ATP2B1 ATPase plasma membrane Ca2+ transporting 1 [Homo sapiens (human)] - Gene

Summary

Official Symbol: ATP2B1provided by HGNC
Official Full Name: ATPase plasma membrane Ca2+ transporting 1provided by HGNC
Primary source: HGNC:HGNC:814
See related: Ensembl:ENSG00000070961 MIM:108731
Gene type: protein coding
RefSeq status: REVIEWED
Organism: Homo sapiens
Lineage: Eukaryota; Metazoa; Chordata; Craniata; Vertebrata; Euteleostomi; Mammalia; Eutheria; Euarchontoglires; Primates; Haplorrhini; Catarrhini; Hominidae; Homo
Also known as: PMCA1; PMCA1kb
Summary: The protein encoded by this gene belongs to the family of P-type primary ion transport ATPases characterized by the formation of an aspartyl phosphate intermediate during the reaction cycle. These enzymes remove bivalent calcium ions from eukaryotic cells against very large concentration gradients and play a critical role in intracellular calcium homeostasis. The mammalian plasma membrane calcium ATPase isoforms are encoded by at least four separate genes and the diversity of these enzymes is further increased by alternative splicing of transcripts. The expression of different isoforms and splice variants is regulated in a developmental, tissue- and cell type-specific manner, suggesting that these pumps are functionally adapted to the physiological needs of particular cells and tissues. This gene encodes the plasma membrane calcium ATPase isoform 1. Alternatively spliced transcript variants encoding different isoforms have been identified. [provided by RefSeq, Jul 2008]
Expression: Ubiquitous expression in brain (RPKM 34.3), bone marrow (RPKM 16.4) and 23 other tissues See more
Orthologs: mouse all
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Genomic context

Location:: 12q21.33

Exon count:: 24

Annotation release	Status	Assembly	Chr	Location
109.20210226	current	GRCh38.p13 (GCF_000001405.39)	12	NC_000012.12 (89588049..89709366, complement)
105.20201022	previous assembly	GRCh37.p13 (GCF_000001405.25)	12	NC_000012.11 (89981826..90103143, complement)

Chromosome 12 - NC_000012.12 Genomic Context describing neighboring genes

Genomic regions, transcripts, and products

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Go to reference sequence details

Genomic Sequence:

Go to nucleotide: Graphics FASTA GenBank

Expression

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See details

Project title: HPA RNA-seq normal tissues
Description: RNA-seq was performed of tissue samples from 95 human individuals representing 27 different tissues in order to determine tissue-specificity of all protein-coding genes
BioProject: PRJEB4337
Publication: PMID 24309898
Analysis date: Wed Apr 4 07:08:55 2018

Bibliography

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GeneRIFs: Gene References Into Functions

What's a GeneRIF?

Crosstalk among Calcium ATPases: PMCA, SERCA and SPCA in Mental Diseases.
Title: Crosstalk among Calcium ATPases: PMCA, SERCA and SPCA in Mental Diseases.
Association of ATP2B1 gene polymorphism with incidence of eclampsia.
Title: Association of ATP2B1 gene polymorphism with incidence of eclampsia.
Association between ATP2B1 gene polymorphism and the onset of cerebral infarction.
Title: Association between ATP2B1 gene polymorphism and the onset of cerebral infarction.
ATP2B1 rs17249754 is positively associated with the risk of developing essential hypertension in patients from Burkina Faso, West Africa.
Title: Positive association between ATP2B1 rs17249754 and essential hypertension: a case-control study in Burkina Faso, West Africa.
No significant association was found between ATP2B1 gene polymorphism and the odds of high blood pressure.
Title: Association between ATP2B1 and CACNB2 polymorphisms and high blood pressure in a population of Lithuanian children and adolescents: a cross-sectional study.
enhanced eNOS activity induced by ATP2B1 gene silencing may be mediated via higher levels of intracellular Ca(2+), and the effect was confirmed to be dependent on the eNOS-calmodulin interaction
Title: ATP2B1 Gene Silencing Increases NO Production Under Basal Conditions Through the Ca2+/calmodulin/eNOS Signaling Pathway in Endothelial Cells.
rs17249754-A (OR = 0.43, p = 0.0002) and rs1401982-G (OR = 0.47, p = 0.0007) were associated with decreased susceptibility of concurrent extra and intracranial stenosis even after Bonferroni correction. These two minor alleles were also significantly associated with less stenotic arteries and moderate-to-severe stenosis.
Title: Association of ATP2B1 common variants with asymptomatic intracranial and extracranial large artery stenosis in hypertension patients.
Cryoelectron microscopy reveals the PMCA1 structure in complex with neuroplastin.
Title: Structure of the human plasma membrane Ca2+-ATPase 1 in complex with its obligatory subunit neuroplastin.
people with the major allele of ATP2B1 rs17249754 are susceptible to hypertension especially in low intake of Ca and high ratio of Na and K
Title: People with the major alleles of ATP2B1 rs17249754 increases the risk of hypertension in high ratio of sodium and potassium, and low calcium intakes.
Crossover analysis and stratified analysis indicated that BMI has a major effect on the development of hypertension, while ATP2B1 variants have a minor effect
Title: Gender-Specific Association of ATP2B1 Variants with Susceptibility to Essential Hypertension in the Han Chinese Population.

