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Q9C5U0

- AHK4_ARATH

UniProt

Q9C5U0 - AHK4_ARATH

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Protein

Histidine kinase 4

Gene

AHK4

Organism
Arabidopsis thaliana (Mouse-ear cress)
Status
Reviewed - Annotation score: 5 out of 5- Experimental evidence at protein leveli

Functioni

Cytokinins (CK) receptor related to bacterial two-component regulators. Binds also the synthetic urea-type cytokinin thiadiazuron, a potent defoliant and herbicide. Functions as a histidine kinase and transmits the stress signal to a downstream MAPK cascade. This protein undergoes an ATP-dependent autophosphorylation at a conserved histidine residue in the kinase core, and a phosphoryl group is then transferred to a conserved aspartate residue in the receiver domain. In the presence of cytokinin, feed phosphate to phosphorelay-integrating histidine phosphotransfer protein (HPt) and activates subsequent cascade. In the absence of cytokinin, removes phosphate from HPt proteins, decreasing the system phosphoload. Involved in meristems establishment in seedlings. Acts as a redundant negative regulator of drought and salt stress responses, and abscisic acid (ABA) signaling in a cytokinin-dependent manner. Required to set vascular asymmetric cell divisions that establish phloem and procambium cell lines. Redundant positive regulator of cytokinin signaling that regulates many development process including seed germination, cell division, seed size, chlorophyll retention during leaf senescence, root repression and shoot promotion. Can interact with isoprenoid-type cytokinins trans-zeatin (tZ and tZR), isopentenyladenine (iP), and isopentenyladenosine (iPR), the meta hydroxylated derivative of benzyladenine m-topolin, buta-2,3-dienyladenine (HA-8), penta-2,3-dienyladenine (HA-1), 4-methyl-penta-2,3-dienyladenine (HA-10), 4-hydroxy-2-butynyladenine (RM1), 2-butynyladenine (RM6), and to a lower extent, with cis-zeatin (cZ), zeatin riboside and dihydrozeatin (DZ). Together with AHK3, involved in the cytokinin-dependent responses to Pi starvation and sucrose stresses. Required for the formation of auxin-transporting vascular tissues in the hypocotyl, and primary and lateral roots, but not in adventitious roots, thus leading to auxin basipetal transport that regulates root development and branching. Involved in alkamides (e.g. N-isobutyl decanamide) and N-acylethanolamides (NAE) signaling that control meristematic activity and differentiation processes during plant development. Prevents the uptake of sulfate by mediating cytokinin-dependent down-regulation of high-affinity sulfate transporters (e.g. SULTR1;1 and SULTR1;2) expression in roots. Together with AHK2, required for growth and reproduction promotion stimulated by the endophytic fungus Piriformospora indica in a trans-zeatin-dependent manner. Required to trigger the phytotoxic effect of the snapdragon (Antirrhinum majus) flowers volatile organic compound (VOC) methyl benzoate (MB). Plays a role in the cytokinin-mediated repression of the iron uptake pathway.28 Publications

Catalytic activityi

ATP + protein L-histidine = ADP + protein N-phospho-L-histidine.
[a protein]-serine/threonine phosphate + H2O = [a protein]-serine/threonine + phosphate.

Enzyme regulationi

Activated by cytokinins to initiate phosphorelay signaling. This cytokinin-mediated activation is repressed by the trans-zeatin antagonists 6-(2-hydroxy-3-methylbenzylamino)purine (PI-55) and 6-(2,5-Dihydroxybenzylamino)purine (LGR-991).3 Publications

pH dependencei

Optimum pH to bind cytokinin is about 7-8.5 at 0 degrees Celsius.2 Publications

Temperature dependencei

Cytokinin-binding is more stable at 0 degrees Celsius than at 20 and 37 degrees Celsius.2 Publications

Sites

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Binding sitei285 – 2851Cytokinin
Binding sitei307 – 3071Cytokinin; via amide nitrogen
Binding sitei317 – 3171Cytokinin

GO - Molecular functioni

  1. cytokine binding Source: TAIR
  2. cytokinin receptor activity Source: TAIR
  3. enzyme binding Source: UniProtKB
  4. osmosensor activity Source: TAIR
  5. phosphoprotein phosphatase activity Source: TAIR
  6. phosphorelay sensor kinase activity Source: InterPro
  7. protein histidine kinase activity Source: TAIR
  8. protein histidine kinase binding Source: UniProtKB
  9. protein kinase binding Source: UniProtKB
  10. transmembrane histidine kinase cytokinin receptor activity Source: UniProtKB

GO - Biological processi

  1. carbohydrate homeostasis Source: TAIR
  2. cellular response to phosphate starvation Source: UniProtKB
  3. cellular response to sucrose stimulus Source: UniProtKB
  4. cytokinin-activated signaling pathway Source: TAIR
  5. defense response to bacterium Source: TAIR
  6. dephosphorylation Source: GOC
  7. embryonic root morphogenesis Source: TAIR
  8. osmosensory signaling pathway Source: TAIR
  9. peptidyl-histidine phosphorylation Source: GOC
  10. phosphorelay signal transduction system Source: UniProtKB
  11. protein phosphorylation Source: TAIR
  12. regulation of meristem development Source: UniProtKB
  13. regulation of seed germination Source: TAIR
  14. regulation of shoot system development Source: TAIR
  15. response to water deprivation Source: TAIR
  16. sulfate transport Source: UniProtKB
Complete GO annotation...

Keywords - Molecular functioni

Developmental protein, Hydrolase, Kinase, Protein phosphatase, Transferase

Keywords - Biological processi

Cytokinin signaling pathway

Enzyme and pathway databases

BioCyciARA:AT2G01830-MONOMER.
ARA:GQT-91-MONOMER.
ARA:GQT-92-MONOMER.

