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Information on EC 2.7.13.3 - histidine kinase and Organism(s) Arabidopsis thaliana and UniProt Accession P49333
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2.7.13.3 histidine kinaseIUBMB Comments
This entry has been included to accommodate those protein-histidine kinases for which the phosphorylation site has not been established (i.e. either the pros- or tele-nitrogen of histidine). A number of histones can act as acceptor.
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Word Map
- 2.7.13.3
- autophosphorylation
- photoreceptors
-
far-red
- seedling
-
phosphorelay
- chromophore
-
perception
-
regulons
- hypocotyls
-
quorum
-
perceived
- pfr
-
chemoreceptor
-
senses
-
light-induced
- chey
-
flagellar
- synechocystis
-
etiolated
- caulobacter
-
photomorphogenesis
-
transmitter
-
quorum-sensing
-
fluence
- biliverdin
-
cryptochrome
- crescentus
-
methyl-accepting
-
dark-grown
- radiodurans
-
transphosphorylation
-
autoinducer
-
histidine-containing
-
chemosensory
-
photomorphogenic
-
extracytoplasmic
- arca
- deinococcus
- phototropins
- tetrapyrrole
- ompc
- c-di-gmp
-
diguanylate
- spo0a
-
four-helix
-
gravitropism
-
photoresponses
-
non-cognate
- xanthus
-
photoisomerization
- synthesis
- medicine
The taxonomic range for the selected organisms is: Arabidopsis thaliana
The enzyme appears in selected viruses and cellular organisms
The enzyme appears in selected viruses and cellular organisms
Synonyms
histidine kinase, sensor kinase, sensor protein, phytochrome a, ethylene receptor, sensor histidine kinase, bacteriophytochrome, ornithine decarboxylase antizyme, chemotaxis protein, hybrid histidine kinase, 
more
topSYNONYM 
ORGANISM
UNIPROT
COMMENTARY 
LITERATURE
AHK5
CKI1
ethylene resistant 1
-
the carboxyl-terminal part of ethylene resistant 1 consists of a histidine kinase domain and a response regulator domain
histidine kinase CKI1
additional information
-
AHK5 belongs to the the canonical HK class of proteins in Arabidopsis thaliana
REACTION TYPE
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
phospho group transfer
-
-
-
-
SYSTEMATIC NAME
IUBMB Comments
ATP:protein-L-histidine N-phosphotransferase
This entry has been included to accommodate those protein-histidine kinases for which the phosphorylation site has not been established (i.e. either the pros- or tele-nitrogen of histidine). A number of histones can act as acceptor.
CAS REGISTRY NUMBER
COMMENTARY
99283-67-7
protein-histidine kinases, EC 2.7.13.1, EC 2.7.13.2, and EC 2.7.13.3 are not distinguished in Chemical Abstracts
SUBSTRATE
PRODUCT
REACTION DIAGRAM
ORGANISM
UNIPROT
COMMENTARY
(Substrate)
(Substrate)
LITERATURE
(Substrate)
(Substrate)
COMMENTARY
(Product)
(Product)
LITERATURE
(Product)
(Product)
Reversibility
r=reversible
ir=irreversible
?=not specified
r=reversible
ir=irreversible
?=not specified
ATP + protein L-histidine
ADP + protein N-phospho-L-histidine
-
canonical histidine kinase activity of the transmitter domain of the ETR1 ethylene receptor from Arabidopsis is not required for signal transmission
-
-
?
additional information
?
-
ETR1 acts as an ethylene receptor
-
-
?
additional information
?
-
ETR1 acts as an ethylene receptor
-
-
?
additional information
?
-
ETR1 acts as an ethylene receptor
-
-
?
additional information
?
-
ETR1 acts as an ethylene receptor
-
-
?
additional information
?
-
ETR1 acts as an ethylene receptor
-
-
?
additional information
?
-
-
histidine kinases are part of two-component signal transduction systems that act to integrate environmental stimuli into a cellular response via a phosphotransfer relay mechanism, e.g. involved in stomatal guard cell response to H2O2, regulation of the physiological function, overview
-
-
?
additional information
?
