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Set Analyses:

VarElect

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GLUL Gene

protein-coding GIFtS: 71
GCID: GC01M182350

Glutamate-Ammonia Ligase

(Previous name: glutamate-ammonia ligase (glutamine synthase))
(Previous symbol: GLNS)

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(According to ¹HGNC, ²Entrez Gene,
³UniProtKB/Swiss-Prot, ⁴UniProtKB/TrEMBL, ⁵OMIM, ⁶GeneLoc, ⁷Ensembl, ⁸DME, ⁹miRBase, ¹⁰fRNAdb, ¹²H-InvDB, ¹³NCBI, ¹⁴NONCODE, and/or ¹⁵RNAdb)
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Aliases
Glutamate-Ammonia Ligase¹ ²		Glutamate-Ammonia Ligase (Glutamine Synthase)¹
GLNS¹ ² ³ ⁵		PIG43²
Glutamine Synthetase¹ ²		PIG59²
Glutamate Decarboxylase² ³		Cell Proliferation-Inducing Protein 59²
Glutamate--Ammonia Ligase² ³		Glutamine Synthase²
GS² ³		Proliferation-Inducing Protein 43²
EC 6.3.1.2³ ⁸		EC 4.1.1.15³

External Ids:	HGNC: 4341¹	Entrez Gene: 2752²	Ensembl: ENSG00000135821⁷	OMIM: 138290⁵	UniProtKB: P15104³

Export aliases for GLUL gene to outside databases

Previous GC identifers: GC01M179878 GC01M177780 GC01M178872 GC01M179592 GC01M179593 GC01M179083 GC01M180617 GC01M153587

(According to Entrez Gene, GeneCards, Tocris Bioscience, Wikipedia's Gene Wiki, PharmGKB,
UniProtKB/Swiss-Prot, and/or UniProtKB/TrEMBL)
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Entrez Gene summary for GLUL Gene:

The protein encoded by this gene belongs to the glutamine synthetase family. It catalyzes the synthesis of

glutamine from glutamate and ammonia. Glutamine is a main source of energy and is involved in cell proliferation,

inhibition of apoptosis, and cell signaling. This gene is expressed during early fetal stages, and plays an

important role in controlling body pH by removing ammonia from circulation. Mutations in this gene are associated

with congenital glutamine deficiency. Several alternatively spliced transcript variants have been found for this

gene.(provided by RefSeq, Oct 2009)

GeneCards Summary for GLUL Gene:

GLUL (glutamate-ammonia ligase) is a protein-coding gene. Diseases associated with GLUL include glutamine deficiency, congenital, and glutathione synthetase deficiency. GO annotations related to this gene include manganese ion binding and identical protein binding.

UniProtKB/Swiss-Prot: GLNA_HUMAN, P15104

Function: This enzyme has 2 functions: it catalyzes the production of glutamine and 4-aminobutanoate

(gamma-aminobutyric acid, GABA), the latter in a pyridoxal phosphate-independent manner (By similarity).

Essential for proliferation of fetal skin fibroblasts

(According to GeneLoc and/or HGNC, and/or
Entrez Gene (NCBI build 37),
and/or miRBase,
Genomic Views according to UCSC (hg19) and Ensembl (release 75), Regulatory elements and Epigenetics data according to QIAGEN, and/or SwitchGear Genomics)
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RefSeq DNA sequence:

NC_000001.11 NT_004487.20 NC_018912.2

Regulatory elements:

Regulatory transcription factor binding sites in the GLUL gene promoter:
ATF-2 c-Jun
Other transcription factors

SwitchGear Promoter luciferase reporter plasmids (see all 2): GLUL promoter sequence

Search Chromatin IP Primers for GLUL

Epigenetics:

DNA Methylation CpG Assay Predesigned for Pyrosequencing in human, mouse, rat GLUL

Genomic Location:
Genomic View: UCSC Golden Path with GeneCards custom track

Entrez Gene cytogenetic band: 1q31 Ensembl cytogenetic band: 1q25.3 HGNC cytogenetic band: 1q31

GLUL Gene in genomic location: bands according to Ensembl, locations according to (and/or Entrez Gene and/or Ensembl if different)
GLUL gene location

GeneLoc information about chromosome 1 GeneLoc Exon Structure

GeneLoc location for GC01M182350: view genomic region (about GC identifiers)

Start:	182,350,839 bp from pter	End:	182,361,341 bp from pter
Size:	10,503 bases	Orientation:	minus strand

(According to ¹UniProtKB, HORDE, ²neXtProt, Ensembl, and/or Reactome, Modification sites according to PhosphoSitePlus, Specific Peptides from DME, RefSeq according to NCBI, PDB rendering according to OCA and/or Proteopedia, Recombinant Proteins from EMD Millipore, R&D Systems, GenScript, Enzo Life Sciences, OriGene, Novus Biologicals, Sino Biological, ProSpec, and/or Cloud-Clone Corp.,
Biochemical Assays by EMD Millipore, R&D Systems, OriGene, GenScript, Cell Signaling Technology, Enzo Life Sciences, and/or Cloud-Clone Corp., Antibodies by EMD Millipore, R&D Systems, Cell Signaling Technology, OriGene, Novus Biologicals, Thermo Fisher Scientific, LSBio, Abcam, and/or Cloud-Clone Corp.)
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UniProtKB/Swiss-Prot: GLNA_HUMAN, P15104 (See protein sequence)

Recommended Name: Glutamine synthetase

Size: 373 amino acids; 42064 Da

Cofactor: Biotin (By similarity)