Names & Taxonomyi

Protein namesi
Recommended name:
Histidine kinase 4 (EC:2.7.13.3)
Alternative name(s):
Arabidopsis histidine kinase 4
Short name:
AtHK4
Cytokinin receptor CYTOKININ RESPONSE 1
Short name:
AtCRE1
Short name:
Cytokinin receptor CRE1
Phosphoprotein phosphatase AHK4 (EC:3.1.3.16)
Protein AUTHENTIC HIS-KINASE 4
Protein ROOT AS IN WOL 1
Protein WOODEN LEG
Gene namesi
Name:AHK4
Synonyms:CRE1, RAW1, WOL
Ordered Locus Names:At2g01830
ORF Names:T23K3.2
OrganismiArabidopsis thaliana (Mouse-ear cress)
Taxonomic identifieri3702 [NCBI]
Taxonomic lineageiEukaryotaViridiplantaeStreptophytaEmbryophytaTracheophytaSpermatophytaMagnoliophytaeudicotyledonsGunneridaePentapetalaerosidsmalvidsBrassicalesBrassicaceaeCamelineaeArabidopsis
ProteomesiUP000006548: Chromosome 2

Organism-specific databases

TAIRiAT2G01830.

Subcellular locationi

Endoplasmic reticulum membrane 1 Publication; Multi-pass membrane protein 1 Publication

Topology

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Topological domaini1 – 124124CytoplasmicSequence AnalysisAdd
BLAST
Transmembranei125 – 14521HelicalSequence AnalysisAdd
BLAST
Topological domaini146 – 429284ExtracellularSequence AnalysisAdd
BLAST
Transmembranei430 – 45021HelicalSequence AnalysisAdd
BLAST
Topological domaini451 – 1080630CytoplasmicSequence AnalysisAdd
BLAST

GO - Cellular componenti

  1. endoplasmic reticulum Source: TAIR
  2. integral component of membrane Source: UniProtKB-KW
  3. plasma membrane Source: UniProtKB
Complete GO annotation...

Keywords - Cellular componenti

Endoplasmic reticulum, Membrane

Pathology & Biotechi

Disruption phenotypei

Reduced sensitivity to cytokinin (mostly in shoots). Narrow vascular cylinder composed mainly of protoxylem cell files, with no apparent metaxylem or phloem. Hypersensitivity to ABA. Strong drought and salinity tolerance only in the presence of CK. Reduced cytokinin repression of several Pi starvation-responsive genes and increased sucrose sensitivity. More rapid germination, reduced requirement for light, and decreased far-red light sensitivity. Reduced sensitivity to N-isobutyl decanamide. Impaired benzyladenine (6-BA)-mediated repression of the iron uptake pathway. Impaired meristematic development in seedlings.12 Publications

Mutagenesis

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Mutagenesisi225 – 2251A → L: Loss of activity. 1 Publication
Mutagenesisi227 – 2271A → L: Loss of activity. 1 Publication
Mutagenesisi273 – 2731Y → E: Loss of activity. 1 Publication
Mutagenesisi279 – 2791M → A: No effect on activity. 1 Publication
Mutagenesisi285 – 2851D → A, E or R: Loss of activity. 1 Publication
Mutagenesisi301 – 3011T → I in wol-1; locked in the phosphoprotein phosphatase active form, retardation of the primary root growth with reduced cell number and exclusive xylem differentiation within the vascular tissue associated with abnormal vascular asymmetric cell divisions, impaired metaxylem and phloem differentiation, and reduced cytokinin-binding ability leading to impaired kinase activity and cyokinin-mediated activation. 6 Publications
Mutagenesisi303 – 3031P → V: Loss of activity. 1 Publication
Mutagenesisi304 – 3041F → A: Loss of activity. 1 Publication
Mutagenesisi306 – 3061L → A: Loss of activity. 1 Publication
Mutagenesisi307 – 3071L → A: Loss of activity. 1 Publication
Mutagenesisi317 – 3171T → A: No effect on activity. 1 Publication
Mutagenesisi343 – 3431G → L: Loss of activity. 1 Publication
Mutagenesisi435 – 4351G → C: Constitutively activated independently of cytokinin. 1 Publication
Mutagenesisi436 – 4361F → S: Constitutively activated independently of cytokinin. 1 Publication
Mutagenesisi447 – 4471M → T: Constitutively activated independently of cytokinin. 1 Publication
Mutagenesisi459 – 4591M → I in wol-3; retardation of the primary root growth, no production of lateral roots and enhanced formation of adventitious roots associated with impaired auxin basipetal transport. 1 Publication
Mutagenesisi471 – 4711V → A: Constitutively activated independently of cytokinin. 1 Publication
Mutagenesisi482 – 4821H → Q: Reduced phosphoprotein phosphatase activity. 2 Publications
Mutagenesisi490 – 4901G → D in cre1-1; impaired histidine-kinase receptor activity and reduced responses to cytokinins, including rapid cell proliferation and shoot formation in tissue culture, repression of sulfate uptake, retardation of the primary root growth, no production of lateral roots and enhanced formation of adventitious roots associated with impaired auxin basipetal transport, as well as reduced cell number within the vascular tissues in roots. 4 Publications
Mutagenesisi493 – 4931G → R in cre1-6; reduced sensitivity to cytokinin. 1 Publication
Mutagenesisi494 – 4941M → L: Constitutively activated independently of cytokinin. 1 Publication
Mutagenesisi529 – 5291L → F in wol-2/raw1; impaired metaxylem and phloem differentiation, and reduced sensitivity to cytokinins. 1 Publication
Mutagenesisi529 – 5291L → F in wol-2; retardation of the primary root growth with reduced cell number and exclusive xylem differentiation within the vascular tissue associated with abnormal vascular asymmetric cell divisions, and impaired cytokinin-binding ability. 1 Publication
Mutagenesisi708 – 7081F → L: No histidine kinase activity, but normal phosphoprotein phosphatase activity. 1 Publication
Mutagenesisi996 – 9961D → N: Cytokinin-mediated autophosphorylation but impaired phosphotransfer to an HPt, abolished phosphoprotein phosphatase activity. 2 Publications
Mutagenesisi1008 – 10081T → I in cre1-4; slightly reduced sensitivity to cytokinin, and impaired cytokinin repression of several Pi starvation-responses. 1 Publication
Mutagenesisi1032 – 10321A → T in cre1-9; impaired cytokinin repression of several Pi starvation-responses. 1 Publication

PTM / Processingi

Molecule processing

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Chaini1 – 10801080Histidine kinase 4PRO_0000398589Add
BLAST

Amino acid modifications

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Modified residuei482 – 4821Phosphohistidine; by autocatalysis1 PublicationPROSITE-ProRule annotation
Modified residuei996 – 99614-aspartylphosphatePROSITE-ProRule annotation

Post-translational modificationi

Autophosphorylated predominantly on His residues. Activation probably requires a transfer of a phosphate group between a His in the transmitter domain and an Asp of the receiver domain.1 Publication

Keywords - PTMi

Phosphoprotein

Proteomic databases

PaxDbiQ9C5U0.
PRIDEiQ9C5U0.

