Motif ID: ACUGCAU

Z-value: 0.465


Mature miRNA associated with seed ACUGCAU:

NamemiRBase Accession
hsa-miR-217 MIMAT0000274



Activity profile for motif ACUGCAU.

activity profile for motif ACUGCAU


Sorted Z-values histogram for motif ACUGCAU

Sorted Z-values for motif ACUGCAU



Network of associatons between targets according to the STRING database.



First level regulatory network of ACUGCAU

PNG image of the network

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Top targets:


Promoter Score Refseq Gene Symbol Gene Name
chr2_-_100721792 1.148 NM_001025108
AFF3
AF4/FMR2 family, member 3
chr10_-_105614952 1.142 NM_014631
SH3PXD2A
SH3 and PX domains 2A
chr2_-_100758970 0.981 NM_002285
AFF3
AF4/FMR2 family, member 3
chr7_-_83824216 0.518 NM_006080
SEMA3A
sema domain, immunoglobulin domain (Ig), short basic domain, secreted, (semaphorin) 3A
chr21_-_36421586 0.451 NM_001754
RUNX1
runt-related transcription factor 1
chr3_-_160283361 0.390 NM_002268
KPNA4
karyopherin alpha 4 (importin alpha 3)
chr3_+_173116243 0.382 NM_014932
NLGN1
neuroligin 1
chr18_-_43652153 0.376 NM_024430
PSTPIP2
proline-serine-threonine phosphatase interacting protein 2
chr8_+_17354562 0.367 NM_001008539
SLC7A2
solute carrier family 7 (cationic amino acid transporter, y+ system), member 2
chr10_-_62149487 0.360 NM_020987
ANK3
ankyrin 3, node of Ranvier (ankyrin G)
chr8_+_17396285 0.352 NM_001164771
NM_003046
SLC7A2

solute carrier family 7 (cationic amino acid transporter, y+ system), member 2

chr1_-_8003224 0.348 NM_001561
TNFRSF9
tumor necrosis factor receptor superfamily, member 9
chr4_+_77870864 0.336 NM_018243
SEPT11
septin 11
chr4_-_120549115 0.320 NM_033437
PDE5A
phosphodiesterase 5A, cGMP-specific
chr5_-_137911317 0.282 NM_004134
HSPA9
heat shock 70kDa protein 9 (mortalin)
chr10_-_61900659 0.279 NM_001149
ANK3
ankyrin 3, node of Ranvier (ankyrin G)
chr10_-_62493282 0.279 NM_001204403
ANK3
ankyrin 3, node of Ranvier (ankyrin G)
chr20_+_62526453 0.265 NM_025219
DNAJC5
DnaJ (Hsp40) homolog, subfamily C, member 5
chr1_+_65613771 0.260 NM_013410
AK4
adenylate kinase 4
chr1_+_169075869 0.259 NM_001677
ATP1B1
ATPase, Na+/K+ transporting, beta 1 polypeptide
chr1_+_65613211 0.255 NM_203464
NM_001005353
AK4

adenylate kinase 4

chr10_-_62332666 0.245 NM_001204404
ANK3
ankyrin 3, node of Ranvier (ankyrin G)
chr10_+_98592658 0.240 NM_032440
LCOR
ligand dependent nuclear receptor corepressor
chr4_-_120549827 0.229 NM_001083
PDE5A
phosphodiesterase 5A, cGMP-specific
chr9_+_112810877 0.202 NM_001004065
NM_001198656
AKAP2

A kinase (PRKA) anchor protein 2

chr16_+_77822400 0.198 NM_020927
VAT1L
vesicle amine transport protein 1 homolog (T. californica)-like
chr5_-_146833150 0.196 NM_001387
DPYSL3
dihydropyrimidinase-like 3
chr10_+_98592015 0.194 NM_001170765
NM_001170766
LCOR

ligand dependent nuclear receptor corepressor

chr5_-_133304274 0.191 NM_020199
C5orf15
chromosome 5 open reading frame 15
chr1_+_151584514 0.189 NM_030918
SNX27
sorting nexin family member 27
chr6_+_126111782 0.186 NM_001122842
NM_001199621
NM_181782
NCOA7


