Motif ID: REST.p3

Z-value: 4.882


Transcription factors associated with REST.p3:

Gene SymbolEntrez IDGene Name
REST 5978 RE1-silencing transcription factor

Activity-expression correlation:

GenePromoterPearsonP-valuePlot
RESTchr4_+_574691220.495.2e-02Click!


Activity profile for motif REST.p3.

activity profile for motif REST.p3


Sorted Z-values histogram for motif REST.p3

Sorted Z-values for motif REST.p3



Network of associatons between targets according to the STRING database.



First level regulatory network of REST.p3

PNG image of the network

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Top targets:


Promoter Score Refseq Gene Symbol Gene Name
chr9_+_134447795 19.795 NM_020064
BARHL1
BarH-like homeobox 1
chr2_-_27384617 14.421 NM_003353
UCN
urocortin
chr3_+_9720509 13.523 NM_153635
CPNE9
copine family member IX
chr8_-_145530639 12.309 NM_031309
SCRT1
scratch homolog 1, zinc finger protein (Drosophila)
chr6_+_110406890 11.072 NM_005284
GPR6
G protein-coupled receptor 6
chr5_-_151284515 10.969 NM_000171
NM_001146040
GLRA1

glycine receptor, alpha 1

chr4_+_158361185 10.352 NM_000826
NM_001083619
NM_001083620
GRIA2


glutamate receptor, ionotropic, AMPA 2


chr5_+_71050745 10.229 NM_004291
CARTPT
CART prepropeptide
chr15_+_65904994 9.975 NM_001031807
SKOR1
SKI family transcriptional corepressor 1
chr7_+_154943466 9.089 NM_001427
EN2
engrailed homeobox 2
chr7_-_19778928 8.682 NM_152774
TMEM196
transmembrane protein 196
chr10_+_26545599 8.601 GAD2
glutamate decarboxylase 2 (pancreatic islets and brain, 65kDa)
chr8_-_57521065 8.543 NM_006211
PENK
proenkephalin
chr19_-_60383280 8.168 NM_003180
SYT5
synaptotagmin V
chr20_+_30019465 7.418 NM_001011718
XKR7
XK, Kell blood group complex subunit-related family, member 7
chr14_-_50631407 7.287 TRIM9
tripartite motif containing 9
chr12_-_3472245 7.271 LOC100129223
hypothetical LOC100129223
chr5_+_33972247 7.188 NM_016568
RXFP3
relaxin/insulin-like family peptide receptor 3
chr8_+_57520915 7.187


chr20_+_44091219 6.990 NM_020708
SLC12A5
solute carrier family 12 (potassium/chloride transporter), member 5
chr20_+_44091265 6.835 SLC12A5
solute carrier family 12 (potassium/chloride transporter), member 5
chr9_-_138236775 6.698 NM_178138
LHX3
LIM homeobox 3
chr3_-_133236447 6.563 NM_130808
CPNE4
copine IV
chr11_+_22316242 6.423 NM_020346
SLC17A6
solute carrier family 17 (sodium-dependent inorganic phosphate cotransporter), member 6
chr11_-_74819165 6.367 NM_001039548
KLHL35
kelch-like 35 (Drosophila)
chr11_-_74818858 6.367 KLHL35
kelch-like 35 (Drosophila)
chr3_-_48675321 6.330 NM_001407
CELSR3
cadherin, EGF LAG seven-pass G-type receptor 3 (flamingo homolog, Drosophila)
chr8_-_91164100 6.328 NM_004929
CALB1
calbindin 1, 28kDa
chr11_-_27698800 6.302 NM_001143805
NM_001143806
NM_170732
BDNF


brain-derived neurotrophic factor


chr11_+_627268 6.194 NM_000797
DRD4
dopamine receptor D4
chr2_+_207215386 6.121 NM_001102659
LOC200726
hCG1657980
chr1_-_149955870 5.910 NM_007185
CELF3
CUGBP, Elav-like family member 3
chr3_+_122109716 5.854 NM_014980
STXBP5L
syntaxin binding protein 5-like
chr20_-_10148137 5.839 LOC100131208
similar to hCG2045825
chr11_-_74818685 5.781 KLHL35
kelch-like 35 (Drosophila)
chr11_-_122357545 5.630 NM_001098169
BSX
brain-specific homeobox
chr17_+_1120607 5.552 NM_001164405
BHLHA9
basic helix-loop-helix family, member a9
chr14_+_92459197 5.440 NM_001275
CHGA
chromogranin A (parathyroid secretory protein 1)
chr14_-_73962557 5.318 NM_001105579
TMEM90A
transmembrane protein 90A
chr17_-_24357014 5.195 NM_001098635
NM_178860
SEZ6