Expressioni

Tissue specificityi

Mostly expressed in roots, specifically in the vascular cylinder and pericycle, and, to a lower extent, in leaves and flowers. Present in seedlings.4 Publications

Developmental stagei

Expressed specifically in the vasculature since the early stages of embryogenesis. At the globular stage of embryogenesis, detected in the four innermost cells, which are the precursors of the vascular tissue. During the heart, torpedo, and nearly mature stages, expressed in the procambium of the cotyledon shoulders, prospective hypocotyl, and embryonic root. In seedlings, mainly localized in meristematic tissues (e.g. shoot apical meristem SAM, root tips, and growing leaf and lateral root primordia), especially in vasculature. Present in all the vasculature and the shoot apical meristem (SAM) of the adult plant. In flowers, localized in carpels and developing ovules. In the root tips, expressed in the central cylinder.3 Publications

Inductioni

Rapidly induced by dehydration. Down-regulated by Pi starvation and induced by cytokinins.2 Publications

Gene expression databases

GenevestigatoriQ9C5U0.

Interactioni

Subunit structurei

Homodimer. Interacts with AHP1, AHP2, AHP3, AHP5, AHK3, AMPD, FBR12, WNK5 and At4g15630.5 Publications

Binary interactionsi

WithEntry#Exp.IntActNotes
AHK3Q9C5U12EBI-1100775,EBI-1100653
AMPDO804522EBI-1100775,EBI-1807679
At1g03430Q67XQ13EBI-1100775,EBI-1100725
At4g15630Q8L8Z12EBI-1100775,EBI-1807704
WNK5Q9SCU52EBI-1100775,EBI-1807651

Protein-protein interaction databases

BioGridi117. 30 interactions.
IntActiQ9C5U0. 29 interactions.
STRINGi3702.AT2G01830.2-P.

Structurei

Secondary structure

1
1080
Legend: HelixTurnBeta strand
Show more details
Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Helixi151 – 19343Combined sources
Turni194 – 1963Combined sources
Beta strandi197 – 1993Combined sources
Helixi204 – 21310Combined sources
Helixi215 – 2173Combined sources
Beta strandi221 – 2288Combined sources
Helixi232 – 2343Combined sources
Helixi235 – 2428Combined sources
Turni249 – 2513Combined sources
Beta strandi259 – 2613Combined sources
Beta strandi263 – 2675Combined sources
Helixi269 – 2746Combined sources
Helixi279 – 2813Combined sources
Helixi283 – 29513Combined sources
Turni307 – 3093Combined sources
Beta strandi312 – 32110Combined sources
Helixi331 – 3366Combined sources
Beta strandi338 – 3469Combined sources
Helixi348 – 35710Combined sources
Helixi362 – 3643Combined sources
Beta strandi365 – 3717Combined sources
Beta strandi379 – 3824Combined sources
Beta strandi385 – 3873Combined sources
Beta strandi395 – 3995Combined sources
Beta strandi408 – 4147Combined sources

3D structure databases

Select the link destinations:
PDBei
RCSB PDBi
PDBji
Links Updated
EntryMethodResolution (Å)ChainPositionsPDBsum
3T4JX-ray1.65A/B149-418[»]
3T4KX-ray1.77A/B149-418[»]
3T4LX-ray1.53A/B149-418[»]
3T4OX-ray1.75A/B149-418[»]
3T4QX-ray2.30A/B149-418[»]
3T4SX-ray1.60A/B149-418[»]
3T4TX-ray1.70A/B149-418[»]
ProteinModelPortaliQ9C5U0.
SMRiQ9C5U0. Positions 150-416, 465-752, 941-1060.
ModBaseiSearch...
MobiDBiSearch...

Miscellaneous databases

EvolutionaryTraceiQ9C5U0.

Family & Domainsi

Domains and Repeats

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Domaini198 – 411214CHASEPROSITE-ProRule annotationAdd
BLAST
Domaini479 – 760282Histidine kinasePROSITE-ProRule annotationAdd
BLAST
Domaini946 – 1071126Response regulatoryPROSITE-ProRule annotationAdd
BLAST

Compositional bias

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Compositional biasi823 – 8264Poly-Ala

Sequence similaritiesi

Contains 1 CHASE domain.PROSITE-ProRule annotation
Contains 1 histidine kinase domain.PROSITE-ProRule annotation
Contains 1 response regulatory domain.PROSITE-ProRule annotation

Keywords - Domaini

Transmembrane, Transmembrane helix

Phylogenomic databases

eggNOGiCOG0784.
InParanoidiQ9C5U0.
KOiK14489.
OMAiNQNRPSF.
PhylomeDBiQ9C5U0.

