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In addition to these conventional sites of mitochondrial ROS generation, it has been recently reported that the branched-chain 2-oxoacid dehydrogenase (BCKDH) complex in mitochondria can produce superoxide and H 2 O 2 at higher rates than complex I from mitochondria (Quinlan et al., 2014)

When ROS levels rise, PUFAs in the sperm membrane are oxidized in a process known as lipid peroxidation, which generates toxic byproducts such as MDA and 4-HNE