nuclear receptor coactivator 7


chr4_-_120548329 0.186 NM_033430
PDE5A
phosphodiesterase 5A, cGMP-specific
chr8_-_119964059 0.181 NM_002546
TNFRSF11B
tumor necrosis factor receptor superfamily, member 11b
chr14_+_33408458 0.180 NM_001164749
NM_001165893
NM_022123
NM_173159
NPAS3



neuronal PAS domain protein 3



chr1_+_145209067 0.177 NM_203458
NOTCH2NL
notch 2 N-terminal like
chr9_+_112887780 0.172 NM_001136562
AKAP2
A kinase (PRKA) anchor protein 2
chr2_+_208576239 0.171 NM_001142300
NM_152523
CCNYL1

cyclin Y-like 1

chr8_+_104512975 0.166 NM_001100117
RIMS2
regulating synaptic membrane exocytosis 2
chr4_-_139163222 0.166 NM_014331
SLC7A11
solute carrier family 7 (anionic amino acid transporter light chain, xc- system), member 11
chr8_-_56685869 0.165 NM_152417
TMEM68
transmembrane protein 68
chr18_+_31158540 0.165 NM_030632
ASXL3
additional sex combs like 3 (Drosophila)
chr21_-_36260972 0.165 NM_001001890
NM_001122607
RUNX1

runt-related transcription factor 1

chr3_-_120461362 0.165 NM_173825
RABL3
RAB, member of RAS oncogene family-like 3
chr16_-_53737713 0.164 NM_001127897
NM_015272
RPGRIP1L

RPGRIP1-like

chr12_+_62654120 0.163 NM_001252078
NM_001252079
NM_006313
USP15


ubiquitin specific peptidase 15


chr15_+_71433787 0.161 NM_024817
THSD4
thrombospondin, type I, domain containing 4
chr7_+_36429388 0.156 NM_018685
ANLN
anillin, actin binding protein
chr10_-_27443309 0.155 NM_001253866
NM_014263
NM_139312
YME1L1


YME1-like 1 (S. cerevisiae)


chr6_+_4776641 0.153 NM_004824
CDYL
chromodomain protein, Y-like
chr6_+_4890225 0.151 NM_001143971
CDYL
chromodomain protein, Y-like
chrX_-_108976509 0.148 NM_004458
NM_022977
ACSL4

acyl-CoA synthetase long-chain family member 4

chr5_-_146889618 0.147 NM_001197294
DPYSL3
dihydropyrimidinase-like 3
chr2_+_99953749 0.145 NM_015904
EIF5B
eukaryotic translation initiation factor 5B
chr5_-_168006413 0.144 NM_024594
PANK3
pantothenate kinase 3
chr6_+_4836331 0.144 NM_001143970
CDYL
chromodomain protein, Y-like
chr8_+_104831415 0.135 NM_014677
RIMS2
regulating synaptic membrane exocytosis 2
chr11_+_109292837 0.134 NM_207645
C11orf87
chromosome 11 open reading frame 87
chr17_+_41052813 0.133 NM_000151
G6PC
glucose-6-phosphatase, catalytic subunit
chr11_+_94277005 0.133 NM_002033
PIWIL4
FUT4
piwi-like 4 (Drosophila)
fucosyltransferase 4 (alpha (1,3) fucosyltransferase, myeloid-specific)
chr7_+_28725597 0.133 NM_001011666
CREB5
cAMP responsive element binding protein 5
chr8_+_1449531 0.132 NM_004745
DLGAP2
discs, large (Drosophila) homolog-associated protein 2
chr11_-_16424391 0.132 NM_001145819
NM_017508
SOX6

SRY (sex determining region Y)-box 6

chr7_+_28452143 0.132 NM_182898
CREB5
cAMP responsive element binding protein 5
chr6_+_7541863 0.130 NM_001008844
NM_004415
DSP

desmoplakin

chr2_+_65216482 0.129 NM_003038
SLC1A4
solute carrier family 1 (glutamate/neutral amino acid transporter), member 4
chr12_-_50101192 0.129 NM_175736
NM_198900
FMNL3

formin-like 3

chr2_+_65215577 0.127 NM_001193493
SLC1A4
solute carrier family 1 (glutamate/neutral amino acid transporter), member 4
chr2_+_228336877 0.126 NM_001135187
NM_001135188
NM_001135189
NM_004504
AGFG1



