BIPOLAR DISORDER GENES Kegg pathway analysis of over 400 bipolar genes
Polygenicblog on the relationships between genes and risk factors
June, 2011: In future, Bipolar genes will be listed at this page, along with the risk factors and KEGG pathways. Links to new genes will be provided, with the NextBio highlights. If you know of any other genes, please Thank you
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The
tables below contain lists of genes reported to be associated with
Bipolar disorder. Negative studies also exist for many of these genes,
but are not reported. Each gene is linked to its ENTREZ gene ID. The links
are designed to collect general data in relation to Bipolar disorder,
rather than specific association data. Protein/protein or transriptional
control interactions are generally culled from ENTREZ gene data unless
specified. , Entrez and OMIM data are provided by the NCBI
GeneCards links are with the kind permission of GeneCards.org.
ENVIRONMENTAL RISK FACTORS ARE INCLUDED HERE
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For genome wide association study see Baum et al, 2008 A2BP1, ABCA6, ABCB11, ABTB1, ANK3, ASTN2, AQP4, BMP6, BRE, c11orf44, C14orf143, C20orf17, C21orf127, CAMK2D, CDH20, CHES1, CNTNAP5, CNTN5, CNTN6, CSMD1, CYP4V2, DGKH, DDX42, DFNB31, DOCK3, EDA, EFNA5, FALZ, FLJ39058, FZD2, GAB2, GALNTL4, GASP, GPR51, GRM1, HK2, JAM3, KCTD8, KLF12, KRN1, LAMA3, LDB2, LLGL2, LRCH1, LY86, MAK, MGC42174, MUC2, NCL, NGL1, NYD-SP26, NXN, OR51F2, PAX7, PHF17, PLA2R1, PLCG2, PLSCR4, PSMF1, PTGIS, PTPRB, PTPRG, PTPRN2, RBMS3, RFC3, RGS17, RNPEPL1, RYR2, SDC2, SGCD, SHOC2, SLC26A7, SLC39A3, SLIT3, SNX27, SKIV2L, SRP54, STAB1, SORCS2, SUPT4H1, TCF7L1, TDP1, TEC, UNQ689, VAV3, VPRBP, VGCNL1: The Wellcome Trust Case Control Consortium. Genome-wide association study.The following genes are those corresponding to the rs SNP numbers defined in the supplementary material of this paper (moderate to strong association) AOF1 AK3L2 AKAP10 C14orf58 CAPN6 CDC25B CMTM8 CSF2RB DFNB31(see above) DPP10 ESRRG FAM126A GABRB1 GRM7 KLHDC1 LAMP3 LOC283547 LOC731264 LRRC7 PALB2 PAX5 PTPRE (PTPRG RNPEPL1 see above) SOX5 SVEP1 SYK SYN3 SYNE1 TDRD9 THRB THSD7A TRDN ZBTB44 ZNF274 ZNF490 ZNF659 ZNF678 |
PI3K/AKT signalling, growth factors and related | ANK3, AKT1, BDNF, BMP6, CABIN1, EGFR, FYN, GSK3B, IGF1, IMPA2, NRG1, PIK3C3, PIK4CA, PIP5K2A, PDLIM5, RGS4, TCF4, WDFY2 |
Adhesion/ Junctions | CNTNAP2, JAM3, NCAM1, NRXN1 |
NMDA and glutamate-related | CIT, DAO, DAOA, DTNBP1, GRIA1, GRID1, GRIN1, GRIN2A, GRIN2B, GRIK4, GRM3, GRM4, GRM7, NOS1, NOSIAP, PCLO, SNAP25, SYN3, SYNJ1 |
Dopaminergic/Serotonergic | COMT, DRD2, DRD3, HTR2A, HTR5A, HTR6, NCKAP5, SLC6A3, SLC6A4,SLC18A1, SLC18A2, MAOA, TDO2, TH |
GABA | GABRA1, GABRB2, GAD1 |
Circadian | CLOCK, NPAS2, NR1D1, TIMELESS, PER3 |
Cytokines | CSF2RB, IL1B, IL1RN, IL6, IL10, TNFA |
Oxidative and other stress | ND4, NDUFV2, MTHFR, MTHFD, MTR, NOS3 |
Endoplasmic reticulum stress | XBP1 |
Miscellaneous | ACE, ABCB1, ABCA13, AGT, APOE, BRD1, C10orf120, CACNA1C, CDKN2A, CHRNA7, DISC1, DPYSL2, FABP7, FBXO38. GLT8D1, GNL3, GPR50, GPR78, hCAP-D3, INTS6, ITIH3, MC5R, MCHR1, MCHR2, MLC1, MMP9, MRCL3, MRLC2, NALCN, NEK4, NPAS3, PB1, PDE4B,PFN4, PPARD, PPP3CC, PSKH1, RELN, S100B, SEMG2, SLIT3, SP4, SST, SYNGR1, TAAR6, TMEM108, TP53I3, TSNAX, TSPAN8, TUBA8, YWHAH, ZNF804A |
Table 2:Growth factors: Phosphatidyl-inositol metabolism and PI3K/AKT1 signaling and tyrosine kinase pathway related genes
Genes
associated with bipolar disorder |
Association
studies |
Primary
role |
Links
to other Bipolar genes |
Expression
changes |
BDNF
brain derived
neurotrophic factor OMIM OMIM
|
A number
of studies suggest association of this gene with bipolar disorder (Geller
et al. 2004;Lohoff et al. 2005;Rybakowski et al. 2003;Neves-Pereira
et al. 2002). Its protein levels are decreased in the hippocampus of
Bipolar disorder patients (Knable et al. 2004). |
Growth
factor activating the PI3K/AKT cascade (Gavalda
et al. 2004) |
Expression
stimulated by NMDA receptor activation (GRIN1,
GRIN2A) and by DRD1(Fang
et al. 2003),(Kuppers
and Beyer 2001)
and DRD2 (Takeuchi
et al. 2002) receptor
stimulation. Controls TH (Zhou
et al. 1998) and SLC6A4 (Mossner
et al. 2000), (Rumajogee
et al. 2002). expression(Zhou
et al. 1994;Zhou et al. 1997). |
Decreased
hippocampal protein levels in schizophrenia and Bipolar disorder (Knable
et al. 2004). Serum
levels decreased during depressive and manic episodes (Cunha
et al. 2006). |
BMP6 bone morphogenetic protein 6 GeneCard | Genome-wide association study Baum et al, 2007 | Bone morphogenesis: Also has trophic and protective effects on neurones | ||
NRG1
Neuregulin
1 OMIM
GeneCard |
This gene,
associated with schizophrenia in many studies (Stefansson
et al. 2002;Stefansson et al. 2003;Williams et al. 2003;Yang et al.
2003)has
also been classified as a susceptibility gene in bipolar disorder (Green
et al. 2005). |
Glial
growth factor |
Activates
the PI3K/AKT pathway (Flores
et al. 2000). Reduces
NMDA currents and increases internalization of GRIN1 (Gu
et al. 2005).(Li
et al. 2003). Released
from cultured Schwann cells by BDNF (Esper
and Loeb 2004). |
Decreased
mRNA expression in prefrontal cortex (Tkachev
et al. 2003). Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006)
|
EGFR epidermal growth factor receptor (erythroblastic leukemia viral (v-erb-b) oncogene homolog, GeneCard | WGA Sklar et al, 2008 | EGF receptor | Activates PI3K/AKT pathway | |
v-akt
murine thymoma viral oncogene homolog 1 14q32.32 |
Weak evidence for association in a Bipolar pedigree (Toyota et al. 2003). |
Survival
factor activated via growth factor stimulation of PI3 kinases. Phosphorylates
and inhibits elements of the apoptotic pathway. |
Phosphatidylinositol
3 phosphate (Product of PIK3C3 activates AKT1 (Franke
et al. 1995) BDNF,
CCL2 activate AKT1
(Selzman et al. 2002;Turner
et al. 1997;Lentzsch et al. 2003).
Angiotensin II
(cf AGT) stimulates AKT1 (Griendling
and Ushio-Fukai 2000;Chiu et al. 2004).
Phosphorylation
targets include GSK3B (inactivated) (Shaw
et al. 1997), NMDA
receptors (GRIN1) activated, NOS3
(activated)(Wu
2002) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
BCR breakpoint
cluster region 22q11.23 High
incidence of Bipolar disorder in 22q11 deletion syndrome (Papolos
et al. 1996) |
BCR has been associated with major depression and bipolar disorder (Hashimoto et al. 2005). |
Serine-threonioneKinase
and GTPase activating protein |
Binds
to PLCG1 and the P85 unit of phosphatidylinositol 3 kinase (Muller
et al. 1992). Tyrosine phosphorylated BCR binds to the adapter
molecule growth factor receptor-bound protein 2 (GRB2) (Ma
et al. 1997) which links the EGF receptor tyrosine kinase to Ras activation
and that of its downstream kinases ERK1 and ERK2. GRB2 is involved in
several growth factor signaling pathways including those mediated by
NGF and BDNF (Araki et al. 2000;Yamada et al. 1999;Qian
et al. 1998) |
Stanley
consortium Combined analysis: Unchanged (Higgs et al. 2006) |
FYN FYN oncogene related to SRC, FGR, YES | Polish study Szczepankiewicz et al, 2009 | Associates with GRIN2A, GRIN2B, NTRK2. BDNF potentiates NMDA responses via FYN Xu et al, 2006 | ||
SYK Spleen tyrosine kinase GeneCard | Wellcome Trust Case control Consortium | Important role in leukocyte signaling pathways and in phagocytic cell activation. | BCR signaling Takata et al, 1994) .Coupled to IL2RB Miyazaki and Tanaguchi, 1996 | |
DGKH diacylglycerol kinase, eta GeneCard | Genome-wide association study Baum et al, 2007
|
Output of the PI signalling cascade | ||
DOK5 Docking protein 5 | Associated with amygdala activation in youths with BD Liu et al, 2010 | Interacts with phosphorylated tyrosins kinases including NTRK2 | ||
Dual
specificity phosphatase 6 |
Association reported in a Korean study (Lee et al. 2006a) |
Binds
to and inactivates erk1 (MAPK3)
and erk2 (MAPK1)
(downstream components of growth factor receptor signaling) (Muda
et al. 1998). |
|
Stanley
consortium Combined analysis: Downregulated (Higgs et al. 2006) |
FAT FAT
tumor suppressor homolog 1 4q35 |
Association reported in an Australian case and family study (Blair et al. 2006) |
Cell-cell
adhesion cadherin involved in cell migration Binds to ENAH,
VASP
and MENA |
Binds
to beta catenin (Cox et al. 2000) |
Stanley
consortium Combined analysis: Unchanged (Higgs et al. 2006) |
GAB2 GRB2-associated binding protein 2 GeneCard | Genome-wide association study Baum et al, 2007 | Adaptor protein transmitting growth factor signals to PI3K/ AKT pathway | Phosphorylated by AKT1 Lynch and Daly, 2002 Binds to PLCG2 Mao et al, 2006 | |
Glycogen
synthase kinase 3 beta 3q13.3
? |
A polymorphism in the promoter region of GSK3B influences the age of onset of bipolar disorder. However, its frequency is not different from that in he normal population (Benedetti et al. 2004b;Benedetti et al. 2004a). Assdociation resticted to female patients in a Polish study(Szczepankiewicz et al. 2006) |
Phosphorylates
and inactivates glycogen synthase.Involved in energy metabolism and
neuronal development. Inhibited
by lithium and valproate (Chen et al. 1999;Klein and Melton 1996) |
Phosphorylated
by AKT1 (Takahashi-Yanaga
et al. 2004).HTR2A
receptors decrease GSK3B phosphorylation (Li et
al. 2004). Wnt activation by GSK3B
inhibition is suppressed by myoinositol (Hedgepeth et al. 1997)
the putative product of IMPA2. BDNF increases
-phosphorylation of GSK3B PDGF phosphorylates
GSK3B via activation of PLCG1 (Fang
et al. 2002). Phosphorylates
KCNQ2(Borsotto et al. 2006) |
No
mRNA expression change in frontal or occipital cortex (Agam et
al. 2003). RNA downregulated in dorsolateral prefrontal cortex (Nakatani
et al. 2006). Protein unmodified in prefrontal cortex (Beasley
et al. 2002) Stanley
consortium Combined analysis: Unchanged (Higgs et al. 2006;Lesort
et al. 1999) |
IGF1 Insulin-like growth factor 1 | Pereira et al, 2011 | |||
IMPA2 inositol(myo)-1(or 4)-monophosphatase 2 OMIM GeneCard 18p11.2 Described as a Bipolar disorder locus in several studies (Detera-Wadleigh et al. 1999;Esterling et al. 1997;McInnes et al. 2001;Mors et al. 1997) |
A polymorphism in this gene associates with bipolar disorder in Palestinian and Norwegian populations (Sjoholt et al. 2004b) . (Sjoholt et al. 2004a). |
Inositol
monophosphatases regenerate inositol from inositol monophosphates (the
breakdown productes of inositol triphosphate) and are a target of lithium,
which inhibits magnesium binding to the enzymes (Hallcher
and Sherman 1980).
High concentrations of lithium repress IMPA2 promoter activity
and expression in HeLa cells (Seelan
et al. 2004). |
|
IMPA2
expression is decreased in B lymphoblast cell lines from bipolar patients
and increased in the temporal cortex in male bipolar subjects (Yoon
et al. 2001) Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
KCNQ2
potassium voltage-gated
channel, KQT-like subfamily, member 2 20q13.3 |
Association reported in a French study (Borsotto et al. 2006) |
- |
Activated
by phosphatidylinositol 4,5 biphosphate, product of PIP5K2A (Zhang
et al. 2003). Phosphorylated by GSK3B, dephosphorylated by PPP2R2C (Borsotto
et al. 2006) |
Stanley
consortium Combined analysis: Unchanged (Higgs et al. 2006) |
KIT v-kit Hardy-Zuckerman 4 feline sarcoma viral oncogene homolog | GWAS USA Scott et al, 2009 | Mast cell growth factor, aka stem cell factor receptor | ||
NTRK2 neurotrophic tyrosine kinase, receptor, type 2 | GWAS Smith et al, 2009 | BDNF receptor | ||
PPP2R2C
protein phosphatase 2 (formerly 2A), regulatory subunit B (PR 52), gamma
isoform 4p16.1 |
Association reported in a French study (Borsotto et al. 2006) |
|
Dephosphorylates
and activates KCNQ2 (Borsotto
et al. 2006) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
PIK3C3 phosphoinositide-3-kinase, class 3 OMIM 602609 GeneCard (VPS34) 18q12.3 Described
as a susceptibility locus in a Canadian study (Maziade
et al. 2005) |
A promoter variant of this gene is associated with with bipolar disorder and schizophrenia (Stopkova et al. 2004a). |
Phosphoinositide
kinase. Converts Phosphatidylinositol
to Phosphatidylinositol 3 phosphate (Volinia
et al. 1995).
