variegated than ftsh2.
• ftsh2 ftsh8  — Completely white seedlings. Restoration of Heterozygocity of FTSH8 allows chloroplast development in the cotyledons.
• fug1-1/var2-2  — Double mutant suppresses leaf variegation due to var2 loss of function. First leaves are smaller with reduced cell size.
• fug1-3/var2-1  — Double mutant suppresses leaf variegation due to var2 loss of function. First leaves are smaller with reduced cell size.
• var2-5  — variegated
• var2-5, svr1-2  — Variegation caused by var2 single mutant is suppressed.
• var2-5;svr1-1  — WT phenotype. The single var2 variegated phenotype is suppresed.

Function-related keywords:

• shoot axis apex ,  leaf lamina base ,  root ,  shoot system ,  plant embryo ,  vascular leaf ,  stamen ,  sepal ,  flower ,  inflorescence flower pedicel ,  cotyledon ,  petiole ,  hypocotyl ,  leaf apex ,  collective leaf structure ,  pollen ,  plant sperm cell ,  pollen tube cell ,  pollen

Literature:

• Mutations in the Arabidopsis VAR2 locus cause leaf variegation due to the loss of a chloroplast FtsH protease.   DOI:  10.1046/j.1365-313x.2000.00738.x ;  PMID:  10849347
• The yellow variegated mutant of Arabidopsis is plastid autonomous and delayed in chloroplast biogenesis.   DOI:  10.1093/jhered/90.1.207 ;  PMID:  9987930
• Identification, characterization, and molecular cloning of a homologue of the bacterial FtsH protease in chloroplasts of higher plants.   DOI:  10.1074/jbc.271.46.29329 ;  PMID:  8910594
• The YELLOW VARIEGATED (VAR2) locus encodes a homologue of FtsH, an ATP-dependent protease in Arabidopsis.   DOI:  10.1093/pcp/pcd067 ;  PMID:  11134419
• A critical role for the Var2 FtsH homologue of Arabidopsis thaliana in the photosystem II repair cycle in vivo.   DOI:  10.1074/jbc.M105878200 ;  PMID:  11717304
• Chloroplast and mitochondrial proteases in Arabidopsis. A proposed nomenclature.   DOI:  10.1104/pp.125.4.1912 ;  PMID:  11299370
• The VAR1 locus of Arabidopsis encodes a chloroplastic FtsH and is responsible for leaf variegation in the mutant alleles.   DOI:  10.1046/j.1365-2443.2002.00558.x ;  PMID:  12167156
• Coordinated regulation and complex formation of yellow variegated1 and yellow variegated2, chloroplastic FtsH metalloproteases involved in the repair cycle of photosystem II in Arabidopsis thylakoid membranes.   DOI:  10.1105/tpc.017319 ;  PMID:  14630971
• The Arabidopsis FtsH metalloprotease gene family: interchangeability of subunits in chloroplast oligomeric complexes.   DOI:  10.1111/j.1365-313x.2003.02014.x ;  PMID:  14996218
• Expression in multigene families. Analysis of chloroplast and mitochondrial proteases.   DOI:  10.1104/pp.104.043299 ;  PMID:  15266057
• New functions of the thylakoid membrane proteome of Arabidopsis thaliana revealed by a simple, fast, and versatile fractionation strategy.   DOI:  10.1074/jbc.M406763200 ;  PMID:  15322131
• Mutations in ClpC2/Hsp100 suppress the requirement for FtsH in thylakoid membrane biogenesis.   DOI:  10.1073/pnas.0402764101 ;  PMID:  15304652
• Developmental and light effects on the accumulation of FtsH protease in Arabidopsis chloroplasts–implications for thylakoid formation and photosystem II maintenance.   DOI:  10.1111/j.1365-313X.2005.02401.x ;  PMID:  15918877
• Functional redundancy of AtFtsH metalloproteases in thylakoid membrane complexes.   DOI:  10.1104/pp.105.061234 ;  PMID:  16040665