Cofactor: Magnesium or manganese (By similarity)

Subunit: Homooctamer and homotetramer. Interacts with PALMD (By similarity)

Developmental stage: Expressed during early fetal stages

2 PDB 3D structures from and Proteopedia for GLUL:

2OJW (3D)

2QC8 (3D)

Secondary accessions: Q499Y9 Q5T9Z1 Q7Z3W4 Q8IZ17

Explore the universe of human proteins at neXtProt for GLUL: NX_P15104

Explore proteomics data for GLUL at MOPED

Post-translational modifications:

Ubiquitinated by ZNRF1 (By similarity)¹

Ubiquitination² at Lys11, Lys14, Lys25, Lys91, Lys95, Lys103, Lys259, Lys268, Lys276, Lys291,
Lys334, Lys372

Modification sites at PhosphoSitePlus

Selected DME Specific Peptides for GLUL (P15104) (see all 24)

GFPGPQG

GTNAEVM

YIWIDGT

PQGPYYC

See GLUL Protein Expression from SPIRE MOPED, PaxDB, and MaxQB

REFSEQ proteins (3 alternative transcripts):

NP_001028216.1 NP_001028228.1 NP_002056.2

ENSEMBL proteins:

ENSP00000356537 ENSP00000344958 ENSP00000307900 ENSP00000398320

Reactome Protein details: P15104

GLUL Human Recombinant Protein Products:

	Browse Purified and Recombinant Proteins at EMD Millipore
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	Browse recombinant and purified proteins available from Enzo Life Sciences
	OriGene Purified Proteins for GLUL OriGene Protein Over-expression Lysate for GLUL OriGene MassSpec for GLUL OriGene Custom Protein Services for GLUL
	GenScript Custom Purified and Recombinant Proteins Services for GLUL
	Novus Biologicals GLUL Proteins Novus Biologicals GLUL Lysates
	Browse Sino Biological Recombinant Proteins Browse Sino Biological Cell Lysates
	ProSpec Recombinant Protein for GLUL
	Browse Proteins at Cloud-Clone Corp.

GLUL Antibody Products:

	EMD Millipore Mono- and Polyclonal Antibodies for the study of GLUL
	Browse R&D Systems for Antibodies
	OriGene Antibodies for GLUL OriGene Custom Antibody Services for GLUL
	Novus Biologicals GLUL Antibodies
	Abcam antibodies for GLUL
	Browse Antibodies at Cloud-Clone Corp.
	ThermoFisher Antibody for GLUL
	LSBio Antibodies in human, mouse, rat for GLUL

GLUL Assay Products:

	Browse Kits and Assays available from EMD Millipore
	OriGene Custom Assay Services for GLUL
	Browse R&D Systems for biochemical assays
	GenScript Custom Assay Services for GLUL
	Browse Enzo Life Sciences for kits & assays
	Browse ELISAs at Cloud-Clone Corp. Browse CLIAs at Cloud-Clone Corp.

(According to HGNC, IUPHAR, InterPro, ProtoNet, UniProtKB, and/or BLOCKS, Sets of similar genes according to GeneDecks)
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5 InterPro protein domains:

IPR008147 Gln_synt_beta

IPR014746 Gln_synth/guanido_kin_cat_dom

IPR008146 Gln_synth_cat_dom

IPR027302 Gln_synth_N_conserv_site

IPR027303 Gln_synth_gly_rich_site

Graphical View of Domain Structure for InterPro Entry P15104

ProtoNet protein and cluster: P15104

UniProtKB/Swiss-Prot: GLNA_HUMAN, P15104

Similarity: Belongs to the glutamine synthetase family

GLUL for domains About GeneDecksing

(According to ¹UniProtKB, Genatlas, LifeMap Discovery™, IUBMB, and/or ²DME, Human phenotypes from GenomeRNAi, Animal models from MGI Mar 06 2013, inGenious Targeting Laboratory, genOway,
transcription factor targeting from QIAGEN and/or HOMER, miRNA Gene Targets from miRTarBase, shRNA from OriGene, siRNAs from OriGene, QIAGEN, microRNA from QIAGEN, SwitchGear Genomics, Gene Editing from DNA2.0, Clones from OriGene, GenScript, Sino Biological, DNA2.0, and Vector BioLabs, Cell Lines from GenScript, ESI BIO, In Situ Hybridization Assays from Advanced Cell Diagnostics, Ontologies according to Gene Ontology Consortium 01 Apr 2014 via Entrez Gene.)
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Molecular Function:

UniProtKB/Swiss-Prot Summary: GLNA_HUMAN, P15104

Function: This enzyme has 2 functions: it catalyzes the production of glutamine and 4-aminobutanoate

(gamma-aminobutyric acid, GABA), the latter in a pyridoxal phosphate-independent manner (By similarity).

