PolygenicPathways Updates

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As of 12/02/2011 all genes have been moved to the risk factor page where genes and environmental factors are shown together.

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Genes associated with schizophrenia are grouped into families relating to their general overall function. The tables link to ENTREZ, OMIM and citations  for each gene. Protein/protein interactions are generally culled from information available from the ENTREZ gene hyperlink unless otherwise referenced.  (See below for chromosomal region linkage/gene association concordance).BIOCAR7TA pathway links are provided for some genes (see Terms and conditions) and Disclaimer . GeneCards links are with the kind permission of GeneCards.org: ENVIRONMENTAL RISK FACTORS ARE INCLUDED HERE

Genes2cognition proteomics: Enhanced Links to genes are provided by

 A large number of the protein products of these genes (DLG2, DPYSL2, GRM3, GRM5, GRIN1, GRIN2A, GRIN2B, GRIN2D, GSK3B, HOMER1,HOMER2, LGI1, MOG, NOS1, NDUFV2, PLA2, PLP1, SLC1A2, STX1A, SYNGR1, YWHAH ) are components of the NMDA receptor complex G2C.These and others (CALR, FYN, GABBR1, GAD1, GNAS, GRIA1, GRIA4, GRM3, GRM7, KIF2, NRXN1, NUMBl, PICK1, PDE4B, PIP5K2A, S100B, SLC25A1, SOD2, MAP6, SYN2, SYN3, XRCC1) are components of the postsynaptic density G2C. ADRA1A, ARCVF, APC, DISC1 , DRPLA, NCAM1, NPTN, PAWR, PAX6, RGS4, ZDHHC8, GRIK3, GRIK4 can also be related to elements of the postsynaptic density or the control of NMDA receptor function (see below). About 25% of the genes so far associated with schizophrenia can be related to this complex.
Genes involved in both Bipolar disorder and schizophrenia
PI3K/AKT signalling, growth factors and related ANK3, AKT1, BDNF, BMP6, CABIN1, EGFR, FYN, GSK3B, 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, SNAP25, SYN3, SYNJ1
Dopaminergic/Serotonergic COMT, DRD2, DRD3, HTR2A, HTR5A, HTR6, 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

 

Entrez Gene Links

General Function and protein/protein interactions

Positive association evidence 
Genes controlling glutamate receptor agonist availability  

DAO D-amino acid oxidase

OMIM GeneCards

Metabolises the NMDA receptor glycine site agonist D-serine (Chumakov et al., 2002)

Associated in Chinese, French, German Israeli and Scottish case studies(Chumakov et al., 2002)(Korostishevsky et al., 2004;Liu et al., 2004;Ma et al., 2006;Schumacher et al., 2004)  

DAOA D-amino acid oxidase activator OMIM

GeneCards

Activates DAO (Chumakov et al., 2002)

Associated in American, Chinese, French, German and Israeli case studies (Addington et al., 2004;Chumakov et al., 2002;Korostishevsky et al., 2004;Schumacher et al., 2004;Wang et al., 2004). .
SRR Serine racemase GeneCards

Converts L-serine to D-Serine: Binds to PICK1 (Fujii et al., 2005

)

Association reported in a Japanese study Morita et al, 2006 
CBS cystathionine-beta-synthase converts homocysteine to cystathionine Russian study Golimbet at al, 2009
KMO kynurenine 3-monooxygenase (kynurenine 3-hydroxylase) Tryptophan metabolism: related to hynurenate and quinolinate: NMDA antagonists and agonists respectively Japanese study Aoyama et al, 2006
MTR 5-methyltetrahydrofolate-homocysteine methyltransferase OMIM GeneCards 5-methyltetrahydrofolate + L-homocysteine = tetrahydrofolate +
L-methionine
Association reported in a Polish case study Kempisty et al, 2007
MTHFD1 methylenetetrahydrofolate dehydrogenase (NADP+ dependent) 1, methenyltetrahydrofolate cyclohydrolase, formyltetrahydrofolate synthetase OMIM GeneCards Multifunctional folate metabolism enzyme Association reported in a Polish case study Kempisty et al, 2007

MTHFR Methylenetetrahydrofolate reductase OMIM 607093 GeneCards

Controls homocysteine metabolism: Homocysteine stimulates the NMDA receptor via an action at the glutamate binding site but inhibits NMDA effects via an antagonist effect at the glycine binding site (Lipton et al., 1997). 

Significant association reported in a meta-analysis of 10 case studies (Muntjewerff et al., 2005).   Homocysteine and NMDA

NAALAD2 N-acetylated alpha-linked acidic dipeptidase 2 GeneCards

Hydrolyses N-acetylaspartylglutamate (NAAG) to glutamate and N-acetyl aspartate. NAAG is both a metabotropic receptor agonist (GRM3) and an NMDA receptor partial agonist (Coyle, 1997)

Situated at the chromosome 11 breakpoint of the chromosome1/11 translocation reported to associate with schizophrenia (Semple et al., 2001) N-acetylaspartate schizophrenia
SLC15A1
solute carrier family 15 (oligopeptide transporter), member 1 (PEPT1) GeneCards
Dipeptide transporter whose substrates include NAAG, the NAALAD2 substrate  (Sala-Rabanal et al, 2006) Association reported in Askenazi Jewish families (Fallin et al., 2005) 

PRODH Proline dehydrogenase

OMIM 606810 GeneCards

Proline can be a precursor for glutamate via PRODH

(Thompson et al., 1985)

Association reported in American Chinese and French family studies  (Li et al., 2004a;Liu et al., 2002a),(Jacquet et al., 2002). 

SLC1A2 solute carrier family 1 (glial high affinity glutamate transporter), member 2 OMIM 600300 GeneCards

Glutamate and cysteine transporter
Expression in astrocytes positively modulated by GRM3 and negatively modulated by GRM5 Aronica et al, 2003. Expression increased by BDNF  (Rodriguez-kern et al, 2003) and by AKT1 in astrocytes (Li et al,2006) 

Association reported in a Japanese case study (Deng et al., 2004b).
SLC1A4 solute carrier family 1 (glutamate/neutral amino acid transporter), member 4 Glutamate transporter Japanese study Deng et al, 2008
SLC1A6 solute carrier family 1 (high affinity aspartate/glutamate transporter), member 6 Glutamate transporter Japanese study Deng et al, 2007
SLC6A5 solute carrier family 6 (neurotransmitter transporter, glycine), member 5 Transports the NMDA receptor co-agonist glycine Japanese study Deng et al, 2008
SLC17A7 solute carrier family 17 (sodium-dependent inorganic phosphate cotransporter), member 7 (VGLUT1) Glutamate transporter Taiwanese study of rare variants Shen et al, 2009
SLC17A6 solute carrier family 17 (sodium-dependent inorganic phosphate cotransporter), member 6 VGLUT2   Taiwan study Shen et al, 2010
Table 2: Genes controlling glutamate packaging and release and presynaptic function  

NOSIAP (CAPON) (also postsynaptic)  OMIM 605551 GeneCards

Binds to NOS1 and synapsins (SYN1, SYN2, SYN3)

Linked in American family and associated in Columbian and Chinese case studies (Brzustowicz et al., 2004;Miranda et al., 2005;Zheng et al., 2005)

CPLX2  Complexin II OMIM 605033 GeneCards

Syntaxin binding protein involved in synaptic vesicle exocytosis

Association reported in a Korean case study (Lee et al., 2005).

DTNBP1 Dysbindin OMIM 607145 GeneCards

Positively regulates glutamate release and upregulates the expression 
of presynaptic proteins SYN1, SNAP25 (Numakawa et al., 2004). Also part of the
biogenesis of organelles complex involved in endosome to lysosome transport. Dysbindin or MUTED knockdown blocks D2 dopamine receptor (DRD2) internalisation, increasing cell surface expression Iizuka et al, 2007

Association reported in multiple case and family studies (Straub et al., 2002;Tang et al., 2003b;van den Oord et al., 2003;Van Den et al., 2003;Williams et al., 2004a).
CMYA5 cardiomyopathy associated 5 Binds to dysbindin GWAS 2010
BLOC1S3 biogenesis of lysosome-related organelles complex-1, subunit 3 GeneCards Components of this complex are involved in the biogenesis of organelles including melanosomes and platelet-dense granules. Includes DTNB1 and MUTED inter alia See HERMANSKY-PUDLAK SYNDROME Irish studyMorris et al, 2007
MUTED Muted GeneCards see above. Dysbindin or MUTED knockdown blocks D2 dopamine receptor (DRD2) internalisation, increasing cell surface expression Iizuka et al, 2007 Irish studyMorris et al, 2007

ENTH Epsin 4 OMIM 607262 GeneCards

Transport of synaptic vesicles (Pimm et al., 2005)

Associated with schizophrenia in a British study (Pimm et al., 2005) and in a chinese family study (Tang et al 2006)
SHANK3 SH3 and multiple ankyrin repeat domains 3 Postsynaptic density protein Binds to SPTAN1 (DISC1 partner) and PAX6 Mutations observed in two families Gauthier et al, 2010

SNAP25 synaptosomal-associated protein, 25kDa

 

Synaptic vesicle and transmitter release Irish study Fanous et al, 2010

SNAP29 OMIM 604202 GeneCards

Syntaxin binding protein involved in glutamate release; Binds to STX1A

Association reported in an American case study (Saito et al., 2001)

SYN2 Synapsin II OMIM 600755 GeneCards

Associated with synaptic vesicles and glutamate release. Binds to CAPON and SYN3 .Expression regulated by DRD1 and DRD2 receptors (Chong et al, 2006)

Association in Chinese and Korean case studies (Chen et al., 2004a;Lee et al., 2005) and in a family based chinese study {8923}

SYN3 Synapsin III OMIM 602705 GeneCards

Associated with synaptic vesicles and glutamate release. Binds to CAPON and SYN2

Association reported in American case studies(Lachman et al., 2005;Porton et al., 2004)
SYT11 Synaptotagmin XI GeneCards Family of memrane trafficking proteins involved in transmitter release Association reported in a Japanese study Inoue at al, 2007

STX1A Syntaxin 1a OMIM 186590 GeneCards

Function in vesicle fusion and transmitter release
Binds to complexins, SLC6A3, SLC6A4, SLC6A5 and SNAP29

Association reported in a Canadian case study (Wong et al., 2004).

SYNGR1 Synaptogyrin 1 OMIM 603925 GeneCards

Associated with presynaptic vesicles.

Nonsense mutation in an Indian family (Verma et al., 2004). Association also revealed in non-familial cases (Verma et al., 2005a)

GABBR1 GABA-B receptor 1

OMIM 603540 GeneCards

Agonists inhibit glutamate release.Binds to ATF4 and ATF5 White at al, 2000(as does DISC1).
Also associates with YWHAH (Couve at al, 2001)

Weakly associated with schizophrenia in Japanese family and European case studies (Imai et al., 2002) (Zai et al., 2005).
GABRG2 gamma-aminobutyric acid (GABA) A receptor, gamma 2   Canadian study Zai et al, 2009

GRM3 Metabotropic glutamate receptor mgluR3 OMIM 601115 GeneCards

Binds to PICK1 . Agonists inhibit glutamate release.

Association reported in Japanese and Chinese case studies (Fujii et al., 2003) (Chen et al., 2005) and in a family based American  study (Egan et al., 2004)

GRM4 Metabotropic glutamate receptor mgluR4  OMIM 604100 GeneCards

Agonists inhibit glutamate release

Association reported in Ashenazi Jewish families (Fallin et al., 2005)
GRM7 glutamate receptor, metabotropic 7 GeneCards Binds to PICK1 and TUBA8 Association reported in a Japanese study Ohtsuki et al, 2008

GRM8 Metabotropic glutamate receptor mgluR8 OMIM 601116 GeneCards

Agonists inhibit glutamate release

Associated with schizophrenia in a Japanese case study (Takaki et al., 2004)
APBA2 amyloid beta (A4) precursor protein-binding, family A, member 2 (X11-like) GeneCards Binds to NRXN1 ( see Entrez gene) Duplication observed in a UK study Kirov et al, 2007
NRXN1 Neurexin 1 GeneCards regulates synapse formation Chubykin et al, 2005 Binds to APBA2 (see Entrez gene) Deletion disrupting NRXN1 observed in a UK study Kirov et al, 2007
Other genes whose products affect glutamate release or uptake BDNF enhances depolarization-evoked glutamate release in cortical neurones (Balkowiec and Katz, 2002). Nicotinic CHRNA7 agonists stimulate glutamate release in the hippocampus and frontal cortex (Liu et al., 2003b;Marchi et al., 2002).CNR1 agonists inhibit glutamate  release in a number of brain regions (Wang, 2003) (Gerdeman and Lovinger, 2001). GRM3, GRM4 and GRM8 receptor activation inhibits glutamate release (DD and Marek, 2002;Guo and Ikeda, 2005). HTR2A agonists stimulate frontal cortical glutamate release (Aghajanian and Marek, 1999), HTR6 receptor antagonism selectively increases glutamate release in the frontal cortex and hippocampus (Dawson et al., 2001). DRD2 receptor activation  inhibits striatal and accumbens glutamate release (Bamford et al, 2004, Kalivas and Duffy, 1997))  HTR7 stimulation inhibits glutamate release in the dorsal raphe (Harsing, Jr. et al., 2004). IL1B and TNFA inhibit glutamate uptake into astrocytes and this effect is blocked by NO synthase inhibitors and mimicked by NO donors (Hu et al., 2000;Ye and Sontheimer, 1996). S100B stimulates glutamate uptake into astrocytes (Tramontina et al, 2006). NPY inhibits hippocampal glutamate release and blocks LTP(Whittaker et al., 1999).
Genes coding for glutamate receptors and scaffold proteins, and related.    
CIT Citron kinase Binds to DISC1, GRIN1, GRIN2D USA study Nicodemus et al, 2010
GRIA1 AMPA receptor subunit
OMIM
138248 GeneCards

AMPA receptor

Binds to PICK1,GRIA4, HOMER1

Association reported in an Italian case study Magri et al, 2006
GRIA3 glutamate receptor, ionotrophic, AMPA 3 GeneCards AMPA receptor Association restricted to females in an italian case study Magri et al, 2008

GRIA4 (AMPA receptor subunit )
OMIM
138246
GeneCards

AMPA receptor: Binds to PICK1 (Xia et al., 1999) and GRIA1

Association demonstrated in a Japanese case study(Makino et al., 2003).
GRIK2 glutamate receptor, ionotropic, kainate 2 Kainate receptor subunit  French study Bah et al, 2004

GRIK3 Kainate receptor

OMIM 138243 GeneCards

Kainate receptor subunit

Association reported in an Italian case study (Begni et al., 2002). 

GRIK4 Kainate receptor

OMIM 600282 GeneCards

Kainate receptor subunit Association with bipolar disorder and schizophrenia  reported in a Scottish case study (Pickard et al., 2006)

GRID1 glutamate receptor, ionotropic, delta 1 GeneCards

 

Association reported in Askenazi Jewish families (Fallin et al., 2005) 
GOPC golgi-associated PDZ and coiled-coil motif containing
Binds to GRID1 and GRID2 GWAS Moskvina et al, 2009

GRIN1 (NMDA receptor subunit NR1) OMIM 138249 GeneCards

Binds to GRIN2A, GRIN2B, DLG2

Association reported in American, Chinese and Italian case studies (Begni et al., 2003;Martucci et al., 2003;Qin et al., 2005; Zhao et al, 2006).

GRIN2A NMDA receptor subunit NR2A OMIM 138253 GeneCards

Binds to GRIN1, DLG2

Association reported in two Japanese case studies (Itokawa et al., 2003b;Iwayama-Shigeno et al., 2005)

GRIN2B NMDA receptor subunit NR2B OMIM 138252 GeneCards

Binds to GRIN1, DLG2

Association reported in Japanese Chinese and Italian case studies. (Di Maria et al., 2004;Miyatake et al., 2002;Ohtsuki et al., 2001))

GRIN2D NMDA receptor subunit NR2D OMIM 602717 GeneCards

Binds to GRIN1

Association reported in a Japanese case study (Makino et al., 2005)

GRM5 Metabotropic glutamate receptor mgluR5 OMIM  604102 GeneCards

GRM5 receptor activation potentiates both AMPA and NMDA receptor mediated responses (Ugolini et al., 1999) (Awad et al., 2000;Doherty et al., 2000) Interacts with RGS4 Schwenst and McGinty, 2007

Association in a British case study (Devon et al., 2001). Located at the Chromosome 11 breakpoint associated with schizophrenia (Semple et al., 2001) .
APC adenomatous polyposis coli (colorectal tumor suppressor)
OMIM
175100 GeneCards

Binds to PSD-95 and GRIN2B and is a component of the NMDA receptor  PSD-95 complex  (Yanai et al, 2000)

Association in a Chinese family study(Cui et al, 2005)

ARVCF armadillo repeat gene deleted in velocardiofacial syndrome OMIM 602269 GeneCards

ARVCF binds to the PDZ domain of erbin (ERBB2IP) (Laura et al., 2002) a protein concentrated in postsynaptic membranes which binds to PSD-95 and to the NRG1 receptor ERBB2 (Huang et al., 2001)

Possible association in a Chinese family study (Xie et al., 2005).

ATXN1 (SCA1): ataxin 1) OMIM 601556 GeneCards

Metabotropic mGluR1 (GRM1), NMDA-like  (GRID1/GRID2) and AMPA receptor GRIA2/3 trafficking is altered in mutant ATXN1 transgenic mice (Skinner et al., 2001). Possible role in ubiquitin-proteasome and chaperone system Davidson et al, 2000  

CAG repeats in this gene have been reported to associate with schizophrenia in American case (Joo et al., 1999)or family (Pujana et al., 1997)studies  (Pujana et al., 1997).Association also reported in Ashkenazi Jewish families (Fallin et al., 2005)
DLG1 discs, large homolog 1 Related to GRIN2A, GRIN2B, GRIA1, GRIK2 Japanes study Sato et al, 2008

DLG2 Chapsyn-110 

OMIM 603583 GeneCards

Postsynaptic scaffold protein; Binds to GRIN1, GRIN2A, GRIN2B, NOS1 and NRG1 receptor ERBB4

DLG2 polymorphisms have not been reported in schizophrenia. However, PSZA11q14 is located in the first intron of and is antisense to DLG2 and has been suggested as a candidate gene for schizophrenia (Polesskaya et al., 2003). 

DRPLA dentatorubral-pallidoluysian atrophy or atrophin-1 OMIM 607462 GeneCards

Interacts with an insulin receptor tyrosine kinase substrate BAIAP2 (Okamura-Oho et al., 1999) which is localised in postsynaptic densities and binds to PSD-95 and DLG2 (Choi et al., 2005).

Possible association of a CAG repeat in one family study(Morris-Rosendahl et al., 1997).
EFNB2 Ephrin B2   Chinese study Zhang et al, 2010

HOMER1 OMIM 604798 GeneCards

Controls the localisation and clustering of Group 1 metabotropic glutamate receptors, GRM1 and GRM5 (Ango et al., 2000;Tadokoro et al., 1999). NMDA and GRM5 receptors are linked via HOMER and Shank (Tu et al., 1999). Gene ID 9456

Borderline association in a British case study (Norton et al., 2003).
HOMER2 Homer hololog 2 Regulates GRM1 GRIA1 and GRIK1 Irish study Gilks et al, 2010

LGI1 leucine-rich, glioma inactivated 1 OMIM 604619 GeneCards

Binds to ADAM22 and potentiates AMPA responses in the hippocampus (Fukata et al, 2006 Association reported in an Askenazi case-parent trio study(Fallin et al., 2005)
NPTN neuroplastin GeneCards Inhibits hippocampal LTP by reducing the surface expression of AMPA receptors (Empson et al, 2006) Saito et al, 2006

PICK1 protein kinase C alpha binding protein OMIM 605926 GeneCards

Controls AMPA receptor clustering (Hanley and Henley, 2005) and binds to a number of AMPA receptor subunits including GRIA1 and GRIA4 (Xia et al., 1999).Also binds to the metabotropic receptor GRM3 (Hirbec et al., 2002) , the dopamine transporter SLC6A3 (Torres et al., 2001), NRG1 receptor ERBB2 (Jaulin-Bastard et al., 2001) and to serine racemase (SRR) an enzyme responsible for the synthesis of the NMDA co-agonist D-Serine (Fujii et al., 2005). Gene ID 2893

Association reported in Taiwanese and Japanese case studies (Fujii et al., 2005;Hong et al., 2004b).

RGS4 regulator of G-protein signalling 4 OMIM  602516 GeneCards

Binds to NRG1 receptor ERBB3 (se Gene data). nhibits signaling via GRM1 and GRM5 (Saugstad et al, 1998) Transcription regulated by PHOX2B. Grillet et al, 2003

Suggested as a susceptibility gene in Indian family, Irish case, Israeli family, and Welsh case studies (Chowdari et al., 2002;Fallin et al., 2005;Morris et al., 2004;Williams et al., 2004b)

ZDHHC8 zinc finger, DHHC domain containing 8 OMIM 608784 GeneCards

Localised to vesicle-like clusters along dendritic shafts occasionally colocalising with PSD-95 and the NMDA receptor subunit GRIN2B (Mukai et al., 2004). Gene ID 29801

Associated in American and Chinese case studies (Chen et al., 2004c;Mukai et al., 2004).

NPAS3 neuronal PAS domain protein 3 OMIM 609430 GeneCards

Altered PSD-95 expression and impaired glutamatergic signalling are observed in NPAS3 knockout mice (Brunskill et al., 2005).

Situated in the breakpoint of a t(9;14)(q34;q13) translocation  in a mother and daughter affected by schizophrenia (Kamnasaran et al., 2003).
PPP1R1B protein phosphatase 1, regulatory (inhibitor) subunit 1B (dopamine and cAMP regulated phosphoprotein, DARPP-32) GeneCards

Dopamine D1 receptor simulation increases phosphorlation: Dephosphorylated by AMPA and NMDA receptor activation Nishi et al, 2002

Association observed in an american family-based study.Meyer-Lindenberg et al, 2007

Genes whose products affect NMDA receptor activation or elements of the postsynaptic scaffold

Protein products associated with the postsynaptic density (Genes to Cognition)

AKT1 is activated by NMDA receptor stimulation and phosphorylates NMDA receptors, potentiating their effects (Sanchez-Perez et al, 2006) . BDNF potentiates NMDA receptor mediated responses via a specific action at GRIN2B) (Levine and Kolb, 2000). In the cortex, BDNF stimulates the expression of AMPA receptors and maintains their elevated activity (Balkowiec and Katz, 2002;Nagano et al., 2003). BDNF increases the expression of a number of PDZ domain proteins, including DLG1, GRIP1 and PICK1 in cortical culture (Jourdi et al., 2003). DISC1 binds to elements of the postsynaptic glutamate receptor scaffold (AKAP9, ACTN2, GRIPAP1, HAPIP, SPTAN1) (Millar et al., 2003). GRM5 receptor activation potentiates both AMPA and NMDA receptor mediated responses (Ugolini et al., 1999) (Awad et al., 2000). NRG1 inhibits NMDA receptor mediated responses in the prefrontal cortex and increases the internalisation of the NR1 subunit (GRIN1) of the NMDA receptor (Gu et al., 2005). PAX6 binds to elements of the postsnaptic scaffold (Cooper and Hanson, 2005)

Dopamine can both potentiate and inhibit NMDA responses in the frontal cortex. The stimulatory effects are mediated by DRD1/DRD5 receptor activation (Wirkner et al., 2004) via a direct coupling of DRD1 to GRIN1 and GRIN2A (Lee et al., 2002).  DRD2 and DRD4 agonists repress NMDA responses in the hippocampus via transactivation of the platelet derived growth factor receptor PDGFRA  (Kotecha et al., 2002) (Ferguson, 2003) and DRD4 agonists inhibit NMDA effects in prefrontal cortex (Wang et al., 2003). EGF enhances NMDA receptor mediated responses in the hippocampus (Abe and Saito, 1992) and facilitates hippocampal LTP (Abe et al., 1991;Ishiyama et al., 1991). IL1B inhibits NMDA receptor mediated synaptic responses in the rat dentate gyrus (Coogan and O'Connor, 1997).

Table 4: Genes coding for downstream signalling effectors and related    

HMBS Hydroxymethylbilane synthase (Invoved in heme synthesis)  OMIM 609806 GeneCards

NOS1 (Boehning and Snyder, 2003) and PTGS2 (Percival et al., 1994) are  heme dependent enzymes. HMBS is part of the heme synthesis pathway.

Association reported in an American case study (Sanders et al., 1993)

NOS1Nitric oxide synthase

OMIM 163731 GeneCards

BIOCARTA

Stimulated by NMDA receptor activation via PSD95 (DLG4 Gene ID 1742) which associates with NMDA and NRG1 receptors ERBB2 and ERBB4: Binds to DLG2 and CAPON:

Association described in Japanese and German case studies and in Askenazi Jewish families (Shinkai et al., 2002) (Fallin et al., 2005) (Reif et al., 2006).    GWAS Moskvina et al, 2009

PLA2G4Aphospholipase A2, group IVA (Cytosolic, calcium-dependent)

OMIM 600522 GeneCards

Cytosolic Phospholipase A2 activity is stimulated by the free radical derivative of nitric oxide and superoxide, peroxynitrite (Guidarelli et al., 2000;Tommasini et al., 2002) and by NMDA receptor activation in the striatum  (L'hirondel et al., 1999).

Associated in American case and family, British family and Chinese family studies(Hudson et al., 1996;Tao et al., 2005b;Wei and Hemmings, 2004a;Yu et al., 2005) .

PLA2G4B OMIM 606088 GeneCards

Associated in a Chinese family (Tao et al., 2005a)

PLA2G4C

OMIM 603602 GeneCards

Associated in a Chinese family study (Tao et al., 2005a)
PLA2G6 phospholipase A2, group VI (cytosolic, calcium-independent)   Brazilian study Junquiera et al, 2004

PPP3CC Calcineurin A gamma

OMIM 114107 GeneCards

NMDA receptor activation stimulates NOS1 activity via calcineurin dephosphorylation of the enzyme.

Association in an American family study (Gerber et al., 2003)

PTGS2 Cyclooxygenase-2

OMIM  600262 GeneCards

Activity and expression are increased by NMDA receptor activation (Adams et al., 1996) (Pepicelli et al., 2002) Binds to TP53

Associated with schizophrenia in a British family study (Wei and Hemmings, 2004a)

SOD2 OMIM 147460 GeneCards

SOD2 activity is inhibited by NMDA receptor activation via nitration releasing superoxide into the extracellular space (Muscoli et al., 2004). Binds to TP53 (Zhao et al, 2005)

Association reported in a Turkish case study (Akyol et al., 2005).
Genes involved in neuritogenesis and plasticity    
ALK (anaplastic lymphoma kinase) pleiotrophin receptor  OMIM 105590 GeneCards

Promotes neurite outgrowth (Motegi et al, 2004).

Association in a Japanese case study Kunugi et al, 2006

CHN2 Chimerin (chimaerin) 2 OMIM 602857 GeneCards

CHN2 is induced by BDNF and its overexpression in cerebellar neurones prevents the inhibitory effects of myelin-associated glycoprotein (MAG) on neurite extension (Mizuno et al., 2004).

Associated with schizophrenia inmale Japanese cases (Hashimoto et al., 2005).

DPYSL2 dihydropyrimidinase-like 2 OMIM 602463 GeneCards

A member of the collapsin response mediator protein family involved in neuronal development and axon guidance (Charrier et al., 2003;Quach et al., 2004a). NMDA receptor activation results in increased calpain cleavage of DPYSL2 and the resulting product reduces the surface expression of the NMDA receptor subunit GRIN2B (Bretin et al, 2006) Binds to DISC1 (Camargo et al, 2006)

Polymorphisms in DPYSL2 have been suggested to reduce susceptibility to schizophrenia in a Japanese case study(Nakata et al., 2003).  Also associated with schizophrenia in an Askenazi Jewish family study (Fallin et al., 2005) and in Caucasian and African Americans (Case study) (Hong et al., 2005)

DISC1

Disrupted in schizophrenia 1

OMIM 605210 GeneCards

Involved in neurite outgrowth and cortical development. Binds to other SZ gene products CIT, FEZ1, PCNT, PDE4B, MLC1,MYT1L, NDEL , NDE1 TRAF3IP1 and to a number of proteins associated with the glutamatergic receptor scaffold (ACTN2 ,AKAP9 , SPTAN1) as well as to DPYSL2 Camargo et al, 2006
Also binds to ATF4 and ATF5 transcription factors.  (Millar et al., 2003) as does GABBR1. ATF4 binds to PTGS2 promoter (Schroer et al, 2002) ATF4 binds to nrf1 and nrf2 (NFE2L1 and NFE2L2 ) . NFE2L2 controls the transcription of genes related to glutathione/ quinone oxidative stress including GCLM, GSTM1 (Chanas et al, 2002) and NQO2 (Wang and Jaiswal, 2006) . NFE2L2 controls the expression of BDNF, CHGB, GABBR1, GSTM1, SYN2 in neurones (Lee et al, 2003) and of GSTM1 and PTGS2 in astrocytes (Lee et al, 2003). NFE2L1 controls the expression of TNF Novotny et al, 1998 and also regulates glutathione function Mhyrstad et al, 2001. The actions of NFE2L2 are inhibited by TP53 Faraonio et al,2006

DISC1 and DISC2, a non-coding sequence antisense to DISC1 are within a chromosomal translocation associated with schizophrenia and psychosis in a Scottish family (Millar et al., 2000). Subsequent American, Finnish and Scottish case studies have confirmed an association of DISC1 with schizophrenia (Blackwood and Muir, 2004;Callicott et al., 2005;Cannon et al., 2005;Hennah et al., 2003;Hodgkinson et al., 2004;Millar et al., 2001;Sachs et al., 2005;Zhang et al., 2006). MAJOR HUB GENE

FEZ1 OMIM 604825 GeneCards

Binds to DISC1: Role in neurite development and axonal outgrowth: Binds to BRD1 (Assmann et al, 2006)

Association in a Japanese case study (Yamada et al., 2004b)
KIF2A
kinesin heavy chain member 2A OMIM 602951 GeneCards
Involved in BDNF-mediated neurite extension (Pfenninger et al, 2003) Association  reported in a chinese family study
(Li et al, 2006)
NDE1 LIS1-interacting protein NUDE1GeneCards Binds to DISC1 Female-specific association in Finnish familiesHennah et al, 2006
NDEL1 nudE nuclear distribution gene E homolog (A. nidulans)-like 1 GeneCards Thiol-activated peptidase (EOPA).Role in microtubule organisation and cell migration. Binds to DISC1 DISC1-dependent association in an American study Burdick et al, 2008

NTNG1 Netrin G1 OMIM 608818 GeneCards

Promotes neurite extension in thalamic and cortical neurones (Nakashiba et al., 2002)

Associated with schizophrenia in case and family based Japanese studies (Aoki-Suzuki et al., 2005;Fukasawa et al., 2004)

NTNG2 Netrin G2 GeneCards

Promotes neurite extension in thalamic and cortical neurones (Nakashiba et al., 2002)

Associated with schizophrenia in a Japanese family study (Aoki-Suzuki et al., 2005)

PLXNA2

Plexin A2 OMIM 601054 GeneCards

Semaphorin receptor involved in axon guidance

Associated with schizophrenia in an american case study (Mah et al., 2006)
PNOC Prepronociceptin GeneCards Precursor for nocipeptin/orphanin. Induced by BDNF in thehippocampus and plays a role in neurite outgrowth Ring et al, 2006 Association reported in an Askenazi case-parent trio study Fallin et al., 2005)

RTN4 reticulon 4 (NOGO) 604475 GeneCards

Potent neurite outgrowth inhibitor acting via RTN4R 

A CAA insertion observed in schizophrenia is in the 3' untranslated region of the gene (Novak et al., 2002). Also associated with schizophrenia in females but not males in a Singaporean case study (Tan et al., 2005). .

RTN4R Reticulon 4 receptor OMIM 605566 GeneCards

RTN4 receptor that also binds to MAG. 

Mutations in the RTN4R receptor were found in 3/120 schizophrenic patients and in 0/300 normal subjects. Two of the schizophrenic patients with a missense mutation were resistant to neuroleptic treatment (Sinibaldi et al., 2004).
Other genes controlling neurite/ dendrite morphology APOE (Qiu et al., 2004), BDNF (Alonso et al., 2004),CHL1 (Demyanenko et al., 2004),  CHN2 (Mizuno et al., 2004),CNR1 (Zhou and Song, 2001), DISC1 (Miyoshi et al., 2003),  DPYSL2 (Quach et al., 2004b), EGF (Mark et al., 1995), FEZ1 (Miyoshi et al., 2003), HOMER  (Sala et al., 2001),IL1B (Gilmore et al., 2004),IL2 (Jiang and Lu, 1998), LTP and NMDA receptor activation (GRIN1, GRIN2A, GRIN2B, GRIN2D)(Collin et al., 1997) , MAG (Domeniconi et al., 2002), NPAS3 (Pieper et al., 2005), NRG1 (Gerecke et al., 2004), NTF3 (Baker et al., 1998), RTN4 and RTN4R, TNFA (Gilmore et al., 2004)
Table 6: Growth factors, tyrosine kinase receptors, phoshoinositide kinase/AKT signaling

AKT1 v-akt murine thymoma viral oncogene homolog 1OMIM 164730

BIOCARTA GeneCards

Growth factor signalling.

The AKT1 pathway is also activated by NMDA receptor stimulation and AKT1 phosphorylates NMDA receptors , potentiating their effects (Sanchez-Perez et al., 2006).

Reported as a schizophrenia susceptibility gene in American and Japanese case and Australian family studies (Emamian et al., 2004) (Ikeda et al., 2004;Schwab et al., 2005).   
GSK3B glycogen synthase kinase 3 beta GeneCards Serine threonine kinase involved in multiple signaling networks Association in a Brazilian Case control study Souza et al, 2008

BDNF

brain derived neurotrophic factor

OMIM 113505 GeneCards

BDNF release is stimulated by NMDA receptor activation in the hippocampus  (Jiang et al., 2005).BDNF enhances depolarization-evoked glutamate release in cortical neurones (Balkowiec and Katz, 2002).BDNF potentiates NMDA receptor mediated responses via a specific action at GRIN2B) (Levine and Kolb, 2000). In the cortex, BDNF stimulates the expression of AMPA receptors and maintains their elevated activity (Balkowiec and Katz, 2002;Nagano et al., 2003). BDNF increases the expression of a number of PDZ domain proteins, including DLG1, GRIP1 and PICK1 in cortical culture (Jourdi et al., 2003). Enhances phosphorylation of GRIN1 (Suen et al, 1997) GRIN2B (Lin et al, 1998) and GRIA1 (Wu et al, 2004). BDNF increases SLC1A2  expression in astrocytes (Rodriguez-kern et al, 2003)

Involved in oligodendrocyte differentiation

Reported association in American case and family, French, Hungarian and Scottish case and  Spanish family studies .(Fanous et al., 2004b;Krebs et al., 2000a;Krebs et al., 2000b;Muglia et al., 2003;Neves-Pereira et al., 2005;Rosa et al., 2006;Szekeres et al., 2003)
BMP6 bone morphogenetic protein 6   Taiwan study Lin et al, 2009

CNTF ciliary neurotrophic factor OMIM 118945 GeneCards

Growth factor: Potent survival factor for neurones astrocytes and oligodendrocytes; Promotes neurite extension. OLIG2 nuclear export in astrocytes is induced by CNTF via phosphorylation of OLIG2 by AKT1 (Setoguchi and Kondo 2004)

Binds to APOE

Null mutation  (Lin and Tsai, 2004;Thome et al., 1996)
DDR1 discoidin domain receptor family, member 1 GeneCards Tyrosine kinase expressed in oligodendrocytes and activated by collagens Franco-Pons et al, 2006 Association reported in a Spanish case study Roig et al, 2007
DGKI diacylglycerol kinase, iota Final step of PI3 kinase pathway: Phosphotylates diacylglycerol to phosphatidic acid GWAS Moskvina et al, 2009
EFNB2 Ephrin B2 Tyrosine kinase Association in a Chinese study Zhang et al, 2010

EGF Epidermal growth factor OMIM 131530 GeneCards

NRG1 inhibits EGF binding to either EGFR or the Neuregulin receptor ERBB2.

Increases expression of SLC1A2 in astrocytes (Zelenaia et al, 2000)

Involved in oligodendrocyte differentiation

Associated in a Finnish case study (Anttila et al., 2004). In one case study the association was conditional on a concurrent TNFA polymorphism (Kampman et al., 2005) In another case study association was only reported in men (Hanninen et al., 2005a) .
EGFR EGF receptor GeneCards EGF/ NRG1, ERBB2, ERBB3 interact Benzel et al, 2007
EGR2 early growth response 2 (Krox-20 homolog, Drosophila) GeneCards Important role in peripheral myelination: Upregulated by NMDA receptor activation during hippocampal long-term potentiation Inokuchi et al, 1996 Yamada et al, 2007
EGR3 early growth response 3 GeneCards Synthesis indiced by BDNF in hippocampus Roberts et al, 2006 by NMDA receptor activation Yamagata et al, 1994 and by NRG1 in muscle Jacobson et al, 2004 Yamada et al, 2007
EGR4 early growth response 4 GeneCards   Yamada et al, 2007
FGF1 fibroblast growth factor 1 (acidic) GeneCards Plays a role in oligodendrocyte myelination (inter alia) Harari et al, 1997 Weak association in a Dutch case-control studyJungerius et al, 2007
FGF14 Fibroblast growth factor 14   Jungerius eyt al, 2007
FGF18 Fibroblast growth factor 18 Binds to FGFR1, 2, 3 and 4 Jungerius eyt al, 2007
FGFR1 fibroblast growth factor receptor 1 (fms-related tyrosine kinase 2, Pfeiffer syndrome) GeneCards FGF1 and FGF2 receptor Weak association in a Dutch case-control studyJungerius et al, 2007
FYN FYN oncogene related to SRC, FGR, YES GeneCards

Associated with BDNF receptor signalling (NTRK2) Iwasaki et al, 1998. Associates with GRIN2A and GRIN2B Takagi et al, 1999 BDNF enhances NMDA currents via FYN Xu et al, 2006. Involved in NCAM1 signalling Beggs et al, 1997

Rybakowski et al, 2007
GDNF Glial derived neurotrophic factor GeneCards Ligand for GFRA1 and 2 Association reported in an Italian study Michelato et al, 2004
GFRA1
GDNF family receptor alpha 1GeneCards
GDNF and neurturin receptor. Possible involvement in oligodendrocyte development (Strelau and Unsicker 1999). Expression regulated by TNF in glial cells (Remy et al, 2003). Involved in the development and migration of cortical GABA neurones (Pozas and Ibanez,2005) Association reported in an Ashkenazi Jewish case-parent trio study
(Fallin et al., 2005).
GFRA2 GDNF family receptor alpha 2 GDNF and neurturin receptor Canadian study Souza et al, 2010
GFRA3 GDNF family receptor alpha 3 Artemin receptor Canadian study Souza et al, 2010
IGF2 insulin-like growth factor 2 (somatomedin A)   Croatian study Pal et al, 2009
LIF leukemia inhibitory factor (cholinergic differentiation factor)   Japanese study Okahisa et al, 2010
METmet proto-oncogene (hepatocyte growth factor receptor)   USA study Burdick et al, 2010
NEUROG1 Neurogenin 1   US study Fanous et al, 2007

NRG1 Neuregulin 1 OMIM 142445 GeneCards

Glial growth factor: NRG1 inhibits NMDA receptor mediated responses in the prefrontal cortex and increases the internalisation  of GRIN1  (Gu et al., 2005). NRG1 receptor ERBB2, binds to the AMPA receptor clustering protein PICK1 (Jaulin-Bastard et al., 2001). A cleaved intracellular domain of NRG1 translocates to the nucleus binds to the PSD-95 (DLG4) promoter (Bao et al, 2004)

Association reported in Chinese, Icelandic, Portugese, Scottish and Welsh  populations (Li et al., 2004b;Norton et al., 2005;Petryshen et al., 2005;Stefansson et al., 2002;Stefansson et al., 2003;Tang et al., 2003a;Williams et al., 2003b;Yang et al., 2003b).(Case and family studies).
NRG2 Neuregulin 2 GeneCards Binds to ERBB3 and ERBB4 Benzel et al, 2007
NRG3 Neuregulin 3 GeneCards Oligodendrocyte growth factor (Carteron et al, 2006): Binds to ERBB4 Association reported in an Ashkenazi Jewish case-parent trio study
(Fallin et al., 2005).
ERBB2 v-erb-b2 erythroblastic leukemia viral oncogene homolog 2, neuro/glioblastoma derived oncogene homolog GeneCards Heterodimerises with EGFR or ERBB3 Benzel et al, 2007
ERBB3 v-erb-b2 erythroblastic leukemia viral oncogene homolog 3 Neuregulin receptor interacting with EGFR and ERBB2 Chinese study Li et al, 2009

ERBB4 Neuregulin receptor OMIM 600543 GeneCards

Binds to NRG1 , DLG2 and PSD-95 (DLG4) . NRG1 induces the expression of NOS1 in cerebellar granule cells via ERBB4 (Krainock and Murphy, 2001b). A cleaved intracellular domain of ERBB4 phosphorylates the p53 activator MDM2 and increases TP53 levels in peripheral cells  (Arasada and Carpenter 2005) 

Interaction between NRG1 and ERBB4 in a UK casestudy (Norton et al., 2005). Association also reported in an Israeli case study(Silberberg et al., 2006).

IMPA2 inositol(myo)-1(or 4)-monophosphatase 2 OMIM 605922 GeneCards

PI signalling and inositol phosphate metabolism

IMPA2 associates with schizophrenia in a Japanese case study (Yoshikawa et al., 2001).
JARID2 jumonji, AT rich interactive domain 2 OMIM GeneCards Negative regulation of cell growth Toyoda et al, 2000 Pedrosa et al,2006
NRN1 Neuritin 1   Australian study Chandler et al, 2010

NTF3 neurotrophin 3 OMIM 162660 GeneCards

Neurotrophin; Ligand for NTRK3: Promotes oligodendrocyte survival

NTF3 polymorphisms associating with schizophrenia have been reported in Japanese case studies (Hattori and Nanko, 1995;Nanko et al., 1994) and in male schizophrenics in England (case study) (Dawson et al., 1995).  A metaanalysis combining case results from Caucasian patients with a Swedish study provided support for an association   between NTF3 and schizophrenia (Jonsson et al., 1997).
NTRK3 neurotrophic tyrosine kinase, receptor, type 3  Receptor for NTF3 GWAS European ancestry Shi et al, 2009
PDGFRB platelet-derived growth factor receptor, beta polypeptide GeneCards Binds to EGFR and FYN Association in a Korean study Kim et al, 2008
PIK3C2G phosphoinositide-3-kinase, class 2, gamma polypeptide GeneCards PI3 kinase: Not fully characterised Weak association in a Dutch case-control studyJungerius et al, 2007

PIK3C3 phosphoinositide-3-kinase, class 3 OMIM 602609 GeneCards

Converts Phosphatidylinositol to Phosphatidylinositol 3 phosphate (Volinia et al. 1995) Involved in EGF receptor endocytosis (Zheleznova et al, 2003)
A promoter variant of PIK3C3 is associated with bipolar disorder and schizophrenia in case studies(Stopkova et al., 2004). Also associated in a family based chinese study (Duan et al 2005)
PI4KA phosphatidylinositol 4-kinase, catalytic, alpha GeneCards Catalyzes the first step in the biosynthesis of phosphatidylinositol 4,5-bisphosphate Association in a Dutch case-control studyJungerius et al, 2007
PIK42B phosphatidylinositol 4-kinase type 2 beta   Scottish study Houlihan et al, 2009
PI4KA phosphatidylinositol 4-kinase, catalytic, alpha (PIK4CA) Synthesis of phosphatidylinositol 4,5-bisphosphate Dutch study Jungerius et al, 2007
PIP4K2A phosphatidylinositol-5-phosphate 4-kinase, type II, alpha   Irish study Thistleton et al, 2010

PIP5K2A

phosphatidylinositol-4-phosphate 5-kinase, type II, alpha OMIM 603140 GeneCards

Metabolises Phosphatidylinositol 3 phosphate to phosphatidylinositol 3,4-biphosphate and phosphatidylinositol 3,4,5-triphosphate (Zhang et al. 1997). Also metabolises 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.Phosphatidylinositol 3,4,5-triphosphate inhibits the GTPase activating effects of RGS4 Ishii et al,2002

Associated with both bipolar disorder and schizophrenia in case studies (Stopkova et al., 2003).
PLCB1 phospholipase C, beta 1 (phosphoinositide-specific) catalyzes the formation of inositol 1,4,5-trisphosphate and diacylglycerol from phosphatidylinositol 4,5-bisphosphate GWAS Moskvina et al, 2009
PTPRZ1 protein tyrosine phosphatase, receptor-type, Z polypeptide 1 OMIM GeneCards Involved in NRG1/ERBB4 signaling Buxbaum et al, 2007 Activates ALK Perez-Pinera et al, 2007 and FYN Pariser et al, 2005 LTP enhanced in knockout miceNiisato et al, 2005 UK case studyBuxbaum et al, 2007
SEMA3D Semaphorin 3D   Japanese study Fuji et al, 2010
SPRY4 sprouty homolog 4 GeneCards Negative regulator of FGF signaling and of the MAPK pathway Association in UK trios Zaharieva et al, 2008
WDFY2 WD repeat and FYVE domain containing 2 Binds to AKT1 GWAS Moskvina et al, 2009
Oligodendrocyte related genes    
ARHGEF10 Rho guanine nucleotide exchange factor (GEF) 10 GeneCards Role in myelination of peripheral nerves Weak association in a Dutch case-control studyJungerius et al, 2007
CNP 2',3'-cyclic nucleotide 3' phosphodiesterase  OMIM 123830 GeneCards

Almost exclusively localised in myelin containing cells

Described as a susceptibility gene in a UK case study (Peirce et al., 2006)

GNPAT glyceronephosphate O-acyltransferase

OMIM 602744 GeneCards

Involved in the synthesis of plasmalogens which are important constrituents of myelin. Abnormal myelinisation is observed in GNPAT deficiency (Szrriha et al, 1997)

 

Polymorphism reported in Taiwan (Lin et al, 2006)
MAG myelin-associated glycoprotein OMIM 159460 GeneCards

Mediates myelin-neuron cell-cell interactions; Inhibits neurite extension via RTN4R
Splicing controlled by QK1. Coimmunoprecipitates with MBP and PLP1.Also binds to FYN and PLCG1 Jaramillo et al, 1994
Arvanitis et al, 2002

 

Polymorphism associated with schizophrenia in a chinese family study (Wan et al., 2005)
MOG myelin oligodendrocyte glycoprotein OMIM 159465 GeneCards

May be involved in completion and maintenance of the myelin sheath and in cell-cell communication. Recognised by HLA-DRB1.

Associated with schizophrenia in a Chinese family study (Liu et al., 2005b).
CSPG5 Neuroglycan C Midkine receptor: Promotes oligodedrocyte growth Ishihara_tanaka et al, 2006 Chinese study So et al, 2010
MYT1L myelin transcription factor 1-like Binds to DISC1 Copy number variation Vrijenhioek et al, 2008

NOTCH4 Notch homolog 4 OMIM 164951 GeneCards

NOTCH signalling in general plays an important role in oligodendrocyte development (Park and Appel, 2003;Popko, 2003) but the role of NOTCH4  has not been specifically examined.Ckeaved by PSEN2

Association reported in African American case and family, Indian family and Chinese mixed pedigree studies (Luo et al., 2004;Prasad et al., 2004;Skol et al., 2003;Wang et al., 2005). Japanese and Chinese studies suggest association only in subsets of early onset patients and/or with negative symptoms (Takahashi et al., 2003) and in a GWAS study Stefanssonnet al, 2009.
OLIG2  
oligodendrocyte lineage transcription factor 2
OMIM 606386 GeneCards
Key transcription factor in oligodendrocyte development (Ligon et al, 2006). Inhibits astrocyte differentiation  and reactive astrocytosis (Cassiani-Ingoni et a, 2006). Transport from nucleus to cytoplasm may be needed in relation to inhibitory effects on astrocyte  differentiation.This export is induced by CNTF via phosphorylation of OLIG2 by AKT1 (Setoguchi and Kondo 2004). See also PAX6 and  PHOX2B interactions with OLIG2 in development Association reported in a UK case study
Georgieva et al, 2006

PAX6paired box gene 6 (aniridia, keratitis) OMIM 607108 GeneCards

Regulates oligodendrocyte development and NRG1 expression (Assimacopoulos et al., 2003;Sun et al., 1998). Binds to HOMER3 ( a partner of GRM5), DYNLL1 (a NOS1 inhibitor) and TRIM11  (Cooper and Hanson 2005) and SHANK3

Promoter polymorphism associates with schizophrenia in a German case study (Stober et al., 1999).

PLP1 OMIM 300401 GeneCards

Proteolipid protein with role in myelinisation. Coimmunoprecipitates with MAG and MBP. Arvanitis et al, 2002

Weak association in males in a family-based chinese study (Qin et al., 2005c)
PSAP Prosaposin GeneCards Myelinotrophic factor Weak association in a Dutch case-control studyJungerius et al, 2007
SOX10 SRY (sex determining region Y)-box 10 OMIM GeneCards Transcription factor with a key role in oligodendrocyte development. Targets include CNTF Ito et al, 2006, ERBB3 Britsch et al, 2001, MBP Wei et al, 2004 TYR Murisier et al, 2007. Regulated by QK1 Aberg et al, 2006 Associated in a Japanese study Maeno et al, 2007
TF Transferrin GeneCards Iron transport: Important role in oligodendrocyte function Connor and Menzies, 1996 Association in the Chinese Han population Qu et al, 2007

QK1 Quaking OMIM 609590 GeneCards

Required for oligodendrocyte differentiation and myelinisation in mice.   Controls the messenger RNA translation or splicing of both myelin basic protein and MAG (Ryder and Williamson, 2004;Wu et al., 2002) Regulates PLP1, MAG, MBP, TF, SOX10, and CDKN1BAberg et al, 2006. Binds to PTPB2 Lim et al, 2006

Associated with schizophrenia in Swedish family-based study (Aberg et al., 2005)

Genes whose products kill oligodendrocytes

AMPA (Yoshioka et al., 1995), kainate (Sanchez-Gomez and Matute, 1999) and NMDA (Salter and Fern, 2005) receptor activation (cf GRIA4, GRIK3. GRIN1, 2A,2B,2D), dopamine (Khorchid et al., 2002),  IL1B (Takahashi et al., 2003a) , LTA)(Plo et al., 1999), Lysophosphatidylcholine  (Fressinaud and Vallat, 1994) a product of PLA2G4,  Nitric oxide (Rosenberg et al., 1999) or peroxynitrite (Mitrovic et al., 1996;Scott et al., 2003) products of NOS1,  TGF beta (Schuster et al., 2002;Schuster et al., 2003), which signals via PCQAP (Kato et al., 2002) TNFA (D'Souza et al., 1996), TP53 (Eizenberg et al., 1995).

Genes whose products protect oligodendrocytes or are involved in their development

CNR1 agonists (Molina-Holgado et al., 2002), CNTF DRD2 and DRD3 receptor activation (Rosin et al., 2005).GRM5 receptor agonists (Deng et al., 2004a). Inhibition of the PI3 kinase cell survival pathway produces apoptosis in oligodendrocytes (Vemuri and McMorris, 1996) (cf AKT1 , PIK3C3, PIP5K2A). BDNF, (Du et al., 2003),  NOTCH signalling (Park and Appel, 2003) (cf NOTCH4 ), NTF3  (Barres et al., 1994;Heinrich et al., 1999;Kumar et al., 1998), NRG1 (Calaora et al., 2001;Fernandez et al., 2000;Flores et al., 2000), PAX6 (Sun et al., 1998), IL2  (Jiang and Lu, 1998), IL10 (Molina-Holgado et al., 2001) S100B (Deloulme et al., 2004). Glutathione (cf SLC1A2, GSTM1) protects against diverse toxic insults(Liu et al., 2002b) (Khorchid et al., 2002;Lee et al., 2004) (Richter-Landsberg and Vollgraf, 1998). Nitric oxide (NOS1) protects oligodendrocytes in conditions of thiol depletion (Rosenberg et al., 1999).
  Dopaminergic system genes    

ALDH3B1 Aldehyde dehydrogenase 3 family, member B1 OMIM 600466 GeneCards

Multiple substrates include catecholamine metabolites.

Association in Chinese case studies (Sun et al., 2005b;Xu et al., 2004):

COMT Catechol-O-methyltransferase OMIM 116790 GeneCards

Methylates catecholamines and catecholamine derived quinones (Zhu, 2004).  

COMT activation also increases the synthesis and release of homocysteine from astrocytes  (Huang et al, 2005) (related to S-adenosylmethionine metabolism)

A meta-analysis of case and family based data suggested a stronger effect in patients of European ancestry  (Glatt et al., 2003a). See (Munafo et al., 2005) for conflicting conclusion.
CHGA Chromogranin A GeneCards Role in tranmitter/hormone secretion in neural and endocrine cells Association in a Japanese case-control Takahashi et al, 2006

CHGB Chromogranin B OMIM 118920 GeneCards

Plays a role in dopamine storage.

Suggested as a susceptibility gene in Japanese and Chinese case studies (Kitao et al., 2000) (Zhang et al., 2002)
DBH Dopamine beta hydroxylase Cinverts dopamine to noradrenaline Croatian study Pal et al, 2009

SLC6A3 DAT1 dopamine transporter OMIM 126455 GeneCards

Binds to PICK1 and STX1A

Reported association in American case and family and Chinese case studies (Blum et al., 1997;Fanous et al., 2004a;Zhang et al., 2003)

DRD2 Dopamine receptor OMIM 126450 IUPHAR GeneCards

Target of most neuroleptics: Dysbindin or MUTED knockdown blocks D2 dopamine receptor (DRD2) internalisation, increasing cell surface expression Iizuka et al, 2007. Agonists attenuate oligodendrocyte damage induced by glutamate or oxidative stress Roisin et al, 2005

Significant association in a meta-analysis of 24 case studies (Glatt et al., 2003b)

DRD3 Dopamine receptor OMIM 126451 IUPHAR GeneCards

Agonists attenuate oligodendrocyte damage induced by glutamate or oxidative stress Roisin et al, 2005

Significant association reported in a meta-analysis of over 30 case studies (Williams et al., 1998)

DRD4 Dopamine receptor OMIM 126452 IUPHAR GeneCards

No overall association in a meta-analysis of 19 worldwide independent samples but significant association reported in Caucasian  subjects especialy in family studies (Lung et al., 2002;Williams et al., 1998)

DRD5 Dopamine receptor OMIM 126453 IUPHAR GeneCards

Association in two British case studies (Williams et al., 1997) (Muir et al., 2001)

KCNN3 small conductance calcium-activated potassium channel, (Alias KCa2.3, SK3, SKCA3, hSK3) OMIM 602983 GeneCards

Controls the firing rate of dopaminergic neurones in the substantia nigra (Wolfart et al., 2001)  

Associated with schizophrenia in Israeli ,Indian, Serbian and Taiwanese case studies (Dror et al., 1999;Ivkovic et al., 2006;Ritsner et al., 2002;Saleem et al., 2000;Tsai et al., 1999a)

MAOA Monoamine oxidase A OMIM 309850 GeneCards

Oxidises monoamines, generating hydrogen peroxide

RCH2NH2 + H2O + O2 = RCHO + NH3 + H2O2

Association reported in British and Swedish case studies (Jonsson et al., 2003;Wei and Hemmings, 1999).
MAOB monoamine oxidase B   Spanish study Carrera et al, 2008
RNLS FAD-dependent amine oxidase (c10orf59)MAOC Metabolises catecholamines Luft, 2005 GWAS African American Shi et al, 2009
NCKAP5 NCK-associated protein 5 Partner binds to dopamine receptors GWAS Wang et al, 2010

PAHPhenylalanine hydroxylase OMIM 261600 GeneCards

L-phenylalanine + tetrahydrobiopterin + O2 = L-tyrosine +
4a-hydroxytetrahydrobiopterin

Weak association reported in an American sib-pair study (Wilcox et al., 2002).
SLC18A1 solute carrier family 18 (vesicular monoamine), member 1 GeneCards Dopamine, noradrenaline and serotonin vesicular uptake Association reported in a Japanese study Richards et al, 2006
SLC18A2 solute carrier family 18 (vesicular monoamine), member 2 GeneCards   Association reported in a spanish studyGutierrez et al, 2007

TH Tyrosine hydroxylase OMIM 191290 GeneCards

L-tyrosine + tetrahydrobiopterin + O2 =
L-DOPA + 4a-hydroxytetrahydrobiopterin

Association reported in French, Japanese , Korean and Welsh case studies  (Kurumaji et al., 2001;Meloni et al., 1995;Pae et al., 2003b;Thibaut et al., 1997;Wei et al., 1995)

CCKAR cholecystokinin A receptor OMIM 118444 GeneCards

Regulates dopamine release

Association reported in Japanese and Spanish case studies and weak (Tachikawa et al., 2000;Wang et al., 2002;Zhang et al., 2000) association reported in an America  case study (Sanjuan et al., 2004)

Glutathione, quinone and oxidative stress related genes

For an interesting history on quinones and schizophrenia see Smythies 1997 .
GCLC glutamate-cysteine ligase, catalytic subunit GeneCards First rate limiting enzyme of glutathione synthesis. Expression controlled by NFE2L2, binding partner of ATF4 (see DISC1, GABBR1): Glutathione depletion impairs synaptic plasticity (LTP) and reduces NMDA receptor-mediated effects Steullet et al, 2006 Association reported in a Swiss study Gysin et al, 2007
GCLM
glutamate-cysteine ligase, modifier subunit
OMIM 600176
GeneCards
First rate limiting enzyme of glutathione synthesis. Expression controlled by NFE2L2, binding partner of ATF4 (see DISC1, GABBR1) Association reported in two case and one family study
Tosic et al, 2006
GSS Glutathione synthase ATP-dependent conversion of gamma-L-glutamyl-L-cysteine to glutathione Swiss Study Tosic et al, 2006

GSTM1 Glutathione transferase M1 OMIM 138350 GeneCards

Cyclised quinones formed from dopamine  or other catecholamines are conjugated by glutathione, a reaction catalysed principally by the mu class of glutathione transferases (Baez et al., 1997) including GSTM1. Expression controlled by NFE2L2, binding partner of ATF4 (see DISC1, GABBR1)

A GSTM1 deletion has been associated with schizophrenia in Japanese and Korean case studies (Harada et al., 2001;Pae et al., 2004).
GSTT1 glutathione S-transferase theta 1 GeneCards   Saadat et al, 2007

HP Haptoglobin OMIM GeneCards

Potent antioxidant: Binds to hemoglobin preventing induction of oxidative stress. Association reported in a Chinese study Wan et al, 2007

MDR1/ABCB1 P-glycoprotein 1 OMIM GeneCards

Involved in NO-related glutathione efflux Watts et al, 2006

 

Association reported in a female Croatian case-study Bozina et al, 2006 GWAS Moskvina et al, 2009
ME2 malic enzyme 2, NAD(+)-dependent, mitochondrial GeneCards

Plays an important role in regenerating reduced glutathione Vogel et al, 1999

Lee et al, 2007 (psychosis and mania)
MSRA Methionine sulfoxide reductase GeneCards Converts methionine sulfoxide to methionine: Possible role in repairing damage to oxidised proteins Costa Rica Family study Walss-Bass et al, 2008

NQO2 NAD(P)H dehydrogenase, quinone 2 OMIM 160998 GeneCards

Quinone reductases play a role in eliminating dopamine-derived quinones (Graumann et al., 2002).Expression controlled by NFE2L2, binding partner of ATF4 (see DISC1, GABBR1)

A deletion in the promoter region of the NQO2 gene is associated with schizophrenia in a Japanese case study (Harada et al., 2003).

S100B Catecholase

OMIM 176990 GeneCards

Converts DOPA to dopachrome, adrenaline to adrenochrome and noradrenaline to noradrenachrome (Senior et al., 2003). Role in neurite extension and astrocytosis and oligodendrocyte development (Deloulme et al., 2004) Binds to TP53

Associated with schizophrenia in a Chinese case study (Liu et al., 2005a)

SOD2 Manganese-dependent superoxide dismutase OMIM 147460 GeneCards

2 superoxide + 2 H(+) <=> O2 + H2O

Binds to TP53 in mitochondria (Zhao et al, 2005)

Association reported in a Turkish case study (Akyol et al., 2005).

TYR Tyrosinase OMIM 606933 GeneCards

L-tyrosine + L-dopa + O(2) <=> L-dopa + dopaquinone + H(2)O. Expression controlled by TP53  (Khlgatian et al, 2002)

Schizophrenia has been linked to a chromosomal translocation at (1:11)(q42.1, q14.3). Genes within the chromosome 11 breakpoint include GRM5, NAALADase II  and TYR (Petit et al., 1999;Semple et al., 2001).

TP53 tumor protein p53 OMIM 191170 GeneCards

Numerous binding partners including PTGS2, SOD2  and S100B. Controls TYR expression (Khlgatian et al, 2002). A cleaved intracellular domain of ERBB4 phosphorylates the p53 activator MDM2 and increases TP53 levels in peripheral cells  (Arasada and Carpenter 2005) . Activated by nitric oxide (Zhou et al, 2004).The actions of NFE2L2 (see DISC1) are inhibited by TP53 Faraonio et al,2006

Associated with schizophrenia in Canadian case and family studies, Chinese and Korean case studies and portugese families (Ni et al., 2005;Park et al., 2004;Yang et al., 2004).
TP53I3 tumor protein p53 inducible protein 3 Binds to TP53:Possible quinone reductase GWAS Moskvina et al, 2009

ND4 NADH dehydrogenase subunit 4 OMIM 516003 GeneCards

Oxidative phosphorylation Complex 1

Reported to associate with schizophrenia, particularly in males in a UK case study(Marchbanks et al., 2003).
NDUFV2
24-kDa subunit of Complex I; NADH dehydrogenase (ubiquinone) flavoprotein 2 (24kD) GeneCards
Oxidative phosphorylation Complex 1 Association reported in a Japanese study (Waskizuka et al, 2006)
UCP2
Uncoupling protein 2 OMIM 601693 GeneCards
Regulates mitochondrial glutathione levels in microglia (de Bilbao et al, 2004. Binds to YWHAZ Assocation reported in a Japanese study (Yasuno et al, 2006)
UCP4 (SLC25A27)
Uncoupling protein 4 GeneCards
Mitochondrial uncoupling  reduces the generation of reactive oxygen species and increases the resistance of cells to oxidative stress (Liu et al, 2006) Assocation reported in a Japanese study (Yasuno et al, 2006)

Other related genes: SLC1A2 mediates the glial uptake of the glutathione precursor cysteine and controls glutathione levels (Chen and Swanson, 2003). Quinones can be formed spontaneously from the oxidation of dopamine or from the enzymatic catalysis of dopamine by PTGS2 (arachidonate or H2O2 + dopamine leading to the formation of aminochrome) (Hastings, 1995). COMT methylation of catecholamines prevents quinone generation and COMT methylates dopamine-derived quinones (Zhu, 2004). 

AKT1, BDNF, CAPON, CCKAR, CHRNA7, CNR1, COMT, DNTBP1, GAD1, GRM3, IL10, MLC1, NOTCH4, NRG1, PRODH, RGS4, RTN4 and TNF, are subject to regulation by hypoxia (Schmidt-Kastner et al, 2006).

Serotonin related    
HTR1A 5-hydroxytryptamine (serotonin) receptor 1A   US study Huang et al, 2004
HTR1B 5-hydroxytryptamine (serotonin) receptor 1B   Croatian Study Pal et al, 2009

HTR2A Serotonin receptor OMIM 182135 IUPHAR GeneCards

Binds to PSD-95
(DLG4)

Significant association in a meta-analysis of 31 case-control association studies (Abdolmaleky et al., 2004).
HTR2C 5-hydroxytryptamine (serotonin) receptor 2C   Croatian Study Pal et al, 2009
HTR4 5-hydroxytryptamine (serotonin) receptor 4   Japanese study Suzuki et al, 2003

HTR5A  Serotonin receptor OMIM 601305 IUPHAR GeneCards

Association reported in Japan (case) and France (family) (Iwata et al., 2001; Dubertret et al 2004)

HTR6 Serotonin receptor OMIM 601109 IUPHAR GeneCards

Association reported in Taiwan (case study)(Tsai et al., 1999b).

HTR7 Serotonin receptor OMIM 182137 IUPHAR GeneCards

Association reported in Japan (case study)(Ikeda et al., 2005)

SLC6A4 Serotonin transporter OMIM 182138 GeneCards

Binds to STX1A

Meta-analysis of 10 separate case studies supports association.
TDO2 tryptophan 2,3-dioxygenase   US Study Miller et al, 2009

TPH1 Tryptophan hydroxylase OMIM 191060 GeneCards

Activated by YWHAH

Association reported in American, Japanese and Taiwanese case studies (Hong et al., 2001;Nolan et al., 2000;Sekizawa et al., 2004;Shinkai et al., 2000)
YWHAE tyrosine 3-monooxygenase/tryptophan 5-monooxygenase activation protein, epsilon polypeptide [Homo sapiens]   Japanese study Ikeda et al, 2008

YWHAH tyrosine 3-monooxygenase/tryptophan 5-monooxygenase-activation protein, eta polypeptide, 14-3-3 eta. OMIM 113508 GeneCards

Family members activate TH and TPH1 (inter alia). Associates with GABBR1 (Couve at al, 2001)

Associates with schizophrenia in a family-based Canadian study  (Wong et al., 2003) and with early onset schizophrenia in a Japane case study (Toyooka et al., 1999) .
GABA-A receptors and other GABAergic (see above for GABBR1)    
GAD1 Glutamate decarboxylase 1 OMIM  605363 GeneCards
GABA synthesis
Associated with childhood-onset schizophrenia in an American family based study (Addington et al., 2005)
GABRA1 gamma-aminobutyric acid (GABA) A receptor, alpha 1 GeneCards GABA-A receptors are formed from multiple permutations of diverse subunits, of which these are members Bollan et al, 2003 Petryshen et al, 2005
GABRA6 gamma-aminobutyric acid (GABA) A receptor, alpha 6 GeneCards Petryshen et al, 2005
GABRB2 gamma-aminobutyric acid (GABA) A receptor, beta 2 GeneCards Lo et al, 2004
GABRP gamma-aminobutyric acid (GABA) A receptor, pi GeneCards Petryshen et al, 2005
Other Receptors, peptides, downstream signalling and neurotransmitter metabolising enzymes    

ACE Angiotensin converting enzyme (synthesises angiotensin II) OMIM 106180 GeneCards

Converts angiotensin 1 to angiotensin 2 

A strong interaction between a low activity COMT genotype and a high activity ACE genotype has been observed in schizophrenic patients non-responsive to neuroleptics. The risk of possessing this combination of polymorphisms is 10 times higher in the schizophrenic non-responsive group (Illi et al., 2003).
ADCYAP1 adenylate cyclase activating polypeptide 1 (pituitary)(PACAP) OMIM GeneCards Enhances NMDA receptor function and increases BDNF expression Yaka et al, 2003 Influences oligodendrocyte proliferation Lelievre et al, 2006. Involved in DISC1 signalingHattori et al, 2007 Hashimoto et al, 2007
ADIPOR2 adiponectin receptor 2   GWAS European ancestry Shi et al, 2009
ADORA1Adenosine A1 receptor Interacts with DRD1 and GRM1 Japanese study Gotoh et al, 2009
ADRA1A adrenergic, alpha-1A-, receptor GeneCards

Adrenoceptor: Modulates NMDA recptors via RGS4

Liu et al, 2006 Binds to Neuregulin receptor ERBB3 Thaminy et al, 2003
Association reported in a Spanish population (Clark et al,2005)
CHAT Choline acetyl transferase GeneCards Acetlycholine synthesis Mancama et al, 2007
CHRFAM7A CHRNA7 (cholinergic receptor, nicotinic, alpha 7, exons 5-10) and FAM7A (family with sequence similarity 7A, exons A-E) fusion   US Study Sincus et al, 2009
CHRM5 cholinergic receptor, muscarinic 5   Canadian study De Luca et al, 2004
CHRNA4 cholinergic receptor, nicotinic, alpha 4   Canadian study De Luca et al, 2006

CHRNA7 Nicotinic receptor alpha subunit OMIM 118511 GeneCards

Stimulates glutamate release

Associated with schizophrenia in American family, French case, South African family and Azorean family studies (Freedman et al., 2001;Houy et al., 2004;Leonard et al., 2002;Raux et al., 2002;Riley et al., 2000;Xu et al., 2001) .
CHRNB2 cholinergic receptor, nicotinic, beta 2 (neuronal)   Canadian study De Luca et al, 2006

CNR1 Cannabinoid receptor 1 OMIM 114610 GeneCards

Activation inhibits glutamate release (Auclair et al., 2000;Wang, 2003). CNR1 receptor stimulation promotes the survival of oligodendrocyte progenitor cells (Molina-Holgado et al., 2002).  

Association reported in French and Japanese case studies (Leroy et al., 2001;Ujike et al., 2002).   
GPR78 Orphan GPCR
OMIM 606921 GeneCards
Orphan receptor Suggestive association in a Scottish case study (Underwood et al,2006
GPR85 G protein-coupled receptor 85 (SREB2) GeneCards Orphan receptor: Overexpression reduces brain weight, increases ventricular size and produces schizophrenia-like behavioural effects in mice Matsumoto et al, 2008 Association in an American family study Matsumoto et al, 2008

HRH2 H2 Histamine receptor

OMIM 142703 GeneCards

Histamine receptor

Associated with schizophrenia in a British case study (Orange et al., 1996).
MCHR1 (GPR24) Melanocyte concentrating hormone receptor 1 OMIM 601751 GeneCards Regulates mesolimbic dopamine signaling Pissios et al, 2008. GRIN1 expression reduced in knockout mice Adamantidis et al, 2005 Association reported in a Faroese/Scottish study
Severinsen et al, 2006
MCHR2 melanin-concentrating hormone receptor 2   US study Miller et al, 2009
MC5R melanocortin 5 receptor   US study Miller et al, 2009

NPY Neuropeptide Y

OMIM 162640 GeneCards

Inhibits glutamate release

Associated with schizophrenia in ta Japanese case study (Itokawa et al., 2003a) . 
OPCML opioid binding protein/cell adhesion molecule-like   GWAS O'Donovan et al, 2008

OPRS1 opioid receptor, sigma 1 OMIM 601978 GeneCards

Regulates oligodendrocyte differentiation Hayash and Su, 2004

Association in a Japanese case study(Ishiguro et al., 1998).
OPRM1 opioid receptor, mu 1   Czech republic Sery et al, 2010

PDE4B Phosphodiesterase 4B OMIM 600127 GeneCards

Binds to DISC1 (Millar et al., 2005). Activated by NMDA receptors during hippocampal LTP (Ahmed and Frey 2003)

Disrupted by a balanced translocation in a subject diagnosed with schizophrenia and a relative with chronic psychiatric illness (Millar et al., 2005)
PDE4D phosphodiesterase 4D, cAMP-specific (phosphodiesterase E3 dunce homolog   Finnish study Tommpo et al, 2009
PDE7B phosphodiesterase 7B   Japanese and UK study Ikeda et al, 2010
PDYN Prodynorphin   Chinese study Zhang et al, 2004
RGS2 regulator of G-protein signaling 2, 24kDa GeneCards Binds to GNAS American study Campbell et al, 2008
RGS5 regulator of G-protein signaling 5 GeneCards   American study Campbell et al, 2008

TAAR6 Trace amine receptor 6 (ex 4) OMIM 608923 GeneCards

Orphan receptor

Association reported in an American study of european and african american pedigrees(Duan et al., 2004)
Adhesion Molecules    
CDH4 Cadherin 4   International schizophrenia consortium O'Dushlaine et al, 2010

CHL1 cell adhesion molecule with homology to L1CAM (close homolog of L1) OMIM 607416 GeneCards

A soluble form of CHL1 promotes neurite extension in cerebellar granule cells (Naus et al., 2004)

Associates with schizophrenia in a Japanese case study (Sakurai et al., 2002).
L1CAM L1 cell adhesion molecule   Japanese studyKuramaji et al, 2001
CNTNAP2 contactin associated protein-like 2 GeneCards Member of the neurexin family of adhesion molecules and receptors. Involved in the clustering of potassium channels in myelinated axons Poliak et al, 2002 Haploinsufficiency in association with epilepsy and schizophrenia in the Amish population Friedman et al, 2007
CNTN1 Contactin 1   International schizophrenia consortium O'Dushlaine et al, 2010
GLG1 golgi glycoprotein 1   International schizophrenia consortium O'Dushlaine et al, 2010
HLA-DQA2   International schizophrenia consortium O'Dushlaine et al, 2010
JAM3 junction adhesion molecule 3 GWAS Moskvina et al, 2009 GWAS Moskvina et al, 2009
NCAM1 Neural cell adhesion molecule 1 GeneCards Promotes assembly of the PSD signaling complex Sytnyk et al, 2006. Stimulates PI3K/AKT signaling (Ditlevsen et al, 2003). TrkB phosphorylation and BDNF signaling is impaired in NCAM knockout mice Muller at al, 2000. Polysialylated-NCAM interacts with BDNF (Vutskits et al, 2001et al. 2001). Oligodendrocytes express GRIN1 and activation of NMDA receptors in these cells increases the expression of polysialylated NCAM1 that is required for oligodendrocyte migration (Wang et al, 1996).Polysialylated by ST8S1A2 Kojima et al, 1996 Sullivan et al, 2006
NRXN1 Neurexin 1   International schizophrenia consortium O'Dushlaine et al, 2010
PDCD1LG2 programmed cell death 1 ligand 2   International schizophrenia consortium O'Dushlaine et al, 2010
Tight junction    
CDC42, CTNNA2, HCLS1, HRAS,
PRKCH, MYH11, MYH15 and PARD3)
In progress International schizophrenia consortium O'Dushlaine et al, 2010
Glycan structures    
MAN2A1,
GALNT2, GALNT13, OGT and XYLT1
In progress International schizophrenia consortium O'Dushlaine et al, 2010

Snares

STX18 and TSNARE1

"" International schizophrenia consortium O'Dushlaine et al, 2010

cell cycle

MAD1L1

"" International schizophrenia consortium O'Dushlaine et al, 2010
Circadian 
CLOCK circadian locomoter output cycles kaput protein; clock OMIM GeneCards

Controls the expression of ATF4 (see DISC1, GABBR1) Igarashi et al, 2007

Circadian control
Association repoted in a Japanese study Takao et al, 2007
NPAS2 neuronal PAS domain protein 2   USA study Mansour et al, 2009
PER3 period homolog 3 OMIM GeneCards Circadian control Association reported in an American study Mansour et al, 2006
TIMELESS OMIM GeneCards Circadian control Association reported in an American study Mansour et al, 2006
Glycolysis and Krebs Cycle    
ENO2 Enolase 2 (gamma neuronal) GeneCards

2-phospho-D-glycerate = phosphoenolpyruvate + H2O

Association reported in a Scandinavian study Olsen et al, 2008
FBP1 fructose-1,6-bisphosphatase 1 GeneCards D-fructose 1,6-bisphosphate + H2O = D-fructose 6-phosphate + phosphate Association reported in a Scandinavian study Olsen et al, 2008

PCK1 phosphoenolpyruvate carboxykinase 1 (soluble) GeneCards

GTP + oxaloacetate = GDP + phosphoenolpyruvate + CO2 Association reported in a Scandinavian study Olsen et al, 2008
Cytokines and immune-related    
BTN2A2 butyrophilin, subfamily 2, member A2   GWAS Shi et al, 2009
BTN3A1 butyrophilin, subfamily 3, member A1   GWAS Shi et al, 2009
BTN3A2 butyrophilin, subfamily 3, member A2   GWAS Shi et al, 2009
C3 complement component 3   Spanish study: Fannanas et al, 1992
C4A complement component 4A (Rodgers blood group)   UK study Rudduck et al, 1985
C4B complement component 4B (Chido blood group)   UK study Rudduck et al, 1985
CFB complement factor B   Spanish study: Fannanas et al, 1992
CCR5 chemokine (C-C motif) receptor 5 GeneCards MCP-2, Mip1-alpha and Mip1-beta chemokine receptor expressed on T-cells and macrophages 32bp deletion associated with late-onset schizophrenia in a Danish study Rasmussen et al, 2006
CD4 CD4 molecule   Belgian study Zamani et al,1994
CLU Clusterin   Chinese study Zhou at al, 2010
CSF2RA colony stimulating factor 2 receptor, alpha, low-affinity (granulocyte-macrophage) OMIM GeneCards  receptor for colony stimulating factor 2, a cytokine which controls the production, differentiation, and function of granulocytes and macrophages. Lencz et al, 2007
CSF2RB colony stimulating factor 2 receptor, beta, low-affinity (granulocyte-macrophage) GeneCards Common beta receptor subunit for IL3, IL5 and GM-CSF signaling Chen et al, 2007

CTLA4 cytotoxic T-lymphocyte-associated protein 4 OMIM 123890 GeneCards

Member of the immunoglobulin superfamily which transmits an inhibitory signal to T cells

Association with schizophrenia has been reported in a Korean case study (Jun et al., 2002a).
CHI3L1 chitinase 3-like 1 (cartilage glycoprotein-39) Alias: GP39, HC-gp39, HCGP-3P, YKL40, YYL-40 GeneCards Inhbits effects of TNF and IL1B in peripheral cells Ling and Recklies,2004. Signals via AKT1 Recklies et al,2002   Zhao et al, 2007
EBF3 Early B-cell factor 3   GWAS O'Donovan et al, 2008
IFNG Interferon gamma   Associated in males but not females in a polish study Paul-Samojedny et al, 2010
IL1A Interleukin 1 alpha   Chinese study Chowdari et al, 2006
IGH@ immunoglobulin heavy locus Immunoglobulins recognize foreign antigens and initiate immune responses Italian study Frezza et al, 2009

IL1B Interleukin 1B OMIM 147720 GeneCards

Cytokine inducing PTGS2 in the CNS.

Toxic effects on oligodendrocytes

Associated with schizophrenia in Italian, German and Finnish case studies (Katila et al., 1999;Meisenzahl et al., 2001;Zanardini et al., 2003)

IL1RN Interleukin 1 receptor antagonist OMIM 147679 GeneCards

Inhibits activity of IL1B

A polymorphism in IL1RA has been reported to exert a protective effect in an Italian case study (Zanardini et al., 2003). Polymorphisms associated with schizophrenia have also been reported in spanish and korean case studies (Kim et al., 2004;Papiol et al., 2005)

IL2: Interleukin 2 OMIM 147680 GeneCards

Cytokine; Stimulates oligodendrocyte proliferation and maturation (Jiang and Lu 1998)

Association reported in a German case study (Schwarz et al., 2005)
IL3 interleukin 3 (colony-stimulating factor, multiple) OMIM GeneCards Cytokine ligand for IL3RA Chen et al, 2007
IL3RA interleukin 3 receptor, alpha (low affinity) OMIM GeneCards IL3 specific component of a heteromeric receptor unit, that includes CSF2RB Lencz et al, 2007 Association in a chinese studySun et al, 2008

IL4: Interleukin 4 OMIM 147780 GeneCards

Cytokine: protects oligodendrocytes ()

Association reported in a German case study (Schwarz et al., 2005)
IL6R Interleukin 6 receptor   Chinese study Sun et al, 2008
IL9 Interleukin 9   US study Chen et al, 2008
IL12B interleukin 12B (natural killer cell stimulatory factor 2, cytotoxic lymphocyte maturation factor 2, p40) GeneCards Cytokineimportant for sustaining a sufficient number of memory/effector Th1 cells to mediate long-term protection to an intracellular pathogen. Associated with schizophreniai in an American case-control study Shirts et al, 2008
IL18 interleukin 18 (interferon-gamma-inducing factor) GeneCards Proinflammatory cytokine inducing interferon-gamma production in T-cells, Binds to IL18 R1 and IL18RAP Associated with schizophrenia and cytomegalovirus seropositivity in an American case-control study Shirts et al, 2008
IL18R1 interleukin 18 receptor 1 GeneCards Il18 cytokine recptor: expression induced by IL12 Associated with schizophrenia and with cytomegalovirus seropositivity in an American case-control study Shirts et al, 2008

IL18RAP interleukin 18 receptor accessory protein GeneCards

Potentiates the IL18 binding activity of IL18R1 (IL1RRP), a ligand binding subunit of the IL18 receptor Associated with schizophrenia and HSV-1 and cytomegalovirus seropositivity in an American case-control study Shirts et al, 2008

LTA Lymphotoxin alpha OMIM 153440 GeneCards

Cytokine: Toxic effects on oligodendrocytes

Associates with schizophrenia in a Korean case study (Jun et al., 2003).

IL10 Interleukin-10 OMIM 124092 BIOCARTA GeneCards

Anti-inflammatory Cytokine: Protective effects on oligodendrocytes

Generally represses effects of TNF and IL1B

Promoter polymorphisms have been associated with schizophrenia in Italy (case) and China (case and family) (Chiavetto et al., 2002) (He et al., 2006;Jun et al., 2002b;Yu et al., 2004).
IL10RA Interleukin 10 receptor GeneCards IL10 receptor signalling via AKT pathway Association reported in an Austrian case study Schosser et al, 2006

TNFA: Tumor necrosis factor alpha OMIM 191160 GeneCards

Proinflammatory cytokine: Expression controlled by NFE2L1 in mice (Prieschl et al, 1998 (see DISC1). Toxic effects on oligodendrocytes ()

Association reported in Chinese case, male Finnish case, German family, Italian case and Singaporean case studies (Boin et al., 2001b;Boin et al., 2001a;Hanninen et al., 2005b;Schwab et al., 2003;Tan et al., 2003;Zhang et al., 2005)
TNFR2 = TNFRSF1B TNF receptor Thabet et al, 2010
TRAF3IP1 TNF receptor-associated factor 3 interacting protein 1  Binds to DISC1 Finnish study (stage 1)Tomppo et al, 2009
HLA-A (HLA-A10) major histocompatibility complex, class I, A GeneCards Immune system In association with chlamydophila infection Fellerhoff et al, 2007
HLA-B major histocompatibility complex, class I, B GeneCards Immune system HLA-B maternal-fetal genotype-matching effect on schizophrenia demonstrated for female offspring : related to birth complications rather than schizophrenia (see environmental risk factors) Palmer et al, 2006
HLA-DRB1 DRB1*04

major histocompatibility complex, class II, DR beta 1

OMIM 142857

HLA-DRB1 recognises peptides generated from myelin oligodendrocyte glycoprotein (MOG) (Weissert et al., 2002). It also recognises peptides derived from the influenza virus (Hennecke and Wiley, 2002;Wedderburn et al., 1995) , from the spirochete Borrelia burgdorferi (Steere et al., 2003;Willett et al., 2004) which causes Lyme disease carried by  Ixodes ticks and from peptides derived from the Rubella virus (Neepom et al, 1997)

The frequency of this allele is positively associated with rheumatoid arthritis (Reveille, 1998)and reduced in schizophrenic patients and their healthy mothers (Wright et al., 1996). The incidences of schizophrenia and rheumatoid arthritis are inversely correlated (Eaton et al., 1992;Gorwood et al., 2004;Vinogradov et al.,
1991). Association also reported in chinese family (Yu et al,2003  and 2 Japanese case studies (Sasaki et al,1999; Arinami et al, 1998 and in Kuwaiti arabs (case study) Heider et al,2000
 
HLADQA1 major histocompatibility complex, class II, DQ alpha 1   GWAS Shi et al, 2009
MICB MHC class I polypeptide-related sequence B OMIM GeneCards  ligand for the NKG2D type II receptor of natural killer cells. Associated with Herpes virus seropositivity and schizophrenia Shirts et al, 2007
PRSS16 protease, serine, 16 (thymus)  Thought to play a role in the alternative antigen presenting pathway used by cortical thymic epithelial cells during the positive selection of T cells. GWAS Shi et al, 2009
TGIF1 TGFB-induced factor homeobox 1 GeneCards Transcriptional repressor of SMAD signaling (part of the TGF beta pathway) and of retinoic acid receptor(RXRalpha) signaling (See Zaharieva et al, 2008 for potential association with SMAD5 or TGFB1. Association reporterd in an American study Chavarria-Siles et al, 2007
Protein sorting and ER stress    
CALR Calreticulin Calcium storage protein in the endoplasmic reticulum Point mutation obsreved in a single schizophrenia patient (1/100) and not in 280 controls Aghajani et al, 2006
HSPA1B HSP70-2 GeneCards Heat shock protein (molecular chaperones that assist other proteins in their folding, transport and assembly into complexes) Association suggested in a Korean study Pae et al, 2005
HSPA1L heat shock 70kDa protein 1-like GeneCards see above Association in a Korean case-control study Kim et al, 2008

XBP1 X-box binding protein 1. OMIM 194355 GeneCards

Endoplasmic reticulum stress

Associated with schizophrenia in Chinese and Japanese case studies (Chen et al., 2004c;Kakiuchi et al., 2004)
WNT Signalling    
DAAM1 dishevelled associated activator of morphogenesis 1   UK study Proitsi et al, 2008
DAAM2 dishevelled associated activator of morphogenesis 2   UK study Proitsi et al, 2008
DKK4 dickkopf homolog 4 GeneCards Role in Wnt signaling Association in chinese trios and a Hong Kong case study Proitsi et al, 2008

FZD3 Frizzled 3 OMIM 606143 GeneCards

Receptor for wingless

Associated with schizophrenia in Japanese case and Chinese case and family studies (Katsu et al., 2003;Yang et al., 2003a;Zhang et al., 2004) .
KREMEN1 kringle containing transmembrane protein 1 Binds to DKK1 Association in a Japanese study Aleksic et al, 2010
SFRP1 secreted frizzled-related protein 1   UK study Proitsi et al, 2008
Extracellular matrix proteins, structural , intracellular transport and related    
CAPZA2 capping protein (actin filament) muscle Z-line, alpha 2   US study Burdick et al, 2010
FLNB Filamin B Binds to actin cytoskeleton UK study Proitsi et al, 2008
ITGA3 integrin, alpha 3 (antigen CD49C, alpha 3 subunit of VLA-3 receptor) GeneCards Component of an integrin receptor interacting with several extracellular matrix proteins. Binds to CALR  Scandinavian case-control Kahler et al, 2008
KIF2A kinesin heavy chain member 2A Microtubule-associated motor protein Chinese study Li et al, 2006
MAP4 microtubule-associated protein 4   Chinese study So et al, 2009
MAP6
Microtubule-associated protein 6 OMIM 601783
GeneCards

Calmodulin-binding and calmodulin-regulated protein involved in microtubule stabilization.

Associated in a Japanese case study
Shimizu et al, 2006
MYH9 myosin, heavy chain 9, non-muscle   Association in a Chinese study Liu et al, 2008
MYO9B Myosin IXb GeneCards Myosin motor Association reported in a Dutch study Jungerius et al, 2007
MYO18B Myosin XVIII b   US study Purcell et al, 2009
MXRA5 matrix-remodelling associated 5   GWAS European ancestry Shi et al, 2009
RELN Reelin GeneCards

Secreted extracellular matrix protein thought to control cell-cell interactions critical for cell positioning and neuronal migration during brain development. Signals via PI3K AKT1 pathway, inter alia Jossin and Goffinet, 2007 Controls the surface expression of NMDA receptors (GRIN1, GRIN2B) Groc et al, 2007 and AMPA receptors Qiu and Weeber, 2007

Associated with working memory in Finnish schizophrenia families Wedenoja et al, 2007 and in GWAS in women Shifman et al, 2008
SPARCL1 SPARC-like 1 (mast9, hevin) GeneCards Extracellular matrix protein localised in postsynaptic terminals and enveloping glia Liveley et al, 2007  

TNXB: tenascin XB

OMIM 600985 GeneCards

Exracellular matrix protein

SNP's within the TNXB locus associate with schizophrenia in a UK family based study (Wei and Hemmings, 2004c)
Retinol Metabolism    

ADH1B alcohol dehydrogenase IB (class I), beta polypeptide OMIM 103720 GeneCards

Multiple substrates include catecholamine metabolites.

Association in a Chinese case study (Xu et al., 2004) and an American family study  (Vawter et al, 2006)
ALDH1A2 aldehyde dehydrogenase 1 family, member A2 Catalyzes the synthesis of retinoic acid (RA) from retinaldehyde. Chinese study Wan et al, 2010
CYP1A2 cytochrome P450, family 1, subfamily A, polypeptide 2

  Turkish study Boke et al, 2007
CYP3A4 cytochrome P450, family 3, subfamily A, polypeptide 4   Chinese study Du et al, 2009
CYP3A5 cytochrome P450, family 3, subfamily A, polypeptide 5   Chinese study Du et al, 2009
RBP1 Retinol binding protein 1, cellular GeneCards Retinol (Vitamin A alcohol) transport Genome-wide UK associationstudy using pooled samples Kirov et al, 2008
Ion Channels    
CACNA1C calcium channel, voltage-dependent, L type, alpha 1C subunit   UK Study Green et al 2009
CACNA1F calcium channel, voltage-dependent, L type, alpha 1F subunit   UK study Wei et al, 2006
CACNG2 calcium channel, voltage-dependent, gamma subunit 2 [ Related to DLG4, GRIA1, GRIA4 and GRIN1 Association in a Chinese study Liu et al, 2008
KCNH2potassium voltage-gated channel, subfamily H (eag-related), member 2   US study Hiffaker et al, 2009
NALCN sodium leak channel, non-selective   GWAS Wang et al, 2010
TRPA1 transient receptor potential cation channel, subfamily A, member 1 Cannabinoid receptor-operated channel Jordt et al, 2004 GWAS African American Shi et al, 2009
Mitochondrial Genes    
ATP6 mitochondrially encoded ATP synthase 6   Japanese study Ueno et al, 2009
CYTB mitochondrially encoded cytochrome b Oxidative phosphorylation Swedish study Lindholm et al, 1997
MT-haplo    
ND2 mitochondrially encoded NADH dehydrogenase 2 Oxidative phosphorylation US study Rollins et al, 2009
ND4 mitochondrially encoded NADH dehydrogenase 4 Oxidative phosphorylation UK study Marchbanks et al, 2003
RNR1 mitochondrially encoded 12S RNA   US study Rollins et al, 2009
Transporters    
ABCA13 ATP-binding cassette, sub-family A (ABC1), member 13   UK study Knight et al, 2009
SLCO6A1 solute carrier organic anion transporter family, member 6A1   GWAS Stefansson et al, 2009
SLC12A2 solute carrier family 12 (sodium/potassium/chloride transporters), member 2
  GWAS USA Potkin et al, 2009
SLC17A1 solute carrier family 17 (sodium phosphate), member 1   GWAS Shi et al, 2009
SLC17A3 solute carrier family 17 (sodium phosphate), member 3   GWAS Shi et al, 2009
Nuclear import export    
IPO5 Importin 5   Chinese study Wang et al, 2010

KPNA3   karyopherin alpha 3 (importin alpha 4) OMIM 601892 GeneCards

Nuclear transport: Involved in the nuclear import of Influenza viral nucleoprotein Melen et al, 2003

Associated in a British family study  (Wei and Hemmings, 2005b)

KPNB3 (RANBP5) GeneCards

OMIM 602008

Nuclear transport : Important role in the nuclear import of Influenza viral RNA polymerase Deng et al, 2006
Associated in a British family study (Wei and Hemmings, 2004c)
RANBP1 RAN binding protein 1 Ran involved in nucleocytoplasmic transport Yudin et al, 2009 USA study Liu et al, 2002
MicroRNA    
MIR-30E microRNA   Xu et al, 2010
MIR-198 MicroRNA (Link is to Sanger Mirbase)   Hansen et al, 2007 (PLOS)
MIR-206 MicroRNA (Link is to Sanger Mirbase)   Hansen et al, 2007 (PLOS)
Miscellaneous    
ACSL6 acyl-CoA synthetase long-chain family member 6   US study Chen et al, 2006
ACSM1 acyl-CoA synthetase medium-chain family member 1 (same as BUCS1) Butanoate metabolism Kegg GWAS Sullivan et al, 2008
ADAM12 ADAM metallopeptidase domain 12
  GWAS O'Donovan et al, 2009
ADSS adenylosuccinate synthase Converts IMP to AMP Zhang et al, 2008
AGA
aspartylglucosaminidase GeneCards

N-glycan degradation : Impaired myelinisation in AGA knockout mice . (Tenhunen et al, 1998)

Linkage in an American family study related to expression change   (Vawter et al, 2006)
AGBL1 ATP/GTP binding protein-like 1 [   GWAS Sullivan et al, 2008
AHI1
Abelson helper integration site 1GeneCards
Required for cerebellar and cortical development Association reported in an Arab-Israeli family (Amman-Zankelstein et al, 2006
AMACR alpha-methylacyl-CoA racemase   Bespalova et al,2010
ANK3 Ankyrin 3  Binds to L1CAM Hortsch et al, 2009 Althanasiu et al, 2010
ANKK1 ankyrin repeat and kinase domain containing 1   French study Dubertret et al, 2010

APOE Apolipoprotein E OMIM 107741 GeneCards

Lipoprotein involved in cholesterol transport. Binds to and potentiates actions of CNTF. Involved in the cerebral uptake of Herpes simplex virus Burgos et al, 2006

Increased frequency of the APOE4 allele reported in a British study (Harrington et al., 1995a) and associated with earlier age of onset an American studies (Arnold et al., 1997;Martorell et al., 2001). A decreased frequency of the APOE2 allele has been associated with early onset schizophrenia in Japan (Kimura et al., 1997) and an increased frequency of APOE3 has been reported in China  (Chen et al., 1999). A Chinese study has also shown that increased frequency of the APOE4 allele and decreased frequency of APOE3 are associated with schizophrenia. This association is modified by date of birth and restricted to individuals born during times of famine (Liu et al., 2003a).
APOL2 Apolipoprotein L,2   Japanese study: Takahishi et al, 2008
AR Androgen receptor   UK study Crow et al, 1993
ARHGAP18 Rho GTPase activating protein 18   US study Potkin et al, 2009
ARHGEF10 Rho guanine nucleotide exchange factor (GEF) 10
  Dutch study: Jungerius et al, 2008
ARRB1 arrestin, beta 1   GWAS Moskvina et al, 2009
ASTN1 Asrotactin 1 GeneCards Role in neuronal migration Adams et al, 2002   Scandinavian case-control Kahler et al, 2008
ASTN2 Astrotactin 2   Copy number variation Vrijenhioek et al, 2008
ATE1 arginyltransferase 1 Involved in ubiquitination GWAS O'Donovan et al, 2009
ATF4 activating transcription factor 4 (tax-responsive enhancer element B67) (CREB2) Binds to DISC1 and GABBR1 . Key transcription factor activated by stress-provoked eif2alpha phosphorylation (see EIS2S1) Rutkowski and Kaufman, 2003 and eif2b page Associated with male patients in a Chinese case control study Qu et al, 2008
ATG9B ATG9 autophagy related 9 homolog B   US study Huffaker et al, 2009
ATM ataxia telangiectasia mutated   Chinese polulation Zhang et al, 2008
ATN1 Atrophin1   German study Morris-Rosendahl et al, 1997
ATXN3 Ataxin 3 Binds to PICK1 Yugoslavian study Culjkovic et al, 2000

B3GAT1 beta-1,3-glucuronyltransferase 1 (Synthesises HNK-1) OMIM 151290 GeneCards

Key enzyme in the biosynthesis of the carbohydrate epitope HNK-1 (human natural killer-1, also known as CD57 and LEU7). HNK-1 is a component of MOG. Mice deficient in HNK-1 show reduced hippocampal LTP (Mikol et al., 1990;Yamamoto et al., 2002) 

 

Located at a chromosomal breakpoint whose translocation (t(6;11)(q14.2;q25) ) segregates with a schizophrenia-like psychosis (Jeffries et al., 2003).
BAK1 BCL2-antagonist/killer 1   UK study Proitsi et al, 2008
BRD1
Bromodomain containing 1 GeneCards
Binds to FEZ1 (Assmann et al, 2006) Association reported in a Danish case control study
Severinsen et al, 2006)
BZRP TSPO translocator protein (18kDa) aka peripheral benzodiazepine receptor GeneCards Important role in controlling cholesterol import into mitochondria and steroid biosynthesis. Fallin et al., 2005

C6orf85=SLC22A23solute carrier family 22, member 23 [

  GWAS Moskvina et al, 2009
C10orf120
  GWAS O'Donovan et al, 2009
C18orf1
  Iceland study Ingason et al, 2007
CBX2 chromobox homolog 2 (Pc class homolog, Drosophila)   GWAS African American Shi et al, 2009
CCDC60 coiled-coil domain containing 60 GeneCards Unknown function  Genome-wide UK associationstudy using pooled samples Kirov et al, 2008
CCDC102B coiled-coil domain containing 102B   GWAS Moskvina et al, 2009
CDC42SE2 CDC42 small effector 2   US study Chen et al, 2006
CDKN2A cyclin-dependent kinase inhibitor 2A (melanoma, p16, inhibits CDK4)   GWAS Moskvina et al, 2009
CENTG2= AGAP1 ArfGAP with GTPase domain, ankyrin repeat and PH domain 1   GWAS European ancestry Shi et al, 2009

CLDN5 Claudin 5 OMIM 602101 GeneCards

Tight junction protein involved in blood-brain barrier regulation

Associated in Chinese  and British family studies (Sun et al., 2004;Wei and Hemmings, 2005)
CLINT1clathrin interactor 1 Intracellular traffic UK study Pimm et al, 2005
CPXM2 carboxypeptidase X (M14 family), member 2   UK GWAS O'Donovan et al, 2009
CTNND2 catenin (cadherin-associated protein), delta 2 (neural plakophilin-related arm-repeat protein)   Copy number variation Vrijenhioek et al, 2008
CREB3L1 cAMP responsive element binding protein 3-like 1   GWAS Moskvina et al, 2009
CTXN3 Cortexin 3 5q23.2   GWAS USA Potkin et al, 2009
CYP2D6 cytochrome P450, family 2, subfamily D, polypeptide 6 [   Spanish study Llerena et al, 2007
DDX31 DEAD (Asp-Glu-Ala-Asp) box polypeptide 31 RNA Helicase GWAS African American Shi et al, 2009
DGCR2 DiGeorge syndrome critical region gene 2   UK study Shifman et al, 2006
DGCR5 DiGeorge syndrome critical region gene 5   UK study Shifman et al, 2006
DGCR6 DiGeorge syndrome critical region gene 6  Binds to GABBR1 US study Liu et al, 2006 GWAS Moskvina et al, 2009
DLX1 distal-less homeobox 1 GeneCards Homeobox gene implcated in TGF beta signaling. Role in the migration of GABA neurones to the cortex Cobos et al, 2007. Negatively regulates OLIG2-dependent oligodendrocyte development Petryniak et al, 2007  Scandinavian case-control Kahler et al, 2008
DNMT3B DNA (cytosine-5-)-methyltransferase 3 beta   Chinese study Zhang et al, 2005
DYM Dymeclin   Japanese study Yazaki et al, 2010
ELP3 elongation protein 3 homolog (S. cerevisiae)   GWAS African American Shi et al, 2009
EML5 echinoderm microtubule associated protein like 5   GWAS European ancestry Shi et al, 2009
ENAH enabled homolog (Drosophila) GeneCards Regulates the actin skeleton Menzies et al, 2004  Scandinavian case-control Kahler et al, 2008
ESD Esterase D   Spanish study Fananas et al, 1990
ESR1 estrogen receptor 1 GeneCards

Interacts with TP53: activated by XBP1(see gene data):Phoshorylated by EGFR activation Marquez et al, 2007

American case control Weickert et al, 2008
EVPL Envoplakin   GWAS Moskvina et al, 2009
FABP7 fatty acid binding protein 7, brain GeneCards Fatty acid binding protein: Target of PAX6 Arai et al, 2005 Association reported in a Japanese study Watanabe et al, 2007
FBXL21 F-box and leucine-rich repeat protein 21 GeneCards F-box proteins are subunits of the "SKP1-cullin-F-box" ubiquitin protein ligase complex. Role in phosphorylation-dependent ubiquitination. Association an an Irish family and case-control study Chen et al, 2008
FBXL7 F-box and leucine-rich repeat protein 7 Ubiquitination see above GWAS Moskvina et al, 2009
FBXO38 F-box protein 38   GWAS Moskvina et al, 2009
FLJ46361   GWAS O'Donovan et al, 2009
FOXP2 Forkhead box P2 OMIM 605317 GeneCards

Role in synaptic plasticity Grozner et al, 2008

Association reported in a Spanish case study (Sanjuan et al, 2006)
FXR1 fragile X mental retardation, autosomal homolog 1   International Schizophrenia Consortium
FXYD2 FXYD domain containing ion transport regulator 2 NA+/K+ ATPase regulator. Association reported in a UK study Choudhury et al, 2007
FXYD6 FXYD domain-containing ion transport regulator 6; phosphohippolin OMIM GeneCards NA+/K+ ATPase regulator. Association reported in a UK study Choudhury et al, 2007
GALNT7
OMIM 605005
GeneCards

O-glycan biosynthesis

Suggestive linkage in an American family study related to expression change   (Vawter et al, 2006)
GC group-specific component (vitamin D binding protein)   UK study Brackenridge et al 1972
GJA8 gap junction protein, alpha 8, 50kDa connexin 50 GeneCards   Ni et al, 2007
GLT8D1 glycosyltransferase 8 domain containing 1   GWAS Moskvina et al, 2009
GNAL guanine nucleotide binding protein (G protein), alpha activating activity polypeptide, olfactory type   Geman study Schwab et al, 1998
GNAO1 guanine nucleotide binding protein (G protein), alpha activating activity polypeptide O   Japanese study Tani et al, 2001
GNAS guanine nucleotide binding protein (G protein), alpha stimulating activity OMIM 139320 GeneCards

G protein

Associated with deficit schizophrenia in an Italian case study (Minoretti et al, 2006)
GNB1L guanine nucleotide binding protein (G protein), beta polypeptide 1-like GeneCards G-protein Association reported in a UK case study and with Bulgarian parent proband trios.Strongest in males Williams et al, 2007
GNB3 guanine nucleotide binding protein (G protein), beta polypeptide 3   German study Zill et al, 2000
GNL3 guanine nucleotide binding protein-like 3 (nucleolar) Binds to TP53 GWAS Moskvina et al, 2009
GTF3C4 general transcription factor IIIC, polypeptide 4, 90kDa   GWAS African American Shi et al, 2009
GPC1 Glypican 1 Involved in polyamine uptake Belting et al, 2003 Polyamines are NMDA receptor ligands Igarashi et al, 2010 Receptor for SLIT3 Liang et al, 1999 GWAS USA Potkin et al, 2009
hCAP-D3 =NCAPd3 non-SMC condensin II complex, subunit D3 Role in mitosis GWAS Moskvina et al, 2009
HDAC3 Histone deacetylase 3   Association in Korean population Kim et al, 2010
HDAC4 Histone deacetylase 4   Association in Korean population Kim et al, 2010
HIST1H2AG histone cluster 1, H2ag   GWAS Shi et al, 2009
HIST1H2AH histone cluster 1, H2ah
  US study Shi et al, 2009
HIST1H2BJ histone cluster 1, H2bj    
HMX3 H6 family homeobox 3   GWAS O'Donovan et al, 2009
HSPA12A heat shock 70kDa protein 12A
  GWAS O'Donovan et al, 2009
HTF9C HpaII tiny fragments locus 9C GeneCards Unknown function Association with attention deficit in a Taiwanese family study Liu et al, 2007
INTS6 integrator complex subunit 6   GWAS Moskvina et al, 2009
ITIH3 inter-alpha (globulin) inhibitor H3   GWAS Moskvina et al, 2009
KLF5 Kruppel-like factor 5 (intestinal) GeneCards zinc finger protein binding the epidermal growth factor response element Association in a Japanese study Yanagi etal , 2008
KLF6 Kruppel-like factor 6   Japanese study Hashimoto et al, 2007
LHPP phospholysine phosphohistidine inorganic pyrophosphate phosphatase   GWAS O'Donovan et al, 2009
LOC220433- RPS4XP16 ribosomal protein S4X pseudogene 16   GWAS Moskvina et al, 2009
LOC730216 Putative Vitamin C transporter : but gene discontinued GWAS Moskvina et al, 2009
LRRTM1 leucine rich repeat transmembrane neuronal 1 GeneCards Uncharacterised overtransmitted paternally to individuals with schizophrenia/schizoaffective disorder Francks et al, 2007
MAMDC1- MDGA2 MAM domain containing glycosylphosphatidylinositol anchor 2   GWAS Moskvina et al, 2009
MAPK8IP2 mitogen-activated protein kinase 8 interacting protein 2   Danish study Severinsen et al, 2006
MAPK14 mitogen-activated protein kinase 14 GeneCards Activated by stress and pro-inflammatory cytokines Association reported in a Scandinavian study Olsen et al, 2008 
MCTP2 multiple C2 domains, transmembrane 2   Scandinavian study Djurovic et al, 2009

MED12 mediator of RNA polymerase II transcription, subunit 12 homolog (Alias HOPA; OPA1; CAGH45; TNRC11; TRAP230; KIAA0192) OMIM 300188 GeneCards

Component of the Mediator Complex, a group of proteins that regulates transcription via nuclear receptor, Wnt and Receptor Tyrosine Kinase pathways Kim et al, 2006

Associated with schizophrenia in two North American case studies (Sandhu et al., 2003;Spinks et al., 2004).
OXT Oxytocin   US Study Souza et al, 2010

PCQAP (MED15) mediator complex subunit 15GeneCards OMIM 607372

A component of TGF/Activin/Nodal/Smad2/3 signal transduction (Kato et al., 2002)

A polymorphic CAG repeat in this gene has been reported to associate with schizophrenia in an Italian case study (De Luca et al., 2003).
MDGA1 MAM domain containing glycosylphosphatidylinositol anchor 1 GeneCards Role in neuronal migration Takeuchi and O'Leary, 2006 Scandinavian case-control Kahler et al, 2008
MEGF10 multiple EGF-like-domains 10 GeneCards Involved in the engulfment of apoptotic cells Suzuki and Nakayama, 2007 Irish case-control Chen et al, 2008
MOV10L1 Moloney leukemia virus 10-like 1, homolog   Danish study Severinsen et al, 2006
MPZL1 Myelin protein zero like 1 GeneCards Concanavalin A receptor Association reported in a Chinese family study  He et al, 2006

MRLC2=MYL12B myosin, light chain 12B, regulatory

  GWAS Moskvina et al, 2009
MRCL3= MYL12A myosin, light chain 12A, regulatory, non-sarcomeric   GWAS Moskvina et al, 2009
MTNR1B melatonin receptor 1B   GWAS Moskvina et al, 2009
NHN1=ZC3H18 zinc finger CCCH-type containing 18   GWAS Moskvina et al, 2009
NEK4 NIMA (never in mitosis gene a)-related kinase 4   GWAS Moskvina et al, 2009
NRGN Neurogranin GeneCards Calmodulin binding protein localised in dendritic spines Association in a Portugese and Brasilian population Ruano et al, 2006 and in a GWAS study Stefanssonnet al, 2009
NUMBL  numb homolog (Drosophila)-like GeneCards Binds to EPS15 a component of EGF  receptor signalling (Salsini et al, 1997) Association reported in Brazilian and Danish subjects (Passos Gregorio et al 2006)
PADI2 peptidyl arginine deiminase, type II Converts arginine residues into citrullines in the presence of calcium ions: Substrates include include myelin basic protein MBP Japanese study Watanabe et al, 2009
PADI4 peptidyl arginine deiminase, type IV   Japanese study Watanabe et al, 2009
PAWR PRKC, apoptosis, WT1, regulator GeneCards Localised in the postsynaptic density . Overexpression attenuates NGF-indiced neuritogenesis Guo et al, 2001 Association reported in female Han Chinese Wang et al, 2008
PCNT Pericentrin Binds to DISC1 Japanese study Anitha et al, 2009
PDCH8 protocadherin 8   UK study Bray et al, 2002

PDLIM5 PDZ and LIM domain 5 OMIM 605904 GeneCards

Cytoplasmic sequestration factor for ID2 (Lasorella and Iavaone 2006). Such effects limit the nuclear entry of transcription factors and prevent their action.

Association in a Japanese case study (Horiuchi et al., 2005)
PEMT phosphatidylethanolamine N-methyltransferase GeneCards Converts phosphatidylethanolamine to phosphatidylcholine via methylation. Plays an important role in the control of homocysteine levels Shields et al, 2005 Association in Han Chinese parent-offspring trios Liu et al, 2007
PGBD1 piggyBac transposable element derived 1

Unknown function

 

Iceland and European GWAS Stephanson et al, 2009

PHOX2B   paired-like homeobox 2b OMIM 603851 GeneCards

Transcription factor involved in the development of noradrenergic neurones. Regulates RGS4 transcription. Grillet et al, 2003.

Associated with schizophrenia in a Japanese case study (Toyota et al., 2004).
PNPO pyridoxamine 5'-phosphate oxidase GeneCards

Rate limiting enzyme for vitamin B6 (pyridoxal 5-phosphate) synthesis . High dose Vitamin B6 reduces elevated homocysteine levels in schizophrenia Miodownick et al, 2007

Association in a Japanese case study Song et al, 2007

KLH1AS (SCA8)

spinocerebellar ataxia 8 OMIM 603680 GeneCards

Non-coding antisense to KLHL1 which regulates oligodendrocyte process elongation Jiang et al, 2007 and neurite outgrowth Seng et al, 2006

Weak association reported in a Canadian study of azorean and swedish families and UK case studies (Vincent et al., 2000).
MAGI3 membrane associated guanylate kinase, WW and PDZ domain containing 3   US study Buxbaum et al, 2007
MMP3 matrix metallopeptidase 3 (stromelysin 1, progelatinase)   Turkish study Kucukali et al, 2009
MMP9 matrix metallopeptidase 9   Polish study Rybakowski et al, 2009
TIMP3 TIMP metallopeptidase inhibitor 3 Inhibits MMP3 and MMP9 Taiwan study Hung et al, 2001
PB1=PBRM1 polybromo 1 [   GWAS Moskvina et al, 2009
PCM1
pericentriolar material 1 GeneCards

Binds to kendrin (PCNT) (Li et al,2001) and NUDEL (Guo et al, 2006), both binding partners of DISC1 Association in a UK family and case study (Gurling et al, 2006)
PFN4 profilin family, member 4   GWAS Moskvina et al, 2009
PGAP1 post-GPI attachment to proteins 1   GWAS Moskvina et al, 2009
PLAA phospholipase A2-activating protein   Athanasiu et al,20010
PNPLA8 patatin-like phospholipase domain containing 8   Chinese study Meng et al, 2010
POM121L2 POM121 membrane glycoprotein-like 2   GWAS Shi et al, 2009
POU3F2 POU class 3 homeobox 2   GWAS USA Potkin et al, 2009
PPAPDC1A phosphatidic acid phosphatase type 2 domain containing 1A
  GWAS O'Donovan et al, 2009
PPARD peroxisome proliferator-activated receptor delta   Chinese study Sun et al, 2008
PRKCA protein kinase C, alpha   UK study Carroll et al, 2009
PSEN2 Presenilin 2 Cleaves NOTCH4 Chinese study Zhang et al, 2009
PSKH1 protein serine kinase H1   GWAS Moskvina et al, 2009
PTBP2 polypyrimidine tract binding protein 2 Binds to QK1 Lim et al, 2006 International Schizophrenia Consortium
PTPN21 protein tyrosine phosphatase, non-receptor type 21   GWAS European ancestry Shi et al, 2009
     
RAPGEF6 Rap guanine nucleotide exchange factor (GEF) 6   US study Chen et al, 2006
RASD2 RASD family, member 2   Association in a Chinese study Liu et al, 2008
RHD Rh blood group, D antigen GeneCards Rhesus blood group, D antigen Maternal-fetal rhesus incomatibility Palmer et al, 2002
RFXDC1=RFX6 regulatory factor X, 6   GWAS Moskvina et al, 2009
ROBO1 roundabout, axon guidance receptor, homolog 1 SLIT receptor GWAS USA Potkin et al, 2009
ROBO2 roundabout, axon guidance receptor, homolog 2 SLIT receptor; Binds to ROBO1 and SLIT3: GWAS USA Potkin et al, 2009
RPS6KA5 ribosomal protein S6 kinase, 90kDa, polypeptide 5   GWAS Moskvina et al, 2009
SLIT3 slit homolog 3 GeneCards SLITS regulate axon guidance and cell migration, and stimulate branching and elongation of sensory axons and cortical dendrites Binds to ROBO2: Binds to GPC1Liang et al, 1999 Association in a Chinese Han population Shi et al, 2004
RPGRIP1L RPGRIP1-like   UK GWAS O'Donovan et al, 2008
RPP21 ribonuclease P/MRP 21kDa subunit   Multinational O'Donovan et al, 2009
RSRC1 arginine/serine-rich coiled-coil 1   US study Potkin et al, 2008
SEMG2 semenogelin II   GWAS Moskvina et al, 2009
SEPSECS Sep (O-phosphoserine) tRNA:Sec (selenocysteine) tRNA synthase   Scottish study Houlihan et al, 2009
SHISA5 shisa homolog 5   Chinese study So et al, 2009
SH3TC2 SH3 domain and tetratricopeptide repeats 2   GWAS Moskvina et al, 2009
SIL1 SIL1 homolog, endoplasmic reticulum chaperone   GWAS O'Donovan et al, 2008
SIRT5 sirtuin (silent mating type information regulation 2 homolog) 5   US study Chowdari et al, 2006
SMARCC1 SWI/SNF related, matrix associated, actin dependent regulator of chromatin, subfamily c, member 1   Chinese study So et al, 2009
SP4 Sp4 transcription factor   US Study Zhou et al, 2009
SREBF1sterol regulatory element binding transcription factor 1 Regulates Lipoprotein receptors and cholssterol synthesis German and scandinavian samples Le Hellard et al, 2008
SREBF2 sterol regulatory element binding transcription factor 2 Regulates cholesterol Homoeostasis German and scandinavian samples Le Hellard et al, 2008

ST8SIA2   (SIAT8B) ST8 alpha-N-acetyl-neuraminide alpha-2,8-sialyltransferase 2 OMIM 602546 GeneCards

Involved in the production of polysialic acid, a modulator of the adhesive properties of neural cell adhesion molecule (NCAM1).

Glycan biosynthesis and metabolism: Ganglioside biosynythesis

Association reported in a Japanese case study (Arai et al., 2005).

SULT4A1 Sulfotransferase 4a1 OMIM 608359 GeneCards

Highly expressed in substantia nigra suggesting a role in catecholamine sulfation (Liyou et al., 2003)

Possible susceptibility gene in an American family study (Brennan and Condra, 2005)
T10 ??   US study Liu et al, 2002
TBP TATA box binding protein Control the initiation of trancription by RNA polymerase Taiwan study Chen et al, 2005
TBX1 T-box 1   UK study Williams et al, 2008
TCERG1L transcription elongation regulator 1-like   GWAS O'Donovan et al, 2008
TCP1 t-complex 1   Chinese study Yang et al, 2004
TGM2 transglutaminase 2 (C polypeptide, protein-glutamine-gamma-glutamyltransferase) GeneCards Catalyses protein cross-linking Association in a British study Bradford et al, 2008
TMEM108 transmembrane protein 108   GWAS O'Donovan et al, 2009
TNIK TRAF2 and NCK interacting kinase   GWAS African American Shi et al, 2009
TRMT2A TRM2 tRNA methyltransferase 2 homolog A   US study Liu et al, 2002
TSKU tsukushi small leucine rich proteoglycan homolog   GWAS Moskvina et al, 2009
TSNAX translin-associated factor X   Scottish study Thomson et al, 2005
TSPAN8 Tetraspanin 8   USA Scholz et al, 2010
TPP2 Tripeptidyl peptidase II GeneCards Involved in the antigen processing of the Influenza virus Guil et al, 2006 Fallin et al., 2005)
TRIM71 tripartite motif-containing 71   GWAS Moskvina et al, 2009
TXNDC5 thioredoxin domain containing 5 (endoplasmic reticulum) Protein-disulfide isomerase induced by hypoxia Taiwanese study Lin et al, 2009

UFD1L ubiquitin fusion degradation 1 like OMIM 601754 GeneCards

Involved in the degradation of ubiquitinated proteins

A polymorphism in the promoter region associates with schizophrenia in an Italian case and family study (De Luca et al., 2001).
UHMK1
U2AF homology motif (UHM) kinase 1 (Kis or Kist)
OMIM 608849 GeneCards
Phosphorylated substrates include myelin basic protein (MBP), stathmin (STMN1) and synapsin 1(SYN1) (Maucuer et al, 1997)  and p27kip1(CDKN1B. Activated by growth factors ( Boehm et al, 2002) Association reported in a UK study
(Puri et al, 2006)
USP46 ubiquitin specific peptidase 46   Japanese study Kushima et al, 2010

MLC1 (WKL1) megalencephalic leukoencephalopathy with subcortical cysts 1 homolog (human) OMIM 605908 GeneCards

Putative transporter: Binds to DISC1(Millar et al., 2003) Part of the dystrophin/glycoprotein complex at astrocytic end feet Boor et al, 2007 Binds to dystrophin glycoprotein complex (DGC) icluding syntrophin, dystrobrevin, Kir4.1 (KCNJ10) and caveolin-1 Lanceotti et al, 2010(Dystrobrevins binds ro DTNBP1). CAV1 binds to ABCB1, EGFR, ERBB2, FYN, ESR1: KCNJ10 binds to DLG1 and DLG4

Associated with familial catatonic schizophrenia (Leegwater et al., 2002) and associates with schizophrenia and bipolar disorder in Indian case and family studies(Meyer et al., 2001;Verma et al., 2005b)
TCF4 Transcription factor 4   International Schizophrenia Consortium
TUBA8 tubulin, alpha 8 OMIM GeneCards Binds to metabotropic receptor GRM7 Saugstad et al, 2002 (Fallin et al., 2005)
VRK2 vaccinia related kinase 2   GWAS Stefansson et al, 2009
XRCC1 X-ray repair complementing defective repair in Chinese hamster cells 1 GeneCards DNA repair Association reported in an Iranian case-control studySaadat et al, 2007
ZBED4 zinc finger, BED-type containing 4   US study Nyegaard et al, 2009

ZNF74 zinc finger protein 74 (Cos52) OMIM 194548 GeneCards

Associated with age of onset in a Japanese case study  (Takase et al., 2001).
ZNF184 Zinc finger protein 184   GWAS Shi et al, 2009
ZNF804A Zinc finger protein 804A   GWAS O'Donovan et al, 2008
ZNF518A Zinc finger protein 518A   GWAS European ancestry Shi et al, 2009
ZNF667 zinc finger protein 667   GWAS Moskvina et al, 2009

Runs of Homozygosity (see Lencz et al, 2007 ) In progress

ROH Genes General function and relation to other genes Association evidence Expression data
roh250 DYNC2H1 dynein, cytoplasmic 2, heavy chain 1 GeneCards Intracellular transport    
roh321 PIK3C3 (vps34) GeneCards vps34 Already implicated (see above)  
roh314

CRHR1 Corticotropin releasing hormone receptor 1 GeneCards

CRH receptor involved in stress responses    
IMP5 intramembrane protease 5 GeneCards Uncharacterised protease    
MAPT Microtubule associated protein tau GeneCards Microtubule related    
STH Saitohin GeneCards Interacts with peroxiredoxin suggesting a role in oxidative stress Gao et al, 2005    
KIAA1267 Uncharacterised    
ARL17 ADP-ribosylation factor-like 17 GeneCards Uncharacterised    
LRRC37A leucine rich repeat containing 37A GeneCards Uncharacterised    
LRRC37A2 Leucine rich repeat containing 37, member A2 GeneCards Uncharacterised    
NSF N-ethylmaleimide-sensitive factor GeneCards Intracellular membtrane fusion and exocytosis of synaptic vesicles (transmitter release). Binds to STX1AHanson et al, 1995 . Regulates AMPA receptor traffic Zhao et al, 2007 via a cascade involving NMDA receptors, nitric oxide and PICK1 Sossa et al, 2007    
WNT3 wingless-type MMTV integration site family, member 3 GeneCards Wnt signaling: regulates adult hippocampal neurogenesis Lie et al, 2005    
WNT9B wingless-type MMTV integration site family, member 9B GeneCards Wnt signaling : Neuronal differentiation    
RPRML reprimo-like GeneCards Uncharacterised    
CDC27 cell division cycle 27 homolog GeneCards Anaphase complex member controlling ubiquitin-mediated proteolysis of B-type cyclins    
       
roh52 CHN1 chimerin (chimaerin) 1 GeneCards Neuronal diacylglycerol-binding protein with Rho GTPase-activating protein domain that inactivates Rac1. Binds to GRIN2A and regulates dendritic spine density Ven de Ven et al, 2005. Increased activity leads to pruning of dendritic spines Buttery et al, 2006 Counters the inhibitory effects of MAG on neutite outgrowth Mizuno et al, 2004    
ATF2 activating transcription factor 2 GeneCards Dimerises with c-jun and posseses histone acetyltransferase activity    
ATP5G3 ATP synthase, H+ transporting, mitochondrial F0 complex, subunit C3 (subunit 9) GeneCards Mitochondrial ATP synthase    
roh15    DUSP12 Dual specificity phosphatase 12 GeneCards Negative regulation of MAP kinases    
ATF6 ctivating transcription factor 6 GeneCards Important role in the endoplasmic reticulum stress pathway . Regulates XBP1 expression Hirota et al, 2006 The ATF6 pathway is blocked by BDNF Chen et al, 2007. See eif2b page    
OLFML2B olfactomedin-like 2B GeneCards Uncharacterised    
NOS1AP nitric oxide synthase 1 (neuronal) adaptor protein GeneCards Binds to NOS1 Already implicated (see above)  
roh129  SGCD sarcoglycan, delta (35kDa dystrophin-associated glycoprotein) GeneCards Component of the dystrophin complex : also plays a role in peripheral myelination Cai et al, 2007    
MRPL22 mitochondrial ribosomal protein L22 GeneCards Mitochondrial protein synthesis    
roh291      GPHN Gephyrin GeneCards Anchors inhibitory (GABA) receptors to the postsynaptic cytoskeleton    
FAM71D (C14orf54) family with sequence similarity 71, member D GeneCards Uncharacterised    
MPP5 membrane protein, palmitoylated 5 (MAGUK p55 subfamily member 5) GeneCards Receptor clustering    
ATP6V1D ATPase, H+ transporting, lysosomal 34kDa, V1 subunit D GeneCards Acidification of intracellular organelles    
EIF2S1 eukaryotic translation initiation factor 2, subunit 1 alpha, 35kDa GeneCards Phosphorylated by four eif2alpha kinases, themselves activated by oxidative stress (hri), viruses (pkr), starvation (gcn2) and endoplasmic reticulum stress (perk): Phosphorylated eif2a activates ATF4, the binding partner of DISC1 and shuts down protein synthesis via eif2b inhibition Carter, 2007: See eif2b page    
PLEK2 pleckstrin 2 GeneCards Cytoskeletal actin organisation Hamaguchi et al, 2007    
roh55            GULP1 GULP, engulfment adaptor PTB domain containing 1 GeneCards Adaptor protein required for engulfment of apoptotic cells by phagocytes    
DIRC1 disrupted in renal carcinoma 1 GeneCards Uncharacterised    
COL3A1 collagen, type III, alpha 1 (Ehlers-Danlos syndrome type IV, autosomal dominant) GeneCards Collagen: Binds to MAG Fahrig et al, 1987    
COL5A2 collagen, type V, alpha 2 GeneCards Collagen    
WDR75 WD repeat domain 75 GeneCards Uncharacterised    
SLC40A1 solute carrier family 40 (iron-regulated transporter), member 1 GeneCards Iron homoeostasis    
ASNSD1 (NS3TP1)asparagine synthetase domain containing 1 GeneCards Uncharacterised    
ANKAR ankyrin and armadillo repeat containing GeneCards Uncharacterised    
OSGEPL1 O-sialoglycoprotein endopeptidase-like 1 GeneCards Uncharacterised endopeptidase    
ORMDL1 ORM1-like 1 GeneCards Uncharacterised    
PMS1 postmeiotic segregation increased 1 GeneCards Putative DNA mismatch repair    
GDF8 MSTN Myostatin GeneCards Negative regulator of muscle growth    
roh173 IMPAD1 inositol monophosphatase domain containing 1 GeneCards Uncharacterised inositol monophosphatase    

Rare structural variants (microdeletions and duplications) belonging to common pathways Walsh et al, 2008 (in progress)

Gene General function and relation to other genes Association evidence
C11orf41

Uncharacterised  
CAV1 caveolin 1, caveolae protein, 22kDa
Scaffolding protein linking integrins to FYN. Associates with EGFR and ERBB2  
DLG2 discs, large homolog 2, chapsyn-110
NMDA and ERBB4 receptor scaffold Already implicated (see above)
ERBB4 v-erb-a erythroblastic leukemia viral oncogene homolog 4
Neuregulin receptor Already implicated (see above)
GRM7 glutamate receptor, metabotropic 7
Metabotropic glutamate receptorbinding to PICK1 and TUBA8 Already implicated (see above)
HIPK3 homeodomain interacting protein kinase 3
Regulates FAS and FADD apoptosis  
LAMA1 laminin, alpha 1
Component of basement membranes  
LARGE like-glycosyltransferase
Binds to dystroglycan  
MAGI2 membrane associated guanylate kinase, WW and PDZ domain containing 2
Binds to atrophin 1 (DRPLA)  
MLL3 myeloid/lymphoid or mixed-lineage leukemia 3
Histone methylation and transcriptional coactivation  
NBPF10 neuroblastoma breakpoint family, member 10
Uncgharcterised  
PHTF2 putative homeodomain transcription factor 2
No data  
PRKAG2 protein kinase, AMP-activated, gamma 2 non-catalytic subunit
Energy-sensing kinase monitoring cellular energy status and functions by inactivating key enzymes involved in regulating de novo biosynthesis of fatty acid and cholesterol.  
PRKCD protein kinase C, delta
Involved in B-cell signaling and the growth and development of multiple cells  
PTK2 PTK2 protein tyrosine kinase 2
Focal adhesion kinase involved in EGFR , ERBB2 ERBB3 and integrin signaling (see gene data). Also involved in NMDA receptor-dependent LTP Yang et al, 2003  
PTPRM protein tyrosine phosphatase, receptor type, M
   
SKP2 S-phase kinase-associated protein 2 (p45)
F-box protein: one of four subunits of ubiquitin protein ligase complexes called SCFs (SKP1-cullin-F-box), which function in phosphorylation-dependent ubiquitination  
SLC12A9 solute carrier family 12 (potassium/chloride transporters), member 9
Potasssium/chloride transport  
SLC1A3 solute carrier family 1 (glial high affinity glutamate transporter), member 3
Glial glutamate transporter  
SLC35F3 solute carrier family 35, member F3
Uncharacterised  
SMARCA2 SWI/SNF related, matrix associated, actin dependent regulator of chromatin, subfamily a, member 2
Members of this family have helicase and ATPase activities and may regulate transcription of certain genes by altering the chromatin structure around those genes.  
STON1-GTF2A1L stoned B/TFIIA alpha/beta like factor
Uncharacterised fusion protein  
TARBP1 TAR (HIV-1) RNA binding protein 1    
TMC4 transmembrane channel-like 4 Uncharacterised  

 

Environmental risk factors.

EPIGENETIC PHENOMENA Wikipedia  
Hypomethylation of leukocyte DNA in males Shimabukuro et al, 2006
Hypermethylation of the Reelin promoter (cortex) Grayson et al, 2005
Hypomethylation of membrane bound COMT promoter (Frontal cortex) Abdolmaleky et al, 2006

 



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Chromosomal regions

I was not able to identify concordant linkage data for chromosomal regions containing ALK (2p23), CTLA4 (2q33), ERBB4 (2q33.3-q34), PMX2B (4p12), FOXP2 (7q31), HTR5A (7q36.1), OPRS1 (9p13.3), B3GAT1 (11q25), NTF3 (12p13), DRPLA (12p13.31), PAH (12q22-q24.2), three genes in the 19q.13 region; PLA2G4C (19q13.3), GRIN2D (19q13.1-qter) and APOE, (19q13.2), S100B (21q22.3)  MED12 (Xq13) or PLP1 (Xq22). 

 

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