• Two types of FtsH protease subunits are required for chloroplast biogenesis and Photosystem II repair in Arabidopsis.   DOI:  10.1105/tpc.105.035071 ;  PMID:  16126834
• AtFtsH6 is involved in the degradation of the light-harvesting complex II during high-light acclimation and senescence.   DOI:  10.1073/pnas.0503472102 ;  PMID:  16157880
• Arabidopsis variegation mutants: new insights into chloroplast biogenesis.   DOI:  10.1093/jxb/erj008 ;  PMID:  16449381
• Recent advances in the study of Clp, FtsH and other proteases located in chloroplasts.   DOI:  10.1016/j.pbi.2006.03.010 ;  PMID:  16603408
• Variegation mutants and mechanisms of chloroplast biogenesis.   DOI:  10.1111/j.1365-3040.2006.01630.x ;  PMID:  17263779
• White leaf sectors in yellow variegated2 are formed by viable cells with undifferentiated plastids.   DOI:  10.1104/pp.107.099002 ;  PMID:  17449646
• The balance between protein synthesis and degradation in chloroplasts determines leaf variegation in Arabidopsis yellow variegated mutants.   DOI:  10.1105/tpc.106.049270 ;  PMID:  17416734
• Quality control of Photosystem II: cleavage and aggregation of heat-damaged D1 protein in spinach thylakoids.   DOI:  10.1016/j.bbabio.2007.05.001 ;  PMID:  17543883
• The E3 ligase AtCHIP ubiquitylates FtsH1, a component of the chloroplast FtsH protease, and affects protein degradation in chloroplasts.   DOI:  10.1111/j.1365-313X.2007.03239.x ;  PMID:  17714429
• Influence of chloroplastic photo-oxidative stress on mitochondrial alternative oxidase capacity and respiratory properties: a case study with Arabidopsis yellow variegated 2.   DOI:  10.1093/pcp/pcn031 ;  PMID:  18296449
• Mutations in SUPPRESSOR OF VARIEGATION1, a factor required for normal chloroplast translation, suppress var2-mediated leaf variegation in Arabidopsis.   DOI:  10.1105/tpc.107.054965 ;  PMID:  18599582
• Transcriptome analyses show changes in gene expression to accompany pollen germination and tube growth in Arabidopsis.   DOI:  10.1104/pp.108.126375 ;  PMID:  18775970
• Activation of the heterotrimeric G protein alpha-subunit GPA1 suppresses the ftsh-mediated inhibition of chloroplast development in Arabidopsis.   DOI:  10.1111/j.1365-313X.2009.03843.x ;  PMID:  19228339
• Identification and characterization of two members of the FtsH gene family in maize (Zea mays L.).   DOI:  10.1007/s11033-009-9691-3 ;  PMID:  19669593
• The variegated mutants lacking chloroplastic FtsHs are defective in D1 degradation and accumulate reactive oxygen species.   DOI:  10.1104/pp.109.146589 ;  PMID:  19767385
• Arrested differentiation of proplastids into chloroplasts in variegated leaves characterized by plastid ultrastructure and nucleoid morphology.   DOI:  10.1093/pcp/pcp127 ;  PMID:  19755395
• Photosynthetic redox imbalance governs leaf sectoring in the Arabidopsis thaliana variegation mutants immutans, spotty, var1, and var2.   DOI:  10.1105/tpc.108.062752 ;  PMID:  19897671
• Reactive oxygen species derived from impaired quality control of photosystem II are irrelevant to plasma-membrane NADPH oxidases.   DOI:  10.4161/psb.5.3.10604 ;  PMID:  20023407
• Comparative transcriptome analysis of green/white variegated sectors in Arabidopsis yellow variegated2: responses to oxidative and other stresses in white sectors.   DOI:  10.1093/jxb/erq075 ;  PMID:  20400527