Submit: New GeneRIF Correction See all GeneRIFs (49)

Phenotypes

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Find tests for this gene in the NIH Genetic Testing Registry (GTR)

Review eQTL and phenotype association data in this region using PheGenI

Associated conditions

Description	Tests
A large-scale genome-wide association study of Asian populations uncovers genetic factors influencing eight quantitative traits. GeneReviews: Not available
Gene-age interactions in blood pressure regulation: a large-scale investigation with the CHARGE, Global BPgen, and ICBP Consortia. GeneReviews: Not available
Genome-wide association studies of serum magnesium, potassium, and sodium concentrations identify six Loci influencing serum magnesium levels. GeneReviews: Not available
Genome-wide association study identifies six new loci influencing pulse pressure and mean arterial pressure. GeneReviews: Not available
Genome-wide association study in Chinese identifies novel loci for blood pressure and hypertension. GeneReviews: Not available
Genome-wide association study in Han Chinese identifies four new susceptibility loci for coronary artery disease. GeneReviews: Not available
Genome-wide association study meta-analysis reveals transethnic replication of mean arterial and pulse pressure loci. GeneReviews: Not available
Genome-wide association study of blood pressure and hypertension. GeneReviews: Not available
Meta-analysis of genome-wide association studies identifies common variants associated with blood pressure variation in east Asians. GeneReviews: Not available

Variation

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See variants in ClinVar

See studies and variants in dbVar

See Variation Viewer (GRCh37.p13)

See Variation Viewer (GRCh38)

HIV-1 interactions

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Protein interactions

Protein	Gene	Interaction	Pubs
Vpu	vpu	HIV-1 Vpu is identified to have a physical interaction with ATPase, Ca++ transporting, plasma membrane 1 (ATP2B1) in human HEK293 and/or Jurkat cell lines by using affinity tagging and purification mass spectrometry analyses	PubMed

Go to the HIV-1, Human Interaction Database

Interactions

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Products	Interactant	Other Gene	Complex	Source	Pubs	Description

General gene information

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Markers

D12S2009 (e-PCR)
- UniSTS:6932

RH66423 (e-PCR)
- UniSTS:10569

G64515 (e-PCR)
- UniSTS:163920

MARC_21823-21824:1025099477:1 (e-PCR)
- UniSTS:268640

WI-7734 (e-PCR)
- UniSTS:10370

RH45012 (e-PCR)
- UniSTS:50850

EST13H11 (e-PCR)
- UniSTS:263102

ATP2B1_4221 (e-PCR)
- UniSTS:463553

WI-18859 (e-PCR)
- UniSTS:54722

D12S1250E (e-PCR)
- UniSTS:151498

Homology

Homologs of the ATP2B1 gene: The ATP2B1 gene is conserved in chimpanzee, Rhesus monkey, dog, cow, mouse, rat, chicken, zebrafish, A.thaliana, and rice.
Orthologs from Annotation Pipeline: 277 organisms have orthologs with human gene ATP2B1
Orthologs

