Apoptosis is the process of cell signalling auto-destruction often called ‘programmed cell death’. Apoptosis is a highly regulated cell signalling process characterized by morphological and biochemical cellular changes.
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The extrinsic or death receptor apoptosis cell signalling pathway is initiated through ligand binding to death receptors on the cell membrane.
The death receptors TNF-R1 (CD120a), TNF-R2 (CD120b), Fas/APO-1 (CD95) and the TNF-related apoptosis-inducing ligand (TRAIL) receptors called DR3, DR4, DR5 and DR6 are subgrouped under the tumor necrosis factor (TNF) receptor superfamily.
This family also includes TWEAK Receptor (CD266 / Fn14) which interacts with its ligand TWEAK (TNF-like weak inducer of apoptosis) (CD255) inducing apoptosis in some tumour cell lines through pathways yet to be fully elucidated.
Structurally, the death receptors are categorised as TNFR associated factors (TRAFs) or death domain homologues which have a cysteine rich extracellular domain and a transmembrane or intracellular death domain.
The apoptosis cell signal is initiated through death receptor oligomerization and via the death domain, is transmitted intracellularly through the recruitment of cytoplasmic adaptor proteins such as Fas associated death domain protein (FADD) and tumour necrosis factor receptor 1-associated death domain protein (TRADD).
The adaptor proteins have a death effector domain which recruits cytosolic proteins e.g. pro-caspase 8 to form the death-inducing signaling complex (DISC) with subsequent activation of the caspase cascade.
One of the earliest features of apoptosis is a morphological change in the plasma membrane. This involves the translocation of the membrane phospholipid phosphatidylserine (PS) during apoptosis from the internal layer to the external layer of the cell membrane.
In the presence of calcium ions, Annexin V has a high specificity and affinity for PS. Thus, the binding of Annexin V to cells with exposed PS provides a very sensitive method for detecting cellular apoptosis.
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Apoptosis of virally-infected cells and primary tumor cells may be induced by cytotoxic T lymphocytes (CTLs) through the release of perforin and granzymes from cytoplasmic cytolytic granules. Further details about this alternative caspase activation pathway are available.
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Dysregulation of apoptosis may lead to pathological conditions. Apoptosis is associated with cancer, autoimmune diseases, transplant rejection, AIDS and neurodegenerative diseases. A greater understanding of the proteins and mechanisms involved in apoptosis will help to develop novel therapeutic approaches to modulate cellular death pathways in the treatment of diseases.
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