• Arabidopsis chloroplast FtsH, var2 and suppressors of var2 leaf variegation: a review.   DOI:  10.1111/j.1744-7909.2010.00980.x ;  PMID:  20666930
• An Arabidopsis pentatricopeptide repeat protein, SUPPRESSOR OF VARIEGATION7, is required for FtsH-mediated chloroplast biogenesis.   DOI:  10.1104/pp.110.164111 ;  PMID:  20935174
• The FtsH protease heterocomplex in Arabidopsis: dispensability of type-B protease activity for proper chloroplast development.   DOI:  10.1105/tpc.110.079202 ;  PMID:  21062893
• A var2 leaf variegation suppressor locus, SUPPRESSOR OF VARIEGATION3, encodes a putative chloroplast translation elongation factor that is important for chloroplast development in the cold.   DOI:  10.1186/1471-2229-10-287 ;  PMID:  21187014
• Identification of nuclear genes encoding chloroplast-localized proteins required for embryo development in Arabidopsis.   DOI:  10.1104/pp.110.168120 ;  PMID:  21139083
• FtsH2 and FtsH5: two homologous subunits use different integration mechanisms leading to the same thylakoid multimeric complex.   DOI:  10.1111/j.1365-313X.2010.04448.x ;  PMID:  21214651
• Fitness analyses of Arabidopsis thaliana mutants depleted of FtsH metalloproteases and characterization of three FtsH6 deletion mutants exposed to high light stress, senescence and chilling.   DOI:  10.1111/j.1469-8137.2011.03684.x ;  PMID:  21438879
• Interplay between N-terminal methionine excision and FtsH protease is essential for normal chloroplast development and function in Arabidopsis.   DOI:  10.1105/tpc.111.087239 ;  PMID:  22010036
• FtsH proteases located in the plant chloroplast.   DOI:  10.1111/j.1399-3054.2011.01548.x ;  PMID:  22121866
• Molecular nature of mutations induced by high-LET irradiation with argon and carbon ions in Arabidopsis thaliana.   DOI:  10.1016/j.mrfmmm.2012.04.010 ;  PMID:  22579628
• Cooperative D1 degradation in the photosystem II repair mediated by chloroplastic proteases in Arabidopsis.   DOI:  10.1104/pp.112.199042 ;  PMID:  22698923
• Possible compensatory role among chloroplast proteases under excess-light stress condition.   DOI:  10.4161/psb.23198 ;  PMID:  23299325
• Modified Clp protease complex in the ClpP3 null mutant and consequences for chloroplast development and function in Arabidopsis.   DOI:  10.1104/pp.113.215699 ;  PMID:  23548781
• Understanding chloroplast biogenesis using second-site suppressors of immutans and var2.   DOI:  10.1007/s11120-013-9855-9 ;  PMID:  23703455
• Genetic interactions reveal that specific defects of chloroplast translation are associated with the suppression of var2-mediated leaf variegation.   DOI:  10.1111/jipb.12078 ;  PMID:  23721655
• Multiple fates of non-mature lumenal proteins in thylakoids.   DOI:  10.1111/tpj.12273 ;  PMID:  23802992
• Proteomic evidence for genetic epistasis: ClpR4 mutations switch leaf variegation to virescence in Arabidopsis.   DOI:  10.1111/tpj.12344 ;  PMID:  24124904
• Proteome and metabolome profiling of cytokinin action in Arabidopsis identifying and up-regulation.   DOI:  10.1093/jxb/ert227 ;  PMID:  24064926
• Phosphorylation of photosystem II core proteins prevents undesirable cleavage of D1 and contributes to the fine-tuned repair of photosystem II.   DOI:  10.1111/tpj.12562 ;  PMID:  24862025