Essential for proliferation of fetal skin fibroblasts

Catalytic activity: ATP + L-glutamate + NH(3) = ADP + phosphate + L-glutamine

Catalytic activity: L-glutamate = 4-aminobutanoate + CO(2)

Induction: By glucocorticoids. Vitamin D and the Wnt signaling pathway inhibit its expression and activity

Genatlas biochemistry entry for GLUL:

L-glutamine synthetase

Enzyme Numbers (IUBMB): EC 6.3.1.2¹ ² EC 4.1.1.15¹

Gene Ontology (GO): Selected molecular function terms (see all 9): About this table

GO ID	Qualified GO term	Evidence	PubMed IDs
GO:0000287	magnesium ion binding	IEA	--
GO:0004351	glutamate decarboxylase activity	IEA	--
GO:0004356	glutamate-ammonia ligase activity	EXP	--
GO:0005515	protein binding	--	--
GO:0005524	ATP binding	IEA	--

GLUL for ontologies About GeneDecksing

Phenotypes:
2 GenomeRNAi human phenotypes for GLUL:

Increased G1 DNA content

Increased cell death HMECs cel

5 MGI mutant phenotypes (inferred from 3 alleles) (MGI details for Glul):

behavior/neurological

growth/size/body

homeostasis/metabolism

mortality/aging

nervous system

GLUL for phenotypes About GeneDecksing

Animal Models:

inGenious Targeting Laboratory: Let us create your new Knockout/Knockin mouse model for GLUL

inGenious Targeting Laboratory: Contact us about creating complex and humanized mouse models for GLUL

genOway customized KO model: permanent, tissue-specific or time-controlled inactivation for GLUL

genOway customized Knockin model: humanization, point mutation, expression monitoring, etc. for GLUL

miRNA Products:		miRTarBase miRNAs that target GLUL: hsa-mir-125a-3p (MIRT038640), hsa-mir-183-5p (MIRT025018), hsa-mir-320b (MIRT036244), hsa-mir-615-3p (MIRT040016), hsa-let-7e-5p (MIRT051664), hsa-mir-186-5p (MIRT021141), hsa-mir-484 (MIRT041957), hsa-mir-185-5p (MIRT045330), hsa-mir-29a-3p (MIRT004021), hsa-mir-296-3p (MIRT038394), hsa-mir-423-5p (MIRT038153), hsa-mir-196a-5p (MIRT048155), hsa-mir-937-3p (MIRT036638), hsa-let-7a-5p (MIRT052399), hsa-mir-320a (MIRT044856), hsa-mir-24-3p (MIRT030540), hsa-mir-149-5p (MIRT045626), hsa-mir-877-5p (MIRT037273), hsa-let-7f-5p (MIRT051362), hsa-mir-877-3p (MIRT036966)

		Block miRNA regulation of human, mouse, rat GLUL using miScript Target Protectors
		Selected qRT-PCR Assays for microRNAs that regulate GLUL (see all 22):
		hsa-miR-140-5p hsa-miR-582-3p hsa-miR-3607-3p hsa-miR-4286 hsa-miR-1285 hsa-miR-340 hsa-miR-3170 hsa-miR-497*

		SwitchGear 3'UTR luciferase reporter plasmid: GLUL 3' UTR sequence

Inhib. RNA Products:		OriGene RNAi products in human, mouse, rat for GLUL
		Predesigned siRNA for gene silencing in human, mouse, rat GLUL

Gene Editing
Products:

DNA2.0 Custom Protein Engineering Service for GLUL

Clone Products:		OriGene clones in human, mouse for GLUL (see all 19) OriGene ORF clones in mouse, rat for GLUL OriGene custom cloning services - gene synthesis, subcloning, mutagenesis, variant library, vector shuttling
		GenScript: all cDNA clones in your preferred vector (see all 3): GLUL (NM_002065)
		Sino Biological Human cDNA Clone for GLUL
		DNA2.0 Custom Codon Optimized Gene Synthesis Service for GLUL
		Vector BioLabs ready-to-use adenovirus/AAV for human, mouse, rat GLUL

Cell Line Products:		GenScript Custom overexpressing Cell Line Services for GLUL
		Browse ESI BIO Cell Lines and PureStem Progenitors for GLUL

In Situ Assay
Products:

Advanced Cell Diagnostics RNAscope RNA in situ hybridization assays for GLUL

(According to UniProtKB, COMPARTMENTS Subcellular localization database, Ontologies according to Gene Ontology Consortium 01 Apr 2014 via Entrez Gene.)
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Subcellular locations from UniProtKB/Swiss-Prot

GLNA_HUMAN, P15104: Cytoplasm. Mitochondrion (By similarity)

Subcellular locations from COMPARTMENTS:

Compartment	Confidence
cytosol	4
chloroplast	3
cytoskeleton	3
endoplasmic reticulum	2
extracellular	2
mitochondrion	2
nucleus	2
plasma membrane	2

About this image

Gene Ontology (GO): Selected cellular component terms (see all 12): About this table

GO ID	Qualified GO term	Evidence	PubMed IDs
GO:0005622	intracellular	--	--
GO:0005737	cytoplasm	IDA	--
GO:0005739	mitochondrion	IEA	--
GO:0005791	rough endoplasmic reticulum	IEA	--
GO:0005829	cytosol	TAS	--

GLUL for ontologies About GeneDecksing

(SuperPaths according to PathCards, Pathways according to R&D Systems, Cell Signaling Technology, KEGG, PharmGKB, BioSystems, Sino Biological, Reactome, Tocris Bioscience, GeneGo (Thomson Reuters), QIAGEN, and/or UniProtKB, Sets of similar genes according to GeneDecks, Interaction Networks according to QIAGEN, and/or STRING, Interactions according to ¹UniProtKB, ²MINT, ³I2D, and/or ⁴STRING, with links to IntAct and Ensembl, Ontologies according to Gene Ontology Consortium 01 Apr 2014 via Entrez Gene).
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SuperPaths for GLUL About (see all 12)

See pathways by source

SuperPath

Contained pathways About

Neurotransmitter uptake and Metabolism In Glial Cells

	Neurotransmitter uptake and Metabolism In Glial Cells			Astrocytic Glutamate-Glutamine Uptake And Metabolism	0.00
	glutamine biosynthesis I	0.00