ArfGAP with FG repeats 1



chr2_+_26568900 0.122 NM_033505
EPT1
ethanolaminephosphotransferase 1 (CDP-ethanolamine-specific)
chr15_-_55562483 0.121 NM_183234
NM_004580
RAB27A

RAB27A, member RAS oncogene family

chr7_-_92465929 0.121 NM_001145306
CDK6
cyclin-dependent kinase 6
chr7_+_134464161 0.119 NM_004342
NM_033138
NM_033157
CALD1


caldesmon 1


chr6_+_126240369 0.118 NM_001199622
NCOA7
nuclear receptor coactivator 7
chr11_+_118307072 0.118 NM_001197104
NM_005933
MLL

myeloid/lymphoid or mixed-lineage leukemia (trithorax homolog, Drosophila)

chr3_-_178789597 0.117 NM_022470
NM_152240
ZMAT3

zinc finger, matrin-type 3

chr7_-_92463211 0.116 NM_001259
CDK6
cyclin-dependent kinase 6
chr3_-_185216545 0.116 NM_080652
TMEM41A
transmembrane protein 41A
chr7_-_23053732 0.115 NM_032581
FAM126A
family with sequence similarity 126, member A
chr11_-_85780105 0.114 NM_001008660
NM_001206946
NM_007166
PICALM


phosphatidylinositol binding clathrin assembly protein


chr15_-_55563091 0.114 NM_183236
RAB27A
RAB27A, member RAS oncogene family
chr8_-_74884142 0.113 NM_001204861
NM_001204862
TCEB1

transcription elongation factor B (SIII), polypeptide 1 (15kDa, elongin C)

chr17_+_27717942 0.112 NM_020791
NM_025142
TAOK1

TAO kinase 1

chr8_-_74884500 0.109 NM_001204857
NM_001204858
NM_001204859
NM_001204860
NM_001204863
NM_001204864
NM_005648
TCEB1






transcription elongation factor B (SIII), polypeptide 1 (15kDa, elongin C)






chr12_+_69201845 0.108 NM_002392
MDM2
Mdm2 p53 binding protein homolog (mouse)
chr12_+_69004614 0.108 NM_001010942
NM_001251917
NM_001251918
NM_001251921
NM_001251922
NM_015646
RAP1B





RAP1B, member of RAS oncogene family





chr6_+_43543877 0.107 NM_006502
POLH
polymerase (DNA directed), eta
chr9_+_112542496 0.105 NM_053016
NM_007203
NM_147150
PALM2
PALM2-AKAP2

paralemmin 2
PALM2-AKAP2 readthrough

chr14_-_24020766 0.104 NM_033400
ZFHX2
zinc finger homeobox 2
chr8_-_116681103 0.104 NM_014112
TRPS1
trichorhinophalangeal syndrome I
chr6_+_138483024 0.104 NM_020340
KIAA1244
KIAA1244
chrX_-_103087106 0.103 NM_016370
RAB9B
RAB9B, member RAS oncogene family
chr7_+_28338939 0.103 NM_182899
CREB5
cAMP responsive element binding protein 5
chr3_+_196594708 0.102 NM_152699
SENP5
SUMO1/sentrin specific peptidase 5
chr4_-_83933961 0.102 NM_001115007
LIN54
lin-54 homolog (C. elegans)
chr6_+_163835668 0.102 NM_006775
NM_206853
NM_206854
NM_206855
QKI



QKI, KH domain containing, RNA binding



chr15_-_55581969 0.100 NM_183235
RAB27A
RAB27A, member RAS oncogene family
chr4_-_69215699 0.100 NM_001031732
NM_133370
YTHDC1

YTH domain containing 1

chr18_+_55102777 0.099 NM_004852
ONECUT2
one cut homeobox 2
chr17_+_29421941 0.098 NM_000267
NM_001042492
NM_001128147
NF1


neurofibromin 1


chr2_+_139259349 0.098 NM_001001664
SPOPL
speckle-type POZ protein-like
chr8_+_99130067 0.098 NM_015029
POP1
processing of precursor 1, ribonuclease P/MRP subunit (S. cerevisiae)
chr7_-_111846384 0.097 NM_014705
DOCK4
dedicator of cytokinesis 4
chr14_-_99737821 0.097 NM_022898
NM_138576
BCL11B

