Heterocyst Metabolism and Development

Abstract

Heterocysts are differentiated cells that are specialized for fixation of N2 in an aerobic environment. In heterocysts in the light, Photosystem I generates ATP, but no photosynthetic production of O2 takes place. Instead, reductant moves into heterocysts from vegetative cells. In return, fixed nitrogen moves from heterocysts to vegetative cells. In neither case is there certainty about the identity of the traffic molecules. Pathways of electron-donation to N2 have been extensively investigated, but their in-vivo importance remains to be critically tested. Nitrogenase in heterocysts is protected from inactivation by O2 by a variety of means, principally by enhanced respiration and by a barrier, the heterocyst envelope, to entry of O2. However, the respiratory apparatus and the biosynthetic processes that result in synthesis of the barrier have been little studied. The detailed mechanisms underlying metabolic, environmental, and developmental control of nitrogenase are under investigation. Studies of heterocyst development are being greatly facilitated by recent advances in the genetics of Anabaena sp. An autoregulated gene, hetR, that is activated shortly after nitrogen-stepdown is critical for the differentiation of heterocysts. Two enigmas remain to be answered: how is it determined which cells will differentiate; and, after differentiation is initiated, what intercellular interactions and intracellular mechanisms regulate the progression of the differentiation process? An evolutionary and biochemical relationship between the processes leading to the formation of heterocysts and akinetes is suggested.