Glutamine synthetase

Glutamine synthetase (GS) (EC 6.3.1.2)^[1] is an enzyme that plays an essential role in the metabolism of nitrogen by catalyzing the condensation of glutamate and ammonia to form glutamine:

Glutamate + ATP + NH₃ → Glutamine + ADP + phosphate

[edit] GS classes

There seem to be three different classes of GS: ^[2] ^[3] ^[4]

Class I enzymes (GSI) are specific to prokaryotes, and are oligomers of 12 identical subunits. ^[5] The activity of GSI-type enzyme is controlled by the adenylation of a tyrosine residue. The adenylated enzyme is inactive ^[6].

Class II enzymes (GSII) are found in eukaryotes and in bacteria belonging to the Rhizobiaceae, Frankiaceae, and Streptomycetaceae families (these bacteria have also a class-I GS). GSII are decamer of identical subunits ^[7]. Plants have two or more isozymes of GSII, one of the isozymes is translocated into the chloroplast.

Class III enzymes (GSIII) has, currently, only been found in Bacteroides fragilis and in Butyrivibrio fibrisolvens. It is a hexamer of identical chains. It is much larger (about 700 amino acids) than the GSI (450 to 470 amino acids) or GSII (350 to 420 amino acids) enzymes.

While the three classes of GS's are clearly structurally related, the sequence similarities are not so extensive.

^ Some evolutionary relationships of the primary biological catalysts glutamine synthetase and RuBisCO. 1987; PubMed: 2900091
^ Evolution of the glutamine synthetase gene, one of the oldest existing and functioning genes. 1993 Apr 1; PubMed: 8096645 Free text
^ Glutamine synthetase II in Rhizobium: reexamination of the proposed horizontal transfer of DNA from eukaryotes to prokaryotes. 1989 Nov; PubMed: 2575672
^ Evolutionary relationships of bacterial and archaeal glutamine synthetase genes. 1994 Jun; PubMed: 7916055
^ GSI picture
^ InterPro:IPR001637 Glutamine synthetase class-I, adenylation site
^ Crystal structure of human glutamine synthetase in complex with ADP and phosphate http://www.rcsb.org/pdb/explore.do?structureId=2OJW

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v • d • e Enzymes: ligases (EC 6)

6.1 - Carbon-Oxygen	Aminoacyl tRNA synthetase

6.2 - Carbon-Sulfur	Succinyl coenzyme A synthetase - Acetyl Co-A synthetase - Long fatty acyl CoA synthetase

6.3 - Carbon-Nitrogen	Glutamine synthetase - Ubiquitin ligase (Von Hippel-Lindau tumor suppressor, UBE3A, Mdm2, Anaphase-promoting complex) - Glutathione synthetase - CTP synthase - Adenylosuccinate synthase - Argininosuccinate synthetase - Holocarboxylase synthetase - GMP synthase - Asparagine synthetase - Carbamoyl phosphate synthetase (I, II) - Gamma-glutamylcysteine synthetase

6.4 - Carbon-Carbon	Carbon-carbon ligases

6.5 - Phosphoric Ester	DNA ligase

v • d • e

glycine→serine→	Serine hydroxymethyltransferase - Serine dehydratase

alanine→	Alanine transaminase

cysteine→	D-cysteine desulfhydrase

threonine→	L-threonine dehydrogenase


HISTIDINE→	Histidine ammonia-lyase - Urocanate hydratase - Formiminotransferase cyclodeaminase

proline→	Proline oxidase - Pyrroline-5-carboxylate reductase - 1-Pyrroline-5-carboxylate dehydrogenase/ALDH4A1 - PYCR1

arginine→	Ornithine aminotransferase - Ornithine decarboxylase - Agmatinase

→alpha-ketoglutarate→TCA	Glutamate dehydrogenase


PHENYLALANINE→tyrosine→	Phenylalanine hydroxylase - Tyrosine aminotransferase - 4-Hydroxyphenylpyruvate dioxygenase - Homogentisate 1,2-dioxygenase - Fumarylacetoacetate hydrolase


asparagine→aspartate→	Asparaginase/Asparagine synthetase - Aspartate transaminase

Other

tyrosine→melanin	Tyrosinase

cysteine+glutamate→glutathione	Gamma-glutamylcysteine synthetase - Glutathione synthetase - Gamma-glutamyl transpeptidase

glycine→creatine	Guanidinoacetate N-methyltransferase - Creatine kinase

glutamate→glutamine	Glutamine synthetase - Glutaminase

proline+lysine→collagen	Prolyl hydroxylase - Lysyl hydroxylase