-
-
the enzyme performs regulatory autophosphorylation
-
-
?
additional information
?
-
CKI1 interacts with proteins AHP2 and AHP3, but CKI1 is incapable of binding to cytokinins
-
-
?
NATURAL SUBSTRATE
NATURAL PRODUCT
REACTION DIAGRAM
ORGANISM
UNIPROT
COMMENTARY
(Substrate)
(Substrate)
LITERATURE
(Substrate)
(Substrate)
COMMENTARY
(Product)
(Product)
LITERATURE
(Product)
(Product)
REVERSIBILITY
r=reversible
ir=irreversible
?=not specified
r=reversible
ir=irreversible
?=not specified
ATP + protein L-histidine
ADP + protein N-phospho-L-histidine
-
canonical histidine kinase activity of the transmitter domain of the ETR1 ethylene receptor from Arabidopsis is not required for signal transmission
-
-
?
additional information
?
-
ETR1 acts as an ethylene receptor
-
-
?
additional information
?
-
ETR1 acts as an ethylene receptor
-
-
?
additional information
?
-
ETR1 acts as an ethylene receptor
-
-
?
additional information
?
-
ETR1 acts as an ethylene receptor
-
-
?
additional information
?
-
ETR1 acts as an ethylene receptor
-
-
?
additional information
?
-
-
histidine kinases are part of two-component signal transduction systems that act to integrate environmental stimuli into a cellular response via a phosphotransfer relay mechanism, e.g. involved in stomatal guard cell response to H2O2, regulation of the physiological function, overview
-
-
?
additional information
?
-
CKI1 interacts with proteins AHP2 and AHP3, but CKI1 is incapable of binding to cytokinins
-
-
?
METALS and IONS
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
INHIBITOR
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
IMAGE
cyanide
-
autokinase activity of the purified ethylene resistant 1 is completely abolished by 2 mM cyanide
ethylene
-
autokinase activity of the purified ethylene resistant 1 is completely abolished by 2 mM ethylene
ORGANISM
COMMENTARY
LITERATURE
UNIPROT
SEQUENCE DB
SOURCE
SOURCE TISSUE
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
SOURCE
additional information
additional information
no expression of CKI1 in seedling, roots, stem, and leaves
additional information
-
spatial distribution of endoplasmic reticulum-localized CKI1 in the syncytial embryo sac is mediated by nuclear position and migration, overview
LOCALIZATION
ORGANISM
UNIPROT
COMMENTARY
GeneOntology No.
LITERATURE
SOURCE
ETR1 of Arabidopsis contains transmembrane domains responsible for ethylene binding and membrane localization
the hybrid histidine kinase is localized in both the cytoplasm and the plasma membrane
the hybrid histidine kinase is localized in both the cytoplasm and the plasma membrane
GENERAL INFORMATION
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
evolution
-
the Arabidopsis thaliana genome encodes eight canonical HKs (AHK1-5, ETR1, ERS1, and CKI1), five canonical AHPs (AHP1-5), and one pseudo-AHP (AHP6) that carries an asparagine instead of the critical histidine residue
malfunction
metabolism
gene CKI1 is part of the genetic pathway consisting of CKI1, AHPs, and type-B ARRs in the regulation of female gametophyte development and vegetative growth
physiological function
additional information
-
structure-function analysis histidine kinase AHK5 bound to its cognate phosphotransfer protein AHP1, overview. AHP1 binds AHK5RD via a prominent hydrogen bond docking ridge and a hydrophobic patch. AHK5RD binds to AHP1-3 with similar, micromolar affinity, consistent with the transient nature of this signaling complex. Architecture of the AHK5RD-AHP1 Interface, overview
malfunction
-
Arabidopsis mutants lacking HK5 show reduced stomatal closure in response to H2O2, overexpression of HK5 results in constitutively less stomatal closure
malfunction
-
genetic modifications of cytokinin-independent1 activity in Arabidopsis cause dysfunction of the two-component signaling pathway and defects in procambial cell maintenance, loss-of-function histidine kinase2 and histidine kinase3 mutants show defects in procambium proliferation and an absence of secondary growth
malfunction
-
ectopic CKI1 expression generates non-propagating seeds with dual fertilized endosperms and no embryos. CKI1 loss-of-function mutants exhibit gametophytic defects resulting in semi-sterility. Cell specification is defective in cki1 mutant embryo sacs. Autonomous endosperms arise from ectopic central cells in the PRC2 fis2 (fertilization independent 2) mutant
