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The TGF-beta/BMP (bone morphogenetic protein) pathway incorporates several signalling pathways that share most, but not all, components of a central signal transduction engine. The general signalling scheme is rather simple: upon binding of a ligand, an activated plasma membrane receptor complex is formed, which passes on the signal towards the nucleus through a phosphorylated receptor-activated SMAD (r-SMAD). In the nucleus, the activated r-SMAD promotes transcription in a complex with a closely-related helper molecule termed the Co-SMAD. However, this simple linear pathway expands into a network when various regulatory components and mechanisms are taken into account. The signalling pathway includes a great variety of different TGF-beta/BMP superfamily ligands and receptors, several types of the r-SMAD, and functionally critical negative feedback loops. The r-SMAD/Co-SMAD can interact with a great number of transcriptional co-activators/co-repressors to regulate positively or negatively effector genes, so that the interpretation of a signal depends on the cell-type and cross talk with other signalling pathways such as Notch, MAPK and Wnt. The pathway plays a number of different biological roles in the control of embryonic and adult cell proliferation and differentiation, and it is implicated in a great number of human diseases.
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