Family and domain databases

Gene3Di1.10.287.130. 1 hit.
3.30.565.10. 2 hits.
InterProiIPR006189. CHASE.
IPR011006. CheY-like_superfamily.
IPR003661. EnvZ-like_dim/P.
IPR003594. HATPase_C.
IPR004358. Sig_transdc_His_kin-like_C.
IPR005467. Sig_transdc_His_kinase_core.
IPR009082. Sig_transdc_His_kinase_dimeric.
IPR001789. Sig_transdc_resp-reg_receiver.
[Graphical view]
PfamiPF03924. CHASE. 1 hit.
PF02518. HATPase_c. 1 hit.
PF00512. HisKA. 1 hit.
PF00072. Response_reg. 1 hit.
[Graphical view]
PRINTSiPR00344. BCTRLSENSOR.
SMARTiSM01079. CHASE. 1 hit.
SM00387. HATPase_c. 1 hit.
SM00388. HisKA. 1 hit.
SM00448. REC. 1 hit.
[Graphical view]
SUPFAMiSSF47384. SSF47384. 1 hit.
SSF52172. SSF52172. 2 hits.
SSF55874. SSF55874. 2 hits.
PROSITEiPS50839. CHASE. 1 hit.
PS50109. HIS_KIN. 1 hit.
PS50110. RESPONSE_REGULATORY. 1 hit.
[Graphical view]

Sequences (2)i

Sequence statusi: Complete.

This entry describes 2 isoformsi produced by alternative splicing. Align

Isoform 1 (identifier: Q9C5U0-1) [UniParc]FASTAAdd to Basket

Also known as: CRE1b

This isoform has been chosen as the 'canonical' sequence. All positional information in this entry refers to it. This is also the sequence that appears in the downloadable versions of the entry.

« Hide

        10         20         30         40         50
MRRDFVYNNN AMFNPLTTHY SSDMNWALNN HQEEEEEPRR IEISDSESLE
60 70 80 90 100
NLKSSDFYQL GGGGALNSSE KPRKIDFWRS GLMGFAKMQQ QQQLQHSVAV
110 120 130 140 150
KMNNNNNNDL MGNKKGSTFI QEHRALLPKA LILWIIIVGF ISSGIYQWMD
160 170 180 190 200
DANKIRREEV LVSMCDQRAR MLQDQFSVSV NHVHALAILV STFHYHKNPS
210 220 230 240 250
AIDQETFAEY TARTAFERPL LSGVAYAEKV VNFEREMFER QHNWVIKTMD
260 270 280 290 300
RGEPSPVRDE YAPVIFSQDS VSYLESLDMM SGEEDRENIL RARETGKAVL
310 320 330 340 350
TSPFRLLETH HLGVVLTFPV YKSSLPENPT VEERIAATAG YLGGAFDVES
360 370 380 390 400
LVENLLGQLA GNQAIVVHVY DITNASDPLV MYGNQDEEAD RSLSHESKLD
410 420 430 440 450
FGDPFRKHKM ICRYHQKAPI PLNVLTTVPL FFAIGFLVGY ILYGAAMHIV
460 470 480 490 500
KVEDDFHEMQ ELKVRAEAAD VAKSQFLATV SHEIRTPMNG ILGMLAMLLD
510 520 530 540 550
TELSSTQRDY AQTAQVCGKA LIALINEVLD RAKIEAGKLE LESVPFDIRS
560 570 580 590 600
ILDDVLSLFS EESRNKSIEL AVFVSDKVPE IVKGDSGRFR QIIINLVGNS
610 620 630 640 650
VKFTEKGHIF VKVHLAEQSK DESEPKNALN GGVSEEMIVV SKQSSYNTLS
660 670 680 690 700
GYEAADGRNS WDSFKHLVSE EQSLSEFDIS SNVRLMVSIE DTGIGIPLVA
710 720 730 740 750
QGRVFMPFMQ ADSSTSRNYG GTGIGLSISK CLVELMRGQI NFISRPHIGS
760 770 780 790 800
TFWFTAVLEK CDKCSAINHM KKPNVEHLPS TFKGMKAIVV DAKPVRAAVT
810 820 830 840 850
RYHMKRLGIN VDVVTSLKTA VVAAAAFERN GSPLPTKPQL DMILVEKDSW
860 870 880 890 900
ISTEDNDSEI RLLNSRTNGN VHHKSPKLAL FATNITNSEF DRAKSAGFAD
910 920 930 940 950
TVIMKPLRAS MIGACLQQVL ELRKTRQQHP EGSSPATLKS LLTGKKILVV
960 970 980 990 1000
DDNIVNRRVA AGALKKFGAE VVCAESGQVA LGLLQIPHTF DACFMDIQMP
1010 1020 1030 1040 1050
QMDGFEATRQ IRMMEKETKE KTNLEWHLPI LAMTADVIHA TYEECLKSGM
1060 1070 1080
DGYVSKPFEE ENLYKSVAKS FKPNPISPSS
Length:1,080
Mass (Da):120,731
Last modified:June 1, 2001 - v1
Checksum:i5950DB968B529401
GO
Isoform 2 (identifier: Q9C5U0-2) [UniParc]FASTAAdd to Basket

Also known as: CRE1a

The sequence of this isoform differs from the canonical sequence as follows:
     1-23: Missing.

Show »
Length:1,057
Mass (Da):117,955
Checksum:iE1381C8685F0D787
GO

Experimental Info

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Sequence conflicti567 – 5671S → G in BAB33311. (PubMed:11234017)Curated
Sequence conflicti1018 – 10181T → A in BAB33311. (PubMed:11234017)Curated

Natural variant

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Natural varianti765 – 7651S → N in strain: cv. Se-0. 1 Publication

Alternative sequence

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Alternative sequencei1 – 2323Missing in isoform 2. 3 PublicationsVSP_039770Add
BLAST