seizure related 6 homolog (mouse)

chr1_+_208178160 4.858 NM_001146261
NM_001146262
NM_001146264
NM_153262
SYT14



synaptotagmin XIV



chr14_-_50631794 4.847 TRIM9
tripartite motif containing 9
chr10_+_26545241 4.841 NM_000818
NM_001134366
GAD2

glutamate decarboxylase 2 (pancreatic islets and brain, 65kDa)

chr4_-_175986952 4.762 NM_001042543
NM_006529
GLRA3

glycine receptor, alpha 3

chr1_+_208178198 4.732 SYT14
synaptotagmin XIV
chr9_+_139153409 4.577 NM_000832
NM_001185090
NM_001185091
NM_007327
NM_021569
GRIN1




glutamate receptor, ionotropic, N-methyl D-aspartate 1




chr14_+_92459244 4.562 CHGA
chromogranin A (parathyroid secretory protein 1)
chr19_-_45416043 4.534 TTC9B
tetratricopeptide repeat domain 9B
chr14_-_104142125 4.427 NM_207379
TMEM179
transmembrane protein 179
chr19_-_45416112 4.199 NM_152479
TTC9B
tetratricopeptide repeat domain 9B
chr6_+_154402135 3.834 NM_000914
NM_001008503
NM_001008504
NM_001008505
NM_001145282
NM_001145283
NM_001145285
NM_001145286
NM_001145284
OPRM1








opioid receptor, mu 1








chr7_-_100595552 3.784 NM_003378
VGF
VGF nerve growth factor inducible
chr7_+_8440541 3.759 NXPH1
neurexophilin 1
chr15_-_70399181 3.739 CELF6
CUGBP, Elav-like family member 6
chr3_-_188870799 3.669 NM_001048
SST
somatostatin
chr15_+_73074882 3.605 NM_001178111
NM_001178112
NM_138967
SCAMP5


secretory carrier membrane protein 5


chr11_+_45863774 3.479 NM_005456
MAPK8IP1
mitogen-activated protein kinase 8 interacting protein 1
chr4_+_8633116 3.425 NM_080819
GPR78
G protein-coupled receptor 78
chr2_-_51112743 3.421 NRXN1
neurexin 1
chr1_+_33124658 3.408 NM_002143
HPCA
hippocalcin
chr12_+_111714096 3.375 RPH3A
rabphilin 3A homolog (mouse)
chr1_-_102234959 3.314 OLFM3
olfactomedin 3
chr3_-_115380539 3.182 NM_000796
NM_033663
DRD3

dopamine receptor D3

chr16_+_66244314 3.123 RLTPR
RGD motif, leucine rich repeats, tropomodulin domain and proline-rich containing
chr5_-_135729062 3.114 NM_001167576
NM_001167577
NM_020389
TRPC7


transient receptor potential cation channel, subfamily C, member 7


chr7_+_8440109 3.107 NM_152745
NXPH1
neurexophilin 1
chr15_-_70399301 2.979 NM_001172684
CELF6
CUGBP, Elav-like family member 6
chr6_+_158877640 2.922 TMEM181
transmembrane protein 181
chr20_-_60228665 2.899 NM_007232
HRH3
histamine receptor H3
chr12_+_111713931 2.835 NM_001143854
NM_014954
RPH3A

rabphilin 3A homolog (mouse)