• Leaf Variegation of Thylakoid Formation1 Is Suppressed by Mutations of Specific σ-Factors in Arabidopsis.   DOI:  10.1104/pp.15.00549 ;  PMID:  25999408
• Mutations in circularly permuted GTPase family genes AtNOA1/RIF1/SVR10 and BPG2 suppress var2-mediated leaf variegation in Arabidopsis thaliana.   DOI:  10.1007/s11120-015-0195-9 ;  PMID:  26435530
• RNA Sequencing Analysis of the msl2msl3, crl, and ggps1 Mutants Indicates that Diverse Sources of Plastid Dysfunction Do Not Alter Leaf Morphology Through a Common Signaling Pathway.   DOI:  10.3389/fpls.2015.01148 ;  PMID:  26734046
• Identification and Roles of Photosystem II Assembly, Stability, and Repair Factors in Arabidopsis.   DOI:  10.3389/fpls.2016.00168 ;  PMID:  26909098
• Thylakoid-Bound FtsH Proteins Facilitate Proper Biosynthesis of Photosystem I.   DOI:  10.1104/pp.16.00200 ;  PMID:  27208291
• and EXECUTER1-mediated signaling is initiated in grana margins and depends on the protease FtsH2.   DOI:  10.1073/pnas.1603562113 ;  PMID:  27303039
• Phenotypic reversion in fas mutants of Arabidopsis thaliana by reintroduction of FAS genes: variable recovery of telomeres with major spatial rearrangements and transcriptional reprogramming of 45S rDNA genes.   DOI:  10.1111/tpj.13257 ;  PMID:  27377564
• Chloroplast Translation Initiation Factors Regulate Leaf Variegation and Development.   DOI:  10.1104/pp.15.02040 ;  PMID:  27535792
• Nitrogen remobilization during leaf senescence: lessons from Arabidopsis to crops.   DOI:  10.1093/jxb/erw365 ;  PMID:  27707774
• D1 fragmentation in photosystem II repair caused by photo-damage of a two-step model.   DOI:  10.1007/s11120-015-0144-7 ;  PMID:  25893898
• FtsH2-Dependent Proteolysis of EXECUTER1 Is Essential in Mediating Singlet Oxygen-Triggered Retrograde Signaling in Arabidopsis thaliana.   DOI:  10.3389/fpls.2017.01145 ;  PMID:  28706530
• Structure of Psb29/Thf1 and its association with the FtsH protease complex involved in photosystem II repair in cyanobacteria.   DOI:  10.1098/rstb.2016.0394 ;  PMID:  28808107
• Balance between Cytosolic and Chloroplast Translation Affects Leaf Variegation.   DOI:  10.1104/pp.17.00673 ;  PMID:  29142022
• FtsH Protease in the Thylakoid Membrane: Physiological Functions and the Regulation of Protease Activity.   DOI:  10.3389/fpls.2018.00855 ;  PMID:  29973948
• The Photosystem II Repair Cycle Requires FtsH Turnover through the EngA GTPase.   DOI:  10.1104/pp.18.00652 ;  PMID:  30131421
• Chloroplast Translation Elongation Factor EF-Tu/SVR11 Is Involved in var2-Mediated Leaf Variegation and Leaf Development in Arabidopsis.   DOI:  10.3389/fpls.2019.00295 ;  PMID:  30915096
• Impaired PSII proteostasis triggers a UPR-like response in the var2 mutant of Arabidopsis.   DOI:  10.1093/jxb/erz151 ;  PMID:  30989223
• Impaired PSII Proteostasis Promotes Retrograde Signaling via Salicylic Acid.   DOI:  10.1104/pp.19.00483 ;  PMID:  31160506
• Phosphorylation of the Chloroplastic Metalloprotease FtsH in Arabidopsis Characterized by Phos-Tag SDS-PAGE.   DOI:  10.3389/fpls.2019.01080 ;  PMID:  31552075
• The chloroplast metalloproteases VAR2 and EGY1 act synergistically to regulate chloroplast development in Arabidopsis.   DOI:  10.1074/jbc.RA119.011853 ;  PMID:  31836664
• EGY3: homologue of S2P protease located in chloroplasts.   DOI:  10.1111/plb.13087 ;  PMID:  31886945