Sequence databases

Select the link destinations:
EMBLi
GenBanki
DDBJi
Links Updated
AJ278528 mRNA. Translation: CAC18521.1.
AJ278529 mRNA. Translation: CAC18522.1.
AJ278530 mRNA. Translation: CAC18523.1.
AB049934 mRNA. Translation: BAB33310.1.
AB049935 mRNA. Translation: BAB33311.1.
AB046871 mRNA. Translation: BAB40776.1.
AC007069 Genomic DNA. Translation: AAD21777.2.
CP002685 Genomic DNA. Translation: AEC05505.1.
CP002685 Genomic DNA. Translation: AEC05506.1.
CP002685 Genomic DNA. Translation: AEC05507.1.
EF598292 Genomic DNA. Translation: ABQ85264.1.
EF598293 Genomic DNA. Translation: ABQ85265.1.
EF598294 Genomic DNA. Translation: ABQ85266.1.
EF598295 Genomic DNA. Translation: ABQ85267.1.
EF598296 Genomic DNA. Translation: ABQ85268.1.
EF598297 Genomic DNA. Translation: ABQ85269.1.
EF598298 Genomic DNA. Translation: ABQ85270.1.
EF598299 Genomic DNA. Translation: ABQ85271.1.
EF598300 Genomic DNA. Translation: ABQ85272.1.
EF598301 Genomic DNA. Translation: ABQ85273.1.
EF598302 Genomic DNA. Translation: ABQ85274.1.
EF598303 Genomic DNA. Translation: ABQ85275.1.
EF598304 Genomic DNA. Translation: ABQ85276.1.
EF598305 Genomic DNA. Translation: ABQ85277.1.
EF598306 Genomic DNA. Translation: ABQ85278.1.
EF598307 Genomic DNA. Translation: ABQ85279.1.
EF598308 Genomic DNA. Translation: ABQ85280.1.
EF598309 Genomic DNA. Translation: ABQ85281.1.
EF598310 Genomic DNA. Translation: ABQ85282.1.
EF598311 Genomic DNA. Translation: ABQ85283.1.
EF598312 Genomic DNA. Translation: ABQ85284.1.
EF598313 Genomic DNA. Translation: ABQ85285.1.
EF598314 Genomic DNA. Translation: ABQ85286.1.
EF598315 Genomic DNA. Translation: ABQ85287.1.
PIRiF84429.
RefSeqiNP_565277.1. NM_126244.2. [Q9C5U0-2]
NP_849925.1. NM_179594.1. [Q9C5U0-1]
NP_973396.1. NM_201667.1. [Q9C5U0-2]
UniGeneiAt.10485.

Genome annotation databases

EnsemblPlantsiAT2G01830.2; AT2G01830.2; AT2G01830. [Q9C5U0-1]
GeneIDi814714.
KEGGiath:AT2G01830.

Keywords - Coding sequence diversityi

Alternative splicing

Cross-referencesi

Sequence databases

Select the link destinations:
EMBLi
GenBanki
DDBJi
Links Updated
AJ278528 mRNA. Translation: CAC18521.1 .
AJ278529 mRNA. Translation: CAC18522.1 .
AJ278530 mRNA. Translation: CAC18523.1 .
AB049934 mRNA. Translation: BAB33310.1 .
AB049935 mRNA. Translation: BAB33311.1 .
AB046871 mRNA. Translation: BAB40776.1 .
AC007069 Genomic DNA. Translation: AAD21777.2 .
CP002685 Genomic DNA. Translation: AEC05505.1 .
CP002685 Genomic DNA. Translation: AEC05506.1 .
CP002685 Genomic DNA. Translation: AEC05507.1 .
EF598292 Genomic DNA. Translation: ABQ85264.1 .
EF598293 Genomic DNA. Translation: ABQ85265.1 .
EF598294 Genomic DNA. Translation: ABQ85266.1 .
EF598295 Genomic DNA. Translation: ABQ85267.1 .
EF598296 Genomic DNA. Translation: ABQ85268.1 .
EF598297 Genomic DNA. Translation: ABQ85269.1 .
EF598298 Genomic DNA. Translation: ABQ85270.1 .
EF598299 Genomic DNA. Translation: ABQ85271.1 .
EF598300 Genomic DNA. Translation: ABQ85272.1 .
EF598301 Genomic DNA. Translation: ABQ85273.1 .
EF598302 Genomic DNA. Translation: ABQ85274.1 .
EF598303 Genomic DNA. Translation: ABQ85275.1 .
EF598304 Genomic DNA. Translation: ABQ85276.1 .
EF598305 Genomic DNA. Translation: ABQ85277.1 .
EF598306 Genomic DNA. Translation: ABQ85278.1 .
EF598307 Genomic DNA. Translation: ABQ85279.1 .
EF598308 Genomic DNA. Translation: ABQ85280.1 .
EF598309 Genomic DNA. Translation: ABQ85281.1 .
EF598310 Genomic DNA. Translation: ABQ85282.1 .
EF598311 Genomic DNA. Translation: ABQ85283.1 .
EF598312 Genomic DNA. Translation: ABQ85284.1 .
EF598313 Genomic DNA. Translation: ABQ85285.1 .
EF598314 Genomic DNA. Translation: ABQ85286.1 .
EF598315 Genomic DNA. Translation: ABQ85287.1 .
PIRi F84429.
RefSeqi NP_565277.1. NM_126244.2. [Q9C5U0-2 ]
NP_849925.1. NM_179594.1. [Q9C5U0-1 ]
NP_973396.1. NM_201667.1. [Q9C5U0-2 ]
UniGenei At.10485.

3D structure databases

Select the link destinations:
PDBei
RCSB PDBi
PDBji
Links Updated
Entry Method Resolution (Å) Chain Positions PDBsum
3T4J X-ray 1.65 A/B 149-418 [» ]
3T4K X-ray 1.77 A/B 149-418 [» ]
3T4L X-ray 1.53 A/B 149-418 [» ]
3T4O X-ray 1.75 A/B 149-418 [» ]
3T4Q X-ray 2.30 A/B 149-418 [» ]
3T4S X-ray 1.60 A/B 149-418 [» ]
3T4T X-ray 1.70 A/B 149-418 [» ]
ProteinModelPortali Q9C5U0.
SMRi Q9C5U0. Positions 150-416, 465-752, 941-1060.
ModBasei Search...
MobiDBi Search...

Protein-protein interaction databases

BioGridi 117. 30 interactions.
IntActi Q9C5U0. 29 interactions.
STRINGi 3702.AT2G01830.2-P.

Chemistry

BindingDBi Q9C5U0.
ChEMBLi CHEMBL6124.