chr1_+_152806874 2.786 NM_000748
CHRNB2
cholinergic receptor, nicotinic, beta 2 (neuronal)
chr15_-_24570010 2.784 NM_021912
GABRB3
gamma-aminobutyric acid (GABA) A receptor, beta 3
chr12_-_97812633 2.763 ANKS1B
ankyrin repeat and sterile alpha motif domain containing 1B
chr18_-_39111243 2.719 NM_020783
SYT4
synaptotagmin IV
chr12_-_29828958 2.684 NM_175861
TMTC1
transmembrane and tetratricopeptide repeat containing 1
chr17_+_77925441 2.624 NM_018949
UTS2R
urotensin 2 receptor
chr20_-_61755145 2.560 NM_015894
STMN3
stathmin-like 3
chr19_-_44427446 2.438 NM_172139
IL28B
interleukin 28B (interferon, lambda 3)
chr1_-_102235377 2.419 NM_058170
OLFM3
olfactomedin 3
chr12_+_117903682 2.411 NM_194286
SRRM4
serine/arginine repetitive matrix 4
chr15_-_70399551 2.379 NM_052840
CELF6
CUGBP, Elav-like family member 6
chr11_-_125278296 2.303 NM_031307
PUS3
pseudouridylate synthase 3
chr7_+_145444385 2.293 NM_014141
CNTNAP2
contactin associated protein-like 2
chr17_+_70178911 2.281 RAB37
RAB37, member RAS oncogene family
chr12_-_29827952 2.196 NM_001193451
TMTC1
transmembrane and tetratricopeptide repeat containing 1
chr17_+_70178850 2.152 NM_175738
RAB37
RAB37, member RAS oncogene family
chr10_+_69990050 2.059 NM_030625
TET1
tet oncogene 1
chr15_-_70399523 2.039 CELF6
CUGBP, Elav-like family member 6
chr19_+_4720144 2.025 C19orf30
chromosome 19 open reading frame 30
chr12_+_46133 2.019 NM_001170738
IQSEC3
IQ motif and Sec7 domain 3
chr6_+_158877455 1.819 NM_020823
TMEM181
transmembrane protein 181
chr10_-_44200458 1.797 NM_000609
NM_001033886
NM_001178134
NM_199168
CXCL12



chemokine (C-X-C motif) ligand 12



chr19_+_45728270 1.792 SPTBN4
spectrin, beta, non-erythrocytic 4
chr1_-_239587090 1.679 NM_002924
RGS7
regulator of G-protein signaling 7
chr2_+_114916368 1.677 NM_020868
DPP10
dipeptidyl-peptidase 10 (non-functional)
chr17_-_3248453 1.660 NM_003553
OR1E1
olfactory receptor, family 1, subfamily E, member 1
chr9_-_73251639 1.635 TRPM3
transient receptor potential cation channel, subfamily M, member 3
chr8_-_145723209 1.627 NM_001024678
LRRC24
leucine rich repeat containing 24
chr5_+_175231098 1.555 NM_001008220
CPLX2
complexin 2
chr10_+_60607232 1.496 NM_001143774
PHYHIPL
phytanoyl-CoA 2-hydroxylase interacting protein-like
chrX_-_47364039 1.492


chr6_+_25760352 1.446 NM_006998
SCGN
secretagogin, EF-hand calcium binding protein
chr3_+_62279707 1.409 C3orf14
chromosome 3 open reading frame 14
chr5_+_175231170 1.393


chr7_+_8439762 1.346 NXPH1
neurexophilin 1
chr5_+_175231210 1.339 CPLX2
complexin 2
chrX_-_47364173 1.334 NM_006950
NM_133499
SYN1

synapsin I

chr16_+_2959362 1.282 PAQR4
progestin and adipoQ receptor family member IV
chr10_-_50640327 1.272 NM_001143996
NM_001143997
NM_018245
OGDHL


oxoglutarate dehydrogenase-like


chr19_+_45728207 1.239 NM_025213
SPTBN4
spectrin, beta, non-erythrocytic 4
chr11_-_1725924 1.239 NM_001170820
LOC402778
CD225 family protein FLJ76511
chr16_+_2959342 1.226 NM_152341
PAQR4
progestin and adipoQ receptor family member IV
chr10_-_102780840 1.223 PDZD7
PDZ domain containing 7
chr18_-_5187240 1.183 NM_001145194
LOC642597
hypothetical protein LOC642597
chr10_-_102780867 1.155 NM_001195263
NM_024895
PDZD7

PDZ domain containing 7

chr2_-_51113040 1.088 NM_001135659
NM_004801
NRXN1

neurexin 1

chr5_-_175969697 1.043 NM_052899
GPRIN1
G protein regulated inducer of neurite outgrowth 1
chr15_+_67009892 0.944 NM_145658
NM_001184780
SPESP1
NOX5
sperm equatorial segment protein 1
NADPH oxidase, EF-hand calcium binding domain 5
chr17_+_39741306 0.929 NM_001144825
NM_001144826
NM_006695
RUNDC3A