malfunction
loss of function of AHK5 also confers tolerance to high salinity, suggesting that AHK5 acts to integrate multiple stress responses. The AHK5 mutant is more resistant to salt stress, the insensitivity of the ahk5-1 mutant in response to salinity is caused by loss of AHK5 function
malfunction
the female gametophytic lethal phenotype of cki1-5 (a null mutant) can be partially rescued by IPT8 or ARR1 (a type-B Arabidopsis RR) driven by a CKI1 promoter. A mutant allele, cki1-8, is cki1-8 is a weak mutant allele with residual CKI1 expression, and can be transmitted through female gametophytes with low frequency, viable homozygous cki1-8 mutant plants that grow larger than wild-type plants, show defective megagametogenesis, and rarely set enlarged seeds. The cki1-8 mutation impairs female gametophyte development, phenotype overview. Mutant cki1-8 responds normally to cytokinin, remarkably smaller siliques are formed in cki1-8 plants than in the wild type
physiological function
-
cytokinin-independent1 and Arabidopsis histidine kinase2 and 3 regulate vascular tissue development in Arabidopsis shoots, the cytokinin-independent activity of CKI1 and cytokinin-induced histidine kinase2 and histidine kinase3 are important for vascular bundle formation in Arabidopsis
physiological function
-
functional HK5 histidine kinase is required for H2O2 responses in stomatal guard cells
physiological function
-
the histidine kinase-related domain of phytochrome A controls the spectral sensitivity and the subcellular distribution of the photoreceptor
physiological function
-
Arabidopsis thaliana endosperm formation requires the cytokinin independent 1 (CKI1) histidine kinase, an activator of the cytokinin signaling pathway, which specifies central cells and restricts egg cell fate. Seeds of flowering plants contain embryos and nutritive endosperm arising from double fertilization of two dimorphic female gametes. The key to specification of only one female gamete for endosperm formation is the regulated localization of the CKI1 histidine kinase by expression control and protein translocation. Dimorphism of the two adjacent gametes is mechanistically established in the syncytial embryo sac by spatially restricted CKI1 expression, followed by translocation of endoplasmic reticulum-localized CKI1 protein via nuclear migration. CKI1-directed specification of the endosperm precursor central cell results in seeds containing an embryo and an endosperm, mechanism, overview
physiological function
cytokinin signaling is mediated by a multiple-step phosphorelay. Key components of the phosphorelay consist of the histidine kinase (HK)-type receptors, histidine phosphotransfer proteins (HP), and response regulators (RRs). Histidine kinase CKI1 acts upstream of histidine phosphotransfer proteins to regulate female gametophyte development and vegetative growth, role of AHP proteins in female gametophyte development, overview
physiological function
the hybrid histidine kinase Arabidopsis histidine kinase 5 (AHK5) mediates stomatal responses to exogenous and endogenous signals in Arabidopsis thaliana, AHK5 integrates abiotic and biotic stimuli in stomatal guard cells through regulation of H2O2 homeostasis. Role for AHK5 in the regulation of survival following challenge by a hemi-biotrophic bacterium and a necrotrophic fungus, as well as in the growth response to salt stress. AHK5 positively regulates salt sensitivity and contributes to resistance to the bacterium Pseudomonas syringae pv. tomato DC3000 and the fungal pathogen Botrytis cinerea, detailed overview. Function of AHK5 in regulating the production of hormones and redox homeostasis. Enzyme AHK5 functions as a negative regulator of root growth inhibition mediated by abscisic acid/ethylene. AHK5 positively regulates salt-induced root growth inhibition
UNIPROT
ENTRY NAME
ORGANISM
NO. OF AA
NO. OF TRANSM. HELICES
MOLECULAR WEIGHT[Da]
SOURCE
SEQUENCE
LOCALIZATION PREDICTION
The reliability class of the localization prediction ranges from 1 (very reliable) to 5 (not reliable).