Proteomic databases

PaxDbi Q9C5U0.
PRIDEi Q9C5U0.

Protocols and materials databases

Structural Biology Knowledgebase Search...

Genome annotation databases

EnsemblPlantsi AT2G01830.2 ; AT2G01830.2 ; AT2G01830 . [Q9C5U0-1 ]
GeneIDi 814714.
KEGGi ath:AT2G01830.

Organism-specific databases

TAIRi AT2G01830.

Phylogenomic databases

eggNOGi COG0784.
InParanoidi Q9C5U0.
KOi K14489.
OMAi NQNRPSF.
PhylomeDBi Q9C5U0.

Enzyme and pathway databases

BioCyci ARA:AT2G01830-MONOMER.
ARA:GQT-91-MONOMER.
ARA:GQT-92-MONOMER.

Miscellaneous databases

EvolutionaryTracei Q9C5U0.

Gene expression databases

Genevestigatori Q9C5U0.

Family and domain databases

Gene3Di 1.10.287.130. 1 hit.
3.30.565.10. 2 hits.
InterProi IPR006189. CHASE.
IPR011006. CheY-like_superfamily.
IPR003661. EnvZ-like_dim/P.
IPR003594. HATPase_C.
IPR004358. Sig_transdc_His_kin-like_C.
IPR005467. Sig_transdc_His_kinase_core.
IPR009082. Sig_transdc_His_kinase_dimeric.
IPR001789. Sig_transdc_resp-reg_receiver.
[Graphical view ]
Pfami PF03924. CHASE. 1 hit.
PF02518. HATPase_c. 1 hit.
PF00512. HisKA. 1 hit.
PF00072. Response_reg. 1 hit.
[Graphical view ]
PRINTSi PR00344. BCTRLSENSOR.
SMARTi SM01079. CHASE. 1 hit.
SM00387. HATPase_c. 1 hit.
SM00388. HisKA. 1 hit.
SM00448. REC. 1 hit.
[Graphical view ]
SUPFAMi SSF47384. SSF47384. 1 hit.
SSF52172. SSF52172. 2 hits.
SSF55874. SSF55874. 2 hits.
PROSITEi PS50839. CHASE. 1 hit.
PS50109. HIS_KIN. 1 hit.
PS50110. RESPONSE_REGULATORY. 1 hit.
[Graphical view ]
ProtoNeti Search...