Gene Ontology Provided by GOA

Function	Evidence Code	Pubs
enables ATP binding	IEA Inferred from Electronic Annotation more info
enables ATP hydrolysis activity	IDA Inferred from Direct Assay more info	PubMed
enables ATPase-coupled cation transmembrane transporter activity	IBA Inferred from Biological aspect of Ancestor more info	PubMed
enables P-type calcium transporter activity	IBA Inferred from Biological aspect of Ancestor more info	PubMed
enables P-type calcium transporter activity involved in regulation of presynaptic cytosolic calcium ion concentration	IDA Inferred from Direct Assay more info	PubMed
enables P-type calcium transporter activity involved in regulation of presynaptic cytosolic calcium ion concentration	IMP Inferred from Mutant Phenotype more info	PubMed
enables PDZ domain binding	IBA Inferred from Biological aspect of Ancestor more info	PubMed
enables calcium ion transmembrane transporter activity	IDA Inferred from Direct Assay more info	PubMed
enables calmodulin binding	IEA Inferred from Electronic Annotation more info
enables metal ion binding	IEA Inferred from Electronic Annotation more info
enables protein binding	IPI Inferred from Physical Interaction more info	PubMed

Process	Evidence Code	Pubs
involved_in aging	IEA Inferred from Electronic Annotation more info
involved_in brain development	IEA Inferred from Electronic Annotation more info
involved_in calcium ion export across plasma membrane	IDA Inferred from Direct Assay more info	PubMed
involved_in cellular calcium ion homeostasis	IBA Inferred from Biological aspect of Ancestor more info	PubMed
involved_in cellular response to corticosterone stimulus	IEA Inferred from Electronic Annotation more info
involved_in cellular response to vitamin D	IEA Inferred from Electronic Annotation more info
involved_in ion transmembrane transport	TAS Traceable Author Statement more info
involved_in negative regulation of cytokine production	ISS Inferred from Sequence or Structural Similarity more info
involved_in negative regulation of cytosolic calcium ion concentration	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in negative regulation of cytosolic calcium ion concentration	ISS Inferred from Sequence or Structural Similarity more info
involved_in neural retina development	IEA Inferred from Electronic Annotation more info
involved_in positive regulation of bone mineralization	ISS Inferred from Sequence or Structural Similarity more info
involved_in positive regulation of calcium ion transport	ISS Inferred from Sequence or Structural Similarity more info
involved_in regulation of blood pressure	ISS Inferred from Sequence or Structural Similarity more info
involved_in regulation of cardiac conduction	TAS Traceable Author Statement more info
involved_in regulation of cellular response to insulin stimulus	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in regulation of cytosolic calcium ion concentration	IBA Inferred from Biological aspect of Ancestor more info	PubMed
involved_in regulation of cytosolic calcium ion concentration	IDA Inferred from Direct Assay more info	PubMed
involved_in regulation of presynaptic cytosolic calcium ion concentration	IEA Inferred from Electronic Annotation more info
involved_in regulation of vascular associated smooth muscle contraction	ISS Inferred from Sequence or Structural Similarity more info
involved_in response to cold	IEA Inferred from Electronic Annotation more info

Component	Evidence Code	Pubs
located_in GABA-ergic synapse	IDA Inferred from Direct Assay more info	PubMed
located_in GABA-ergic synapse	IMP Inferred from Mutant Phenotype more info	PubMed
located_in apical plasma membrane	IEA Inferred from Electronic Annotation more info
located_in basolateral plasma membrane	ISS Inferred from Sequence or Structural Similarity more info
located_in cytoplasmic side of plasma membrane	IEA Inferred from Electronic Annotation more info
located_in dendritic spine membrane	IEA Inferred from Electronic Annotation more info
located_in extracellular exosome	HDA more info	PubMed
located_in glutamatergic synapse	IDA Inferred from Direct Assay more info	PubMed
located_in glutamatergic synapse	IMP Inferred from Mutant Phenotype more info	PubMed
located_in immunological synapse	ISS Inferred from Sequence or Structural Similarity more info
is_active_in integral component of plasma membrane	IBA Inferred from Biological aspect of Ancestor more info	PubMed
located_in integral component of presynaptic active zone membrane	IEA Inferred from Electronic Annotation more info
located_in integral component of presynaptic membrane	ISS Inferred from Sequence or Structural Similarity more info
located_in integral component of synaptic vesicle membrane	ISS Inferred from Sequence or Structural Similarity more info
is_active_in intracellular membrane-bounded organelle	IBA Inferred from Biological aspect of Ancestor more info	PubMed
located_in membrane	HDA more info	PubMed
located_in membrane raft	IEA Inferred from Electronic Annotation more info
located_in neuronal cell body membrane	IEA Inferred from Electronic Annotation more info
located_in nucleoplasm	IDA Inferred from Direct Assay more info
located_in plasma membrane	IDA Inferred from Direct Assay more info	PubMed
located_in plasma membrane	TAS Traceable Author Statement more info