RUN domain containing 3A


chr6_+_153113624 0.923 NM_003381
NM_194435
VIP

vasoactive intestinal peptide

chr2_+_131230252 0.907 NM_001105194
NM_001105195
FAM123C

family with sequence similarity 123C

chr3_+_52787478 0.878 NM_001166434
ITIH1
inter-alpha (globulin) inhibitor H1
chr1_+_152806924 0.859 CHRNB2
cholinergic receptor, nicotinic, beta 2 (neuronal)
chr15_-_29950070 0.854 OTUD7A
OTU domain containing 7A
chrX_-_152794478 0.832 L1CAM
L1 cell adhesion molecule
chr11_-_6633427 0.827 NM_003737
DCHS1
dachsous 1 (Drosophila)
chr10_+_102780974 0.813 NM_030971
SFXN3
sideroflexin 3
chr1_-_239587007 0.781 RGS7
regulator of G-protein signaling 7
chr11_-_1725662 0.742 LOC402778
CD225 family protein FLJ76511
chr16_+_24175173 0.728 CACNG3
calcium channel, voltage-dependent, gamma subunit 3
chr8_+_69027143 0.698 NM_024870
NM_025170
PREX2

phosphatidylinositol-3,4,5-trisphosphate-dependent Rac exchange factor 2

chr8_+_69026895 0.675 PREX2
phosphatidylinositol-3,4,5-trisphosphate-dependent Rac exchange factor 2
chr15_-_29950188 0.668 OTUD7A
OTU domain containing 7A
chr17_-_3283884 0.652 NM_003554
OR1E2
olfactory receptor, family 1, subfamily E, member 2
chr8_+_69027277 0.646 PREX2
phosphatidylinositol-3,4,5-trisphosphate-dependent Rac exchange factor 2
chr19_-_50429306 0.641 NM_138568
EXOC3L2
exocyst complex component 3-like 2
chr8_+_80685859 0.630 STMN2
stathmin-like 2
chr1_-_33108876 0.618 NM_001171940
NM_153756
FNDC5

fibronectin type III domain containing 5

chr5_+_175231144 0.600 CPLX2
complexin 2
chr16_+_24174975 0.586 CACNG3
calcium channel, voltage-dependent, gamma subunit 3
chr12_+_112280757 0.574 PLBD2
phospholipase B domain containing 2
chr3_+_62280435 0.546 NM_020685
C3orf14
chromosome 3 open reading frame 14
chr1_-_151390050 0.499 NM_001014291
SPRR2G
small proline-rich protein 2G
chrX_-_152794592 0.498 NM_000425
NM_001143963
NM_024003
L1CAM


L1 cell adhesion molecule


chr11_-_123030091 0.483 SCN3B
sodium channel, voltage-gated, type III, beta
chrX_-_48943582 0.441 NM_003179
SYP
synaptophysin
chr18_+_55718171 0.423 NM_021127
PMAIP1
phorbol-12-myristate-13-acetate-induced protein 1
chr20_+_5839973 0.415 NM_001819
CHGB
chromogranin B (secretogranin 1)
chr5_+_141283550 0.407 NM_001142603
NM_014773
KIAA0141

KIAA0141

chr6_+_35881048 0.393 NM_182548
LHFPL5
lipoma HMGIC fusion partner-like 5
chr20_+_35445350 0.322 SRC
v-src sarcoma (Schmidt-Ruppin A-2) viral oncogene homolog (avian)
chr12_+_112280753 0.295 NM_001159727
NM_173542
PLBD2

phospholipase B domain containing 2

chr17_-_3546236 0.282 NM_002561
NM_175080
NM_175081
P2RX5-TAX1BP3
P2RX5


P2RX5-TAX1BP3 read-through transcript
purinergic receptor P2X, ligand-gated ion channel, 5