Nutrient regulated and inhibited by glucose or amino acid deprivation.
Involved in the trafficking of EGF and PDGF receptors (Futter
et al. 2001;Siddhanta et al. 1998) |
Product
metabolised by PIP5K2A |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
PIK4CA
phosphatidylinositol 4-kinase, catalytic, alpha polypeptide 22q11.21. High incidence
of Bipolar disorder in 22q11 deletion syndrome (Papolos
et al. 1996) |
Suggestive association in an American study (Saito et al. 2003) |
Converts
Phosphatidylinositol to Phosphatidylinositol 4 phosphate: Role in EGF
receptor trafficking and degradation (Minogue
et al. 2006). |
- |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
PLCG1 20q12-q13.1 Region
linked to psychotic bipolar disorder (Park
et al. 2004) |
Moderate evidence for association in Canadian and Norwegian patients (Lovlie et al. 2001) |
Catalyzes
the formation of inositol 1,4,5-trisphosphate and diacylglycerol from
phosphatidylinositol 4,5-bisphosphate, one of the products of PIP5K2A. |
Bind to
BCR (Muller
et al. 1992) GNAZ (Bartlett
and Hendry 1997) GRIN2A (Gurd
and Bissoon 1997a),
GRIN2B (Gurd
and Bissoon 1997b)
inhibited by SYNJ1 (Ahn
et al. 1998) DRD1
activates PLCG1
via PKA (Yu et
al. 1996) . The
NRG1 receptor ERBB2 binds to and tyrosine phosphorylates PLCG1 (Peles
et al. 1991). Binds
to GRB2 (Pei
et al. 1997) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
PLCG2 phospholipase C, gamma 2 (phosphatidylinositol-specific) GeneCard | Genome-wide association study Baum et al, 2007 | see above | Binds to GAB2 Mao et al, 2006 | |
PIP5K2A phosphatidylinositol-4-phosphate 5-kinase, type II, alpha OMIM GeneCard 10p12.2 Region
linked to Bipolar disorder in a Genome-wide linkage scan (NIMH) (Cheng
et al. 2006) |
Modest association of polymorphisms with bipolar disorder and schizophrenia (Stopkova et al. 2003) |
Metabolises
the PIK3C3 product Phosphatidylinositol 3 phosphate to phosphatidylinositol
3,4-biphosphate and phosphatidylinositol 3,4,5-triphosphate (Zhang
et al. 1997). Also
metabolises the PIK4CA product phosphatidylinositol-4-phosphate,
and phosphatidylinositol-5-phosphate (Rameh
et al. 1997) (products
of phosphatidylinositol 4- and 5-kinases to phosphatidylinositol-4,5-bisphosphate (Loijens
et al. 1996) and is
thus capable of generating multiple second messengers. |
Substrate
generated by PIK3C3, and PIK4CA Products metabolised by
SYNJ1. PIP5K2A
product PI 4,5
P2, inhibits ADRBK2 (Onorato
et al. 1995) and RGS4 (Ishii
et al. 2005). |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
PTPRE | GWAS Moskvina et al, 2009 | |||
RGS4 regulator of G-protein signalling 4 See Review
of genome scan studies (Levinson
2005) |
Suggestive association in a Brazilian study (Cordeiro et al. 2005) |
GTPase-activating
protein |
GTPase
activating property is inhibited by phosphatidylinositol 3,4,5-trisphosphate,
the product of PIP5K2A (Ishii
et al. 2005). |
Stanley
consortium Combined analysis: Downregulated (Higgs
et al. 2006) |
SHOC2 soc-2 suppressor of clear homolog (C. elegans) GeneCard | Genome-wide association study Baum et al, 2007 | Involved in the transduction of growth factor to MAPK signaling Rodriguez-Viciana et al, 2006 |
||
21q22.2 Region linked to bipolar
disorder (Detera-Wadleigh
et al. 1999;McQuillin et al. 2005) |
Rare mutations
of this gene have been observed in a small number of bipolar patients (Stopkova
et al. 2004b;Saito et al. 2001). |
Multifunctional
enzyme that can remove the 5 terminal phosphates of several phosphoinositides
and inositol polyphosphates including Phosphatidylinositol (4,5) biphosphate,
Phosphatidylinositol (3,4,5) triphosphate and Inositol triphosphate.
It can also convert Phosphatidylinositol (3) P, Phosphatidylinositol
(4)P and Phosphatidylinositol(3,5)P2 into Phosphatidylinositol (Johenning
et al. 2004). It can
thus be considered as the reverse equivalent of PIK3C3,
PIK4CA and PIK5K2A. |
Associates
with and Inhibits PLCG1 (Ahn
et al. 1998). Binds
to GRB2 (McPherson
et al. 1994). |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
TCF4 (E2-2,
ITF2, SEF2, SEF2-1, SEF2-1A, SEF2-1B) 18q12-q21described
as a susceptibility locus in a Canadian study (Maziade
et al. 2005) |
A number of groups have reported association of CAG repeats within the CTG18.1 locus with Bipolar disorder (Del Favero et al. 2002;Jin et al. 2001;Parikh et al. 1999;Lindblad et al. 1998). The gene lying within this locus has the symbol TCF4. |
Transcription
factor |
Activated
by beta catenin CTNNB1 (Kolligs
et al. 2002;Zhai et al. 2002) a
target of GSK3B. Controls TH expression (Yoon
and Chikaraishi 1994) |
Stanley
consortium Combined analysis: Downregulated (Higgs
et al. 2006) |
PDLIM5
PDZ and LIM domain
5: (enigma
homolog; enigma-like LIM domain protein)
OMIM 605904
GeneCard
4q22 |
Associated with bipolar disorder in a Japanese case study (Kato et al. 2005) |
Acts
as a cytoplasmic retention factor for ID2
(inhibitor of DNA binding 2) a growth inhibitory gene that prevents
the action of basic loop helix transcription factors by retaining them
in the cytosol (Lasorella
and Iavarone 2006) |
ID2 is
an inhibitor of basic helix-loop-helix transcription factors, including
TCF4 (Langlands
et al. 1997) |
Decreased
expression in prefrontal cortex and lymphocytes (Iwamoto
et al. 2004) |
VAV3 vav 3 oncogene OMIM GeneCard | Genome-wide association study Baum et al, 2007 | Interacts with tyrosine kinase receptors whose activation phosphorylates VAV3 leading to its association with the receptor Zeng et al, 2000. Phosphatidylinositol 3,4,5-trisphosphate accumulation (product of PIP5K2A) recruits Vav2 and Vav3 which activate RAC1 pathways initiating neurite outgrowth in PC12 cells Aoki et al, 2005 | ||
WDFY2 WD repeat and FYVE domain containing 2 | GWAS Moskvina et al, 2009 | Binds to AKT1 | ||
Genes
whose products affect BDNF expression or release |
Increase:
NMDA receptors (GRIN1, GRIN2A, GRIN2B) (Marini
et al. 1998); DRD1
and DRD2 receptor activation (Fang
et al. 2003) (Kuppers
and Beyer 2001) DRD2 (Takeuchi
et al. 2002), GABA
receptor activation (GABRA1/GABRA5)(Development) (Marty
et al. 1996;Berninger et al. 1995).
The GRM3/GRM5 (Lee
et al. 2006b);
HTR2A receptor
activation (Meller
et al. 2002). HTR2A
receptor stimulation (Vaidya
et al. 1997;Vaidya et al. 1999);
TNF (Meeuwsen
et al. 2003;Bayas et al. 2002)
Inhibit:
IL1B (Barrientos
et al. 2004), GABA
receptor activation (GABRA1, GABRA5) (Adult) (Zafra
et al. 1991). |
Genes
associated with bipolar disorder |
Association
studies |
Primary
role |
Links
to other Bipolar genes |
Expression
changes |
PTPRB protein tyrosine phosphatase, receptor type, B GeneCard | Genome-wide association study Baum et al, 2007 | |||
PTPRE protein tyrosine phosphatase, receptor type, E GeneCard | WGA Wellcome Trust Case Control Consortium | |||
PTPRG protein tyrosine phosphatase, receptor type, G GeneCard | Genome-wide association study Baum et al, 2007 AND WGA Wellcome Trust Case Control Consortium | |||
PTPRN2 protein tyrosine phosphatase, receptor type, N polypeptide 2 GeneCard | Genome-wide association study Baum et al, 2007 |
Table 3: Glutamatergic neurotransmission,
receptors and signaling
Genes
associated with bipolar disorder |
Association
studies |
Primary
role |
Links
to other Bipolar genes |
Expression
changes |
DAOD-amino
acid oxidase OMIM 124050
GeneCard
12q24 Described
as a susceptibility locus in Quebec (Shink
et al. 2005) : 12q23-24
region linked to Bipolar disorder in UK
studies (Glaser
et al. 2005b) |
Suggestive association in Askenazi Jewish case-parent trios (Fallin et al. 2005). Associated in a german case study (Schumacher et al. 2004) |
DAO
is responsible for the catabolism of D-amino acids including D-serine (Nagata
1992), an endogeous
glycine-site ligand for the NMDA receptor (Mothet
et al. 2000). |
DAO catabolises
D-serine (Nagata
1992) a ligand for
the NMDA receptor (Mothet
et al. 2000) (GRIN1,
GRIN2A, GRIN2B) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
D-amino
acid oxidase activatorOMIM 607408
13q34 Susceptibility
locus (Craddock et al. 2005) |
D-amino
acid oxidase (DAO) and an activator protein (DAOA, initially
described as G72) are both associated with schizophrenia (Chumakov
et al. 2002) and DAOA has also been linked with bipolar
disorder (Chen et al. 2004;Addington et al. 2004;Hattori et al. 2003). |
D-amino
acid oxidase |
Activates
DAO |
Stanley
consortium Combined analysis: Unchanged (Higgs et al. 2006) |
GRIA1 glutamate receptor, ionotropic, AMPA 1 GeneCard | Association in an American family-based study Kerner et al, 2008 | Glutamatergic receptor | ||
GRID1
glutamate receptor, ionotropic, delta 1
GeneCard |
Associated
with Bipolar disorder in a study of Ashenazi Case-parent trios (Fallin
et al. 2005) |
Glutamate
receptor |
- |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
GRIK4 Kainate receptor |
Association
reported for both schizophrenia and bipolar disorder in a scottish case
study (Pickard
et al. 2006) |
High-affinity
Kainate receptor |
- |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
GRIK5 glutamate receptor, ionotropic, kainate 5 | Spanish study Gratacos et al, 2009 | |||
GRIN1(NMDA receptor subunit NR1) Region
linked to early onset bipolar disorder (Faraone
et al. 2006) |
Polymorphisms
in the GRIN1 promoter (Mundo
et al. 2003)
have been associated with bipolar
disorder. |
|
Trophic
concentrations of NMDA stimulate the PI3kinase/AKT
signaling cascade (Zhu et al. 2002;Lafon-Cazal et al. 2002), (Perkinton
et al. 2002) (Manabe and Lipton 2003)
and Toxic
concentrations of NMDA phosphorylate and inactivate AKT1
and decrease the phosphorylation of GSK3B (Luo
et al. 2003); Binds to DRD1 (Fiorentini et al. 2003) Favaron
et al. 1993). Oligodendrocytes express GRIN1 and activation of
NMDA receptors in these cells increases their expression of polysialylated
NCAM1 that is required for oligodendrocyte migration (Wang
et al. 1996). Expression regulated by BDNF (Schratt
et al. 2004) |
RNA downregulated
in dorsolateral prefrontal cortex (Nakatani
et al. 2006). No change
in protein expression on orbitofrontal cortex or dentate hilus.(Toro
and Deakin 2005) Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
GRIN2A NMDA receptor subunit NR2A |
Polymorphisms in the GRIN2A promoter (Itokawa et al. 2003) have been associated with bipolar disorder. |
Glutamate |
PLCG1
is directly coupled to GRIN2A (Gurd
and Bissoon 1997a).
Expresion regulated by BDNF (Margottil
and Domenici 2003). |
No change
in anterior cingulate cortex. (Woo
et al. 2004). Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
GRIN2B NMDA receptor subunit NR2B |
(Martucci et al. 2006) |
|
Binds
to PLCG1 (Gurd
and Bissoon 1997b);
BDNF activates
NMDA receptors via a GRIN2B
dependent mechanism (Levine
and Kolb 2000) |
No change
in expression (Martucci
et al. 2006) Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
CAMK2D calcium/calmodulin-dependent protein kinase (CaM kinase) II delta OMIM GeneCard | Genome-wide association study Baum et al, 2007 | Involved in glutamate-related synaptic plasticity | Binds to GRIN2B Robison et al, 2005. Regulates BDNF expression Kamata et al, 2006. Controls circadian elements via PER1 Nomura et al, 2003 | |
CAMKK2 calcium/calmodulin-dependent protein kinase kinase 2, beta | Canadian study Barden et al, 2006 | |||
CIT 12q24 |
Association reported in an American study (Lyons-Warren et al. 2005) |
Plays
a role in cytokinesis and in the control of dendritic morphology (Di
Cunto et al. 2003) |
Binds
to GRIN1 and is a component of the NMDA receptor adhesion complex (Husi
et al. 2000). Binds
to DISC1 (Ozeki
et al. 2003) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
DFNB31 deafness, autosomal recessive 31 GeneCard | Genome-wide association study Baum et al, 2007 GWAS Moskvina et al, 2009 | PDZ domain molecule that may be involved in glutamte receptor scaffolding | Binds to CASK a protein that associates with GRIP1 and RGS4 Hong and Hsueh, 2006 | |
Disrupted in schizophrenia 1 1q42.1 1q42 described
a s a susceptibility locus in a Scottish study (Macgregor
et al. 2004) |
Association reported in an American study (Hodgkinson et al. 2004) and an American family study (Maeda et al. 2006). |
Involved
in neurite outgrowth and cortical development. |
Binds
to CIT (Ozeki et al. 2003) , MLC1 (Millar et
al. 2003) and DPYSL2 (Camargo
et al, 2006). Binding partner ATF4 is involved in endoplasmic
reticulum stress and controls GCH1 transcription (Kapatos
et al. 2000). Binds to elements of the glutamate receptor scaffold (see (Carter
2006)) and TENC1 (Millar
et al. 2003)a negative regulator of PI3K/AKT signalling (Hafizi
et al. 2005). Binding partner MGAT3
is involved in N-glycans biosythesis, as is ALG9 |
Decreased
mRNA expression in lymphoblasts; Correlated with manic symptomatology (Maeda
et al. 2006) . Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
Weakly Associated in a study of Ashkenazi case-parent trios (Fallin et al. 2005) |
Nitric oxide synthesis |
Neuronal nitric oxide
synthase stimulated by NMDA receptor activation |
Increased
expression in hippocampus (Benes
et al. 2006) |
|
NOSIAP (CAPON) 1q23.3 Region linked to bipolar
disorder (Levinson
2005) |
Suggested implication based on the identification of a variant with increased expression (Xu et al. 2005a) |
Nitric oxide synthesis |
Binds
to NOS1 |
Increased
expression in both schizophrenia and bipolar disorder (dorsolateral
prefrontal cortex) correlated with genotype (Xu
et al. 2005a). Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
Endothelial nitric
oxide synthase 7q36 Region
linked to Bipolar disorder in a European study (Etain
et al. 2006) |
Associated
with bipolar disorder in a german study (Reif
et al. 2005). |
Endothelial Nitric
oxide synthase |
Phosphorylated
by AKT1 (Wu
2002).