General protein information

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Preferred Names: plasma membrane calcium-transporting ATPase 1

Names: ATPase, Ca++ transporting, plasma membrane 1; plasma membrane calcium pump

NP_001001323.1

EC 7.2.2.10

NP_001353449.1

EC 7.2.2.10

NP_001353450.1

EC 7.2.2.10

NP_001353451.1

EC 7.2.2.10

NP_001353452.1

EC 7.2.2.10

NP_001353453.1

EC 7.2.2.10

NP_001353454.1

EC 7.2.2.10

NP_001353455.1

EC 7.2.2.10

NP_001353456.1

EC 7.2.2.10

NP_001353457.1

EC 7.2.2.10

NP_001353458.1

EC 7.2.2.10

NP_001353459.1

EC 7.2.2.10

NP_001353460.1

EC 7.2.2.10

NP_001353461.1

EC 7.2.2.10

NP_001673.2

EC 7.2.2.10

XP_016874846.1

EC 7.2.2.10

XP_024304759.1

EC 7.2.2.10

XP_024304761.1

EC 7.2.2.10

NCBI Reference Sequences (RefSeq)

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RefSeqs maintained independently of Annotated Genomes

These reference sequences exist independently of genome builds. Explain

These reference sequences are curated independently of the genome annotation cycle, so their versions may not match the RefSeq versions in the current genome build. Identify version mismatches by comparing the version of the RefSeq in this section to the one reported in Genomic regions, transcripts, and products above.

Genomic

NG_029485.2 RefSeqGene

Range

4988..126305

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GenBank, FASTA, Sequence Viewer (Graphics)

mRNA and Protein(s)

NM_001001323.2 → NP_001001323.1 plasma membrane calcium-transporting ATPase 1 isoform 1a

See identical proteins and their annotated locations for NP_001001323.1

Status: REVIEWED

Description

Transcript Variant: This variant (1), as well as variant 6, encodes isoform 1a, also known as hPMCA1a.

Source sequence(s)

AC009522, AC025034, AC068641

Consensus CDS

CCDS41817.1

UniProtKB/Swiss-Prot

P20020

Related

ENSP00000352054.3, ENST00000359142.7

Conserved Domains (2) summary

TIGR01517
Location:15 → 1064

ATPase-IIB_Ca; plasma-membrane calcium-translocating P-type ATPase

pfam12424
Location:1103 → 1142

ATP_Ca_trans_C; Plasma membrane calcium transporter ATPase C terminal
NM_001366520.1 → NP_001353449.1 plasma membrane calcium-transporting ATPase 1 isoform 1b

Status: REVIEWED

Description

Transcript Variant: This variant (3), as well as variants 2, 4, and 5, encodes isoform 1b.

Source sequence(s)

AC009522, AC025034, AC068641

Conserved Domains (2) summary

TIGR01517
Location:15 → 1064

ATPase-IIB_Ca; plasma-membrane calcium-translocating P-type ATPase

pfam12424
Location:1103 → 1149

ATP_Ca_trans_C; Plasma membrane calcium transporter ATPase C terminal
NM_001366521.1 → NP_001353450.1 plasma membrane calcium-transporting ATPase 1 isoform 1b

Status: REVIEWED

Description

Transcript Variant: This variant (4), as well as variants 2, 3, and 5, encodes isoform 1.

Source sequence(s)

AC009522, AC025034, AC068641

Related

ENSP00000392043.3, ENST00000428670.8

Conserved Domains (2) summary

TIGR01517
Location:15 → 1064

ATPase-IIB_Ca; plasma-membrane calcium-translocating P-type ATPase

pfam12424
Location:1103 → 1149

ATP_Ca_trans_C; Plasma membrane calcium transporter ATPase C terminal
NM_001366522.1 → NP_001353451.1 plasma membrane calcium-transporting ATPase 1 isoform 1b

Status: REVIEWED

Description

Transcript Variant: This variant (5), as well as variants 2, 3, and 4, encodes isoform 1b.

Source sequence(s)

AC009522, AC025034, AC068641

Conserved Domains (2) summary

TIGR01517
Location:15 → 1064

ATPase-IIB_Ca; plasma-membrane calcium-translocating P-type ATPase

pfam12424
Location:1103 → 1149

ATP_Ca_trans_C; Plasma membrane calcium transporter ATPase C terminal
NM_001366523.1 → NP_001353452.1 plasma membrane calcium-transporting ATPase 1 isoform 1a

Status: REVIEWED

Description

Transcript Variant: This variant (6), as well as variant 1, encodes isoform 1a.

Source sequence(s)

AC009522, AC025034, AC068641

Conserved Domains (2) summary

TIGR01517
Location:15 → 1064

ATPase-IIB_Ca; plasma-membrane calcium-translocating P-type ATPase

pfam12424
Location:1103 → 1142

ATP_Ca_trans_C; Plasma membrane calcium transporter ATPase C terminal
NM_001366524.1 → NP_001353453.1 plasma membrane calcium-transporting ATPase 1 isoform 2

Status: REVIEWED

Description

Transcript Variant: This variant (7), as well as variant 8, encodes isoform 2.

Source sequence(s)

AC009522, AC025034, AC068641

Conserved Domains (2) summary

TIGR01517
Location:15 → 1064

ATPase-IIB_Ca; plasma-membrane calcium-translocating P-type ATPase

pfam12424
Location:1103 → 1178

ATP_Ca_trans_C; Plasma membrane calcium transporter ATPase C terminal
NM_001366525.1 → NP_001353454.1 plasma membrane calcium-transporting ATPase 1 isoform 2

Status: REVIEWED

Description

Transcript Variant: This variant (8), as well as variant 7, encodes isoform 2.

Source sequence(s)

AC009522, AC025034, AC068641

Conserved Domains (2) summary

TIGR01517
Location:15 → 1064

ATPase-IIB_Ca; plasma-membrane calcium-translocating P-type ATPase

pfam12424
Location:1103 → 1178

ATP_Ca_trans_C; Plasma membrane calcium transporter ATPase C terminal
NM_001366526.1 → NP_001353455.1 plasma membrane calcium-transporting ATPase 1 isoform 3

Status: REVIEWED

Description

Transcript Variant: This variant (9), as well as variant 10, encodes isoform 3.

Source sequence(s)

AC009522, AC025034, AC068641

Conserved Domains (2) summary

TIGR01517
Location:15 → 1064

ATPase-IIB_Ca; plasma-membrane calcium-translocating P-type ATPase

pfam12424
Location:1103 → 1142

ATP_Ca_trans_C; Plasma membrane calcium transporter ATPase C terminal
NM_001366527.1 → NP_001353456.1 plasma membrane calcium-transporting ATPase 1 isoform 3

Status: REVIEWED

Description

Transcript Variant: This variant (10), as well as variant 9, encodes isoform 3.

Source sequence(s)

AC009522, AC025034, AC068641

Conserved Domains (2) summary

TIGR01517
Location:15 → 1064

ATPase-IIB_Ca; plasma-membrane calcium-translocating P-type ATPase

pfam12424
Location:1103 → 1142

ATP_Ca_trans_C; Plasma membrane calcium transporter ATPase C terminal
NM_001366528.1 → NP_001353457.1 plasma membrane calcium-transporting ATPase 1 isoform 4

Status: REVIEWED

Source sequence(s)

AC009522, AC025034, AC068641

Conserved Domains (2) summary

pfam12424
Location:1067 → 1106

ATP_Ca_trans_C; Plasma membrane calcium transporter ATPase C terminal

cl27747
Location:15 → 1023

E1-E2_ATPase; E1-E2 ATPase
NM_001366529.1 → NP_001353458.1 plasma membrane calcium-transporting ATPase 1 isoform 5

Status: REVIEWED

Source sequence(s)

AC009522, AC025034, AC068641

Conserved Domains (2) summary

TIGR01517
Location:15 → 1023

ATPase-IIB_Ca; plasma-membrane calcium-translocating P-type ATPase

pfam12424
Location:1067 → 1106

ATP_Ca_trans_C; Plasma membrane calcium transporter ATPase C terminal
NM_001366530.1 → NP_001353459.1 plasma membrane calcium-transporting ATPase 1 isoform 6

Status: REVIEWED

Source sequence(s)

AC009522, AC025034, AC068641

Conserved Domains (2) summary

TIGR01517
Location:15 → 998

ATPase-IIB_Ca; plasma-membrane calcium-translocating P-type ATPase

pfam12424
Location:1037 → 1076

ATP_Ca_trans_C; Plasma membrane calcium transporter ATPase C terminal
NM_001366531.1 → NP_001353460.1 plasma membrane calcium-transporting ATPase 1 isoform 7

Status: REVIEWED

Description

Transcript Variant: This variant (14), as well as variant 15, encodes isoform 7.

Source sequence(s)

AC009522, AC025034, AC068641

Conserved Domains (2) summary

pfam12424
Location:916 → 955

ATP_Ca_trans_C; Plasma membrane calcium transporter ATPase C terminal

cl27747
Location:15 → 877

E1-E2_ATPase; E1-E2 ATPase
NM_001366532.1 → NP_001353461.1 plasma membrane calcium-transporting ATPase 1 isoform 7

Status: REVIEWED

Description

Transcript Variant: This variant (15), as well as variant 14, encodes isoform 7.

Source sequence(s)

AC009522, AC025034, AC068641

Conserved Domains (2) summary

pfam12424
Location:916 → 955

ATP_Ca_trans_C; Plasma membrane calcium transporter ATPase C terminal

cl27747
Location:15 → 877

E1-E2_ATPase; E1-E2 ATPase
NM_001682.3 → NP_001673.2 plasma membrane calcium-transporting ATPase 1 isoform 1b

See identical proteins and their annotated locations for NP_001673.2

Status: REVIEWED

Description

Transcript Variant: This variant (2) lacks a segment in the coding region, which leads to a frameshift, compared to variant 1. The resulting isoform (1b) contains a longer and distinct C-terminus compared to isoform 1a. Variants 2, 3, 4, and 5 all encode the same isoform (1b).

Source sequence(s)

AC009522, AC025034, AC068641

Consensus CDS

CCDS9035.1

UniProtKB/Swiss-Prot

P20020

UniProtKB/TrEMBL

A0A024RBC7

Related

ENSP00000261173.2, ENST00000261173.6

Conserved Domains (2) summary

TIGR01517
Location:15 → 1064

ATPase-IIB_Ca; plasma-membrane calcium-translocating P-type ATPase

pfam12424
Location:1103 → 1149

ATP_Ca_trans_C; Plasma membrane calcium transporter ATPase C terminal

RefSeqs of Annotated Genomes: Homo sapiens Updated Annotation Release 109.20210226

The following sections contain reference sequences that belong to a specific genome build. Explain

Reference GRCh38.p13 Primary Assembly

Genomic

NC_000012.12 Reference GRCh38.p13 Primary Assembly

Range

89588049..89709366 complement

Download

GenBank, FASTA, Sequence Viewer (Graphics)

mRNA and Protein(s)

XM_024448991.1 → XP_024304759.1 plasma membrane calcium-transporting ATPase 1 isoform X4

Conserved Domains (2) summary

TIGR01517
Location:15 → 1023

ATPase-IIB_Ca; plasma-membrane calcium-translocating P-type ATPase

pfam12424
Location:1067 → 1113

ATP_Ca_trans_C; Plasma membrane calcium transporter ATPase C terminal
XM_017019357.2 → XP_016874846.1 plasma membrane calcium-transporting ATPase 1 isoform X3

Conserved Domains (2) summary

TIGR01517
Location:15 → 1023

ATPase-IIB_Ca; plasma-membrane calcium-translocating P-type ATPase

pfam12424
Location:1067 → 1142

ATP_Ca_trans_C; Plasma membrane calcium transporter ATPase C terminal
XM_024448993.1 → XP_024304761.1 plasma membrane calcium-transporting ATPase 1 isoform X6

Conserved Domains (2) summary

pfam12424
Location:1067 → 1106

ATP_Ca_trans_C; Plasma membrane calcium transporter ATPase C terminal

cl27747
Location:15 → 1023

E1-E2_ATPase; E1-E2 ATPase