B-cell CLL/lymphoma 11B (zinc finger protein)

chr7_+_28475233 0.097 NM_004904
CREB5
cAMP responsive element binding protein 5
chr2_-_172291227 0.096 NM_024770
METTL8
methyltransferase like 8
chr12_-_31479120 0.096 NM_001135811
NM_021238
NM_001135812
FAM60A


family with sequence similarity 60, member A


chr5_-_171433876 0.094 NM_012300
NM_033644
NM_033645
FBXW11


F-box and WD repeat domain containing 11


chr1_+_218458628 0.094 NM_016052
RRP15
ribosomal RNA processing 15 homolog (S. cerevisiae)
chr8_+_90769974 0.094 NM_003821
RIPK2
receptor-interacting serine-threonine kinase 2
chr6_-_119031230 0.092 NM_001178035
CEP85L
centrosomal protein 85kDa-like
chr14_+_32546445 0.092 NM_001030055
NM_001173
ARHGAP5

Rho GTPase activating protein 5

chrX_+_49832214 0.092 NM_000084
CLCN5
chloride channel 5
chr5_+_149109814 0.091 NM_001172698
NM_133263
PPARGC1B

peroxisome proliferator-activated receptor gamma, coactivator 1 beta

chr8_+_99129488 0.091 NM_001145860
NM_001145861
POP1

processing of precursor 1, ribonuclease P/MRP subunit (S. cerevisiae)

chr17_+_72733350 0.090 NM_001006638
NM_001163990
RAB37

RAB37, member RAS oncogene family

chr7_+_134576150 0.090 NM_033139
NM_033140
CALD1

caldesmon 1

chr17_+_72732958 0.088 NM_001163989
RAB37
RAB37, member RAS oncogene family
chr6_-_57086204 0.087 NM_183227
RAB23
RAB23, member RAS oncogene family
chr9_+_110045516 0.086 NM_002874
RAD23B
RAD23 homolog B (S. cerevisiae)
chr2_+_70142152 0.086 NM_001202513
NM_001202514
NM_002357
MXD1


MAX dimerization protein 1


chr14_+_32798478 0.086 NM_004274
AKAP6
A kinase (PRKA) anchor protein 6
chr3_-_24536176 0.085 NM_000461
NM_001128176
NM_001128177
NM_001252634
THRB



thyroid hormone receptor, beta



chr1_+_167691156 0.083 NM_001146191
NM_003953
NM_024569
MPZL1


myelin protein zero-like 1


chr1_+_205538111 0.083 NM_181644
MFSD4
major facilitator superfamily domain containing 4
chr17_-_36105005 0.082 NM_000458
NM_001165923
HNF1B

HNF1 homeobox B

chr12_-_25403824 0.082 NM_004985
NM_033360
KRAS

v-Ki-ras2 Kirsten rat sarcoma viral oncogene homolog

chr1_+_26438267 0.080 NM_001243533
NM_152835
PDIK1L

PDLIM1 interacting kinase 1 like

chr6_+_37787266 0.080 NM_021943
ZFAND3
zinc finger, AN1-type domain 3
chr11_-_86666211 0.079 NM_012193
FZD4
frizzled family receptor 4
chr4_+_39699663 0.079 NM_001111112
NM_001111113
NM_005339
UBE2K


ubiquitin-conjugating enzyme E2K


chr9_-_23826062 0.079 NM_004432
ELAVL2
ELAV (embryonic lethal, abnormal vision, Drosophila)-like 2 (Hu antigen B)
chr5_+_176730812 0.079 NM_013237
PRELID1
PRELI domain containing 1
chrX_+_11129405 0.079 NM_001122608
NM_001171991
NM_005333
HCCS


holocytochrome c synthase


chr14_+_29236171 0.079 NM_005249
FOXG1
forkhead box G1
chr8_-_82598517 0.078 NM_001144878
NM_001144879
NM_005536
IMPA1


inositol(myo)-1(or 4)-monophosphatase 1


chr6_-_57087018 0.076 NM_016277
RAB23
RAB23, member RAS oncogene family
chr6_+_64281910 0.076 NM_003463
PTP4A1
protein tyrosine phosphatase type IVA, member 1
chr2_+_110371887 0.075 NM_023016
ANKRD57
ankyrin repeat domain 57
chr20_-_48530275 0.075 NM_001135773
SPATA2
spermatogenesis associated 2
chr3_+_50712671 0.075 NM_004947
DOCK3
dedicator of cytokinesis 3
chr11_-_75062650 0.075 NM_004041
NM_020251
ARRB1

arrestin, beta 1

chr4_-_159592979 0.075 NM_001008393
C4orf46
chromosome 4 open reading frame 46
chr12_-_93323088 0.073 NM_003566
EEA1
early endosome antigen 1
chr11_-_85780899 0.072 NM_001206947
PICALM
phosphatidylinositol binding clathrin assembly protein
chr8_+_76452202 0.072 NM_004133
HNF4G
hepatocyte nuclear factor 4, gamma
chr4_-_76598611 0.071 NM_203504
NM_203505
G3BP2

GTPase activating protein (SH3 domain) binding protein 2

chr4_+_40058507 0.071 NM_018177
N4BP2
NEDD4 binding protein 2
chr3_+_147127151 0.070 NM_003412
ZIC1
Zic family member 1
chr1_+_203444882 0.070 NM_002725
NM_201348
PRELP

proline/arginine-rich end leucine-rich repeat protein

chr4_-_76598289 0.069 NM_012297
G3BP2
GTPase activating protein (SH3 domain) binding protein 2
chr8_+_92082423 0.069 NM_016023
OTUD6B
OTU domain containing 6B
chr20_-_48532067 0.069 NM_006038
SPATA2
spermatogenesis associated 2
chr12_-_109251333 0.069 NM_001161330
NM_018984
SSH1

slingshot homolog 1 (Drosophila)

chr10_-_118765087 0.069 NM_001127211
NM_018330
KIAA1598

KIAA1598

chr6_+_126102278 0.068 NM_001199619
NM_001199620
NCOA7

nuclear receptor coactivator 7

chr4_-_83931973 0.068 NM_001115008
NM_194282
LIN54

lin-54 homolog (C. elegans)

chr11_+_118477212 0.068 NM_015157
PHLDB1
pleckstrin homology-like domain, family B, member 1
chr4_+_72052354 0.068 NM_001098484
NM_001134742
SLC4A4

solute carrier family 4, sodium bicarbonate cotransporter, member 4

chr4_-_146100793 0.067 NM_001102653
OTUD4
OTU domain containing 4
chr10_+_86088156 0.067 NM_018999
FAM190B
family with sequence similarity 190, member B
chr12_+_99038918 0.067 NM_001160
NM_013229
NM_181861
NM_181868
NM_181869
APAF1




apoptotic peptidase activating factor 1




chr16_-_9057250 0.066 NM_003470
USP7
ubiquitin specific peptidase 7 (herpes virus-associated)
chr9_-_23821477 0.065 NM_001171197
ELAVL2
ELAV (embryonic lethal, abnormal vision, Drosophila)-like 2 (Hu antigen B)
chr11_+_118478305 0.065 NM_001144758
NM_001144759
PHLDB1

pleckstrin homology-like domain, family B, member 1

chr20_+_43514340 0.065 NM_003404
NM_139323
YWHAB

tyrosine 3-monooxygenase/tryptophan 5-monooxygenase activation protein, beta polypeptide

chr5_+_71403040 0.065 NM_005909
MAP1B
microtubule-associated protein 1B
chr5_-_60458228 0.065 NM_001048249
C5orf43
chromosome 5 open reading frame 43
chr3_-_16306283 0.065 NM_001047434
NM_206831
DPH3

DPH3, KTI11 homolog (S. cerevisiae)

chr16_+_7382750 0.064 NM_145891
NM_145892
NM_145893
RBFOX1


RNA binding protein, fox-1 homolog (C. elegans) 1


chr5_+_135468522 0.062 NM_001001419
NM_001001420
NM_005903
SMAD5


SMAD family member 5


chr10_+_25464289 0.062 NM_020752
GPR158
G protein-coupled receptor 158
chr9_-_23821842 0.062 NM_001171195
ELAVL2
ELAV (embryonic lethal, abnormal vision, Drosophila)-like 2 (Hu antigen B)
chrX_-_129402753 0.061 NM_017666
ZNF280C
zinc finger protein 280C
chr2_-_95825262 0.061 NM_032788
ZNF514
zinc finger protein 514
chr15_-_62352565 0.060 NM_001018088
NM_017684
NM_018080
NM_020821
VPS13C



vacuolar protein sorting 13 homolog C (S. cerevisiae)



chr4_+_57774035 0.059 NM_005612
REST
RE1-silencing transcription factor
chr2_-_60780404 0.059 NM_018014
NM_022893
NM_138559
BCL11A


B-cell CLL/lymphoma 11A (zinc finger protein)


chr9_+_33025207 0.058 NM_001539
DNAJA1
DnaJ (Hsp40) homolog, subfamily A, member 1
chr5_+_149151502 0.058 NM_001172699
PPARGC1B
peroxisome proliferator-activated receptor gamma, coactivator 1 beta
chr10_-_7453412 0.057 NM_001029880
SFMBT2
Scm-like with four mbt domains 2
chr2_+_166428845 0.057 NM_024969
CSRNP3
cysteine-serine-rich nuclear protein 3
chr6_-_13814546 0.057 NM_001031713
CCDC90A
coiled-coil domain containing 90A
chr7_+_7222142 0.057 NM_020156
C1GALT1
core 1 synthase, glycoprotein-N-acetylgalactosamine 3-beta-galactosyltransferase, 1
chr14_+_100705101 0.056 NM_003403
YY1
YY1 transcription factor
chr2_-_202507348 0.056 NM_152388
TMEM237
transmembrane protein 237
chr21_-_37432789 0.056 NM_017438
SETD4
SET domain containing 4
chr9_+_115983807 0.056 NM_001859
SLC31A1
solute carrier family 31 (copper transporters), member 1
chr8_-_30706532 0.055 NM_031271
TEX15
testis expressed 15
chrX_+_152599544 0.055 NM_001080485
ZNF275
zinc finger protein 275
chr1_-_212004113 0.055 NM_014873
LPGAT1
lysophosphatidylglycerol acyltransferase 1
chr4_+_57775064 0.055 NM_001193508
REST
RE1-silencing transcription factor
chr7_+_91875411 0.055 NM_019004
ANKIB1
ankyrin repeat and IBR domain containing 1
chr1_-_146644045 0.054 NM_005399
PRKAB2
protein kinase, AMP-activated, beta 2 non-catalytic subunit
chr6_-_7313352 0.054 NM_003144
SSR1
signal sequence receptor, alpha
chr16_-_12009804 0.054 NM_001130006
NM_002094
GSPT1

G1 to S phase transition 1

chr12_-_48499415 0.054 NM_014554
SENP1
SUMO1/sentrin specific peptidase 1
chr12_+_65218351 0.054 NM_015279
TBC1D30
TBC1 domain family, member 30
chr18_+_67068134 0.053 NM_152721
DOK6
docking protein 6
chr16_-_12010518 0.052 NM_001130007
GSPT1
G1 to S phase transition 1
chr4_-_23891608 0.051 NM_013261
PPARGC1A
peroxisome proliferator-activated receptor gamma, coactivator 1 alpha

Gene Ontology Analysis

Gene overrepresentation in process category:

enrichment   p-value GO term description
2.13 2.29e-04 GO:0033036 macromolecule localization
2.35 1.26e-03 GO:0045184 establishment of protein localization
2.11 4.93e-03 GO:0008104 protein localization
1.46 4.95e-03 GO:0044260 cellular macromolecule metabolic process
2.27 5.60e-03 GO:0015031 protein transport
2.32 6.31e-03 GO:0046907 intracellular transport
1.77 1.20e-02 GO:0006464 protein modification process
1.73 1.54e-02 GO:0043412 macromolecule modification
1.18 1.57e-02 GO:0009987 cellular process
1.92 2.70e-02 GO:0051641 cellular localization
1.99 2.73e-02 GO:0051649 establishment of localization in cell
1.38 3.72e-02 GO:0043170 macromolecule metabolic process

Gene overrepresentation in compartment category:

enrichment   p-value GO term description
1.36 1.89e-03 GO:0005634 nucleus
1.17 5.06e-03 GO:0044424 intracellular part
1.15 9.61e-03 GO:0005622 intracellular

Gene overrepresentation in function category:

enrichment   p-value GO term description
1.17 3.30e-04 GO:0005488 binding
1.27 9.34e-03 GO:0005515 protein binding
1.60 1.19e-02 GO:0000166 nucleotide binding