Publicationsi

« Hide 'large scale' publications
  1. "A novel two-component hybrid molecule regulates vascular morphogenesis of the Arabidopsis root."
    Maehoenen A.P., Bonke M., Kauppinen L., Riikonen M., Benfey P.N., Helariutta Y.
    Genes Dev. 14:2938-2943(2000) [PubMed] [Europe PMC] [Abstract]
    Cited for: NUCLEOTIDE SEQUENCE [MRNA] (ISOFORM 2), TISSUE SPECIFICITY, DEVELOPMENTAL STAGE, FUNCTION, MUTAGENESIS OF THR-301.
    Strain: cv. Columbia.
    Tissue: Root.
  2. Cited for: NUCLEOTIDE SEQUENCE [MRNA] (ISOFORMS 1 AND 2), FUNCTION, ENZYME REGULATION, MUTAGENESIS OF HIS-482; GLY-490 AND ASP-996.
    Strain: cv. Wassilewskija.
    Tissue: Seedling.
  3. "Novel family of sensor histidine kinase genes in Arabidopsis thaliana."
    Ueguchi C., Koizumi H., Suzuki T., Mizuno T.
    Plant Cell Physiol. 42:231-235(2001) [PubMed] [Europe PMC] [Abstract]
    Cited for: NUCLEOTIDE SEQUENCE [MRNA] (ISOFORM 2), TISSUE SPECIFICITY.
  4. Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
    Strain: cv. Columbia.
  5. The Arabidopsis Information Resource (TAIR)
    Submitted (APR-2011) to the EMBL/GenBank/DDBJ databases
    Cited for: GENOME REANNOTATION.
    Strain: cv. Columbia.
  6. "The genetic architecture of shoot branching in Arabidopsis thaliana: a comparative assessment of candidate gene associations vs. quantitative trait locus mapping."
    Ehrenreich I.M., Stafford P.A., Purugganan M.D.
    Genetics 176:1223-1236(2007) [PubMed] [Europe PMC] [Abstract]
    Cited for: NUCLEOTIDE SEQUENCE [GENOMIC DNA] OF 722-983, VARIANT ASN-765.
    Strain: cv. Ag-0, cv. An-1, cv. Br-0, cv. C24, cv. Ct-1, cv. Cvi-1, cv. Edi-0, cv. Ga-0, cv. Kas-2, cv. Kin-0, cv. Landsberg erecta, cv. Ll-0, cv. Lz-0, cv. Ms-0, cv. Mt-0, cv. Nd-1, cv. Nok-3, cv. Oy-0, cv. Se-0, cv. Sorbo, cv. Tsu-1, cv. Van-0, cv. Wa-1 and cv. Wassilewskija.
  7. "The Arabidopsis sensor His-kinase, AHk4, can respond to cytokinins."
    Suzuki T., Miwa K., Ishikawa K., Yamada H., Aiba H., Mizuno T.
    Plant Cell Physiol. 42:107-113(2001) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, INTERACTION WITH AHP1; AHP2; AHP3 AND AHP5.
  8. "The AHK4 gene involved in the cytokinin-signaling pathway as a direct receptor molecule in Arabidopsis thaliana."
    Ueguchi C., Sato S., Kato T., Tabata S.
    Plant Cell Physiol. 42:751-755(2001) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, DISRUPTION PHENOTYPE.
  9. "The Arabidopsis AHK4 histidine kinase is a cytokinin-binding receptor that transduces cytokinin signals across the membrane."
    Yamada H., Suzuki T., Terada K., Takei K., Ishikawa K., Miwa K., Yamashino T., Mizuno T.
    Plant Cell Physiol. 42:1017-1023(2001) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, ACTIVATION BY CYTOKININS, MUTAGENESIS OF THR-301.
  10. "CREam of cytokinin signalling: receptor identified."
    Schmuelling T.
    Trends Plant Sci. 6:281-284(2001) [PubMed] [Europe PMC] [Abstract]
    Cited for: REVIEW.
  11. "His-Asp phosphorelay signal transduction in higher plants: receptors and response regulators for cytokinin signaling in Arabidopsis thaliana."
    Oka A., Sakai H., Iwakoshi S.
    Genes Genet. Syst. 77:383-391(2002) [PubMed] [Europe PMC] [Abstract]
    Cited for: REVIEW.
  12. "Characterization of the ARR15 and ARR16 response regulators with special reference to the cytokinin signaling pathway mediated by the AHK4 histidine kinase in roots of Arabidopsis thaliana."
    Kiba T., Yamada H., Mizuno T.
    Plant Cell Physiol. 43:1059-1066(2002) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, MUTAGENESIS OF GLY-490.
  13. "Mutations at CRE1 impair cytokinin-induced repression of phosphate starvation responses in Arabidopsis."
    Franco-Zorrilla J.M., Martin A.C., Solano R., Rubio V., Leyva A., Paz-Ares J.
    Plant J. 32:353-360(2002) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, DISRUPTION PHENOTYPE, INDUCTION, MUTAGENESIS OF GLY-493; THR-1008 AND ALA-1032.
  14. "Two-component signal transduction pathways in Arabidopsis."
    Hwang I., Chen H.-C., Sheen J.
    Plant Physiol. 129:500-515(2002) [PubMed] [Europe PMC] [Abstract]
    Cited for: GENE FAMILY, NOMENCLATURE.
  15. "Histidine kinase homologs that act as cytokinin receptors possess overlapping functions in the regulation of shoot and root growth in Arabidopsis."
    Nishimura C., Ohashi Y., Sato S., Kato T., Tabata S., Ueguchi C.
    Plant Cell 16:1365-1377(2004) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, DISRUPTION PHENOTYPE, TISSUE SPECIFICITY, DEVELOPMENTAL STAGE.
  16. "Two cytokinin receptors of Arabidopsis thaliana, CRE1/AHK4 and AHK3, differ in their ligand specificity in a bacterial assay."
    Spichal L., Rakova N.Y., Riefler M., Mizuno T., Romanov G.A., Strnad M., Schmuelling T.
    Plant Cell Physiol. 45:1299-1305(2004) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION.
  17. "Interallelic complementation at the Arabidopsis CRE1 locus uncovers independent pathways for the proliferation of vascular initials and canonical cytokinin signalling."
    de Leon B.G.-P., Zorrilla J.M.F., Rubio V., Dahiya P., Paz-Ares J., Leyva A.
    Plant J. 38:70-79(2004) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, MUTAGENESIS OF THR-301 AND LEU-529.
  18. "A novel regulatory pathway of sulfate uptake in Arabidopsis roots: implication of CRE1/WOL/AHK4-mediated cytokinin-dependent regulation."
    Maruyama-Nakashita A., Nakamura Y., Yamaya T., Takahashi H.
    Plant J. 38:779-789(2004) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, MUTAGENESIS OF GLY-490.
  19. Cited for: FUNCTION, TISSUE SPECIFICITY, DISRUPTION PHENOTYPE.
  20. "A live cell hormone-binding assay on transgenic bacteria expressing a eukaryotic receptor protein."
    Romanov G.A., Spichal L., Lomin S.N., Strnad M., Schmuelling T.
    Anal. Biochem. 347:129-134(2005) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, BIOPHYSICOCHEMICAL PROPERTIES.
  21. "Interaction between phosphate-starvation, sugar, and cytokinin signaling in Arabidopsis and the roles of cytokinin receptors CRE1/AHK4 and AHK3."
    Franco-Zorrilla J.M., Martin A.C., Leyva A., Paz-Ares J.
    Plant Physiol. 138:847-857(2005) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, DISRUPTION PHENOTYPE.
  22. "Topolins and hydroxylated thidiazuron derivatives are substrates of cytokinin O-glucosyltransferase with position specificity related to receptor recognition."
    Mok M.C., Martin R.C., Dobrev P.I., Vankova R., Ho P.S., Yonekura-Sakakibara K., Sakakibara H., Mok D.W.
    Plant Physiol. 137:1057-1066(2005) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION.
