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The mutant also had substantially depleted intracellular levels of key energy pathway intermediates, including adenosine and adenosine monophosphate (AMP), L-carnitine and propionylcarnitine, pantothenate, -ketoglutaric acid, and -nicotinamide adenine dinucleotide (NAD) and nicotinamide (which drive generation of adenosine triphosphate (ATP) in the mitochondria), as well as the urea cycle intermediates ornithine and argininosuccinic acid (Extended Data Fig