Metabolism

Metabolism

0.38

Metabolic pathways

0.38

Transmission across Chemical Synapses

Transmission across Chemical Synapses

0.72

Neuronal System

0.68

Circadian entrainment

GABAergic synapse

0.53

Glutamatergic synapse

0.39

Metabolism of amino acids and derivatives

Metabolism of amino acids and derivatives

Amino acid synthesis and interconversion (transamination)

0.00

Pathways by source See SuperPaths
Show all pathways

1 BioSystems Pathway for GLUL

glutamine biosynthesis I

2 Reactome Pathways for GLUL

	Astrocytic Glutamate-Glutamine Uptake And Metabolism
	Amino acid synthesis and interconversion (transamination)

Selected Kegg Pathways (Kegg details for GLUL) (see all 8):

	Alanine, aspartate and glutamate metabolism
	Arginine and proline metabolism
	Glyoxylate and dicarboxylate metabolism
	Nitrogen metabolism
	Metabolic pathways

GLUL for pathways About GeneDecksing

Pathway & Disease-focused RT² Profiler PCR Arrays including GLUL:

Nephrotoxicity in human mouse rat

GABA & Glutamate in human mouse rat

Necrosis in human mouse rat

Interactions:

Search GeneGlobe Interaction Network for GLUL

STRING Interaction Network Preview (showing 5 interactants - click image to see 25)

Selected Interacting proteins for GLUL (P15104^{1, 2, 3} ENSP00000307900⁴) via UniProtKB, MINT, STRING, and/or I2D (see all 187)

Interactant		Interaction Details
GeneCard	External ID(s)	Interaction Details
SKIL	P12757²^,³	MINT-61661 I2D: score=3
GLUL	P15104¹^,²	EBI-746653,EBI-746653 MINT-68068
NUDT18	Q6ZVK8²^,³, ENSP00000307852⁴	MINT-68171 I2D: score=5 STRING: ENSP00000307852
PLEKHF2	Q9H8W4²^,³, ENSP00000322373⁴	MINT-66879 I2D: score=5 STRING: ENSP00000322373
PIAS1	O75925³, ENSP00000249636⁴	I2D: score=3 STRING: ENSP00000249636

About this table

Gene Ontology (GO): Selected biological process terms (see all 15): About this table

GO ID	Qualified GO term	Evidence	PubMed IDs
GO:0001504	neurotransmitter uptake	TAS	--
GO:0006536	glutamate metabolic process	--	--
GO:0006538	glutamate catabolic process	TAS	--
GO:0006542	glutamine biosynthetic process	TAS	--
GO:0006807	nitrogen compound metabolic process	--	--

GLUL for ontologies About GeneDecksing

(Chemical Compounds according to UniProtKB, Enzo Life Sciences, EMD Millipore, Tocris Bioscience, ApexBio, HMDB, BitterDB, and/or Novoseek, Ligands according to IUPHAR, and Drugs according to DrugBank, Enzo Life Sciences, and/or PharmGKB)
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Browse Small Molecules at EMD Millipore

Enzo Life Sciences drugs & compounds for GLUL

Browse compounds at ApexBio

Browse Tocris compounds for GLUL (GLNA)

2 HMDB Compounds for GLUL About this table

Compound	Synonyms	CAS #	PubMed Ids
L-Glutamic acid	(2S)-2-Aminopentanedioate (see all 49)	56-86-0	--
L-Glutamine	(2S)-2,5-diamino-5-oxopentanoic acid (see all 31)	56-85-9	--

9 DrugBank Compounds for GLUL About this table

Compound	Synonyms	CAS #	Type	Actions	PubMed Ids
L-Glutamine	(S)-2,5-diamino-5-oxopentanoic acid (see all 4)	56-85-9	target	product of	16484281 16326016 16547651 16382338 16530878
L-Glutamic Acid	(2S)-2-Aminopentanedioic acid (see all 15)	56-86-0	target	--	16973920 16673373 16876704 17284842
L-Methionine	(S)-2-amino-4-(methylthio)butanoic acid (see all 6)	63-68-3	enzyme	inhibitor	6127334 4144717 4308004
Ceftriaxone	Cefatriaxone (see all 4)	73384-59-5	enzyme	inducer	10696140 20423712
Diazoxide	--	364-98-7	enzyme	inhibitor	7926271
Dihydroergotoxine	--	11032-41-0	enzyme	inducer	6117619
Ethyl biscoumacetate	Pelentan (see all 2)	548-00-5	enzyme	inhibitor	7195738
Pegvisomant	GH (see all 6)	218620-50-9	enzyme	inhibitor	1970314
Picrotoxin	--	124-87-8	enzyme	inhibitor	840320

Selected Novoseek inferred chemical compound relationships for GLUL gene (see all 95) About this table