Nitric oxide
affects AKT1, SLC6A3, SLC6A4, GRIN2A, GRIN2B, HTR2A and TH
function (Yasukawa
et al. 2005) (Nozik-Grayck
et al. 2002) (Choi
et al. 2000).(Kuhn
and Geddes 2003;Park et al. 2002)(Bryan-Lluka
et al. 2004;Mishra et al. 2002).
NOS3 knockout decreases cerebral BDNF expression (Chen
et al. 2005) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
GRM1 glutamate receptor, metabotropic 1 OMIM GeneCard | Genome-wide association study Baum et al, 2007 | Linked to inositol triphosphate receptor ITPR1 via HOMER1 Tu et al, 1998 | ||
GRM3 Metabotropic glutamate receptor mgluR3 OMIM 601115 GeneCard
7q21.1-q21.2 Region
linked to Bipolar disorder in a Genome-wide linkage scan (NIMH) (Cheng
et al. 2006) |
Association reported in a case-parent trio study in Askemazi Jews (Fallin et al. 2005) |
- |
MGlur2/3
antagonism suppresses stress-induced increases in frontal cortex BDNF
expresion (Lee
et al. 2006b) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
GRM4 Metabotropic glutamate receptor mgluR4 |
Association
reported in a case-parent trio study in Askemazi Jews (Fallin
et al. 2005) |
Metabotropic
glutamate receptor |
- |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
GRM7 glutamate receptor, metabotropic 7 GeneCard | Wellcome Trust Case control Consortium | Knockout increases hippocampal BDNF levels Mitsukawa et al, 2006 | ||
PCLO Piccolo | Kleinman, 2011 | |||
SNAP25 synaptosomal-associated protein, 25kDa | French study Etain et al, 2009 | |||
SYN3 Synapsin III GeneCard | Wellcome Trust Case control Consortium | Glutamate release | Binds to NOS1AP | |
VAPA VAMP (vesicle-associated membrane protein)-associated protein A, 33kDa | US study Lohoff et al, 2008 |
Genes
associated with bipolar disorder |
Association
studies |
Primary
role |
Links
to other Bipolar genes |
Expression
changes |
CCL2 17q11.2-q12 |
A polymorphism in this gene was shown to influence the symptomatology of bipolar disorder in a Korean population (Pae et al. 2004a). |
Chemokine |
Stimulates
the PI3K/AKT pathway (Jones
et al. 2003) (Selzman
et al. 2002). TNF
upregulates CCL2 expression (Murao
et al. 2000) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
CMTM8 CKLF-like MARVEL transmembrane domain containing 8 GeneCard | Wellcome Trust Case control Consortium | Chemokine-like. Regulates EGF receptor signaling Jin et al, 2005 | ||
CSF2RB colony stimulating factor 2 receptor, beta, low-affinity (granulocyte-macrophage) GeneCard | Wellcome Trust Case control Consortium | Beta chain of the high affinity receptor for IL-3, IL-5 and CSF. | ||
EDA ectodysplasin A OMIM GeneCard | Genome-wide association study Baum et al, 2007 | TNF family related cytokine | - | - |
IFNG Interferon gamma | Italian study Clerici et al, 2009 | |||
IL1B Interleukin 1B OMIM 147720 GeneCard 2q14 |
Association reported in a Spanish study (Papiol et al. 2004) |
Cytokine Activates
PKR (EIF2AK2)
Williams,
2001 |
BDNF
stimulates IL1B secretion from macrophages (Asami
et al. 2006). IL1B
suppresses the learning-associated increases in hippocampal BDNF
expression (Barrientos
et al. 2004) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
IL1RN
Interleukin
1 receptor antagonist |
Association reported in a Spanish study (Papiol et al. 2004) |
IL1B inhibitor |
Social
isolation decreases hippocampal BDNF mRNA, an effect blocked
by IL1RN infusion (Barrientos
et al. 2003) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
IL2RB 22q13.1 Region
linked to bipolar disorder (see review of genome scans (Levinson
2005)) |
Association reported in a case-parent trio study in Askenazi Jews (Fallin et al. 2005) |
IL2 receptor |
IL-2 Activates the PI3K pathway in T cells and myelin (Benczik and Gaffen 2004). Associated with phosphoinosite kinase PIK3R1 (Migone et al. 1998;Chakraborty et al. 2003) |
Increased
levels of serum IL2 receptors often reported in bipolar disorder (Tsai
et al. 2003). Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
IL6 Interleukin 6 | Modifies age of onset in an Italian study Clerici et al, 2009 | |||
IL10 Interleukin 10 | Italian study Clerici et al, 2009 | |||
TNFA:
Tumor necrosis factor alpha OMIM 191160 6p21.3 Folate-sensitive
fragile site(Tastemir
|
A polymorphism in this gene has been associated with both bipolar disorder and major depression in a Korean population (Pae et al. 2004b;Jun et al. 2003) |
Cytokine |
Stimulates
the PI3K/AKT pathway (Marchetti
et al. 2004). Inhibits
AKT1 activity by increasing the production of C2 ceramide
which maintains AKT1 in a dephosphorylated active state (Teruel
et al. 2001).
Stimulates the expression of BDNF and CCL2
in human astrocytes (Meeuwsen
et al. 2003) and of
GCH1 in C6 glioma cells (D'Sa
et al. 1996).Both
TNF and IL1B stimulate the activity of the brain serotonin
transporter SLC6A4 via p38 mitogen-activated protein kinase
activation (Zhu
et al. 2006) |
Increased
expression in hippocampus (Benes
et al. 2006) Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
Table
4: Downstream targets of BDNF or PI3K/AKT signaling
Genes
associated with bipolar disorder |
Association
studies |
Primary
role |
Links
to other Bipolar genes |
Expression
changes |
GTP cyclohydrolase
1 (dopa-responsive dystonia) 14q22.1-q22.2 Region
linked to Bipolar disorder (Kealey
et al. 2005) |
Associated
with bipolar disorder in an Irish study (Kealey
et al. 2005). |
Tetrahydrobiopterin
synthesis |
Stimulated
by growth factors via the PI3 kinase pathway (Bauer
et al. 2002). Tetrahydrobiopterin
is a cofactor for TH TPH and NOS3: regulated by TNFA (D'Sa
et al. 1996)and ATF4(Kapatos
et al. 2000), binding
partner of DISC1(Stegenga
et al. 1996;Hirayama and Kapatos 1995) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
glucose-6-phosphate
dehydrogenase Xq28 OMIM MAFD2 |
Association reported in Italian studies (Bocchetta 2003;Bocchetta et al. 1999). |
D-glucose
6-phosphate + NADP+ = D-glucono-1,5-lactone 6-phosphate + The NADPH
so formed plays and important role in the maintenance of reduced glutathione (Fujii
1995). |
Expression
stimulated by growth factors (Stanton
et al. 1991). PDGF results
in the rapid release of G6PD that is attenuated by PI3K
or PLCG1 inhibition (Tian
et al. 1994).
G6PD expression
is controlled by sterol regulatory element binding protein
SREBp1a (Amemiya-Kudo
et al. 2002) whose
transcription is in turn controlled by AKT1 (Porstmann
et al. 2005). |
G6PD
deficiency has been associated with Bipolar disorder (Bocchetta
2003;Bocchetta et al. 1999) Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
SLC25A4
(ANT1) Adenine
nucleotide translocator 4q35 Identified
as a susceptibility locus in American (NIMH) and Australian studies (Willour
et al. 2003;Blair et al. 2002) |
An ANT1
mutation associated with Bipolar disorder has been described in a family
with dominantly inherited progressive external ophthalmoplegia (Siciliano
et al. 2003). |
Mitochondrial
Adenine nucleotide translocator Binds
to BAX and modifies the mitochondrial |
Activated
by serotonin (HTR2B)
receptor stimulation via the PI3K/AKT pathway (Nebigil
et al. 2003). Facilitates
glutamate transport (Buck
et al. 2003) |
Stanley
consortium Combined analysis: Downregulated (Higgs
et al. 2006) |
Somatostatin 3q28 ? |
Weak association
reported in a Japanese study (Nakatani
et al. 2006) |
- |
Activates
SSTR5. Expression synthesis or release promoted by BDNF
in hypothalamic, cortical and hippocampal neurones (Givalois
et al. 2006;Villuendas et al. 2001;Marty and Onteniente 1999).
In SH-SY5Y human neuroblastoma
cells, which express all SSTR isoforms, somatostatin inhibits erk1/2
and RAC activity and indirectly suppresses the PDGF-activated PI3 kinase
pathway via inhibition of RAC, a downstream target of the PI3 kinase (Pola
et al. 2003).
Somatostatin
is generally antiproliferative. |
RNA downregulated
in dorsolateral prefrontal cortex (Nakatani
et al. 2006) Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006). |
Somatostatin
receptor 5 16p13.3 |
Polymorphisms
in the gene coding for this receptor are associated with bipolar disorder (Nyegaard
et al. 2002). |
Growth
suppressive |
Forms
heterodimers with DRD2 (Rocheville
et al. 2000) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
DPYSL2dihydropyrimidinase-like
2 OMIM 602463
GeneCard Bipolar locus (see
meta-analysis review) (Levinson
2005) |
Weakly Associated in a study of Ashkenazi case-parent trios (Fallin et al. 2005) |
Involved in axonal
outgrowth. |
Phosphorylated
by GSK3B (Yoshimura
et al. 2005). |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
Table 5: Genes
whose products activate the PI3K/AKT1 pathway
Genes
associated with bipolar disorder |
Association
studies |
Primary
role |
Links
to other Bipolar genes |
Expression
changes |
ALOX12 17p13.1 Region
linked to bipolar disorder (see review of genome scans (Levinson
2005)) |
An ALOX12
polymorphism associates with Bipolar disorder in a Brazilian population (Fridman
et al. 2003). |
Arachidonate
metabolism Arachidonate
+ O2 -> 12(S)HPETE |
12(S)
HETE activates the AKT1 pathway in epidermal cancer
cells (Szekeres
et al. 2002) and in
prostate cancer cells (Pidgeon
et al. 2002) and is
also linked to stimulation of PI3 kinase (Szekeres
et al. 2000;Szekeres et al. 2002).
Stimulated by AGT) (Zhu
et al. 2000),NMDA
receptor activation (GRIN1, GRIN2A) and noradrenaline (Wolfe
et al. 1990). ALOX12
inhibition prevents oligodendrocyte cell death induced by glutathione
inhibition (Wang
et al. 2004). |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
AGT 1q42-q43
GeneCard
1q42 identified as a susceptibility locus in a Scottish study (Macgregor
et al. 2004) |
Associated with Bipolar disorder in a Brazilian population (Meira-Lima et al. 2000). |
Cleaved
by renin to Angiotensin I which is cleaved by ACE to Angiotensin II |
Angiotensin
stimulates AKT1 activity and promotes neurite extension
in hypothalamic neurones (Yang
et al. 2002). PLCG1
is stimulated by angiotensin II (AGT) in smooth muscle cells (Marrero
et al. 1994). Stimulates
TH expression (Kim
et al. 1996). |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
Apolipoprotein E OMIM 107741 GeneCard 19q13.2 Region
linked to bipolar disorder (see review of genome scans (Levinson
2005)) |
Early onset bipolar disorder patients possess an increased frequency of the APOE4 allele (Bellivier et al. 1997). |
Cholesterol
transport |
APOE
activates the PI3K/AKT pathway in neuro2A cells (Laffont
et al. 2002). APOE
3 and APOE4 also acutely activate GSK3B
in human SH-SY5Y neuroblastoma cells and APOE4 subsequently
inactivates GSK3B. APOE4 also stimulated
AKT1 activity in these cells (Cedazo-Minguez
et al. 2003). Apolipoprotein
E4 also inhibits phosphatidylinositol 4-kinase (PI4K) and phosphatidylinositol
4-phosphate 5-kinase (PIP5K) activity in the rat cerebral cortex (Zambrzycka
and Kacprzak 2003).
