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both in apo form and in the complex forms with substrate and product. Protein consists of four domains. of which three domains have essentially the same fold. The catalytic cleft is formed by four loops from each domains. The N-acetyl group of the substrate is recognized by V370 and N389 in domain 3, from which the substrate specificity arises. Substrate rotates about 180Â° on the axis linking C4 and the midpoint of the C5-O5 bond in the reaction