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Under certain conditions, protons can re-enter the mitochondrial matrix without contributing to ATP synthesis.
This process is known as proton leak or mitochondrial uncoupling and is due to the facilitated diffusion of protons into the matrix.
The process results in the unharnessed potential energy of the proton electrochemical gradient being released as heat.
The process is mediated by a proton channel called thermogenin, or UCP1.
Thermogenin is a 33kDa protein first discovered in 1973.
Thermogenin is primarily found in brown adipose tissue, or brown fat, and is responsible for non-shivering thermogenesis.
Brown adipose tissue is found in mammals, and is at its highest levels in early life and in hibernating animals.
In humans, brown adipose tissue is present at birth and decreases with age.

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