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JAK3 is a protein coding gene. The protein encoded by this gene is a member of the Janus kinase (JAK) family of tyrosine kinases involved in cytokine receptor-mediated intracellular signal transduction. Other members of the Janus family include JAK1, JAK2 and TYK2. JAKs are conserved in domain structure, with an N-terminal FERM domain and a Src homology 2 (SH2)-like domain, followed by a pseudokinase domain (JAK homology 2, JH2) and a tyrosine kinase domain (JH1). They are relatively large kinases of approximately 1150 amino acids with apparent molecular weights of 120-130 kDa. It is predominantly expressed in immune cells and transduces a signal in response to its activation via tyrosine phosphorylation by interleukin receptors. Non-receptor tyrosine kinase is also involved in various processes, such as cell growth, development, or differentiation.
JAK3 mediates essential signaling events in both innate and adaptive immunity, and plays a crucial role in hematopoiesis during T-cells development. In the cytoplasm, it plays a pivotal role in signal transduction via its sole association with type I receptors sharing the common subunit gamma (γc), such as IL2R, IL4R, IL7R, IL9R, IL15R and IL21R, which regulate development and activation of lymphoid lineage cells. In IL-2 receptor (IL-2R) signaling, as well as cytokine signaling, when ligand binds to extracellular domain of cell surface receptors β or γ chains, JAK1 and JAK3 phosphorylates specific tyrosine residues of receptor subunits on the cytoplasmic tails, creating docking sites for signal transducers and activators of transcription (STATs) proteins. Then, for example, STAT5A and STAT5B are recruited. Subsequently, JAK3 phosphorylates the STATs proteins. Phosphorylated STATs then form homodimer or heterodimers, and translocate to the nucleus to activate transcription of specific target genes in a cytokine-specific fashion.
|Type||Subgroup||Cytokine Receptor||JAK Kinase|
|type I receptors||homodimeric||EPO, TPO, GH, G-CSF||JAK2|
|uses common beta chain (CSF2RΒ)||IL-3, IL-5, GM-CSF||JAK2|
|uses gp130 chain||IL-6, IL-11||JAK1, JAK2, Tyk2|
|uses common gamma chain (γc)||IL-2, IL-4, IL-7, IL-9, IL-15, IL-21||JAK1, JAK3|
|type II receptors||IFN-α, IFN-β, IFN-γ||JAK1, JAK2, Tyk2|
Table 1 Some cytokine receptors and their involvement with JAK kinases （O'Shea JJ et. al, 2005)
Mutations in JAK3 gene are associated with autosomal SCID (severe combined immunodeficiency disease). JAK3 mutations are also found in patients of T-cell acute lymphoblastic leukemia (T-ALL), acute megakaryoblastic leukemia, T-cell prolymphocytic leukemia, and juvenile myelomonocytic leukemia and natural killer T-cell lymphoma (NK/T-lymphoma). JAK plays a central role in the control of lymphopoiesis. The pseudokinase domain (JAK homology 2, JH2) of JAK3 shows significant sequence homology with classical protein kinases but several conserved functional residues or motifs are missing or altered. JAK JH2s is of particular interest as approximately half of clinical JAK3 mutations cluster into it. Recent studies have focused on tyrosine kinases as potential targets for the treatment of Rheumatoid arthritis (RA), which is a chronic, systemic disease characterized by persistent inflammatory synovitis. Among them, inhibitors specific for the JAK family are, so far, the most efficacious in treating RA. Selective JAK3 inhibition should be sufficient for immunosuppression without causing effects outside the immune system.
JAK3 Inhibition Mechanism (Warren J Leonard et al., 2016)
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