The reliability class of the localization prediction ranges from 1 (very reliable) to 5 (not reliable). ETR1_ARATH
738
3
82566
Swiss-Prot
other Location (Reliability: 2)
PDB
SCOP
CATH
UNIPROT
ORGANISM
MOLECULAR WEIGHT
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
SUBUNIT
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
additional information
high affinity binding is observed between AHK1 monomers resulting in the formation of the dimer present at physiological conditions. A high affinity interaction is also observed between AHK1 and AHP2 transfer protein
POSTTRANSLATIONAL MODIFICATION
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
no phosphoprotein
no automodification of AHK1 with ATP is observed
additional information
-
the enzyme performs regulatory autophosphorylation
CRYSTALLIZATION (Commentary)
ORGANISM
UNIPROT
LITERATURE
hanging drop vapor diffusion method, using 2.54 M ammonium sulfate, 15.9% (v/v) glycerol, and 0.1 M MES buffer pH 5.05
-
puriifed kinase AHK5RD in complex with AHP1, X-ray diffraction structure determination and analysis at 1.95 A resolution
-
structures of free, Mg2+-bound, and beryllofluoridated CKI1RD (a stable analogue of the labile phosphorylated form) are identical and similar to the active state of receiver domains of bacterial response regulator. Mg2+ binding and beryllofluoridation alter the conformational equilibrium of the beta3-alpha3 loop close to the phosphorylation site. Mutations that perturbe the conformational behavior of the beta3-alpha3 loop while keeping the active-site aspartate intact result in suppression of CKI1 function. The beta3-alpha3 loop may directly interact with the ATP-binding site of the CKI1 histidine kinase domain
PROTEIN VARIANTS
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
D1050A
mutation changes conformation of the beta3-alpha3 loop in solution and shifts residue F1102 to the inactive orientation
D1050E
mutation changes conformation of the beta3-alpha3 loop in solution and shifts residue F1102 to the inactive orientation
additional information
additional information
-
construction of ahk5 T-DNA insertion mutants, Arabidopsis mutants lacking AHK5 show reduced stomatal closure in response to H2O2, which is reversed by complementation with the wild type gene. Overexpression of AHK5 results in constitutively less stomatal closure. Abiotic stimuli that generate endogenous H2O2, such as darkness, nitric oxide and the phytohormone ethylene, also show reduced stomatal closure in the ahk5 mutants. However, ABA causes closure, dark adaptation induces H2O2 production and H2O2 induces NO synthesis in mutants, phenotype, overview
additional information
-
generation of CKI1 loss-of-function mutants. Transformation of egg cell to central cell by ectopic CKI1 retains female gametophyte organization
additional information
generation of the ahk5-1 mutant containing a T-DNA insertion in the receiver domain of the AHK5 histidine kinase. The AHK5 mutant is more resistant to salt stress. Two complemented lines, PAHK5-AHK5/ahk5-1-1 and PAHK5-AHK5/ahk5-1-4, are generated by complementation of the ahk5-1 mutant with full-length AHK5 including 3205 bases upstream of the ATG start codon
additional information
-
generation of the ahk5-1 mutant containing a T-DNA insertion in the receiver domain of the AHK5 histidine kinase. The AHK5 mutant is more resistant to salt stress. Two complemented lines, PAHK5-AHK5/ahk5-1-1 and PAHK5-AHK5/ahk5-1-4, are generated by complementation of the ahk5-1 mutant with full-length AHK5 including 3205 bases upstream of the ATG start codon
