Abstract
Photosystem II is the chlorophyll-containing enzyme in which the very first chemical energy storing reaction of photosynthesis occurs, when water is split into molecular oxygen and hydrogen equivalents at a catalytic center composed of four Mn ions and one Ca2+. All the oxygen in the atmosphere is derived from this reaction, and without it the biosphere, as we know it, would not exist. Indeed, the appearance of this catalytic center ∼3 billion years ago gave rise to the "big bang of evolution". Thus, understanding the structure and functioning of this metal cluster is a major topic in science, and here I discuss it in terms of research that has occurred over the past 12 years dating back to when it was first proposed to be a Mn3CaO4 cubane with the fourth Mn attached to cubane by one of its oxo bridging bonds. Since then, a number of novel properties have emerged for this metalloprotein with implications for its catalytic mechanism and evolutionary origin.
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