The isoforms were not specified. |
Increased
protein levels in the caudate putamen and brodmann area 9 , decreased
levels in Brodmann Area 10.(Digney
et al. 2005). Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
CALCA |
A polymorphism in this gene has been associated with bipolar disorder in a Swedish study (Buervenich et al. 2001). |
|
Stimulates
PI3K/AKT signaling (Parameswaran
et al. 2000) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
CHRNA7 Nicotinic receptor alpha 7 subunit OMIM 118511 GeneCard |
Association reported in a Taiwanese study (Hong et al. 2004) |
Nicotinic
receptor |
Binds
to the P85 subunit of phosphatidylinositol 3-kinase (PIK3R1). Nicotine
stimulation results in AKT1 phosphorylation (Kihara
et al. 2001). |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
DTNBP1 Dysbindin 6p22.3 |
Associated with bipolar disorder in subset of psychotic patients (Raybould et al. 2005). Weakly Associated in a study of Ashkenazi case-parent trios (Fallin et al. 2005) |
Overexpression
reduces glutamate release (Numakawa
et al. 2004) |
Dysbindin
overexpression in primary cortical neurones increases AKT1 phosphorylation
and protects neurones via stimulation of the PI3K/AKT cascade (Numakawa
et al. 2004) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
P2RX7
purinergic receptor P2X, ligand-gated ion channel, 7 Described
as a susceptibility locus in Quebec (Shink
et al. 2005) : 12q23-24
region linked to Bipolar disorder in UK
studies (Glaser
et al. 2005b) |
Association suggested in a Canadian Case study (Barden et al. 2006) |
Purinergic
receptor forming a pore/channel. |
Phosphorylated
by ADRBK2 (Feng
et al. 2005). Stimulates
AKT phosphorylation in astrocytes via diverse routes (Jacques-Silva
et al. 2004). Astrocytic
receptors control glutamate release(Duan
et al. 2003). IL1B
increases astrocytic P2RX7 expression and P2RX7 stimulation increases
IL1B release (Narcisse
et al. 2005) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
Genes
associated with bipolar disorder |
Association
studies |
Primary
role |
Links
to other Bipolar genes |
Expression
changes |
ASTN2 Astrotactin GeneCard | Genome-wide association study Baum et al, 2007 | Astrotactin may guide migrating neurones along glial paths Zheng et al, 1996 | - | - |
CDH12 cadherin 12, type 2 (N-cadherin 2) | UK study Hamshere et al, 2009 | |||
CDH20 cadherin 20, type 2 OMIM GeneCard | Genome-wide association study Baum et al, 2007 | Cadherins are calcium-dependent adhesive proteins that mediate cell-to-cell interaction | - | - |
CNTNAP2 contactin associated protein-like 2 | Irish study O'Dushlaine et al, 2010 | |||
CNTN5 contactin 5 GeneCard | Genome-wide association study Baum et al, 2007 | Glycosylphosphatidylinositol -anchored neuronal membrane protein: Acts as a cell adhesion molecule | ||
CNTN6 Contactin 6 GeneCard | Genome-wide association study Baum et al, 2007 | see above | ||
CNTNAP5 contactin associated protein-like 5 GeneCard | Genome-wide association study Baum et al, 2007 | Neurexin family member. Neurexins act as cell adhesion molecules and receptors. | - | - |
DOCK3 dedicator of cytokinesis 3 GeneCard | Genome-wide association study Baum et al, 2007 | Regulates N-cadherin nediated cell adhesion Chen et al, 2005 | - | - |
DOCK9 Dedicator of cytokinesis 9 GeneCard | Association described in an American family study Detera-Wadleigh et al, 2007 | Binds to the rho GTPase cdc42 Meller et al, 2002 | ||
DSCAM Down syndrome cell adhesion molecule GeneCard | Association reported in American and Japanese population Amano et al, 2008 | Involved in neuronal self-avoidance Gao, 2007 | ||
JAM3 junctional adhesion molecule 3 OMIM GeneCard | Genome-wide association study Baum et al, 2007 | Localised in tight junctions in epithelial cells | - | - |
LAMA3 laminin, alpha 3 GeneCard | Genome-wide association study Baum et al, 2007 | Basement membrane component | Binds to heparin and promotes cell adhesion via syndecan SDC2 Utani et al, 2001 | |
NCAM1 11q23.1
? |
An NCAM1 polymorphism is associated with Bipolar disorder in the Japanese population (Arai et al. 2004). |
NCAM1
is a neural adhesion molecule and is its own neuronal receptor and substrate
ligand. NCAM1/NCAM1 binding stimulates
axonal growth and neuritic sprouting. This effect is mediated via interaction
with tyrosine kinase receptors, which are activated by NCAM1
mediated adhesion. |
Stimulates
PI3K/AKT signaling via FGFR1 interactions (Kiselyov
et al. 2003). TrkB
phosphorylation and BDNF signaling is impaired in NCAM knockout
mice. Polysialylated-NCAM interacts with BDNF (Vutskits
et al. 2001). Oligodendrocytes
express GRIN1 and activation of NMDA receptors in these cells
increases their expression of polysialylated NCAM1 that is required
for oligodendrocyte migration (Wang
et al. 1996). |
Increased protein expression
of a splice variant (NCAM-VASE) in the hippocampus (Vawter
et al. 1998). Stanley
consortium Combined analysis: Downregulated (Higgs
et al. 2006) |
NRXN1 Neurexin 1 | Irish study O'Dushlaine et al, 2010 | |||
SDC2 Syndecan 2 (heparan sulfate proteoglycan ) GeneCard | Genome-wide association study Baum et al, 2007 | Review Essner et al, 2005 | - | - |
Table 6: Genes involved in vesicular
or protein traffic, glycosylation and endoplasmic reticulum stress associated
genes
Genes
associated with bipolar disorder |
Association
studies |
Primary
role |
Links
to other Bipolar genes |
Expression
changes |
ALG9 |
Disrupted by a translocation breakpoint segregating with Bipolar disorder in a small family (Baysal et al. 2002) |
A deficiency
of ALG9 causes an accumulation of lipid-linked-GlcNAc
(2) Man (6) and -GlcNAc(2)Man(8) containing structures and defects in
N-glycosylation (Frank
et al. 2004b).(GlcNac
= N-acetylglucoseamine; Man = mannose) This
enzyme is involved in glycosylation, a post-translational protein modification
involving a multitude of targets. The N-linked
glycosylation pathway necessitates the assembly of an oligosaccharide
core with a dolichylpyrophosphate lipid carrier and the transfer of
this assembly to nascent polypeptide chains. ALG9 is
involved in this assembly (Frank
et al. 2004b). |
Glycosylation
is involved in protein folding and processing. Defects in type 1 glycosylation
are associated with increased endoplasmic reticulum stress(Lecca
et al. 2005;Freeze 2002;Shang et al. 2002).
Deletion in yeast
results in the hypoglycosylation of secreted proteins (Burda
et al. 1996). |
- |
FKBP5 FK506 binding protein 5 | Family-based association Willour et al, 2008. | Involved in protein sorting/folding immunomodulation and glucocorticoid receptor regulation | ||
GALNTL4 UDP-N-acetyl-alpha-D-galactosamine:polypeptide N-acetylgalactosaminyltransferase-like 4 GeneCard | Genome-wide association study Baum et al, 2007 | |||
GASP GPRASP1 G protein-coupled receptor associated sorting protein 1 GeneCard | Genome-wide association study Baum et al, 2007 | GPCR soring including dopamine receptor DRD2 Bartlett et al, 2005 | - | - |
HIP1R 12q24 12q23-24
region linked to Bipolar disorder in UK
studies (Glaser
et al. 2005b) |
This gene was isolated from Bipolar pedigrees showing decreased age of onset with successive generations (Provencal et al. 2004). |
Actin
and phospholipid binding protein: Plays a role in growth factor receptor
trafficking (Hyun
et al. 2004). |
Binds
to phosphatidylinositol 3,4-bisphosphate the product of PIP5K2A
and phosphatidylinositol 3,5-bisphosphate (Hyun
et al. 2004) a product
of PIP5K3 (Ikonomov
et al. 2002). Binds
to dynein light chain (DYNLL1)
a protein involved in intracellular transport. DYNLL1 also binds to
and inhibits the activity of NOS1 (Jaffrey
and Snyder 1996)and
to the BDNF receptor NTRK2 (Yano
et al. 2001) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
HSPA4 heat shock 70kDa protein 4 | Korean study Pae et al, 2009 | |||
HSPA5 9q33-q34.1 9q34 described
as a locus for early onset Bipolar disorder (Faraone
et al. 2006) |
Association observed in Japanese case and family study (Kakiuchi et al. 2005) |
Folding
and assembly of proteins in the endoplasmic reticulum. |
Binds
to three key proteins involved in ER stress response.
(PERK (Gene symbol = EIF2AK3),
IRE1 alpha (gene symbol ERN1)
and the transcription factor ATF6)
and its saturation by unfolded proteins releases these stress response
activators (Xu
et al. 2005b).
ATF6 controls the expression of XBP1 which is spliced
by IRE1 alpha to produce a highly active transcription factor (Yoshida
et al. 2001;Lee et al. 2002) |
No change
in protein levels in temporal cortex (Bown
et al. 2000). Stanley
consortium Combined analysis: Downregulated (Higgs
et al. 2006) |
HSP90B1 heat shock protein 90kDa beta (Grp94), member 1 12q24.2-q24.3 GeneCard | Association in a Japanese case control and american bipolar family trio set Kakiuchi et al, 2007 | Molecular chaperone involved in endoplasmic reticulum stress Little at al, 1994 | ||
KIF13A
kinesin family member 13A 6p23 ? |
Weakly
Associated in a study of Ashkenazi case-parent
trios (Fallin
et al. 2005) |
Motor protein transporting
the mannose-6-phosphate receptor (Nakagawa
et al. 2000) |
- |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
NAPG
N-ethylmaleimide-sensitive factor attachment protein, gamma 18p11.22
. |
Association reported in an American case study (Weller et al. 2006) |
Play a
role in intracellular membrane fusion and vesicular trafficking |
|
Stanley
consortium Combined analysis: Downregulated (Higgs
et al. 2006) |
SNX27 sorting nexin family member 27 GeneCard | Genome-wide association study Baum et al, 2007 | Involved in endocytosis of plasma membrane receptors and protein trafficking | Involved in HTR4 serotonin receptor sorting in mice Joubert et al, 2004 | |
SORCS2 sortilin-related VPS10 domain containing receptor 2 OMIM GeneCard | Genome-wide association study Baum et al, 2007 | Vacuolar protein sorting (by homology) | ||
SYBL1
Synaptobrevin-like 1 (VAMP7, VAMP-7, TI-VAMP) Xq28 and
Yq12 OMIM MAFD2 |
Association reported in a German population and in male Americans (Muller et al. 2002;Saito et al. 2000) |
In PC12
cells SYBL1 mediates the vesicular transport from endosomes
to lysosomes and thus assists in the breakdown of endocytosed protein
products. An antibody to SYBL1 inhibits the breakdown
of EGF (Advani
et al. 1999) in PC12
cells. SYBL1 is thus likely to participate in the breakdown
of a large number of endocytosed macromolecules including other growth
factors that are processed in this way. SYBL1 also plays a role
in neurite and dendrite outgrowth in PC12
cells and hippocampal neurones (Martinez-Arca
et al. 2000, 2001) |
- - |
RNA upregulated
in dorsolateral prefrontal cortex (Nakatani
et al. 2006). Stanley
consortium Combined analysis: Downregulated (Higgs
et al. 2006) |
WFS1 4p16 |
Polymorphisms in this gene have been associated with both bipolar disorder and major depression (Koido et al. 2004;Furlong et al. 1999). |
- |
Endoplasmic
reticulum cation channel activated by inositol 1,4,5 triphosphate, the
product of PLCG1 (Osman
et al. 2003). Transcription
controlled by XBP1 (Kakiuchi
et al. 2006). |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
XBP1 X-box binding protein 1. OMIM 194355 GeneCard 22q12 OMIM MAFD1 |
Association reported in Japan and Taiwan (Hou et al. 2004;Kakiuchi et al. 2003) |
ER
stress |
Regulates WFS1 transcription (Kakiuchi et al. 2006). Controls expression of DNAJC3 (p58IPK), DNAJB9 (ERDJ4), DNAJB11 (HEDJ), EDEM.protein disulfide isomerase-P5 and SERP1 (RAMP4) (Lee at al, 2003). |
RNA upregulated
in dorsolateral prefrontal cortex (Nakatani
et al. 2006). In lymphoblastoid
cells, XBP1-dependent transcription activity of the Bipolar associated
allele was reduced and in the cells with this allele, induction of XBP1
expression after ER stress was markedly reduced. Valproate, one
of three mood stabilizers, rescued the impaired response by inducing
ATF6, the controller of XBP1(Kakiuchi
et al. 2003). Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
Also involved
in traffic: AKT1 (receptor endocytosis (Clague
and Urbe 2001)), PIK3C3 (EGF
and PDGF receptors) (Futter
et al. 2001); PIK4CA
(EGF receptors (Minogue
et al. 2006); SYNJ1 (synaptic
vesicle recycling and endocytosis (Cremona
et al. 1999;Singer-Kruger et al. 1998).
Table
7: Circadian genes
CLOCK
is a transcription factor involved in the control of circadian rhythms. It forms
a heterodimer with ARNTL (aryl hydrocarbon receptor nuclear translocator-like,
also known as BMAL-1), which together control the expression of the period gene
(PER1) (Gekakis et al.
1998).
Period and TIMELESS associate with each other and translocate
into the nucleus where they switch off the CLOCK-ARNTL
mediated transcription of the period gene (Sangoram
et al. 1998). The association
between period and timeless is controlled by phosphorylation of the two proteins
by casein kinase 1 epsilon (CSNK1E) that phosphorylates period and GSK3B,
which phosphorylates timeless (Harms
et al. 2003;Martinek et al. 2001).
CSNKIE also phosphorylates cryptochromes 1 and 2 (CRY1, CRY2) when they are
associated with period and also phosphorylates and activates ARNTL (Eide
et al. 2002)
Genes
associated with bipolar disorder |
Association
studies |
Primary
role |
Links
to other Bipolar genes |
Expression
changes |
ARNTL
11p15.
Identified as a linkage region in an American study (NIMH) (McInnis
et al. 2003) |
Association reported in two American studies (Nievergelt et al. 2006) (Mansour et al. 2006). |
Circadian
rhythm |
Forms
heterodimer with CLOCK. |
RNA upregulated
in dorsolateral prefrontal cortex (Nakatani
et al. 2006). Stanley
consortium Combined analysis: Downregulated (Higgs
et al. 2006) |
BHLHB2 basic helix-loop-helix domain containing, class B, 2 GeneCard | Associaton reported in an American family study Shi et al 2008 | represses Clock/ARBTL-induced transactivation of the mouse Per1 promoter through direct protein-protein interactions with Bmal1 and/or competition for E-box elements. | ||
CRY2 Cryptochrome 2 | Sjoholm et al, 2010 | |||
CSNK1E casein kinase 1, epsilon GeneCard | Association in an American study Shi et al 2008 | Phosphorylates ARNTL Eide et al 2002 | ||
NPAS2 neuronal PAS domain protein 2 | USA Mansour et al, 2009 | |||
PER3 1p36.23 |
Association
reported in an American study (Nievergelt
et al. 2006). |
Circadian
rhythm |
- |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
4q12 Region
linked toBipolar disorder in UK Irish Sib-pair Genome screen (Lambert
et al. 2005) |
CLOCK has been associated with the recurrence of Bipolar depression in one study (Benedetti et al. 2003) see also Shi et al 2008 |
Circadian
rhythm |
Forms
heterodimer with ARNTL. GSK3B
controls elements in the circadian cycle (Harms
et al. 2003) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
GPR50
Orphan receptor Xq28 OMIM MAFD2 |
This receptor
has been associated with Bipolar disorder and major depression particularly
in women (Thomson
et al. 2004). |
Dimerises
with melatonin receptors MT1 and MT2 and inhibits activity of MT1 (Levoye
et al. 2006) |
Melatonin
plays a key role in circadian control. Activation of the AKT
pathway has been implicated in the neuroprotective effects of melatonin
in stroke models (Kilic
et al. 2005) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
NR1D1 nuclear receptor subfamily 1, group D, member 1 (Rev-ErbAalpha) | Sardinian study Severino et al, 2009 | |||
RFX4
regulatory factor X, 4 12q24 12q23-24
region linked to Bipolar disorder in UK
studies (Glaser
et al. 2005b) |
Association reported in a British study (Glaser et al. 2005a) |
Associated
with two other similar transcription factors RFX2 and RFX3: Light-induced
and localised in the suprachiasmatic nucleus (Araki
et al. 2004). Involved
in cortical development in mice (Zarbalis
et al. 2004). |
- |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
? |
Weak association reported in an American study (Mansour et al. 2006) |
- |
Phosphorylated
by GSK3B (Iitaka
et al. 2005) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
Table 8: Mitochondrial
and energy-related genes
Genes
associated with bipolar disorder |
Association
studies |
Primary
role |
Links
to other Bipolar genes |
Expression
changes |
MTND1 ND4 NADH dehydrogenase subunit 4 OMIM OMIM GeneCard Mitochondrial
DNA |
Mutation
leading to reduced activity observed in a Japanese population -(Munakata
et al. 2004) |
Subunit
of Mitochondrial complex 1 |
- |
- |
ND4L mitochondrially encoded NADH 4L | Rollins et al, 2009 | |||
NDUFAB1 NADH dehydrogenase (ubiquinone) 1, alpha/beta subcomplex, 1, 8kDa | GWAS WTCCC | |||
NDUFV2 18p11.31-p11.2 Described
as a Bipolar disorder locus in several studies (Detera-Wadleigh
et al. 1999;Esterling et al. 1997;McInnes et al. 2001;Mors et al. 1997) |
NDUFV2 polymorphisms are associated with Bipolar disorder and schizophrenia in Japanese and American populations (Washizuka et al. 2004;Washizuka et al. 2003) |
Subunit
of Mitochondrial complex 1 |
- |
Increased
mRNA expression in DLPFC (Nakatani
et al. 2006). Decreasesd
expression in lymphoblastoid cell lines isolated from bipolar disorder
patients (Washizuka
et al. 2005). Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
Table 9: Channels
and transporters
Genes
associated with bipolar disorder |
Association
studies |
Primary
role
|
Links
to other Bipolar genes |
Expression
changes |
ABCA6 ATP-binding cassette, sub-family A (ABC1), member 6 GeneCard | Genome-wide association study Baum et al, 2007 | Putative role in lipid cholesterol-related transport | - | - |
ATP-binding cassette, sub-family B (MDR/TAP), member 1 |
Turkish study Turgut et al, 2009 | |||
MDR1/ABCB1 P-glycoprotein 1 OMIM GeneCards | Turkish study Turgut et al, 2009 | Drug transporter Involved in NO-related glutathione efflux Watts et al, 2006 | ||
ABCB11 ATP-binding cassette, sub-family B (MDR/TAP), member 11 GeneCard | Genome-wide association study Baum et al, 2007 | Major canalicular bile salt export pump in man. | - | - |
ATP1A1 ATPase, Na+/K+ transporting, alpha 1 polypeptide | Goldstein et al, 2009 | |||
ATP1A2 ATPase, Na+/K+ transporting, alpha 2 polypeptide | Goldstein et al, 2009 | |||
ATP1A3 19q13.31 Region
linked to bipolar disorder (see review of genome scans (Levinson
2005)) |
A polymorphism in this gene has been associated with bipolar disorder in one study (Mynett-Johnson et al. 1998a). |
Sodium/Potassium
ATPase |
Would
indirectly affect most transport activity e.g. serotonin (SLC6A4),
and dopamine (SLC6A3) transporters. Sodium potassium
ATPase activity is inhibited by 12-HETE, the product of ALOX12
and arachidonic acid mediated inhibition of the renal sodium pump in
young rats is attenuated by ALOX12 inhibition (Li
et al. 2000). |
A meta-analysis
of multiple studies suggesting sodium pump dysfunction concluded that
lymphocyte activity is decreased in bipolar patients (isoform not specified) (Looney
and el Mallakh 1997).
Stanley consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
CACNA1B calcium channel, voltage-dependent, N type, alpha 1B subunit | WTCCC Moskvina et al,2009 | |||
CACNA1C calcium channel, voltage-dependent, L type, alpha 1C subunit GeneCard | WGA Sklar et al, 2008 GWAS Moskvina et al, 2009 | Binds to RYR2 Mouton et al, 2001 | ||
CACNB2 calcium channel, voltage-dependent, beta 2 subunit | Chinese GWAS Lee et al, 2010 | |||
KCNJ6 GIRK2 potassium inwardly-rectifying channel, subfamily J, member 6 | UK study Hamshere et al, 2009 | |||
KCTD12 potassium channel tetramerisation domain containing 12 | Chinese GWAS Lee et al, 2010 | |||
RYR2 ryanodine receptor 2 OMIM GeneCard | Genome-wide association study Baum et al, 2007 | Calcium release channel: Binds to CACNA1C | ||
SCN8A sodium channel, voltage gated, type VIII, alpha subunit | Chinese study Wang et al, 2008 | |||
Potassium/chloride
transporter (ACCPN, DKFZP434D2135, KCC3, KCC3A, KCC3B) |
Association
reported in an American case study (Meyer
et al. 2005)
|
Overexpression
is growth promoting in fibroblasts (Shen
et al. 2001) |
Expression
is regulated by the nitric oxide/cGMP/Protein kinase G cascade in endothelial
cells (NOS3)(Di
Fulvio et al. 2001).
Activated by osmotic and oxidative stress (Piechotta
et al. 2002) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
SLC22A16 solute carrier family 22 (organic cation/carnitine transporter), member 16 | US study Fan et al, 2010 | |||
SLC26A7 solute carrier family 26, member 7 sulphate anion transporter OMIM GeneCard | Genome-wide association study Baum et al, 2007 | chloride, sulfate, and oxalate transport. | - | - |
SLC39A3 solute carrier family 39 (zinc transporter), member 3 GeneCard | Genome-wide association study Baum et al, 2007 | Zinc transporter | - | - |
TRPM2 Region
linked to bipolar disorder (McQuillin
et al. 2005) |
Association reported in an American study(Xu et al. 2006) |
Calcium-permeable
cation channel regulated by free intracellular ADP-ribose. The encoded
protein is activated by oxidative stress and confers susceptibility
to cell death (Kaneko
et al. 2006). Also
activated by cyclic ADP ribose (Togashi
et al. 2006) |
Binds
to the inositol 1,4,5 triphosphate (PLCG1 product) channel ITPR3 (Tang
et al. 2001). |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
VGCNL1 NALCN sodium leak channel, non-selective GeneCard | Genome-wide association study Baum et al, 2007 | - | - | - |
Genes
associated with bipolar disorder |
Association
studies |
Primary
role
|
Links
to other Bipolar genes |
Expression
changes |
COMT
Catechol-O-methyltransferase
OMIM 116790
22q11.21 High incidence
of Bipolar disorder in 22q11 deletion syndrome (Papolos
et al. 1996) |
Several
studies have reported association of the low-activity allele of COMT
with bipolar disorder (Rotondo
et al. 2002;Mynett-Johnson et al. 1998b;Kirov et al. 1998;Li et al.
1997). COMT
is a susceptibily gene for both bipolar disorder and schizophrenia (Shifman
et al. 2004). |
Catalyzes
the transfer of a methyl group from S-adenosylmethionine to catecholamines,
including dopamine, adrenaline, and noradrenaline, forming 3-methoxytyramine,
metanephrine and normetanephrine respectively. |
Would
affect dopamine metabolism. COMT activation also increases homocysteine
synthesis and release from astrocytes (derived from S-adenosylmethionine) (Huang
et al. 2005) |
No change
in dorsolateral profrontal cortex mRNA expression (Tunbridge
et al. 2004). Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
DDC 7p11 ? |
A Danish study has linked this gene to bipolar disorder (Borglum et al. 2003;Borglum et al. 1999) |
DOPA decarboxylase
catalyses the formation of dopamine from l-DOPA but can also synthesise
5-HT from 5-hydroxytyptophan, tryptamine from tryptophan, tyramine from
tyrosine and phenylethylamine from pheylalanine. It is expressed in
both serotoninergic and dopaminergic neurones (Tison
et al. 1991) and in
astrocytes (Li
et al. 1992). |
|
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
DRD1 5q35.1 5q31-5q35
linked to both schizophrenia and psychosis in bipolar disorder (Sklar
et al. 2004) |
The dopamine receptors DRD1, DRD2 and DRD3 have each been associated with bipolar disorder (Ni et al. 2002;Massat et al. 2002;Chiaroni et al. 2000;Li et al. 1999;Staner et al. 1998;Manki et al. 1996). |
Positively
coupled to adenylate cyclase (Kebabian
1997) |
Increases
neuronal BDNF expression (Fang
et al. 2003),(Kuppers
and Beyer 2001) :
Activates AKT1 (Brami-Cherrier
et al. 2002). Activates
PLCG1 via PKA (Yu
et al. 1996) ;
In LTK and HEK293
cells dopamine D1 receptor stimulation (DRD1) decreases
PLCG1 cortical membrane expression and activity (Yu
et al. 1996). |
Decreased
mRNA in hippocampal area CA2 (Pantazopoulos
et al. 2004). Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
DRD2 Dopamine receptor OMIM 126450 GeneCard 11q23 - ? |
Negatively
coupled to adenylate cyclase (Huff
1996) |
Stimulates
the PI3K/AKT pathway in PC12 or substantia nigra cells via transactivation
of the EGF receptor (Nair
and Sealfon 2003).
In cortical neurones,
DRD2 receptor stimulation also protects neurones from
the toxic effects of glutamate via the PI3K/AKT pathway (Kihara
et al. 2002). Increases
BDNF expression in NG108-15 cells (Takeuchi
et al. 2002) Activates
AKT1 (Brami-Cherrier
et al. 2002) Coupled
to GNAZ (Obadiah
et al. 1999) which
links to PLCG1 (Tran
et al. 2002). In the
mouse striatum amphetamine or Dopamine transporter knockdown inactivate
AKT1 and activate GSK3B, effects that
are blocked by D2 receptor antagonism or by lithium (Beaulieu
et al. 2004).Heterodimerises
with SSTR5 (Rocheville
et al. 2000). |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
|
DRD3 Dopamine receptor |
Negatively
coupled to adenylate cyclase (Kuzhikandathil
et al. 2004). |
Stimulates
the PI3K/AKT pathway via transactivation of the EGF receptor (Beom
et al. 2004). Coupled
to GNAZ (Obadiah
et al. 1999) which
links to PLCG1 (Tran
et al. 2002). Binds
to GRB2 (Oldenhof
et al. 2001). BDNF
increases the expression of DRD3 in the developing brain
and maintains its expression in adulthhood (Guillin
et al. 2001;Guillin et al. 2003;Sokoloff et al. 2002) |
Decreased
expression in lymphocytes (Vogel
et al. 2004). Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
|
NCKAP5 NCK-associated protein 5 | GWAS Wang et al, 2010 | |||
SLC6A3 DAT1 dopamine transporter OMIM 126455 GeneCard 5p15.3 5p15 shows
suggestive evidence for linkage related to psychosis in bipolar disorder
(NIMH) (Cheng et al.
2006) |
Two studies have suggested association of the dopamine transporter to bipolar disorder (Greenwood et al. 2001;Waldman et al. 1997) |
Dopamine
transport |
Activity
controlled by PI3 kinase pathway (Carvelli
et al. 2002). The
PI3 kinase pathway controls dopamine uptake in synaptosomes and in appropriately
transfected HEK-293 cells. Inhibition of phosphatidylinositol (PI) 3-kinase
by LY294002 induces internalization of the human transporter and reduces
transport capacity. Insulin, which activates this pathway, stimulated
dopamine uptake, an effect also blocked by LY294002. Increased dopamine
uptake was also observed in HEK-293 following transfection with constitutively
active PI3 kinase (murine PI 3-kinase p110alpha (PIK3CA) (Carvelli
et al. 2002). In the
mouse striatum amphetamine or Dopamine transporter knockdown inactivate
AKT1 and activate GSK3B effects that
are blocked by DRD2 receptor antagonism or by lithium (Beaulieu
et al. 2004). |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
TH
Tyrosine
hydroxylase OMIM 191290
11p15.5 |
TH has been weakly associated with bipolar disorder in French and Spanish studies (Perez, I et al. 1995;Leboyer et al. 1990) |
Catecholamine
synthesis (L-DOPA) |
Phosphatidylinositol
4-phosphate, the product of PIK3C3 and phosphatidylinositol 4,5-bisphosphate,
the product of PIP5K2A, stimulate TH activity
in vitro (Morita
et al. 1986). BDNF
controls expression (Zhou
et al. 1998). Expression
controlled by TCF4 (Yoon
and Chikaraishi 1994) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
MAOA Monoamine oxidase A OMIM 309850 GeneCard Xp11.3 Suggestive
susceptibility locus in an NIMH pedigree (Zandi
et al. 2003) |
A number of studies have linked this gene to Bipolar disorder (Kirov et al. 1999) |
Monoamine
oxidase. The gene product oxidises dopamine, noradrenaline and serotonin,
and other amines, generating H2O2 in the process (Youdim
and Finberg 1991)
|
A
number of studies have shown that the apoptotic effects of catecholamines
can be related to the generation of H2O2 from
MAO (Bianchi
et al. 2003) H2O2
decreases the expression of TrkB (BDNF receptor) at
the cell surface (Olivieri
et al. 2003). H2O2
oxidises BDNF (Jensen
et al. 2000). H2O2
derived from dopamine oxidises PLCG1 (Kim
et al. 2002). |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
ADRBK2 22q12.1 OMIM MAFD1 |
A single nucleotide polymorphism of this gene has been associated with bipolar disorder (Barrett et al. 2003). |
Beta
adrenergic receptor kinase 2 |
Inhibited
by PI 4,5 P2, product of PIP5K2A (Onorato
et al. 1995) Phosphorylates
and inhibits DRD1 (Tiberi
et al. 1996) and P2RX7 (Feng
et al. 2005) |
Decreased
protein expression in lymphocyte cell lines isolated from bipolar patients (Niculescu,
III et al. 2000).
Stanley consortium Combined analysis: Downregulated (Higgs
et al. 2006) |
GNAZ 22q11.2 High incidence
of Bipolar disorder in 22q11 deletion syndrome (Papolos
et al. 1996) |
GNAZ has been associated with bipolar disorder in an American study (Saito et al. 1999). |
guanine
nucleotide binding protein |
Binds
to PLCG1 (Bartlett
and Hendry 1997) DRD2,
DRD3 receptors
signal via GNAZ (Obadiah
et al. 1999) |
Reduced
expression in olfactory neuroepithelial cells isolated from bipolar
patients (McCurdy
et al. 2006) . Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
ADCY8 adenylate cyclase 8 (brain) GeneCard | Associated with bipolar disorder in a family-based American study Zandi et al, 2008 | |||
ADCY9 16p13.3 Region
linked to Bipolar disorder in a Genome-wide linkage scan (NIMH) (Cheng
et al. 2006) |
ADCY9 has been shown to be weakly associated with bipolar disorder in a family based study (Toyota et al. 2002). |
This form
of adenylate cyclase is coupled to GalphaS linked receptors including
the beta-2 adrenoceptor (Small
et al. 2003). It is
inhibited by calcineurin (Paterson
et al. 1995). |
DRD2
receptor stimulation inhibits Galpha(s) stimulation of ADCY9
and Galpha(s) stimulation of ADCY9
is reduced by blocking ADCY9 glycosylation (Cumbay
and Watts 2004). SSTR5
also signals via Galpha(s) and inhibits adenylate cyclase in in cho
cells although the subtype of adenylate cyclase was not specifically
identified (Carruthers
et al. 1999) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
SLC18A1 8p21.3 |
Associated reported in an American study (Lohoff et al, 2006) | Vesicular monoamine transporter. A splice variant of unknown function, that does not take up serotonin, is localised in the endoplasmic reticulum (Essand et al, 2005). | - | - |
SLC18A2 solute carrier family 18 (vesicular monoamine), member 2 | Spanish study Gutierrez et al, 2006 |
Genes
associated with bipolar disorder |
Association
studies |
Primary
role
|
Links
to other Bipolar genes |
Expression
changes |
HTR2A
Serotonin
receptor OMIM 182135
13q14-q21 13q14
described as a combined locus for both schizophrenia and Bipolar disorder
in a Canadian study (Maziade
et al. 2005). 13q21
also described in an american study (NIMH) (McInnis
et al. 2003) |
Three studies have reported association with Bipolar disorder (Ranade et al. 2003;Bonnier et al. 2002;Chee et al. 2001). |
Serotonin |
Stimulates
BDNF expression (Meller
et al. 2002). (Vaidya
et al. 1997;Vaidya et al. 1999).
Activates AKT1 in PC12 cells (Johnson-Farley
et al. 2005) . Postsynaptic
HTR2A receptors are markedly reduced in BDNF conditional
knockout mice (Rios
et al. 2006). Serotonin
regulates GSK3B activity in the cortex, hippocampus
and striatum in vivo and its phosphoylation at serine9 is increase by
the combined administration of fenfluramine and clorgyline or by the
serotonin uptake blockers fluoxetine or imipramine. 5HT1A receptors
mediate increases and 5HT2 (HTR2A) receptors mediate decreases
in GSK3B phosphorylation (Li
et al. 2004).
|
Reduced
hippocampal mRNA expression (Lopez-Figueroa
et al. 2004) (Knable
et al. 2004). Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
HTR3A 11q23.1 |
One study
has reported association of a polymorphism in HTR3A with bipolar
disorder (Niesler
et al. 2001).
|
Serotonin
receptor/channel |
Assembly
and cell surface expression controlled by HSPA5 (Boyd
et al. 2002). Serotonin
receptor/channel: HTR3 receptors
activate phospholipase D (Khan
and Hichami 1999;Khan and Poisson 1999)whose
product Phosphatidic acid is a potent inhibitor
of PI3 kinase (Lauener
et al. 1995). |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
HTR3B 11q23.1 ? |
A 3 base
pair deletion in the gene coding HTR3B is under-represented in
a sample of Bipolar disorder patients (Frank
et al. 2004a). |
Serotonin
receptor/channel |
- |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
HTR4 5q31-q33 5q33 linked
to Bipolar disorder in a European study (Etain
et al. 2006) |
This receptor
has been associated with Bipolar disorder in a Japanese study (Ohtsuki
et al. 2002).
|
Activates
adenylate cyclase (Dumuis
et al. 1989). In hippocampal
neurones, HTR4 activation increases neuronal excitability
via a protein kinase A mediated inhibition of calcium-dependent potassium
channels (Torres
et al. 1995). |
This receptor
also activates the ERK kinases ERK1 (MAPK3) and ERK2 (MAPK1) via Ras
and protein kinase A in HEK-293 cells (Norum
et al. 2003). cAMP
generated by 5HT-4 receptor activation also activates RAP1 via the guanine
nucleotide-exchange factor Epac1 (RAPGEF3) and this leads to activation
of the small GTPase Rac (Maillet
et al. 2003). RAPGEF3
is able to activate PI3 kinases in other models (Sable
et al. 1997;Filippa et al. 1999;Mei et al. 2002) but
the effects of HTR4 receptor stimulation have not been
specifically examined. |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
HTR5A 7q36.1 7q36 linked
to Bipolar disorder in a European study (Etain
et al. 2006) |
A polymorphism
associating with bipolar disorder has been reported in one study (Birkett
et al. 2000). |
Localised
in the suprachiasmatic nucleus. Activation can produce phase shifts
of the circadian clock (Sprouse
et al. 2004)Negatively
coupled to adenylate cyclase (Nelson
2004). In C6 glioma
cells HTR5A activation increases the accumulation of
inositol 1,4,5 triphosphate and inhibit ADP ribosyl cyclase activity
via G-protein linked pathways (Noda
et al. 2003).Cyclic
ADP ribose, the product of this enzyme increases ryanodine-sensitive
intracellular calcium mobilisation (Meszaros
et al. 1993).
|
Cyclic
ADPribose activates TRPM2 (Togashi
et al. 2006) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
HTR6Serotonin receptor OMIM 601109 GeneCard 1p35-36
identified as a susceptibility locus in a European study (Schumacher
et al. 2005) |
A limited
association between bipolar disorder and this receptor has been observed
in one study (Vogt
et al. 2000). |
Positively
coupled to adenylate cyclase (Grimaldi
et al. 1998). |
- |
Stanley
consortium Combined analysis: Downregulated (Higgs
et al. 2006) |
SLC6A4
Serotonin
transporter OMIM 182138 17q11.1-q12 Region
linked to bipolar disorder (see review of genome scans (Levinson
2005) |
Numerous studies support association of this transporter to unipolar, bipolar and seasonal affective disorders (Lotrich and Pollock 2004) |
Serotonin
transporter |
BDNF
increases expression in embryonic rat raphe (Rumajogee
et al. 2002). Synaptosomal
serotonin transport is inhibited by tetanus toxin an effect mediated
by tyrosine kinase receptor activation and a subsequent increase in
PLCG1 phosphorylation (Najib
et al. 2000). |
Increased
frontal cortex mRNA expression (Sun
et al. 2001). Reduced
affinity for [3H]citalopram in molecular layer of the hippocampus (Dean
et al. 2003). Reduced
imipramine binding in platelets of unaffected relatives of bipolar patients
(Kd and Bmax) (Leboyer
et al. 1999) Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
TDO2 tryptophan 2,3-dioxygenase | US study Miller et al, 209 | |||
TPH2 Tryptophan hydroxylase 2 GeneCard | Association reported in Canadian case and family study Harvey et al, 2007 and in a Swedish population Van den Bogaert et al, 2006 | Converts tryptophan to 5-hydroxytryptophan. Rate limiting enzyme of serotonin synthesis. |
Genes
associated with bipolar disorder |
Association
studies |
Primary
role
|
Links
to other Bipolar genes |
Expression
changes |
GABRA1 5q34-q35 |
(Horiuchi et al. 2004) |
GABA receptor |
PDGF receptors
inactivate GABA receptors (Valenzuela
et al. 1995) in rat
hippocampal neurones and independently of their subunit composition
via activation of PLCG1.
In contrast, BDNF promotes the cell surface clustering
of GABAA receptors via activation of PLCG1 and subsequent
increases in intracellular calcium (Mizoguchi
et al. 2003). |
No change
in immunocytochemical study in prefrontal cortex (Ishikawa
et al. 2004) Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
GABRA4 gamma-aminobutyric acid (GABA) A receptor, alpha 4 | U K WTCCC study Craddock et al, 2010 | |||
GABRA5 15q11.2-q12 15q11
decribed a s a susceptibility locvus in a Canadian study(Maziade
et al. 2005) |
(De bruyn et al. 1996;Otani et al. 2005;Papadimitriou et al. 1998) U K WTCCC study Craddock et al, 2010 |
GABA |
In developing
neurones the chloride gradient is such as to enable GABA (GABRA1,
GABRA5) to promote excitation rather than inhibition. GABA-related
depolarisation can activate calcium channels and activate downstream
signaling cascades. In the developing hypothalamus
GABA and muscimol activate the MAPK/CREB pathway and increase the expression
of BDNF (Obrietan
et al. 2002). Similar
effects have been observed in hippocampal culture where the effects
of GABA switch from increasing to decreasing BDNF expression
during maturation, in parallel with the switch of GABA actions from
excitation to inhibition (Marty
et al. 1996;Berninger et al. 1995)
The excitatory effects of GABA also contribute to the trophic effects
of this transmitter in developing oligodendrocytes (Wang
et al. 2003). In the
adult brain, where GABA is inhibitory, GABA-A receptor activation reduces
BDNF and NGF expression in the hippocampus (Zafra
et al. 1991). |
Stanley
consortium Combined analysis: Downregulated (Higgs
et al. 2006) |
GABRA1 | WTCCC | |||
GABRB1 gamma-aminobutyric acid (GABA) A receptor, beta 1 GeneCard | Wellcome Trust Case control Consortium | - | - | - |
GABRB2 gamma-aminobutyric acid (GABA) A receptor, beta 2 | US family-based study Perlis et al, 2008 | |||
GABBR2 gamma-aminobutyric acid (GABA) B receptor, 2 (GPR51) OMIM GeneCard | Genome-wide association study Baum et al, 2007 | GABAB receptor inhibiting glutamate release | Binds to ATF4 and ATF5 (binding partners of DISC1) | |
GABRB3 gamma-aminobutyric acid (GABA) A receptor, beta 3 | U K WTCCC study Craddock et al, 2010 | |||
GABRR1 gamma-aminobutyric acid (GABA) receptor, rho 1 | U K WTCCC study Craddock et al, 2010 | |||
GABRR3 | WTCCC | |||
Suggested
susceptibility locus in german and danish studies (Cichon
et al. 2001;Ewald et al. 2003) |
Weak association
reported in a Danish study (Lundorf
et al. 2005) |
GABA
synthesis |
Transcription
controlled by BDNF in rat striatal neurones (Mizuno
et al. 1994) |
Reduced
hippocampal protein (Knable
et al. 2004), Decreased
number of GAD67-positive neurones in Brodmann area 9 (Veldic
et al. 2005) and in
cerebellum (Fatemi
et al. 2005). Reduced
density of GRIN2A receptor bearing GAD67-positive neurones in
layer 2 of cingulate cortex (Woo
et al. 2004). Reduced
mRNA expression in prefrontal cortex (Guidotti
et al. 2000).Reduced
mRNA density in CA4 hippocampal region (Heckers
et al. 2002). Stanley
consortium Combined analysis: Downregulated (Higgs
et al. 2006) |
CHRNB3 | Hartz et al, 2011 |
Genes
associated with bipolar disorder |
Association
studies |
Primary
role
|
Links
to other Bipolar genes |
Expression
changes |
MTHFR Methylenetetrahydrofolate reductase 1p36.3 Region
linked to bipolar disorder (see review of genome scans (Levinson
2005)) |
Associated
with Bipolar disorder in Polish males (Kempisty
et al. 2006) |
5-methyltetrahydrofolate
+ NAD(P)(+) <=> 5,10-methylenetetrahydrofolate + NAD(P)H Product
is a cosubstrate for homocysteine remethylation to methionine. |
Homocysteine
is a glutamate site agonist/ glycine site antagonist at NMDA receptors (Lipton
et al. 1997). Homocysteine
is growth suppressive in endothelial cells via inhibition of the PI3K/AKT
pathway (Zhang
et al. 2005). Exerts
cytotoxicity via multiple effects on NMDA, PI3K/AKT, oxidative stress
and endoplasmic reticulum stress-related pathways and effects on HSPA5,
TNFA, inter alia (Sharma
et al. 2006) |
High serum
levels of homocysteine in young male bipolar disorder patients (Levine
et al. 2005). Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
MTR 5-methyltetrahydrofolate-homocysteine methyltransferase OMIM GeneCard | Association described in a Polish study Kempisty et al, 2007 | |||
MTHFD1 methylenetetrahydrofolate dehydrogenase (NADP+ dependent) 1, methenyltetrahydrofolate cyclohydrolase, formyltetrahydrofolate synthetase OMIM GeneCard | Association described in a Polish study Kempisty et al, 2007 | - | - |
Genes
associated with bipolar disorder |
Association
studies |
Primary
role
|
Links
to other Bipolar genes |
Expression
changes |
FZD2 Frizzled-2 GeneCard | Genome-wide association study Baum et al, 2007 | Wnt signaling via cGMP phosphodiesterase Wang, 2004 | ||
HMG2L1 high-mobility group protein 2-like 1 GeneCard | Potash et al, 2007 | Negative regulator of Wnt signaling Yamada et al, 2007 | ||
NXN Nucleoredoxin GeneCard | Genome-wide association study Baum et al, 2007 | Interacts with dishevelled and inhibits Wnt signaling Funato et al, 2006 | ||
WNT2B wingless-type MMTV integration site family, member 2B | US study Zandi et al, 2008 | |||
WNT7A wingless-type MMTV integration site family, member 7A | US study Zandi et al, 2008 |
APH1B anterior pharynx defective 1 homolog B | Part of gamma secretase complex degrading APP | |
APP amyloid beta (A4) precursor protein | ||
CLDN5 Claudin 5 | Tight junction component | |
CNTNAP2 contactin associated protein-like 2 | Already associated | |
COMT Catechol-O-methyl transferase | Already associated | |
GNB1L guanine nucleotide binding protein (G protein), beta polypeptide 1-like | ||
GRM7 glutamate receptor, metabotropic 7 | Already associated | |
LARGE like-glycosyltransferase | ||
MED15 mediator complex subunit 15 | ||
NRXN1 Neurexin 1 | Already associated | |
PARK2 Parkinson disease (autosomal recessive, juvenile) 2, parkin | ||
PI4KA phosphatidylinositol 4-kinase, catalytic, alpha | ||
PTPRD protein tyrosine phosphatase, receptor type, D | ||
RCAN2 regulator of calcineurin 2 | ||
RTN4R Reticulon 4 (NOGO) receptor | reticulon 4 (NOGO) , oligodendrocyte myelin glycoprotein and myelin-associated glycoprotein receptor | |
SNAP29 synaptosomal-associated protein, 29kDa | Neurotransmitter release | |
TXNIP thioredoxin interacting protein | ||
UFD1L ubiquitin fusion degradation 1 like | degradation of ubiquitinated proteins | |
ZDHHC8 zinc finger, DHHC-type containing 8 | ||
ZNF74 zinc finger protein 74 |
Miscellaneous
genes
Genes
associated with bipolar disorder |
Association
studies |
Primary
role
|
Links
to other Bipolar genes |
Expression
changes |
A2BP1 ataxin 2-binding protein 1 (aka FOX-1) GeneCard | Genome-wide association study Baum et al, 2007 UK study Hamshere et al, 2009 | Neuronal splicing regulator Underwood et al, 2005 | ||
ABCA13 ATP-binding cassette, sub-family A (ABC1), member 13 | Knight et al, 2009 | |||
ABTB1 ankyrin repeat and BTB (POZ) domain containing 1 GeneCard | Genome-wide association study Baum et al, 2007 | |||
ACE angiotensin converting enzyme | Turkish study Kucukali et al, 2010 | |||
AK3L2 adenylate kinase 3-like 2 GeneCard | Wellcome Trust Case control Consortium | Uncharacterised | ||
AKAP10 A kinase (PRKA) anchor protein 10 GeneCard | Wellcome Trust Case control Consortium | Protein Kinase A anchoring protein with RGS domains, a PKA-binding motif, and a PDZ (PSD95/Dlg/ZO1)-binding motif | ||
AMTN (UNQ689) Amelotin GeneCard | Genome-wide association study Baum et al, 2007 | Tooth enamel protein | ||
ANK3 ankyrin 3, node of Ranvier (ankyrin G) GeneCard | Genome-wide association study Baum et al, 2007 | Found at the axonal initial segment and nodes of Ranvier of neurons in the central and peripheral nervous systems .Regulates PI3 kinase signalling via binding to PIK3R1 Ignatiuk et al, 2006 | ||
ANKRD35 | Moskvina et al, 2009 | |||
AOF1 amine oxidase (flavin containing) domain 1 GeneCard | Wellcome Trust Case control Consortium | Uncharacterised | ||
ATP6V0D1 |
GWAS Moskvina et al, 2009 | |||
AQP4 Aquaporin 4 GeneCard | Genome-wide association study Baum et al, 2007 | Brain-expressed water channel | ||
AUTS2 autism susceptibility candidate 2 | UK study Hamshere et al, 2009 | |||
B3GALT5 UDP-Gal:betaGlcNAc beta 1,3-galactosyltransferase, polypeptide 5 | UK study Hamshere et al, 2009 | |||
BCL2 | Uemara et al, 2011 | |||
BDKRB2 bradykinin receptor B2 | Spanish study Gratacos et al, 2009 | |||
BRCA2 breast cancer 2, early onset | GWAS Tesli et al, 2010 | |||
BRD1 bromodomain containing 1 GeneCard 22q13.33: |
Association suggested in a Danish study (Severinsen et al, 2006) | Unknown | - | - |
BRE brain and reproductive organ-expressed (TNFRSF1A modulator) GeneCard | Genome-wide association study Baum et al, 2007 | Modulates TNF effects via interaction with TNFRSF1A Gu et al, 1998. Inhibits mitochondrial apoptosis pathway Li et al, 2004 | ||
BTBD16 BTB (POZ) domain containing 16 | GWAS , USA Smith et al, 2009 | |||
BPTF (FALZ) bromodomain PHD finger transcription factor GeneCard | Genome-wide association study Baum et al, 2007 | Apoptosis inducing Strachan et al, 2005. Binds to KEAP1 Strachan et al, 2004 a cytosolic binding partner of NFE2L2. | ||
BSN Bassoon (presynaptic cytomatrix protein) | UK study Hamshere et al, 2009 | |||
CABIN1
calcineurin binding protein 1 22q11.23 : |
Weakly
Associated in a study of Ashkenazi case-parent
trios (Fallin
et al. 2005) |
Calcineurin inhibitor |
- |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
PPP3CC protein phosphatase 3 (formerly 2B), catalytic subunit, gamma isoform GeneCard | Association reported in a French study Mathieu et al, 2008 | Inhibited by CABIN1 Esau et al, 2001 | ||
C10orf120 | GWAS Moskvina et al, 2009 | |||
C14orf48 | GWAS Moskvina et al, 2009 | |||
C15orf53 | GWAS Ferreira et al, 2008 | |||
CAPN6 Calpain 6 GeneCard | Wellcome Trust Case control Consortium | Uncharacterised calpain | ||
CAPN10 | GWAS Moskvina et al, 2009 | |||
CDC25B cell division cycle 25 homolog B GeneCard | Wellcome Trust Case control Consortium | CDC phosphatase | Binds to 14-3-3 proteins including YWHAH | |
CDKN2A cyclin-dependent kinase inhibitor 2A (melanoma, p16, inhibits CDK4) | GWAS Moskvina et al, 2009 | |||
CHMP1B chromatin modifying protein 1B | Kato 2007 | |||
CHRM2 cholinergic receptor, muscarinic 2 | Cannon et al, 2010 | |||
CHRNA7 (dup)cholinergic receptor, nicotinic, alpha 7 | Taiwanese study Hong et al, 2004 | |||
CMTM8 | GWAS Moskvina et al, 2009 | |||
CREB1 cAMP responsive element binding protein 1 | Canadian study Mamdani et al, 2008 | |||
CPB2 | GWAS Moskvina et al, 2009 | |||
CSMD1 CUB and Sushi multiple domains 1 GeneCard | Genome-wide association study Baum et al, 2007 | Regulates complement activation and inflammation in the developing CNS Kraus et al, 2006 | ||
CTNNA2 catenin (cadherin-associated protein), alpha 2 | GWAS Scott et al, 2009 | |||
CUTL2
cut-like 2 transcription factor 12q24.11-q24.12 |
Association
reported in a UK case-control study (Glaser
et al. 2005b) |
Transcription
factor |
Binds
to and inbibits NCAM promoter activity (Tissier-Seta
et al. 1993;Valarche et al. 1993) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
CYP4V2 cytochrome P450, family 4, subfamily V, polypeptide 2 OMIM GeneCard | Genome-wide association study Baum et al, 2007 | Cytochrome p450 substrate uncharacterised | ||
DCTN5 Dynactin 5 | GWAS WTCCC | |||
DGCR8
DiGeorge syndrome critical region gene 8 High incidence
of Bipolar disorder in 22q11 deletion syndrome (Papolos
et al. 1996) |
Weakly
Associated in a study of Ashkenazi case-parent
trios (Fallin
et al. 2005) |
Involved in microRNA
biosynthesis |
- |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
DDX42 DEAD (Asp-Glu-Ala-Asp) box polypeptide 42 GeneCard | Genome-wide association study Baum et al, 2007 | RNA chaperone Uhlmann-Schiffler et al, 2006 | ||
DIO2 deiodinase, iodothyronine, type II | Chinese study He et al, 2009 | |||
DIS3L2 DIS3 mitotic control homolog (S. cerevisiae)-like 2 FLJ36974; MGC42174 GeneCard | Genome-wide association study Baum et al, 2007 | Uncharacterised | ||
DPY19L3
dpy-19-like 3 (C. elegans) |
GWAS , USA Smith et al, 2009 | |||
DPP10 dipeptidyl-peptidase 10 GeneCard | Wellcome Trust Case control Consortium Moskvina et al, 2009 | Inactive dipeptidyl peptidase homolog that binds to potassium channels Li et al, 2006 | ||
EFNA5 Ephrin A5 GeneCard | Genome-wide association study Baum et al, 2007 | Tyrosine kinase receptor involved in axon guidance. Signals via PI3 kinase and SRC Wong et al, 2004 | ||
ESRRG estrogen-related receptor gamma GeneCard | Wellcome Trust Case control Consortium | Estrogen receptor | - | - |
EXTL1 exostoses (multiple)-like 1 | GWAS analysis Torkamani et al, 2008 | alpha 1,4- N-acetylglucosaminyltransferase involved in chain elongation of heparan sulfate and possibly heparin | ||
FABP3 | GWAS Moskvina et al, 2009 | |||
FABP7 fatty acid binding protein 7, brain | Japan Imayama et al, 2010 | |||
FAM109A
family with sequence similarity 109, member A (FLJ32356) |
Association
reported in a UK case-control study (Glaser
et al. 2005b) |
Unknown
function |
- |
- |
FAM115A | GWAS Moskvina et al, 2009 | |||
FAM126A (Hyccin) family with sequence similarity 126, member A GeneCard | Wellcome Trust Case control Consortium | Involved in myelination and downregulated by beta-catenin | ||
FBXO38 F-box protein 38 | GWAS Moskvina et al, 2009 | |||
FLVCR2 (C14ORF58) feline leukemia virus subgroup C cellular receptor family, member 2 GeneCard | Wellcome Trust Case control Consortium | |||
FOXN3 (CHES1) forkhead box N3 /chechpoint suppressor 1 GeneCard | Genome-wide association study Baum et al, 2007 | |||
GLT8D1 glycosyltransferase 8 domain containing 1 | GWAS Moskvina et al, 2009 | |||
GNL3 guanine nucleotide binding protein-like 3 (nucleolar) | US GWAS Scott et al, 2009 GWAS Moskvina et al, 2009 | |||
Association
reported in female Scottish patients in bipolar disorder and schizophrenia (Underwood
et al. 2006) |
Orphan
receptor |
- |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
|
GPR109B G protein-coupled receptor 109B HM74 | Miller et al, 2009 | |||
HK2 Hexokinase-2 GeneCard | Genome-wide association study Baum et al, 2007 | phosphorylates glucose to glucose-6-phosphate, first step in glycolysis | ||
hCAP-D3 =NCAPd3 non-SMC condensin II complex, subunit D3 | GWAS Moskvina et al, 2009 | |||
HOXA3 | GWAS Moskvina et al, 2009 | |||
HSST3 NDST3 N-deacetylase/N-sulfotransferase (heparan glucosaminyl) 3 | GWAS analysis Torkamani et al, 2008 | |||
HS6ST1
heparan sulfate 6-O-sulfotransferase 1 |
GWAS analysis Torkamani et al, 2008 | |||
HS6ST3
heparan sulfate 6-O-sulfotransferase 3 |
GWAS analysis Torkamani et al, 2008 | |||
INTS6 integrator complex subunit 6 | GWAS Moskvina et al, 2009 | |||
ITIH1 inter-alpha (globulin) inhibitor H1 | USA GWAS Scott et al, 2009 | |||
ITIH3 inter-alpha (globulin) inhibitor H3 | USA GWAS Scott et al, 2009 GWAS Moskvina et al, 2009 | |||
JAM3 junction adhesion molecule 3 | GWAS Moskvina et al, 2009 | |||
MAK male germ cell-associated kinaseGeneCard | Genome-wide association study Baum et al, 2007 | serine/threonine kinase | ||
MCHR1
(GPR24) 22q13.2 |
Suggested as a susceptibility gene in Faroese and Scottish populations (Severinsen et al. 2006). |
Melanocyte-concentrating
hormone receptor |
- |
Stanley
consortium Combined analysis: Downregulated (Higgs
et al. 2006) |
MCHR2 melanin-concentrating hormone receptor 2 | Miller et al, 2009 | |||
NCAN Neurocan | Cichon et al, 2011 GWAS | |||
NCKAP5 (NAP5) NCK-associated protein 5 | GWAS Smith et al, 2009 | |||
KIAA0960 | GWAS Moskvina et al, 2009 | |||
KCTD8 potassium channel tetramerisation domain containing 8 GeneCard | Genome-wide association study Baum et al, 2007 | Uncharacterised | ||
KLF12 Kruppel-like factor 12 AP-2 repressor GeneCard | Genome-wide association study Baum et al, 2007 | Represses transcription factor AP-2alpha (TFAP2A | ||
KLK8 kallikrein-related peptidase 8 Neuropsin GeneCard | Izumi et al, 2008 | Involved in LTP Tamura et al, 2006 : Activated by NMDA receptors Matsumoto-Miyai et al, 2003. Promotes oligodendrocyte cell death demyelination and axonal degeneration Terayama et al, 2007: Modifies neurite outgrowth Oka et al, 2002 | ||
KLHDC1 kelch domain containing 1 GeneCard | Wellcome Trust Case control Consortium | Uncharacterised | ||
KRN1 KRTAP5-9 keratin associated protein 5-9 OMIM | Genome-wide association study Baum et al, 2007 | Hair Keratin | ||
LAMP3 lysosomal-associated membrane protein 3 GeneCard | GWAS Moskvina et al, 2009 | |||
LDB2 Lim domain binding 2 GeneCard | Genome-wide association study Baum et al, 2007 | |||
LLGL2 lethal giant larvae homolog 2 GeneCard | Genome-wide association study Baum et al, 2007 | The lethal (2) giant larvae protein of Drosophila plays a role in asymmetric cell division, epithelial cell polarity, and cell migration | ||
LOC283547 | Wellcome Trust Case control Consortium | Uncharacterised | ||
LOC728267 | GWAS Moskvina et al, 2009 | |||
LOC731264=LOC730018 | Wellcome Trust Case control Consortium | Uncharacterised | ||
LRCH1 leucine-rich repeats and calponin homology (CH) domain containing 1 GeneCard | Genome-wide association study Baum et al, 2007 | Uncharacterised | ||
LRRC4C (NGL1)leucine rich repeat containing 4C GeneCard | Genome-wide association study Baum et al, 2007 | Binding partner for Netrin G1. Involved in axon guidance | ||
LRRC7 leucine rich repeat containing 7 Densin-180 GeneCard | Wellcome Trust Case control Consortium | Synaptic protein linked to PSD95 Ohtakara et al, 2002 | ||
LRRC36 | GWAS Moskvina et al, 2009 | |||
LY86 Lymphocyte antigen 86 GeneCard | Genome-wide association study Baum et al, 2007 | |||
MC5R melanocortin 5 receptor | US study Miller et al, 2009 | |||
MCM3APA | GWAS Moskvina et al, 2009 | |||
MCTP1 multiple C2 domains, transmembrane 1 | GWAS Scott et al, 2009 | |||
MLC1
(WKL1) megalencephalic
leukoencephalopathy with subcortical cysts 1 homolog (human) OMIM 605908 22q13.33
|
Associated with bipolar disorder and schizophrenia in an Indian population (Verma et al. 2005b) |
Putative
transporter expressed in astrocytes, Bergmann glia and ependymal cells (Schmitt
et al. 2003) |
Binds
to DISC1 (Millar
et al. 2003) |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
MMP9 matrix metallopeptidase 9 (gelatinase B, 92kDa gelatinase, 92kDa type IV collagenase) | Rybakowski et al, 2009 | |||
MPPE1 metallophosphoesterase 1 | Lohoff et al, 2010 | |||
MRCL3 =MYL12A myosin, light chain 12A, regulatory, non-sarcomeric | GWAS Moskvina et al, 2009 | |||
MRLC2= MYL12B | GWAS Moskvina et al, 2009 | |||
MUC2 mucin 2, oligomeric mucus/gel-forming GeneCard | Genome-wide association study Baum et al, 2007 | Mucus protein | ||
MYO5B myosin VB GeneCard | WGA Sklar et al, 2008 | Myosin | ||
NEK4 NIMA (never in mitosis gene a)-related kinase 4 | GWAS USA Scott et al, 2009 GWAS Moskvina et al, 2009 | |||
NCL Nucleolin GeneCard | Genome-wide association study Baum et al, 2007 | nucleolar phosphoprotein involved in the synthesis and maturation of ribosomes | ||
NF1X nuclear factor I/X (CCAAT-binding transcription factor) | GWAS USA Scott et al, 2009 | |||
NPAS3 neuronal PAS domain protein 3 | Scottish study Pickard et al, 2008 | |||
NR2E1 nuclear receptor subfamily 2, group E, member 1 GeneCard | Associaton in a Canadian study (females) Kumar et al, 2008 | |||
NR3C1 nuclear receptor subfamily 3, group C, member 1 (glucocorticoid receptor) | Dutch study Spijker et al, 2009 | |||
NSMCE2 non-SMC element 2, MMS21 homolog | US family study Zandi et al 2008 | DNA repair | ||
NYD-SP26 GeneCard | Genome-wide association study Baum et al, 2007 | Unknown function | - | - |
OR51F2 olfactory receptor, family 51, subfamily F, member 2 | Genome-wide association study Baum et al, 2007 | Orphan olfactory receptor | - | - |
OTX2 orthodenticle homeobox 2 14q21-q22 GeneCard | Sabunciyan et al, 2007 | Controls dopamine and serotonin neuronal development. Regulates NCAM1 expression Nguyen et al, 1999 | ||
PALB2 partner and localizer of BRCA2 GeneCard | Wellcome Trust Case control ConsortiumGWAS WTCCC GWAS Moskvina et al, 2009 |
Putative tumor suppressor | ||
PAX5 Paired box 5 GeneCard | Wellcome Trust Case control Consortium | Regulates B cell development Sxhebesta et al, 2007 as well as that of midbrain dopamine neurones Simon et al, 2003 | ||
PAX7 paired box 7 GeneCard | Genome-wide association study Baum et al, 2007 | Transcription factor | ||
PB1= PBRM1 polybromo 1 | GWAS Moskvina et al, 2009 | Binds to CDKN1A promoter Xia et al, 2008 | ||
PDE4B phosphodiesterase 4B, cAMP-specific (phosphodiesterase E4 dunce homolog, | Norwegian study Kahler at al, 2010 (weak) | |||
PHF17 PHD finger protein 17 (JADE1) GeneCard | Genome-wide association study Baum et al, 2007 | Promotes apoptosis | ||
PLA2G1B 12q23-q24.1 |
PLA2G1B has been associated with bipolar disorder in two studies (Dawson et al. 1995). |
Phospholipid
and lysophospholipid metabolism Secretory
phospholipase A2's, including the pancreatic form can also synthesise
lysophosphatidic acid from phosphatidic acid (Eder
et al. 2000) |
Phosphatidic
and lysophosphatidic acid are potent inhibitors of PI3 kinase (Lauener
et al. 1995). |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
PLA2R1 phospholipase A2 receptor 1, 180kDa GeneCard | Genome-wide association study Baum et al, 2007 | Secretory PLA2 receptor Ancian et al, 1995 (see above) | - | - |
PLSCR4 phospholipid scramblase 4 (aka cell growth inhibiting protein 43) GeneCard | Genome-wide association study Baum et al, 2007 | |||
PFN4 Profilin 4 | GWAS Moskvina et al, 2009 | Binds to phosphatidic acid and PI3P Behnen et al, 2009 | ||
PPARD peroxisome proliferator-activated receptor delta GeneCard | Zandi et al, 2008 | Multi faceted nuclear receptor See Wikipedia | ||
POLE2 | GWAS Moskvina et al, 2009 | |||
PPIL5 | GWAS Moskvina et al, 2009 | |||
PSKH1 protein serine kinase H1 | GWAS Moskvina et al, 2009 | |||
PSMF1 proteasome (prosome, macropain) inhibitor subunit 1 (PI31) GeneCard | Genome-wide association study Baum et al, 2007 | Proteasome inhibitor modifying antigen processing Zaiss et al, 2002 | ||
PTGIS prostaglandin I2 (prostacyclin) synthase GeneCard | Genome-wide association study Baum et al, 2007 | Prostacyclin synthesis | ||
RASGRP1 RAS guanyl releasing protein 1 (calcium and DAG-regulated) | GWAS Ferreira et al, 2008 | |||
RBMS3 RNA binding motif, single stranded interacting protein GeneCard | Genome-wide association study Baum et al, 2007 | RNA binding protein | ||
RELN Reelin | USA Goes et al, 2010 | |||
RFC3 replication factor C (activator 1) 3, 38kDa GeneCard | Genome-wide association study Baum et al, 2007 | Binds to PCNA and acts as an accesory protein for DNA replication | ||
RGS5 regulator of G-protein signaling 5 | GWAS Smith et al, 2009 | |||
RGS17 regulator of G-protein signaling 17 GeneCard | Genome-wide association study Baum et al, 2007 | GTPase activating proteins: turn off G-proteins and negatively regulate the signaling of G-protein coupled receptors (GPCRs) | ||
RNPEPL1 arginyl aminopeptidase (aminopeptidase B)-like 1 GeneCard | Genome-wide association study Baum et al, 2007 and Wellcome Trust Case control Consortium | Uncharacterised aminopeptidase | ||
ROR1 receptor tyrosine kinase-like orphan receptor 1 | GWAS Smith et al, 2009 | |||
RORB RAR-related orphan receptor B retinoid related | US study McGrath et al, 2009 | |||
RSRC2 | GWAS Moskvina et al, 2009 | |||
S100B S100 calcium binding protein B | Irish study Roche et al, 2007 | |||
SEMG2 semenogelin II | GWAS Moskvina et al, 2009 | |||
SGCD sarcoglycan, delta (35kDa dystrophin-associated glycoprotein) OMIM GeneCard | Genome-wide association study Baum et al, 2007 | Part of a dystrophin complex bridging the sarcolemmal cytoskeleton and the extracellular matrix | - | - |
SIGLEC6 sialic acid binding Ig-like lectin 6 | GWAS Moskvina et al, 2009 | |||
SKIV2L superkiller viralicidic activity 2-like (S. cerevisiae) GeneCard | Genome-wide association study Baum et al, 2007 | Associates with the exosome, a structure involved in RNA degradation. Putative antiviral activity by blocking translation of poly(A) deficient mRNAs | ||
SLIT3 Slit homolog 3 GeneCard | Genome-wide association study Baum et al, 2007 | Regulate axonal guidance and cell migration | ||
SLITRK2 SLIT and NTRK-like family, member 2 | GWAS Smith et al, 2009 | |||
Slynar (AY070435)
12q24.3 12q23-24 region linked to Bipolar disorder in UK studies (Glaser et al. 2005b) |
Associated in Danish and UK subjects (Kalsi et al, 2006) | Unknown function | - | - |
SOX5 SRY (sex determining region Y)-box 5 GeneCard | Wellcome Trust Case control Consortium | Transcription factor | ||
SP4 Sp4 transcription factor | US studyZhou et al, 2009 | |||
SPR Sepiapterin reductase | New Zealand study McHugh et al, 2009 | |||
STAB1 stabilin 1 GeneCard | Genome-wide association study Baum et al, 2007 | Scavenger receptor expressed on macrophages . Endocytoses low density lipoprotein, Gram-positive and Gram-negative bacteria, and advanced glycosylation end products. | ||
ST3GAL1 ST3 beta-galactoside alpha-2,3-sialyltransferase 1 GeneCard | Associated with bipolar disorder in an American famili-based study Zandi et al, 2008 US family-based study Perlis et al, 2008 | Golgi-localised enzyme catalyzing transfer of sialic acid from CMP-sialic acid to galactose-containing substrates. | ||
SRP54 signal recognition particle 54kDa GeneCard | Genome-wide association study Baum et al, 2007 | Part of a protein-RNA complex that recognizes and transports specific proteins to the endoplasmic reticulum | ||
SUPT4H1 suppressor of Ty 4 homolog 1 (S. cerevisiae) GeneCard | Genome-wide association study Baum et al, 2007 | Chromatin protein | ||
SVEP1 sushi, von Willebrand factor type A, EGF and pentraxin domain containing 1 GeneCard | Wellcome Trust Case control Consortium GWAS Moskvina et al, 2009 | Osteogenesis ? | ||
SYNE1 spectrin repeat containing, nuclear envelope 1 GeneCard | Wellcome Trust Case control Consortium | Element of the nuclear membrane and actin cytoskeleton | Binds to DISC1 Morris et al, 2003 | |
SYNGR1 22q13.1 |
Associated
with both schizophrenia and bipolar disorder in India (Verma et
al. 2005a) |
Membrane
protein asociated with presynaptic vesicles |
- |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
TAAR6 trace amine associated receptor 6 | Korean study Pae et al, 2008 | |||
TACR1 tachykinin receptor 1 | US family-based study Perlis et al, 2008 | |||
TCF7L1 transcription factor 7-like 1 (T-cell specific, HMG-box) GeneCard | Genome-wide association study Baum et al, 2007 | Binds to beta-catenin | ||
TDP1 tyrosyl-DNA phosphodiesterase 1 GeneCard | Genome-wide association study Baum et al, 2007 | DNA repair | ||
TDRD9 tudor domain containing 9 GeneCard | Wellcome Trust Case control Consortium | Uncharacterised | ||
TEC tec protein tyrosine kinase GeneCard | Genome-wide association study Baum et al, 2007 | Non-receptor tyrosine kinase important in T-cell receptor signaling | ||
THRB Thyroid hormone receptor beta (erythroblastic leukemia viral (v-erb-a) oncogene homolog 2, avian) GeneCard | Wellcome Trust Case control Consortium | nuclear hormone receptor for triiodothyronine | ||
THS7DA thrombospondin, type I, domain containing 7A GeneCard | Wellcome Trust Case control Consortium | |||
TLE4 | GWAS Moskvina et al, 2009 | |||
TMEM10 | GWAS Moskvina et al, 2009 | |||
TOM1 target of myb1 GeneCard | Potash et al, 2007 | translocation of growth factor receptor complexes to the lysosome for degradation (putative) | Negative refulator of IL1B and TNF signaling Yamakami and Yokosawa, 2004 | |
TP53I3 | GWAS Moskvina et al, 2009 | |||
TRDN Triadin GeneCard | Wellcome Trust Case control Consortium | Intracellular calcium release | ||
TSPAN8 tetraspanin 8 GeneCard | WGA Sklar et al, 2008 | |||
TSNAX translin-associated factor X 1q42.1 GeneCard | Weak association in Scottish males Thompson et al, 2005 | Complexes with translin forming a single and double-stranded DNA binding protein Gupta et al, 2005Regulating axonal regenerationSchroer et al, 2007 | ||
VPRBP Vpr (HIV-1) binding protein GeneCard | Genome-wide association study Baum et al, 2007 | Binds to vpr (HIV protein). Acts as a receptor for cul-4-DDB1 ubiquitin ligase LeRouzic et al, 2007 | ||
TUBA8
Tubulin alpha 8 22q11.1 High incidence
of Bipolar disorder in 22q11 deletion syndrome (Papolos
et al. 1996) |
Association
reported in a case-parent trio study in Askenazi Jews (Fallin
et al. 2005) |
Tubulin |
Binds
to GRM7 |
Stanley
consortium Combined analysis: Unchanged (Higgs
et al. 2006) |
VIP Vasoactive intestinal polypeptide | Spanish study Soria et al, 2010 | |||
YWHAH |
Weakly Associated in a study of Ashkenazi case-parent trios (Fallin et al. 2005) |
- |
Activates
TH and TPH1.
Binds to phosphorylated TPH1 (Ichimura et al. 1995). Binds to and
inhibits the AKT1 activator PDPK1 (Sato
et al. 2002). |
Stanley
consortium Combined analysis: Downregulated (Higgs et al. 2006) |
ZADH1 | GWAS Moskvina et al, 2009 | |||
ZBTB44 zinc finger and BTB domain containing 44 GeneCard | Wellcome Trust Case control Consortium | |||
ZNF274 zinc finger protein 274 GeneCard | Wellcome Trust Case control Consortium | Nucleolar zinc finger Yano et al, 2000 | ||
ZNF284 | GWAS Moskvina et al, 2009 | |||
ZNF490 zinc finger protein 490 GeneCard | Wellcome Trust Case control Consortium | |||
ZNF659 zinc finger protein 659 GeneCard | Wellcome Trust Case control Consortium | |||
ZNF678 zinc finger protein 678 GeneCard | Wellcome Trust Case control Consortium | |||
ZNF804A zinc finger protein 804A | GWAS Williams et al, 2010 | |||
C11orf44 =FLJ39058 |
Genome-wide association study Baum et al, 2007 |
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