  23. Cited for: FUNCTION, PHOSPHORYLATION AT HIS-482, MUTAGENESIS OF THR-301; HIS-482; PHE-708 AND ASP-996, DISRUPTION PHENOTYPE, DEVELOPMENTAL STAGE.
  24. "Analysis of protein interactions within the cytokinin-signaling pathway of Arabidopsis thaliana."
    Dortay H., Mehnert N., Buerkle L., Schmuelling T., Heyl A.
    FEBS J. 273:4631-4644(2006) [PubMed] [Europe PMC] [Abstract]
    Cited for: INTERACTION WITH AHP1; AHP2; AHP3; AHP5 AND AHK3.
  25. "Biochemical characteristics and ligand-binding properties of Arabidopsis cytokinin receptor AHK3 compared to CRE1/AHK4 as revealed by a direct binding assay."
    Romanov G.A., Lomin S.N., Schmuelling T.
    J. Exp. Bot. 57:4051-4058(2006) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, BIOPHYSICOCHEMICAL PROPERTIES.
  26. "Arabidopsis cytokinin receptor mutants reveal functions in shoot growth, leaf senescence, seed size, germination, root development, and cytokinin metabolism."
    Riefler M., Novak O., Strnad M., Schmuelling T.
    Plant Cell 18:40-54(2006) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, DISRUPTION PHENOTYPE.
  27. "Cytokinin receptors are required for normal development of auxin-transporting vascular tissues in the hypocotyl but not in adventitious roots."
    Kuroha T., Ueguchi C., Sakakibara H., Satoh S.
    Plant Cell Physiol. 47:234-243(2006) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, DISRUPTION PHENOTYPE, MUTAGENESIS OF MET-459 AND GLY-490.
  28. "The floral volatile, methyl benzoate, from snapdragon (Antirrhinum majus) triggers phytotoxic effects in Arabidopsis thaliana."
    Horiuchi J., Badri D.V., Kimball B.A., Negre F., Dudareva N., Paschke M.W., Vivanco J.M.
    Planta 226:1-10(2007) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION.
  29. "Identification of amino acid substitutions that render the Arabidopsis cytokinin receptor histidine kinase AHK4 constitutively active."
    Miwa K., Ishikawa K., Terada K., Yamada H., Suzuki T., Yamashino T., Mizuno T.
    Plant Cell Physiol. 48:1809-1814(2007) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, MUTAGENESIS OF THR-301; GLY-435; PHE-436; MET-447; VAL-471 AND MET-494.
  30. "Functional analysis of AHK1/ATHK1 and cytokinin receptor histidine kinases in response to abscisic acid, drought, and salt stress in Arabidopsis."
    Tran L.S., Urao T., Qin F., Maruyama K., Kakimoto T., Shinozaki K., Yamaguchi-Shinozaki K.
    Proc. Natl. Acad. Sci. U.S.A. 104:20623-20628(2007) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, INDUCTION, DISRUPTION PHENOTYPE.
  31. Cited for: FUNCTION, DISRUPTION PHENOTYPE.
  32. "Mechanism-based inhibitors of cytokinin oxidase/dehydrogenase attack FAD cofactor."
    Kopecny D., Sebela M., Briozzo P., Spichal L., Houba-Herin N., Masek V., Joly N., Madzak C., Anzenbacher P., Laloue M.
    J. Mol. Biol. 380:886-899(2008) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION.
  33. "Toward an interaction map of the two-component signaling pathway of Arabidopsis thaliana."
    Dortay H., Gruhn N., Pfeifer A., Schwerdtner M., Schmuelling T., Heyl A.
    J. Proteome Res. 7:3649-3660(2008) [PubMed] [Europe PMC] [Abstract]
    Cited for: INTERACTION WITH AHP2; AMPD; WNK5 AND AT4G15630.
  34. "The role of auxins and cytokinins in the mutualistic interaction between Arabidopsis and Piriformospora indica."
    Vadassery J., Ritter C., Venus Y., Camehl I., Varma A., Shahollari B., Novak O., Strnad M., Ludwig-Mueller J., Oelmueller R.
    Mol. Plant Microbe Interact. 21:1371-1383(2008) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION.
  35. "Cytokinins negatively regulate the root iron uptake machinery in Arabidopsis through a growth-dependent pathway."
    Seguela M., Briat J.-F., Vert G., Curie C.
    Plant J. 55:289-300(2008) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, DISRUPTION PHENOTYPE.
  36. "The purine derivative PI-55 blocks cytokinin action via receptor inhibition."
    Spichal L., Werner T., Popa I., Riefler M., Schmuelling T., Strnad M.
    FEBS J. 276:244-253(2009) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, ENZYME REGULATION.
  37. "STIMPY mediates cytokinin signaling during shoot meristem establishment in Arabidopsis seedlings."
    Skylar A., Hong F., Chory J., Weigel D., Wu X.
    Development 137:541-549(2010) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, DISRUPTION PHENOTYPE.
  38. "Cytokinin receptor antagonists derived from 6-benzylaminopurine."
    Nisler J., Zatloukal M., Popa I., Dolezal K., Strnad M., Spichal L.
    Phytochemistry 71:823-830(2010) [PubMed] [Europe PMC] [Abstract]
    Cited for: ENZYME REGULATION.
  39. "The cytokinin receptors of Arabidopsis are located mainly to the endoplasmic reticulum."
    Wulfetange K., Lomin S.N., Romanov G.A., Stolz A., Heyl A., Schmuelling T.
    Plant Physiol. 156:1808-1818(2011) [PubMed] [Europe PMC] [Abstract]
    Cited for: SUBCELLULAR LOCATION.
    Strain: cv. Columbia.
  40. "The Arabidopsis eukaryotic translation initiation factor eIF5A-2 regulates root protoxylem development by modulating cytokinin signaling."
    Ren B., Chen Q., Hong S., Zhao W., Feng J., Feng H., Zuo J.
    Plant Cell 25:3841-3857(2013) [PubMed] [Europe PMC] [Abstract]
    Cited for: INTERACTION WITH FBR12 AND AHP1.
  41. "Structural basis for cytokinin recognition by Arabidopsis thaliana histidine kinase 4."
    Hothorn M., Dabi T., Chory J.
    Nat. Chem. Biol. 7:766-768(2011) [PubMed] [Europe PMC] [Abstract]
    Cited for: X-RAY CRYSTALLOGRAPHY (1.53 ANGSTROMS) OF 149-418 IN COMPLEX WITH CYTOKININS OR THIADIAZURON, SUBUNIT, MUTAGENESIS OF ALA-225; ALA-227; TYR-273; MET-279; ASP-285; THR-301; PRO-303; PHE-304; LEU-306; LEU-307; THR-317 AND GLY-343.

Entry informationi

Entry nameiAHK4_ARATH
AccessioniPrimary (citable) accession number: Q9C5U0
Secondary accession number(s): A5YY60
, A5YY75, Q9C5T8, Q9C5T9, Q9FDZ3, Q9SIT0
Entry historyi
Integrated into UniProtKB/Swiss-Prot: October 5, 2010
Last sequence update: June 1, 2001
Last modified: November 26, 2014
This is version 97 of the entry and version 1 of the sequence. [Complete history]
Entry statusiReviewed (UniProtKB/Swiss-Prot)
Annotation programPlant Protein Annotation Program

Miscellaneousi

Keywords - Technical termi

3D-structure, Complete proteome, Reference proteome

Documents

  1. Arabidopsis thaliana
    Arabidopsis thaliana: entries and gene names
  2. PDB cross-references
    Index of Protein Data Bank (PDB) cross-references
  3. SIMILARITY comments
    Index of protein domains and families

External Data

Dasty 3