chr17_-_7095622 0.266 NM_001143775
CTDNEP1
CTD nuclear envelope phosphatase 1
chr5_+_141283647 0.241 KIAA0141
KIAA0141
chr19_+_38867273 0.222 NM_022467
CHST8
carbohydrate (N-acetylgalactosamine 4-0) sulfotransferase 8
chr19_+_51804260 0.155 CALM3
calmodulin 3 (phosphorylase kinase, delta)
chr19_-_50618599 0.139 ERCC1
excision repair cross-complementing rodent repair deficiency, complementation group 1 (includes overlapping antisense sequence)
chr6_-_32121784 0.138 NM_032470
TNXB
tenascin XB
chr1_+_100777044 0.112 GPR88
G protein-coupled receptor 88
chr11_-_27677755 0.110 NM_170734
BDNF
brain-derived neurotrophic factor
chrX_+_152891184 0.100 NM_003492
TMEM187
transmembrane protein 187
chr1_-_20707103 0.096 MUL1
mitochondrial E3 ubiquitin protein ligase 1
chr1_-_20707196 0.072 NM_024544
MUL1
mitochondrial E3 ubiquitin protein ligase 1
chr10_+_73225533 0.066 NM_001171933
NM_001171934
CDH23

cadherin-related 23

chr8_-_94046891 0.054 NM_001191036
C8orf83
chromosome 8 open reading frame 83
chr11_-_72170533 0.054 STARD10
StAR-related lipid transfer (START) domain containing 10
chr2_-_96174502 0.049 DUSP2
dual specificity phosphatase 2
chr22_-_16636748 0.042 NM_197966
BID
BH3 interacting domain death agonist
chr11_+_64565118 0.025 SAC3D1
SAC3 domain containing 1
chr11_+_62136788 0.016 NM_000327
ROM1
retinal outer segment membrane protein 1

Gene Ontology Analysis

Gene overrepresentation in process category:

enrichment   p-value GO term description
8.12 1.76e-11 GO:0007268 synaptic transmission
7.56 1.94e-11 GO:0019226 transmission of nerve impulse
7.56 1.94e-11 GO:0035637 multicellular organismal signaling
3.52 2.70e-09 GO:0003008 system process
3.91 3.22e-09 GO:0050877 neurological system process
4.92 1.32e-07 GO:0007267 cell-cell signaling
5.57 7.80e-07 GO:0007186 G-protein coupled receptor protein signaling pathway
4.02 1.25e-06 GO:0007154 cell communication
7.09 7.19e-06 GO:0007610 behavior
5.84 9.59e-06 GO:0046903 secretion
10.72 2.17e-05 GO:0031644 regulation of neurological system process
2.10 4.50e-05 GO:0023052 signaling
10.66 1.15e-04 GO:0051969 regulation of transmission of nerve impulse
1.80 2.04e-04 GO:0032501 multicellular organismal process
5.03 4.97e-04 GO:0055082 cellular chemical homeostasis
4.84 1.96e-03 GO:0006873 cellular ion homeostasis
11.06 2.24e-03 GO:0007626 locomotory behavior
5.56 3.23e-03 GO:0032940 secretion by cell
19.84 3.34e-03 GO:0051952 regulation of amine transport
7.11 3.43e-03 GO:0055074 calcium ion homeostasis
3.95 3.97e-03 GO:0048878 chemical homeostasis
6.85 4.63e-03 GO:0072507 divalent inorganic cation homeostasis
4.47 4.69e-03 GO:0050801 ion homeostasis
4.15 4.70e-03 GO:0019725 cellular homeostasis
9.70 5.36e-03 GO:0003001 generation of a signal involved in cell-cell signaling
9.70 5.36e-03 GO:0023061 signal release
5.22 5.89e-03 GO:0044057 regulation of system process
2.28 7.50e-03 GO:0065008 regulation of biological quality
9.18 7.70e-03 GO:0050804 regulation of synaptic transmission
26.69 9.23e-03 GO:0014073 response to tropane
26.69 9.23e-03 GO:0042220 response to cocaine
24.47 1.33e-02 GO:0060078 regulation of postsynaptic membrane potential
48.93 1.71e-02 GO:0010155 regulation of proton transport
22.59 1.85e-02 GO:0050433 regulation of catecholamine secretion
10.13 1.91e-02 GO:0001505 regulation of neurotransmitter levels
21.75 2.16e-02 GO:0045776 negative regulation of blood pressure
6.53 2.16e-02 GO:0006874 cellular calcium ion homeostasis
1.96 2.29e-02 GO:0051234 establishment of localization
4.97 2.34e-02 GO:0051046 regulation of secretion
6.32 2.73e-02 GO:0072503 cellular divalent inorganic cation homeostasis
6.29 2.84e-02 GO:0003013 circulatory system process
6.29 2.84e-02 GO:0008015 blood circulation
4.80 3.10e-02 GO:0055080 cation homeostasis
40.04 3.32e-02 GO:0032225 regulation of synaptic transmission, dopaminergic
1.93 4.05e-02 GO:0006810 transport
11.84 4.33e-02 GO:0007269 neurotransmitter secretion
36.70 4.41e-02 GO:0032413 negative regulation of ion transmembrane transporter activity
8.73 4.45e-02 GO:0006836 neurotransmitter transport
8.56 4.96e-02 GO:0008217 regulation of blood pressure