additional information
the T-DNA insertion is transmitted through female gametophytes at an extremely low frequency in CKI1/cki1-8 plants, while cki1-6 mutation, which carries a T-DNA at a more distal location in the CKI1 promoter than cki1-8 mutatin does not appear to be capable of being transmitted through female gametophytes. The cki1-8 mutation impairs female gametophyte development, cki1-8 mutant phenotype during reproductive development and vegetative growth. Mutant phenotypes, overview
PURIFICATION (Commentary)
ORGANISM
UNIPROT
LITERATURE
HiTrap Zn2+-chelating column chromatography, Superdex 75 gel filtration
-
recombinant protein, 1% (w/v) FosCholine-16 is most efficient in solubilizing the recombinant histidine kinase from the bacterial membranes
CLONED (Commentary)
ORGANISM
UNIPROT
LITERATURE
ectopic CKI1 expression in all nuclei of the female gametophyte, under the pAKV promoter using the pOp/LhG4 transactivation system, generates non-propagating seeds with dual fertilized endosperms and no embryos, phenotypes, detailed overview
-
expressed in Escherichia coli strains BL21(DE3), C41 (DE3), and C43 (DE3)
-
gene ahk5, transient expression of GST-tagged full-length AHK5 in Arabidopsis thaliana protoplasts and Nicotiana benthamiana leaf cells the fusion protein in the cytoplasm, independent of whether the GFP tag is fused to the N-terminus or C-terminus of AHK5, expression profile of AHK5 in different tissues and cell types by semi-quantitative RT-PCR
-
overexpression of nonreceptor-type HK gene cytokinin independent1 (CKI1) activates cytokinin signaling, quantitative RT-PCR expression analysis
REF.
AUTHORS
TITLE
JOURNAL
VOL.
PAGES
YEAR
ORGANISM (UNIPROT)
PUBMED ID
SOURCE
Chang, C.; Kwok, S.F.; Bleecker, A.B.; Meyerowitz, E.M.
Arabidopsis ethylene-response gene ETR1: similarity of product to two-component regulators
Science
262
539-544
1993
Arabidopsis thaliana (P49333)
Chen, Y.F.; Randlett, M.D.; Findell, J.L.; Schaller, G.E.
Localization of the ethylene receptor ETR1 to the endoplasmic reticulum of Arabidopsis
J. Biol. Chem.
277
19861-19866
2002
Arabidopsis thaliana (P49333)
Rodriguez, F.I.; Esch, J.J.; Hall, A.E.; Binder, B.M.; Schaller, G.E.; Bleecker, A.B.
A copper cofactor for the ethylene receptor ETR1 from Arabidopsis
Science
283
996-998
1999
Arabidopsis thaliana (P49333)
Schaller, G.E.; Bleecker, A.B.
Ethylene-binding sites generated in yeast expressing the Arabidopsis ETR1 gene
Science
270
1809-1811
1995
Arabidopsis thaliana (P49333), Arabidopsis thaliana
Schaller, G.E.; Ladd, A.N.; Lanahan, M.B.; Spanbauer, J.M.; Bleecker, A.B.
The ethylene response mediator ETR1 from Arabidopsis forms a disulfide-linked dimer
J. Biol. Chem.
270
12526-12530
1995
Arabidopsis thaliana (P49333)
Theologis, A.; Ecker, J.R.; Palm, C.J.; et al.
Sequence and analysis of chromosome 1 of the plant Arabidopsis thaliana
Nature
408
816-820
2000
Arabidopsis thaliana (P49333)
Wang, W.; Hall, A.E.; O'Malley, R.; Bleecker, A.B.
Canonical histidine kinase activity of the transmitter domain of the ETR1 ethylene receptor from Arabidopsis is not required for signal transmission
Proc. Natl. Acad. Sci. USA
100
352-357
2003
Arabidopsis thaliana
Desikan, R.; Horak, J.; Chaban, C.; Mira-Rodado, V.; Witthoeft, J.; Elgass, K.; Grefen, C.; Cheung, M.K.; Meixner, A.J.; Hooley, R.; Neill, S.J.; Hancock, J.T.; Harter, K.
