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Three-Dimensional Organization of Higher-Plant Chloroplast Thylakoid Membranes Revealed by Electron Tomography

^a Electron Microscopy Unit, Weizmann Institute of Science, Rehovot 76100, Israel
^b Department of Biological Chemistry, Weizmann Institute of Science, Rehovot 76100, Israel
^c Department of Biological Chemistry, Hebrew University of Jerusalem, Jerusalem 91904, Israel
^d Department of Plant Sciences, Weizmann Institute of Science, Rehovot 76100, Israel

¹ To whom correspondence should be addressed. E-mail ziv.reich{at}weizmann.ac.il; fax 972-8-934-6010.

The light-harvesting and energy-transducing functions of the chloroplast are performed within an intricate lamellar system of membranes, called thylakoid membranes, which are differentiated into granum and stroma lamellar domains. Using dual-axis electron microscope tomography, we determined the three-dimensional organization of the chloroplast thylakoid membranes within cryo-immobilized, freeze-substituted lettuce (Lactuca sativa) leaves. We found that the grana are built of repeating units that consist of paired layers formed by bifurcations of stroma lamellar sheets, which fuse within the granum body. These units are rotated relative to each other around the axis of the granum cylinder. One of the layers that makes up the pair bends upwards at its edge and fuses with the layer above it, whereas the other layer bends in the opposite direction and merges with the layer below. As a result, each unit in the granum is directly connected to its neighbors as well as to the surrounding stroma lamellae. This highly connected morphology has important consequences for the formation and function of the thylakoid membranes as well as for their stacking/unstacking response to variations in light conditions.

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