Compound	-log (P-Val)	Hits	PubMed IDs for Articles with Shared Sentences (# sentences)
glutamine	96.3	1584	10681395 (8), 10943675 (8), 11589877 (8), 9288153 (8) (see all 99)
methionine sulfoximine	93.5	62	14714472 (2), 20060132 (2), 12390519 (1), 12840174 (1) (see all 38)
glutamate	81.3	432	19728998 (7), 20064572 (6), 20373878 (5), 12390519 (4) (see all 99)
ammonium	73.2	84	8563633 (5), 16173463 (3), 19464754 (3), 14757761 (3) (see all 33)
gamma-glutamyl phosphate	71	5	8563633 (3), 10664863 (1)
phosphinothricin	67	24	17888468 (6), 16510221 (3), 16504520 (1), 12723887 (1) (see all 7)
alpha-ketoglutarate	64.3	5	18421771 (1), 8971000 (1), 1360177 (1), 17477312 (1)
2',3'-cyclic nucleotide	58.3	2	1979258 (1), 10366697 (1)
carbamoyl phosphate	54.2	13	1976588 (5), 17518723 (1), 1687030 (1), 8587121 (1) (see all 5)
nitrate	52.3	38	7534371 (2), 9244251 (1), 18419739 (1), 1967601 (1) (see all 23)

GLUL for compounds About GeneDecksing

(Secondary structures according to fRNAdb,
GenBank/EMBL/DDBJ Accessions according to
Unigene (Build 236 Homo sapiens; Apr 25 2013) or GenBank,
RefSeq according to Entrez Gene,
DOTS (version 10), and/or AceView, transcript ids from Ensembl with links to UCSC,
Conferences by KenesGroup, exon structure from GeneLoc, alternative splicing isoforms according to ASD and/or ECgene,
siRNAs from OriGene, QIAGEN, shRNA from OriGene, microRNA from QIAGEN, SwitchGear Genomics,
Tagged/untagged cDNA clones from OriGene, GenScript, DNA2.0, Vector BioLabs, Primers from OriGene, and/or QIAGEN )
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REFSEQ mRNAs for GLUL gene (3 alternative transcripts):

NM_001033044.2 NM_001033056.2 NM_002065.5

Unigene Cluster for GLUL:

Glutamate-ammonia ligase

Hs.518525 [show with all ESTs]

Unigene Representative Sequence: NM_002065

12 Ensembl transcripts including schematic representations, and UCSC links where relevant:

ENST00000331872(uc010pnt.2 uc001gpd.2) ENST00000491322 ENST00000339526

ENST00000463851 ENST00000461447 ENST00000484996 ENST00000489818 ENST00000475808

ENST00000480604 ENST00000462444 ENST00000311223(uc001gpb.2 uc001gpc.2)

ENST00000417584(uc001gpa.2)

Congresses - knowledge worth sharing:
European Congress of Clinical Microbiology and Infectious Diseases (ECCMID) 10 - 13 May 2014

miRNA Products:		Block miRNA regulation of human, mouse, rat GLUL using miScript Target Protectors
		Selected qRT-PCR Assays for microRNAs that regulate GLUL (see all 22):
		hsa-miR-140-5p hsa-miR-582-3p hsa-miR-3607-3p hsa-miR-4286 hsa-miR-1285 hsa-miR-340 hsa-miR-3170 hsa-miR-497*

		SwitchGear 3'UTR luciferase reporter plasmid: GLUL 3' UTR sequence

Inhib. RNA Products:		OriGene RNAi products in human, mouse, rat for GLUL
		Predesigned siRNA for gene silencing in human, mouse, rat GLUL

Clone Products:		OriGene clones in human, mouse for GLUL (see all 19) OriGene ORF clones in mouse, rat for GLUL OriGene custom cloning services - gene synthesis, subcloning, mutagenesis, variant library, vector shuttling
		GenScript: all cDNA clones in your preferred vector (see all 3): GLUL (NM_002065)
		DNA2.0 Custom Codon Optimized Gene Synthesis Service for GLUL
		Vector BioLabs ready-to-use adenovirus/AAV for human, mouse, rat GLUL

Primer Products:		OriGene qPCR primer pairs and template standards for GLUL OriGene qSTAR qPCR primer pairs in human, mouse for GLUL
		Pre-validated RT² qPCR Primer Assay in human, mouse, rat GLUL
		QuantiTect SYBR Green Assays in human, mouse, rat GLUL
		QuantiFast Probe-based Assays in human, mouse, rat GLUL

Additional mRNA sequence:

AK122784.1 AK301600.1 AY486122.1 AY513283.1 AY947343.1 BC010037.2 BC011700.2 BC011852.2

BC018992.1 BC031964.1 BC051726.1 BC127883.1 BX537384.1 S70290.1 X59834.1 Y00387.2

Selected DOTS entries (see all 59):

DT.97789039 DT.95297120 DT.91909750 DT.95176992 DT.100687848 DT.91707765 DT.100687841 DT.100687844

DT.92475794 DT.95075484 DT.100687842 DT.102837867 DT.85104875 DT.91813488 DT.100687839 DT.91875449

DT.95075478 DT.100873067 DT.75153967 DT.97860944 DT.121426398 DT.100040371 DT.100687840 DT.100687850

Selected AceView cDNA sequences (see all 1446):

AA779772 AA418870 AI688954 BU726821 BQ008649 BF939525 BQ188827 BX382320

CD677430 BU685432 CB117590 CB132724 BG056775 BQ009780 BQ691536 BP377716

BQ690979 AA418869 BF339029 T08012 BM697627 BU543600 BF724702 F03096

GeneLoc Exon Structure

Selected Alternative Splicing Database (ASD) splice patterns (SP) for GLUL (see all 20) About this scheme

ExUns:	1a	·	1b	·	1c	·	1d	^	2	^	3	^	4a	·	4b	^	5	^	6a	·	6b	^	7	^	8a	·	8b	·	8c	·	8d	^	9	^	10a	·	10b	^	11a	·	11b	^	12a	·	12b	·	12c	^	13a	·	13b	·
SP1:									-		-		-		-				-		-		-										-				-		-								-		-
SP2:									-		-		-		-		-		-		-		-										-				-		-								-		-
SP3:																			-		-		-										-				-		-								-		-
SP4:													-		-		-		-		-		-										-				-		-								-		-
SP5:									-		-		-		-		-		-		-		-						-		-		-		-		-		-		-		-		-		-		-		-

ExUns:	13c	·	13d	·	13e	·	13f	^	14a	·	14b	·	14c	^	15a	·	15b	·	15c	·	15d
SP1:					-		-
SP2:					-		-
SP3:					-		-
SP4:					-		-
SP5:					-		-

ECgene alternative splicing isoforms for GLUL

(RNA expression data according to H-InvDB, NONCODE, miRBase, and RNAdb, Expression images according to data from BioGPS, Illumina Human BodyMap, and CGAP SAGE, Sets of similar genes according to GeneDecks, in vivo and in vitro expression data from LifeMap Discovery™, Protein expression images according to data from SPIRE ¹MOPED, ²PaxDb, and ³MaxQB, plus additional links to SOURCE, and/or BioGPS, and/or UniProtKB,
PCR Arrays from QIAGEN, Primers from OriGene, and/or QIAGEN, In Situ Hybridization Assays from Advanced Cell Diagnostics)
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GLUL expression in normal human tissues (normalized intensities)
See probesets specificity/sensitivity at GeneAnnot
About this imageBioGPS <intensity>^2/3
CGAP TAG: TACAGTATGT

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GLUL expression in embryonic tissues and stem cells About this table

Data from LifeMap, the Embryonic Development and Stem Cells Database
selected tissues (see all 27) fully expand


Brain (Nervous System) fully expand to see all 7 entries
Cerebral Cortex
Astrocyte-like cells

Eye (Sensory Organs) fully expand to see all 2 entries
Mature Muller Glia Cells Inner Nuclear Layer
Retina-like cells

Liver (Hepatobiliary System) fully expand to see all 2 entries
Perivenous Hepatocytes Liver Lobule

Thymus (Hematopoietic System) fully expand to see all 2 entries
Double Negative 2 Thymocytes Thymus
Thymus

Testis (Reproductive System)
Leydig Cells Testis Interstitium

GLUL Protein expression data from MOPED¹, PaxDb² and MaxQB³:

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GLUL Protein Expression

SOURCE GeneReport for Unigene cluster: Hs.518525

Pathway & Disease-focused RT² Profiler PCR Arrays including GLUL:

Nephrotoxicity in human mouse rat

GABA & Glutamate in human mouse rat

Necrosis in human mouse rat

Primer Products:		OriGene qPCR primer pairs and template standards for GLUL OriGene qSTAR qPCR primer pairs in human, mouse for GLUL
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(Orthologs according to ^1,2HomoloGene (²older version, for species not in ¹newer version), ³euGenes, ⁴SGD , ⁵MGI Mar 06 2013, with possible further links to Flybase and/or WormBase, and/or ⁶Ensembl pan taxonomic compara , Gene Trees according to Ensembl and TreeFam)
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This gene was present in the common ancestor of eukaryotes.

Orthologs for GLUL gene from Selected species (see all 30) About this table

Organism	Taxonomic classification	Gene	Description	Human Similarity	Orthology Type	Details
mouse (Mus musculus)	Mammalia	Glul^{1 , 5}	glutamate-ammonia ligase (glutamine synthetase)^{1, 5}	89.45(n)¹ 94.64(a)¹		1 (65.43 cM)⁵ 14645¹ NM_008131.3¹ NP_032157.2¹ 153899944⁵
chicken (Gallus gallus)	Aves	GLUL¹	glutamate-ammonia ligase	78.91(n) 88.74(a)		396489 NM_205493.1 NP_990824.1
lizard (Anolis carolinensis)	Reptilia	GLUL⁶	glutamate-ammonia ligase	88(a)	1 ↔ 1	GL343712.1(207755-222114)
African clawed frog (Xenopus laevis)	Amphibia	glul-prov²	glutamate-ammonia ligase (glutamine synthase)	76.31(n)		BC054153.1
zebrafish (Danio rerio)	Actinopterygii	AY423006.1²	--	83.48(n)		AY423006.1
fruit fly (Drosophila melanogaster)	Insecta	Gs2³ Gs1¹	glutamate-ammonia ligase³ Glutamine synthetase 1¹	60(a) (best of 2)³ 60.77(n)¹ 61.05(a)¹		10B11³ 33172¹ NM_001169368.2¹ NP_001162839.1¹
worm (Caenorhabditis elegans)	Secernentea	Y105C5B.28³ gln-1¹	gln-1¹	64(a) (best of 5)³ 59.13(n)¹ 60(a)¹		IV(16168058-16170839)³ 180875¹ NM_076611.3¹ NP_509012.2¹
baker's yeast (Saccharomyces cerevisiae)	Saccharomycetes	GLN1(YPR035W)⁴ GLN1¹	Glutamine synthetase (GS), synthesizes glutamine from more⁴ GLN1¹	57.12(n)¹ 57.02(a)¹		16(642208-643320)⁴ 856147^{1, 4} NP_015360.2¹ NP_015360.1⁴
thale cress (Arabidopsis thaliana)	eudicotyledons	GSR2¹	GSR2	56.76(n) 56.76(a)		842935 NM_105291.3 NP_176794.1
rice (Oryza sativa)	Liliopsida	Os.7879²	Oryza sativa shoot GS2 mRNA for chloroplastic glutamine more	73.63(n)		AK099252.1

ENSEMBL Gene Tree for GLUL (if available)
TreeFam Gene Tree for GLUL (if available)

(Paralogs according to ¹HomoloGene,
²Ensembl, and ³SIMAP, Pseudogenes according to ⁴Pseudogene.org Build 68)
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Paralogs for GLUL gene

3 SIMAP similar genes for GLUL using alignment to 2 protein entries: GLNA_HUMAN (see all proteins):

PIG59 DKFZp434M0813 PIG43

GLUL for paralogs About GeneDecksing

Selected Pseudogenes.org Pseudogenes for GLUL (see all 6)
PGOHUM00000242168 PGOHUM00000242513 PGOHUM00000239935 PGOHUM00000240465 PGOHUM00000235814

(SNPs/Variants according to the ¹NCBI SNP Database, ²Ensembl, ³PupaSUITE, ⁴UniProtKB, and DNA2.0, Linkage Disequilibrium by HapMap, Structural Variations(CNVs/InDels/Inversions) from the Database of Genomic Variants, Mutations from the Human Gene Mutation Database (HGMD), the Human Cytochrome P450 Allele Nomenclature Database, and the Locus Specific Mutation Databases (LSDB), Blood group antigen gene mutations by BGMUT, Resequencing Primers, Cancer Mutation PCR Arrays and Assays, and Copy Number PCR Arrays from QIAGEN)
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SNP ID	Valid	Clinical significance	Chr 1 pos	Sequence	#	AA Chg	Type	More	#	Allele freq	Pop	Total sample	More
Genomic Data					Transcription Related Data				Allele Frequencies
				--	--	--		--	--
VAR_026560⁴		Congenital systemic glutamine deficiency (CSGD)⁴	--	see VAR_026560	2	R C	mis⁴		0	--	--	--	--
VAR_026561⁴		Congenital systemic glutamine deficiency (CSGD)⁴	--	see VAR_026561	2	R C	mis⁴		0	--	--	--	--
rs80358215^1,2	C	pathogenic¹	188905975(-)	`ATCGTC/TGCCCC`	6	R C	mis¹		0	--	--	--	--
rs80358214^1,2	C	pathogenic¹	188906026(-)	`GCATAA/C/TGCATT`	9	S R C	mis¹		0	--	--	--	--
rs78353586^1,2	C	--	153590513(+)	`GACATA/GTGTCA`	3	--	int¹		0	--	--	--	--
rs3028837^1,2	C	--	182351630(-)	`GTGAT-/TCA/ TCATTTAAGAA`	6	--	cds¹		1		CSA	2
rs56354856^1,2	C	--	182351631(+)	`TCTTA-/AAATG A/AATGAATCACT`	6	--	cds¹		0	--	--	--	--
rs71785633^1,2	C	--	182354175(+)	`ATGACAT/-ATGTC`	3	--	int¹		1		CSA	2
rs56029235^1,2	C	--	182358541(+)	`ATATA-/TTTT TTTAG`	3	--	int¹		1		NA	2
rs112035237^1,2	C	--	182358689(+)	`AGACTTG/-TCAAG`	3	--	int¹		1		CSA	2

HapMap Linkage Disequilibrium report for GLUL (182350839 - 182361341 bp)

Structural Variations
Database of Genomic Variants (DGV) 1 variation for GLUL: About this table

Variant ID	Type	Subtype	PubMed ID
esv22717	CNV	Loss	19812545

Human Gene Mutation Database (HGMD): GLUL

Site Specific Mutation Identification with PCR Assays

SeqTarget long-range PCR primers for resequencing GLUL

DNA2.0 Custom Variant and Variant Library Synthesis for GLUL

(in which this Gene is Involved, According to MalaCards, OMIM, UniProtKB, the University of Copenhagen DISEASES database, Conferences by KenesGroup, Novoseek, Genatlas, GeneTests, GAD, HuGE Navigator, and/or TGDB.)
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OMIM gene information: 138290
OMIM disorders: 610015
UniProtKB/Swiss-Prot: GLNA_HUMAN, P15104

Congenital systemic glutamine deficiency (CSGD) [MIM:610015]: Rare developmental disorder with severe

brain malformation resulting in multi-organ failure and neonatal death. Glutamine is largely absent from affected

patients serum, urine and cerebrospinal fluid. Note=The disease is caused by mutations affecting the gene

represented in this entry

Selected diseases for GLUL (see all 63):
About MalaCards

glutamine deficiency, congenital glutathione synthetase deficiency hyperinsulinism-hyperammonemia syndrome retinal detachment

reye syndrome paranoid schizophrenia hepatic encephalopathy acute porphyria

brain edema hyperinsulinism porphyria metabolic acidosis

acute liver failure familial adenomatous polyposis proliferative vitreoretinopathy oligodendroglioma

toxic encephalopathy febrile seizures brain disease temporal lobe epilepsy

3 diseases from the University of Copenhagen DISEASES database for GLUL:

Hepatic encephalopathy Toxic encephalopathy Hepatocellular carcinoma

GLUL for disorders About GeneDecksing

Congresses - knowledge worth sharing:
The International Symposium on Pneumococci and Pneumococcal Diseases (ISPPD) 9 - 13 March 2014
Alzheimer's & Parkinson's Diseases Congress (ADPD) 18 - 22 March 2015

Selected Novoseek inferred disease relationships for GLUL gene (see all 56) About this table

Disease	-log (P-Val)	Hits	PubMed IDs for Articles with Shared Sentences (# sentences)
hyperammonemia	67.3	8	12020613 (2), 15841446 (1), 1352200 (1), 1677585 (1)
hepatic encephalopathy	43.2	2	10564534 (1), 12020613 (1)
hepatocellular carcinoma	39.5	13	1346587 (3), 18282277 (2), 10952242 (2), 20525558 (1) (see all 6)
brain diseases	32.4	3	18062818 (1), 9166739 (1), 11004532 (1)
gliosis	30.3	2	8570019 (1), 19183851 (1)
liver failure	29.5	5	10564534 (1), 17299991 (1), 11549067 (1), 16382338 (1)
alzheimers disease	27.9	26	10733015 (5), 10520940 (2), 9166739 (2), 11746426 (2) (see all 12)
neurodegeneration	27.8	3	20107048 (2), 14681883 (1)
epilepsy temporal lobe	26.3	6	16095760 (1), 14723991 (1), 19170755 (1), 18226171 (1) (see all 5)
reye syndrome	25.9	1	10199339 (1)

Genetic Association Database (GAD): GLUL
Human Genome Epidemiology (HuGE) Navigator: GLUL (3 documents)

Export disorders for GLUL gene to outside databases

(in PubMed. Associations of this gene to articles via ¹Entrez Gene, ²UniProtKB/Swiss-Prot, ³HGNC, ⁴GAD, ⁵PharmGKB, ⁶HMDB, ⁷DrugBank, ⁸UniProtKB/TrEMBL, ⁹ Novoseek, and/or ¹⁰fRNAdb)
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PubMed articles for GLUL gene, integrated from 10 sources (see all 557):
(articles sorted by number of sources associating them with GLUL)

Utopia: connect your pdf to the dynamic
world of online information

cDNA sequence of the long mRNA for human glutamine synthase. (PubMed id 1681907)^{1, 2, 3} van den Hoff M.J.B....Lamers W.H. (Biochim. Biophys. Acta 1991)
Sequence of a human glutamine synthetase cDNA. (PubMed id 2888076)^{1, 2, 3} Gibbs C.S.... Wilson R.H. (Nucleic Acids Res. 1987)
Wnt and steroid pathways control glutamate signalling by regulating glutamine synthetase activity in osteoblastic cells. (PubMed id 18555765)^{1, 2, 9} Olkku A. and Mahonen A. (Bone 2008)
Glutamine synthetase is essential for proliferation of fetal skin fibroblasts. (PubMed id 18662667)^{1, 2, 9} Vermeulen T.... Haeberle J. (Arch. Biochem. Biophys. 2008)
Overexpression of glutamine synthetase in human primary liver cancer. (PubMed id 7909780)^{1, 2, 9} Christa L....Lasserre C. (Gastroenterology 1994)
An association study between polymorphisms in five genes in glutamate and GABA pathway and paranoid schizophrenia. (PubMed id 15642443)^{1, 4, 9} Zhang B....Shen Y. (Eur. Psychiatry 2005)
Pharmacogenetics of antipsychotic response in the CATIE trial: a candidate gene analysis. (PubMed id 19156168)^{1, 4} Need A.C....Goldstein D.B. (Eur. J. Hum. Genet. 2009)
Association analysis of the glutamic acid decarboxylase 2 and the glutamine synthetase genes (GAD2, GLUL) with schizophrenia. (PubMed id 19125103)^{1, 4} Arai S....Fukumaki Y. (Psychiatr. Genet. 2009)
The DNA sequence and biological annotation of human chromosome 1. (PubMed id 16710414)^{1, 2} Gregory S.G.... Bentley D.R. (Nature 2006)
Congenital glutamine deficiency with glutamine synthetase mutations. (PubMed id 16267323)^{1, 2} Haeberle J.... Koch H.G. (N. Engl. J. Med. 2005)

(in PubMed, OMIM, and NCBI Bookshelf)
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		AND	OR
Aliases
Disorders
Free Text

Query String		PubMed OMIM NCBI Bookshelf
	(Note: In FireFox, select the above section and copy using Ctrl-C)

(According to Entrez Gene, HGNC, AceView, euGenes, Ensembl, miRBase, ECgene, Kegg, and/or H-InvDB)
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Entrez Gene: 2752	HGNC: 4341	AceView: GLUL	Ensembl:ENSG00000135821	euGenes: HUgn2752
ECgene: GLUL	Kegg: 2752	H-InvDB: GLUL

(According to HUGE)
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(According to PharmGKB, ATLAS, HORDE, IMGT, LEIDEN, UniProtKB/Swiss-Prot, UniProtKB/TrEMBL, and/or others, e.g. Wikipedia and GeneReviews, via UniProtKB/Swiss-Prot)
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Name	Description
PharmGKB entry for GLUL	Pharmacogenomics, SNPs, Pathways
Wikipedia	http://en.wikipedia.org/wiki/Glutamine_synthetase

Intellectual Property for GLUL gene
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Patent Information for GLUL gene:
Search GeneIP for patents involving GLUL

GeneCards and IP:
Japan Patent Office Licenses GeneCards European Patent Office Licenses GeneCards Improving the IP Search

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GLUL Gene

Glutamate-Ammonia Ligase

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familial adenomatous polyposis
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Porphyria Acute Intermittent
sepsis
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Occlusion Middle Cerebral Artery
Hepatic Encephalopathy
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GLUL Gene

Glutamate-Ammonia Ligase

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