The histidine kinase AHK5 integrates endogenous and environmental signals in Arabidopsis guard cells
PLoS ONE
3
e2491
2008
Arabidopsis thaliana
Klumpler, T.; Pekarova, B.; Marek, J.; Borkovcova, P.; Janda, L.; Hejatko, J.
Cloning, purification, crystallization and preliminary X-ray analysis of the receiver domain of the histidine kinase CKI1 from Arabidopsis thaliana
Acta Crystallogr. Sect. F
65
478-481
2009
Arabidopsis thaliana
Voet-van-Vormizeele, J.; Groth, G.
Ethylene controls autophosphorylation of the histidine kinase domain in ethylene receptor ETR1
Mol. Plant
1
380-387
2008
Arabidopsis thaliana
Hejatko, J.; Ryu, H.; Kim, G.T.; Dobesova, R.; Choi, S.; Choi, S.M.; Soucek, P.; Horak, J.; Pekarova, B.; Palme, K.; Brzobohaty, B.; Hwang, I.
The histidine kinases cytokinin-independent1 and Arabidopsis histidine kinase2 and 3 regulate vascular tissue development in Arabidopsis shoots
Plant Cell
21
2008-2021
2009
Arabidopsis thaliana
Mueller, R.; Fernandez, A.P.; Hiltbrunner, A.; Schaefer, E.; Kretsch, T.
The histidine kinase-related domain of Arabidopsis phytochrome A controls the spectral sensitivity and the subcellular distribution of the photoreceptor
Plant Physiol.
150
1297-1309
2009
Arabidopsis thaliana
Yuan, L.; Liu, Z.; Song, X.; Johnson, C.; Yu, X.; Sundaresan, V.
The CKI1 histidine kinase specifies the female gametic precursor of the endosperm
Dev. Cell
37
34-46
2016
Arabidopsis thaliana, Arabidopsis thaliana Col-0
Bauer, J.; Reiss, K.; Veerabagu, M.; Heunemann, M.; Harter, K.; Stehle, T.
Structure-function analysis of Arabidopsis thaliana histidine kinase AHK5 bound to its cognate phosphotransfer protein AHP1
Mol. Plant
6
959-970
2013
Arabidopsis thaliana
Pham, J.; Liu, J.; Bennett, M.H.; Mansfield, J.W.; Desikan, R.
Arabidopsis histidine kinase 5 regulates salt sensitivity and resistance against bacterial and fungal infection
New Phytol.
194
168-180
2012
Arabidopsis thaliana (Q3S4A7), Arabidopsis thaliana, Arabidopsis thaliana Col-0 (Q3S4A7)
Deng, Y.; Dong, H.; Mu, J.; Ren, B.; Zheng, B.; Ji, Z.; Yang, W.C.; Liang, Y.; Zuo, J.
Arabidopsis histidine kinase CKI1 acts upstream of histidine phosphotransfer proteins to regulate female gametophyte development and vegetative growth
Plant Cell
22
1232-1248
2010
Arabidopsis thaliana (O22267), Arabidopsis thaliana Col-0 (O22267)
Otrusinova, O.; Demo, G.; Padrta, P.; Jasenakova, Z.; Pekarova, B.; Gelova, Z.; Szmitkowska, A.; Kaderavek, P.; Jansen, S.; Zachrdla, M.; Klumpler, T.; Marek, J.; Hritz, J.; Janda, L.; Iwai, H.; Wimmerova, M.; Hejatko, J.; Zidek, L.
Conformational dynamics are a key factor in signaling mediated by the receiver domain of a sensor histidine kinase from Arabidopsis thaliana
J. Biol. Chem.
292
17525-17540
2017
Arabidopsis thaliana (O22267), Arabidopsis thaliana
EXTERNAL LINKS (specific for EC-Number 2.7.13.3)
Transporter Classification Database (TCDB): 9.B.33.1.1, 9.B.238.3.5, 9.B.33.1.2, 2.A.21.9.1, 2.A.21.9.2, 2.A.123.2.13
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