• The Phytotoxicity of Meta-Tyrosine Is Associated With Altered Phenylalanine Metabolism and Misincorporation of This Non-Proteinogenic Phe-Analog to the Plant's Proteome.   DOI:  10.3389/fpls.2020.00140 ;  PMID:  32210982
• Autocatalytic Processing and Substrate Specificity of Arabidopsis Chloroplast Glutamyl Peptidase.   DOI:  10.1104/pp.20.00752 ;  PMID:  32663165
• Arabidopsis CHLOROPHYLLASE 1 protects young leaves from long-term photodamage by facilitating FtsH-mediated D1 degradation in photosystem II repair.   DOI:  10.1016/j.molp.2021.04.006 ;  PMID:  33857689
• VAR2/AtFtsH2 and EVR2/BCM1/CBD1 synergistically regulate the accumulation of PSII reaction centre D1 protein during de-etiolation in Arabidopsis.   DOI:  10.1111/pce.14368 ;  PMID:  35610189
• The Arabidopsis NLP7-HB52/54-VAR2 pathway modulates energy utilization in diverse light and nitrogen conditions.   DOI:  10.1016/j.cub.2022.10.024 ;  PMID:  36332616
• Cooperative role of AtRsmD and AtRimM proteins in modification and maturation of 16S rRNA in plastids.   DOI:  10.1111/tpj.16135 ;  PMID:  36752655
• Chloroplast SRP54 and FtsH protease coordinate thylakoid membrane-associated proteostasis in Arabidopsis.   DOI:  10.1093/plphys/kiad199 ;  PMID:  36994815
• Mutations in the Arabidopsis VAR2 locus cause leaf variegation due to the loss of a chloroplast FtsH protease.   DOI:  10.1046/j.1365-313x.2000.00738.x ;  PMID:  10849347
• Identification, characterization, and molecular cloning of a homologue of the bacterial FtsH protease in chloroplasts of higher plants.   DOI:  10.1074/jbc.271.46.29329 ;  PMID:  8910594
• The YELLOW VARIEGATED (VAR2) locus encodes a homologue of FtsH, an ATP-dependent protease in Arabidopsis.   DOI:  10.1093/pcp/pcd067 ;  PMID:  11134419
• Proteome map of the chloroplast lumen of Arabidopsis thaliana.   DOI:  10.1074/jbc.M108575200 ;  PMID:  11719511
• Chloroplast and mitochondrial proteases in Arabidopsis. A proposed nomenclature.   DOI:  10.1104/pp.125.4.1912 ;  PMID:  11299370
• Proteomic study of the Arabidopsis thaliana chloroplastic envelope membrane utilizing alternatives to traditional two-dimensional electrophoresis.   DOI:  10.1021/pr034025j ;  PMID:  12938931
• In-depth analysis of the thylakoid membrane proteome of Arabidopsis thaliana chloroplasts: new proteins, new functions, and a plastid proteome database.   DOI:  10.1105/tpc.017814 ;  PMID:  14729914
• The Arabidopsis thaliana chloroplast proteome reveals pathway abundance and novel protein functions.   DOI:  10.1016/j.cub.2004.02.039 ;  PMID:  15028209
• New functions of the thylakoid membrane proteome of Arabidopsis thaliana revealed by a simple, fast, and versatile fractionation strategy.   DOI:  10.1074/jbc.M406763200 ;  PMID:  15322131
• High light response of the thylakoid proteome in arabidopsis wild type and the ascorbate-deficient mutant vtc2-2. A comparative proteomics study.   DOI:  10.1104/pp.106.080150 ;  PMID:  16648217
• Membrane proteomic analysis of Arabidopsis thaliana using alternative solubilization techniques.   DOI:  10.1021/pr060525b ;  PMID:  17432890
• Sorting signals, N-terminal modifications and abundance of the chloroplast proteome.   DOI:  10.1371/journal.pone.0001994 ;  PMID:  18431481
• Quantitative proteomics of a chloroplast SRP54 sorting mutant and its genetic interactions with CLPC1 in Arabidopsis.   DOI:  10.1104/pp.108.124545 ;  PMID:  18633119

Sequences: