Motif ID: ACAGUAU

Z-value: 0.883

Mature miRNA associated with seed ACAGUAU:

NamemiRBase Accession
mmu-miR-144 MIMAT0000156



Activity profile for motif ACAGUAU.

activity profile for motif ACAGUAU


Sorted Z-values histogram for motif ACAGUAU

Sorted Z-values for motif ACAGUAU



Network of associatons between targets according to the STRING database.



First level regulatory network of ACAGUAU

PNG image of the network

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Top targets:


Promoter Score Refseq Gene Description
chr8_+_24521967 6.073 NM_013834
Sfrp1
secreted frizzled-related protein 1
chr2_-_7002344 4.756 NM_001110229
NM_001110230
NM_001160292
Celf2


CUGBP, Elav-like family member 2


chr1_+_179374783 4.732 NM_001012330
Zfp238
zinc finger protein 238
chr2_-_84265422 4.635 NM_018782
Calcrl
calcitonin receptor-like
chr3_-_50247306 4.284 NM_011990
Slc7a11
solute carrier family 7 (cationic amino acid transporter, y+ system), member 11
chr6_+_117118552 2.932 NM_001012477
NM_013655
NM_021704
Cxcl12


chemokine (C-X-C motif) ligand 12


chr18_+_39152776 2.886 NM_175164
Arhgap26
Rho GTPase activating protein 26
chr7_+_121558695 2.853 NM_011055
Pde3b
phosphodiesterase 3B, cGMP-inhibited
chr14_+_34736772 2.602 NM_145741
Gdf10
growth differentiation factor 10
chr18_-_47528483 2.465 NM_018744
Sema6a
sema domain, transmembrane domain (TM), and cytoplasmic domain, (semaphorin) 6A
chr2_-_156904966 2.460 NM_001164663
9830001H06Rik
RIKEN cDNA 9830001H06 gene
chr15_+_99223622 2.437 NM_001171035
Tmbim6
transmembrane BAX inhibitor motif containing 6
chr2_-_79296732 2.364 NM_010894
Neurod1
neurogenic differentiation 1
chr11_-_69419419 2.298 NM_013415
Atp1b2
ATPase, Na+/K+ transporting, beta 2 polypeptide
chr4_+_97248556 2.128 NM_001122952
Nfia
D130039L10Rik
nuclear factor I/A
RIKEN cDNA D130039L10 gene
chr5_-_135223110 2.098 NM_007925
Eln
elastin
chr2_+_138104219 2.086 NM_145534
Btbd3
BTB (POZ) domain containing 3
chr4_+_97444312 2.041 NM_001122953
NM_010905
Nfia

nuclear factor I/A

chr17_-_84587278 2.008 NM_001001806
Zfp36l2
zinc finger protein 36, C3H type-like 2
chr10_+_56097078 1.942 NM_010288
Gja1
gap junction protein, alpha 1
chr11_+_114536748 1.838 NM_053273
Ttyh2
tweety homolog 2 (Drosophila)
chr4_-_53172748 1.826 NM_013454
Abca1
ATP-binding cassette, sub-family A (ABC1), member 1
chr15_+_99223377 1.733 NM_001171034
NM_001171036
NM_026669
Tmbim6


transmembrane BAX inhibitor motif containing 6


chr12_-_111934580 1.716 NM_010480
Hsp90aa1
heat shock protein 90, alpha (cytosolic), class A member 1
chr2_+_158492854 1.686 NM_001159662
Ppp1r16b
protein phosphatase 1, regulatory (inhibitor) subunit 16B
chr1_-_166388431 1.663 NM_009721
Atp1b1
ATPase, Na+/K+ transporting, beta 1 polypeptide
chr10_-_20444654 1.641 NM_013875
Pde7b
phosphodiesterase 7B
chr3_+_84756090 1.582 NM_080428
Fbxw7
F-box and WD-40 domain protein 7
chr6_-_113991945 1.516 NM_009723
Atp2b2
ATPase, Ca++ transporting, plasma membrane 2
chr4_-_82151180 1.476 NM_001113209
NM_001113210
NM_008687
Nfib


nuclear factor I/B


chr8_+_98226914 1.434 NM_001195006
NM_145602
Ndrg4

N-myc downstream regulated gene 4

chr12_+_86814822 1.399 NM_010234
Fos
FBJ osteosarcoma oncogene
chr1_+_6720087 1.324 NM_173868
St18
suppression of tumorigenicity 18
chr3_+_60305108 1.320 NM_020007
Mbnl1
muscleblind-like 1 (Drosophila)
chr5_-_124482223 1.294 NM_177876
Vps37b
vacuolar protein sorting 37B (yeast)
chr6_-_113841356 1.274 NM_001036684
Atp2b2
ATPase, Ca++ transporting, plasma membrane 2
chr4_+_11118855 1.258 NM_009830
Ccne2
cyclin E2
chr18_+_58038278 1.243 NM_009194
Slc12a2
solute carrier family 12, member 2
chr10_-_10802102 1.220 NM_001114333
NM_016976
Grm1

glutamate receptor, metabotropic 1

chr2_-_90420626 1.219 NM_008982
Ptprj
protein tyrosine phosphatase, receptor type, J
chr10_+_4424140 1.129 NM_001161822
NM_019958
Rgs17

regulator of G-protein signaling 17

chr2_+_103406466 1.113 NM_178890
Abtb2
ankyrin repeat and BTB (POZ) domain containing 2
chr14_+_20570444 1.109 NM_008695
Nid2
nidogen 2
chr10_+_99478457 1.086 NM_013598
Kitl
kit ligand
chr18_+_69504145 1.085 NM_013685
Tcf4
transcription factor 4
chr10_-_30522912 1.079 NM_172495
Ncoa7
nuclear receptor coactivator 7
chr15_+_100701068 1.055 NM_001077499
Scn8a
sodium channel, voltage-gated, type VIII, alpha
chr18_+_36440748 1.051 NM_008989
Pura
purine rich element binding protein A
chr6_+_113257130 1.044 NM_030178
Brpf1
bromodomain and PHD finger containing, 1
chr16_-_11254052 1.034 NM_001130008
NM_146066
Gspt1

G1 to S phase transition 1

chr15_+_66409094 1.013 NM_001081409
Phf20l1
PHD finger protein 20-like 1
chr19_+_28909559 0.999 NM_009199
Slc1a1
solute carrier family 1 (neuronal/epithelial high affinity glutamate transporter, system Xag), member 1
chrX_-_154036606 0.923 NM_172307
Mbtps2
membrane-bound transcription factor peptidase, site 2
chr16_+_33380860 0.910 NM_011749
Zfp148
zinc finger protein 148
chr5_+_144383313 0.908 NM_011182
Cyth3
cytohesin 3
chr15_+_80541675 0.907 NM_144812
Tnrc6b
trinucleotide repeat containing 6b
chr6_-_90760116 0.905 NM_001134384
Iqsec1
IQ motif and Sec7 domain 1
chr17_+_86567003 0.883 NM_011104
Prkce
protein kinase C, epsilon
chr2_+_112295120 0.864 NM_133749
2900064A13Rik
RIKEN cDNA 2900064A13 gene
chr8_+_106625524 0.843 NM_009868
Cdh5
cadherin 5
chr8_+_113245378 0.825 NM_198625
Mtss1l
metastasis suppressor 1-like
chr12_-_76836549 0.817 NM_030750
Sgpp1
sphingosine-1-phosphate phosphatase 1
chr7_+_56501558 0.811 NM_175272
Nav2
neuron navigator 2
chr12_-_12948513 0.811 NM_008709
Mycn
v-myc myelocytomatosis viral related oncogene, neuroblastoma derived (avian)
chr8_+_23587151 0.807 NM_011394
Slc20a2
solute carrier family 20, member 2
chr18_-_23197156 0.797 NM_001161483
Nol4
nucleolar protein 4
chr16_-_10314010 0.796 NM_007929
Emp2
epithelial membrane protein 2
chr2_+_31428244 0.788 NM_001033389
Fubp3
far upstream element (FUSE) binding protein 3
chr19_+_32832002 0.766 NM_008960
Pten
phosphatase and tensin homolog
chr3_-_107320881 0.750 NM_172271
Slc6a17
solute carrier family 6 (neurotransmitter transporter), member 17
chr14_-_103381744 0.745 NM_177715
Kctd12
potassium channel tetramerisation domain containing 12
chr2_-_167318313 0.729 NM_170756
Spata2
spermatogenesis associated 2
chr14_-_17407701 0.727 NM_011243
Rarb
retinoic acid receptor, beta
chr19_+_27291507 0.725 NM_001161420
NM_013703
Vldlr

very low density lipoprotein receptor

chr9_-_31019392 0.715 NM_001102455
NM_001102456
NM_009691
Aplp2


amyloid beta (A4) precursor-like protein 2


chr15_+_99869071 0.708 NM_001159361
Dip2b
DIP2 disco-interacting protein 2 homolog B (Drosophila)
chr11_-_18918923 0.706 NM_001193271
NM_010789
Meis1

Meis homeobox 1

chr6_+_128312987 0.694 NM_008021
Foxm1
forkhead box M1
chr2_+_166631011 0.692 NM_001085495
Arfgef2
ADP-ribosylation factor guanine nucleotide-exchange factor 2 (brefeldin A-inhibited)
chr2_-_12223518 0.662 NM_001001309
Itga8
integrin alpha 8
chr2_+_181501718 0.660 NM_001171616
NM_008665
Myt1

myelin transcription factor 1

chr10_-_36858671 0.653 NM_008538
Marcks
myristoylated alanine rich protein kinase C substrate
chr2_+_61884596 0.652 NM_033552
Slc4a10
solute carrier family 4, sodium bicarbonate cotransporter-like, member 10
chr5_+_143105467 0.637 NM_178398
Wipi2
WD repeat domain, phosphoinositide interacting 2
chr14_+_120674886 0.627 NM_175341
NM_207515
Mbnl2

muscleblind-like 2

chr3_+_146067605 0.621 NM_138744
Ssx2ip
synovial sarcoma, X breakpoint 2 interacting protein
chr7_+_73984620 0.618 NM_001033877
Adamts17
a disintegrin-like and metallopeptidase (reprolysin type) with thrombospondin type 1 motif, 17
chr16_-_37385024 0.616 NM_001114611
NM_001114612
NM_001114613
NM_172440
Stxbp5l



syntaxin binding protein 5-like



chr17_-_10512217 0.608 NM_001159516
NM_001159517
NM_021881
Qk


quaking


chr2_-_6805937 0.580 NM_001110228
NM_001160293
NM_010160
Celf2


CUGBP, Elav-like family member 2


chr17_-_79754427 0.579 NM_026514
Cdc42ep3
CDC42 effector protein (Rho GTPase binding) 3
chr4_-_128483355 0.575 NM_001081098
Zfp362
zinc finger protein 362
chr17_+_31701700 0.569 NM_016670
Pknox1
Pbx/knotted 1 homeobox
chr12_+_3806955 0.567 NM_007872
Dnmt3a
DNA methyltransferase 3A
chr9_-_75257067 0.567 NM_027418
Mapk6
mitogen-activated protein kinase 6
chr8_+_90723069 0.561 NM_001164497
NM_001164498
NM_001164499
Papd5


PAP associated domain containing 5


chr18_+_35989376 0.560 NM_133687
Cxxc5
CXXC finger 5
chr18_-_23200150 0.554 NM_199024
Nol4
nucleolar protein 4
chr6_+_113027481 0.547 NM_028385
Setd5
SET domain containing 5
chr9_+_32503600 0.546 NM_001038642
NM_011808
Ets1

E26 avian leukemia oncogene 1, 5' domain

chr19_+_21727795 0.543 NM_146096
Fam108b
family with sequence similarity 108, member B
chr13_-_99660903 0.540 NM_001048267
Tnpo1
transportin 1
chr3_-_9610062 0.526 NM_133218
Zfp704
zinc finger protein 704
chr6_-_35489875 0.524 NM_008098
Mtpn
myotrophin
chr15_+_38837839 0.516 NM_008056
Fzd6
frizzled homolog 6 (Drosophila)
chr18_+_5591869 0.511 NM_011546
Zeb1
zinc finger E-box binding homeobox 1
chr10_-_30375572 0.503 NM_001111267
Ncoa7
nuclear receptor coactivator 7
chr6_+_54222814 0.499 NM_023543
Chn2
chimerin (chimaerin) 2
chr7_+_35904310 0.496 NM_007678
Cebpa
CCAAT/enhancer binding protein (C/EBP), alpha
chr9_-_14419410 0.493 NM_001081395
Amotl1
angiomotin-like 1
chr7_+_94732677 0.488 NM_001081414
NM_001143834
Grm5

glutamate receptor, metabotropic 5

chr15_-_93105514 0.488 NM_001033275
Gxylt1
glucoside xylosyltransferase 1
chrX_+_96237919 0.484 NM_199018
Stard8
START domain containing 8
chrX_+_65931715 0.482 NM_008031
Fmr1
fragile X mental retardation syndrome 1 homolog
chr4_-_91038729 0.478 NM_010486
NM_207686
Elavl2

ELAV (embryonic lethal, abnormal vision, Drosophila)-like 2 (Hu antigen B)

chr13_+_75844906 0.478 NM_138953
Ell2
elongation factor RNA polymerase II 2
chr6_+_38383918 0.466 NM_177185
Ubn2
ubinuclein 2
chr7_+_89484203 0.464 NM_001190374
Adamtsl3
ADAMTS-like 3
chr18_-_35374663 0.464 NM_178005
Lrrtm2
leucine rich repeat transmembrane neuronal 2
chr2_+_156439426 0.460 NM_146128
Dlgap4
discs, large homolog-associated protein 4 (Drosophila)
chr18_+_13130739 0.458 NM_008378
Impact
imprinted and ancient
chr15_+_96117952 0.448 NM_175251
Arid2
AT rich interactive domain 2 (ARID, RFX-like)
chr16_+_32608981 0.446 NM_011638
Tfrc
transferrin receptor
chr8_-_72963464 0.437 NM_001004062
Crtc1
CREB regulated transcription coactivator 1
chr6_+_88148657 0.436 NM_008090
Gata2
GATA binding protein 2
chr9_-_50817077 0.432 NM_178710
Sik2
salt inducible kinase 2
chr1_+_106665395 0.424 NM_011800
Cdh20
cadherin 20
chr7_+_107855215 0.423 NM_178764
Fam168a
family with sequence similarity 168, member A
chr15_+_58720783 0.419 NM_175226
Tatdn1
Rnf139
TatD DNase domain containing 1
ring finger protein 139
chr9_+_114640266 0.417 NM_026036
Cmtm6
CKLF-like MARVEL transmembrane domain containing 6
chr18_+_23573908 0.416 NM_010087
NM_207650
Dtna

dystrobrevin alpha

chr14_+_73952438 0.412 NM_011506
Sucla2
succinate-Coenzyme A ligase, ADP-forming, beta subunit
chr11_+_32542845 0.410 NM_134015
Fbxw11
F-box and WD-40 domain protein 11
chr4_-_126106749 0.409 NM_153402
Eif2c3
eukaryotic translation initiation factor 2C, 3
chr9_+_13553574 0.400 NM_023858
Mtmr2
myotubularin related protein 2
chr9_-_88376937 0.400 NM_019666
NM_019796
Syncrip

synaptotagmin binding, cytoplasmic RNA interacting protein

chr2_+_69735301 0.397 NM_001081548
NM_177783
Ubr3

ubiquitin protein ligase E3 component n-recognin 3

chr5_+_139728268 0.395 NM_026269
Get4
golgi to ER traffic protein 4 homolog (S. cerevisiae)
chr11_-_118947424 0.393 NM_007625
Cbx4
chromobox homolog 4 (Drosophila Pc class)
chr16_-_85173944 0.392 NM_001198823
NM_001198824
NM_001198825
NM_001198826
NM_007471
App




amyloid beta (A4) precursor protein




chr6_+_119429620 0.387 NM_133940
Fbxl14
F-box and leucine-rich repeat protein 14
chr16_-_4559511 0.386 NM_031182
Tcfap4
transcription factor AP4
chr7_-_35174519 0.384 NM_025948
Lsm14a
LSM14 homolog A (SCD6, S. cerevisiae)
chr3_-_78949796 0.381 NM_001099624
Rapgef2
Rap guanine nucleotide exchange factor (GEF) 2
chr4_+_130219255 0.379 NM_001159603
NM_001159604
NM_001159605
NM_001159606
NM_030722
Pum1




pumilio 1 (Drosophila)




chr11_+_59761707 0.367 NM_138675
Med9
mediator of RNA polymerase II transcription, subunit 9 homolog (yeast)
chr14_+_19103655 0.367 NM_023526
Nkiras1
NFKB inhibitor interacting Ras-like protein 1
chr9_+_59598963 0.364 NM_173018
Myo9a
myosin IXa
chr7_+_149135647 0.361 NM_029426
Brsk2
BR serine/threonine kinase 2
chr9_-_107134182 0.355 NM_153413
Dock3
dedicator of cyto-kinesis 3
chr4_+_147822690 0.353 NM_020009
Mtor
mechanistic target of rapamycin (serine/threonine kinase)
chr15_-_58046428 0.351 NM_026346
Fbxo32
F-box protein 32
chr2_+_59450080 0.343 NM_198294
Tanc1
tetratricopeptide repeat, ankyrin repeat and coiled-coil containing 1
chr17_-_12511525 0.343 NM_011948
Map3k4
mitogen-activated protein kinase kinase kinase 4
chr10_+_82448214 0.340 NM_021439
Chst11
carbohydrate sulfotransferase 11
chr18_-_4352916 0.338 NM_007746
Map3k8
mitogen-activated protein kinase kinase kinase 8
chr6_+_135312948 0.336 NM_010128
Emp1
epithelial membrane protein 1
chr11_+_20101587 0.331 NM_008996
Rab1
RAB1, member RAS oncogene family
chr7_-_123182257 0.331 NM_001025559
Sox6
SRY-box containing gene 6
chr6_-_148779856 0.323 NM_001081113
Ipo8
importin 8
chr3_+_138405156 0.317 NM_019571
Tspan5
tetraspanin 5
chr1_-_178143971 0.315 NM_001195816
Pld5
phospholipase D family, member 5
chr10_+_85763156 0.311 NM_011595
Timp3
tissue inhibitor of metalloproteinase 3
chr5_+_18732863 0.309 NM_001170746
Magi2
membrane associated guanylate kinase, WW and PDZ domain containing 2
chr2_-_66278870 0.302 NM_018733
Scn1a
sodium channel, voltage-gated, type I, alpha
chr2_+_127951773 0.299 NM_009754
NM_207680
NM_207681
Bcl2l11


BCL2-like 11 (apoptosis facilitator)


chr7_-_74517624 0.299 NM_001033713
Mef2a
myocyte enhancer factor 2A
chr19_+_3851772 0.298 NM_001025566
NM_013490
Chka

choline kinase alpha

chr5_-_74464390 0.274 NM_177561
Usp46
ubiquitin specific peptidase 46
chr5_+_15440508 0.262 NM_001110843
NM_001110844
NM_001110845
NM_001110846
NM_009784
Cacna2d1




calcium channel, voltage-dependent, alpha2/delta subunit 1




chr15_+_100766701 0.261 NM_011323
Scn8a
sodium channel, voltage-gated, type VIII, alpha
chr5_+_43624701 0.259 NM_001177379
NM_175937
Cpeb2

cytoplasmic polyadenylation element binding protein 2

chr7_+_56214442 0.258 NM_001111016
Nav2
neuron navigator 2
chr16_+_94791713 0.255 NM_007890
Dyrk1a
dual-specificity tyrosine-(Y)-phosphorylation regulated kinase 1a
chr1_-_58642994 0.255 NM_172513
Fam126b
family with sequence similarity 126, member B
chr18_+_69505374 0.254 NM_001083967
Tcf4
transcription factor 4
chr16_-_17891608 0.254 NM_001109750
NM_010048
Dgcr2

DiGeorge syndrome critical region gene 2

chr1_+_176431933 0.254 NM_019445
Fmn2
formin 2
chrX_-_138825032 0.253 NM_001033600
NM_019477
NM_207625
Acsl4


acyl-CoA synthetase long-chain family member 4


chr12_+_8680904 0.252 NM_001160221
NM_001160222
Pum2

pumilio 2 (Drosophila)

chr2_-_75542552 0.249 NM_010902
Nfe2l2
nuclear factor, erythroid derived 2, like 2
chr5_+_147043131 0.245 NM_153599
Cdk8
cyclin-dependent kinase 8
chr7_-_81699435 0.244 NM_001038643
NM_023908
Slco3a1

solute carrier organic anion transporter family, member 3a1

chr12_-_52792851 0.240 NM_001172098
NM_052973
Strn3

striatin, calmodulin binding protein 3

chr5_-_151992724 0.226 NM_001163493
NM_146258
Stard13

StAR-related lipid transfer (START) domain containing 13

chr12_-_113146259 0.226 NM_011625
Ppp1r13b
protein phosphatase 1, regulatory (inhibitor) subunit 13B
chr5_-_108561512 0.222 NM_028876
Tmed5
transmembrane emp24 protein transport domain containing 5
chr9_+_21001148 0.219 NM_019798
Pde4a
phosphodiesterase 4A, cAMP specific
chr11_-_95171521 0.217 NM_001195003
NM_001195004
NM_177619
Myst2


MYST histone acetyltransferase 2


chr17_+_28046196 0.217 NM_181413
Anks1
ankyrin repeat and SAM domain containing 1
chr5_-_98459943 0.215 NM_133738
Antxr2
anthrax toxin receptor 2
chr6_-_134742645 0.215 NM_001048054
NM_130447
Dusp16

dual specificity phosphatase 16

chr4_+_132567420 0.211 NM_146155
Ahdc1
AT hook, DNA binding motif, containing 1
chr2_+_177927788 0.211 NM_001007154
Phactr3
phosphatase and actin regulator 3
chr2_+_156547014 0.206 NM_001042487
Dlgap4
discs, large homolog-associated protein 4 (Drosophila)
chr8_-_112261607 0.205 NM_001080930
Atxn1l
ataxin 1-like
chr1_-_17087821 0.204 NM_020604
Jph1
junctophilin 1
chr14_+_28051224 0.203 NM_027871
Arhgef3
Rho guanine nucleotide exchange factor (GEF) 3
chr8_-_125075222 0.203 NM_001037298
Fam38a
family with sequence similarity 38, member A
chr7_+_117265568 0.203 NM_009516
Wee1
WEE 1 homolog 1 (S. pombe)

Gene Ontology Analysis

Gene overrepresentation in process category:

enrichment p-value GO term description
3.51 1.64e-04 GO:0048731 system development
3.01 8.73e-04 GO:0007275 multicellular organismal development
3.15 8.98e-04 GO:0048856 anatomical structure development
3.78 1.82e-03 GO:0048513 organ development
2.76 3.31e-03 GO:0032502 developmental process
2.32 2.77e-02 GO:0019222 regulation of metabolic process
2.50 4.77e-02 GO:0006139 nucleobase, nucleoside, nucleotide and nucleic acid metabolic process
4.15 4.90e-02 GO:0010646 regulation of cell communication

Gene overrepresentation in compartment category:

enrichment p-value GO term description
4.41 6.78e-04 GO:0000267 cell fraction
4.69 2.56e-03 GO:0005626 insoluble fraction
2.42 3.64e-03 GO:0005886 plasma membrane
2.37 4.91e-03 GO:0071944 cell periphery
11.09 5.72e-03 GO:0016324 apical plasma membrane
4.42 1.05e-02 GO:0005624 membrane fraction
8.35 2.16e-02 GO:0045177 apical part of cell

Gene overrepresentation in function category:

enrichment p-value GO term description
18.53 5.83e-03 GO:0042626 ATPase activity, coupled to transmembrane movement of substances
17.93 6.62e-03 GO:0043492 ATPase activity, coupled to movement of substances
8.18 7.68e-03 GO:0022804 active transmembrane transporter activity
17.02 8.12e-03 GO:0015399 primary active transmembrane transporter activity
17.02 8.12e-03 GO:0015405 P-P-bond-hydrolysis-driven transmembrane transporter activity
4.55 1.13e-02 GO:0022857 transmembrane transporter activity
83.30 2.36e-02 GO:0005391 sodium:potassium-exchanging ATPase activity
23.59 2.59e-02 GO:0015662 ATPase activity, coupled to transmembrane movement of ions, phosphorylative mechanism
4.47 3.18e-02 GO:0022891 substrate-specific transmembrane transporter activity
11.83 3.30e-02 GO:0003690 double-stranded DNA binding
69.42 3.46e-02 GO:0004115 3',5'-cyclic-AMP phosphodiesterase activity
69.42 3.46e-02 GO:0008556 potassium-transporting ATPase activity
3.83 4.10e-02 GO:0005102 receptor binding
20.17 4.11e-02 GO:0042625 ATPase activity, coupled to transmembrane movement of ions