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Dysregulated p16 binds cyclin dependent kinase 4 (cdk4), which results in cytoplasmic sequestration of cdk4. Binding between nuclear cyclin D1 and cdk2 results in an inert complex but serves to compete with the binding of cdk2 to cyclins A and E. This may explain why there is increased expression of nuclear cyclin D1 in tumour buds whereas proliferation is decreased.