Gene overrepresentation in compartment category:

enrichment   p-value GO term description
8.70 1.27e-09 GO:0044456 synapse part
6.84 1.62e-08 GO:0045202 synapse
17.07 4.22e-08 GO:0008021 synaptic vesicle
26.10 8.07e-08 GO:0030672 synaptic vesicle membrane
6.12 9.16e-07 GO:0043005 neuron projection
4.29 4.45e-06 GO:0016023 cytoplasmic membrane-bounded vesicle
4.21 6.07e-06 GO:0031988 membrane-bounded vesicle
4.04 1.20e-05 GO:0031410 cytoplasmic vesicle
13.82 1.44e-05 GO:0030665 clathrin coated vesicle membrane
9.18 1.78e-05 GO:0030136 clathrin-coated vesicle
3.91 2.03e-05 GO:0031982 vesicle
6.81 2.55e-05 GO:0030659 cytoplasmic vesicle membrane
6.58 3.68e-05 GO:0012506 vesicle membrane
4.39 3.79e-05 GO:0030054 cell junction
7.50 1.17e-04 GO:0030135 coated vesicle
10.22 1.49e-04 GO:0030662 coated vesicle membrane
5.65 1.84e-04 GO:0044433 cytoplasmic vesicle part
3.41 3.39e-04 GO:0042995 cell projection
6.87 2.83e-03 GO:0043025 neuronal cell body
6.68 3.48e-03 GO:0044297 cell body
2.16 3.83e-03 GO:0044459 plasma membrane part
6.22 5.79e-03 GO:0030425 dendrite
1.44 6.06e-03 GO:0016020 membrane
1.52 6.99e-03 GO:0044425 membrane part
8.90 7.51e-03 GO:0019717 synaptosome
6.95 8.77e-03 GO:0045211 postsynaptic membrane
1.56 1.35e-02 GO:0031224 intrinsic to membrane
1.56 1.63e-02 GO:0016021 integral to membrane
25.91 2.51e-02 GO:0032589 neuron projection membrane
73.39 3.52e-02 GO:0032590 dendrite membrane
1.64 3.79e-02 GO:0005886 plasma membrane

Gene overrepresentation in function category:

enrichment   p-value GO term description
14.28 7.79e-05 GO:0005230 extracellular ligand-gated ion channel activity
18.74 1.01e-04 GO:0005231 excitatory extracellular ligand-gated ion channel activity
27.96 1.51e-03 GO:0005184 neuropeptide hormone activity
8.23 3.14e-03 GO:0015276 ligand-gated ion channel activity
8.23 3.14e-03 GO:0022834 ligand-gated channel activity
4.25 7.84e-03 GO:0005216 ion channel activity
8.90 7.91e-03 GO:0005179 hormone activity
4.18 9.02e-03 GO:0022838 substrate-specific channel activity
33.88 1.13e-02 GO:0016594 glycine binding
4.05 1.18e-02 GO:0015267 channel activity
4.04 1.21e-02 GO:0022803 passive transmembrane transporter activity
97.85 1.86e-02 GO:0001591 dopamine receptor activity, coupled via Gi/Go
7.17 2.63e-02 GO:0043176 amine binding
4.40 2.93e-02 GO:0022836 gated channel activity
2.46 3.20e-02 GO:0005215 transporter activity
73.39 3.71e-02 GO:0016933 extracellular-glycine-gated ion channel activity
73.39 3.71e-02 GO:0016934 extracellular-glycine-gated chloride channel activity
3.05 4.58e-02 GO:0005509 calcium ion binding