The paraveinal mesophyll of soybean leaves in relation to assimilate transfer and compartmentation

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Abstract
The paraveinal mesophyll (PVM) is a unique and specialized, one-cell-thick tissue spanning the vascular bundles at the level of the phloem in soybean (Glycine max) (L.) Merr.) leaves. Its position within the leaf dictates that all photosynthate produced in the palisade and spongy mesophyll must pass through this specialized layer enroute to the phloem. Symplastic continuity, via plasmodesmata, exists between the PVM and bundle sheath, palisade parenchyma and spongy mesophyll. During leaf ontogeny the PVM is the first tissue to differentiate and at maturity these cells are six to eight times larger than other mesophyll cells, are highly vacuolate, and are interconnected by tubular arms. The PVM undergoes several unique structural and metabolic modifications during leaf development. The PVM cytoplasm, in vegetative plants, is dense, enriched in rough endoplasmic reticulum and dictyosomes, but contains few, small starch-free chloroplasts and few microbodies. Unlike the tonoplast of mesophyll cells, the tonoplast of the PVM is unusually thick and dense-staining. During leaf development the vacuoles of PVM cells accumulate a glycoprotein derived from the dictyosomes which reacts with the protein staining reagents, mercuric bromophenol blue and sulfaflavine, and is degraded by Pronase. Both the vacuolar material and tonoplast are also stained by phosphotungstic acid, which at low pH is relatively selective for glycoprotein. A unique role of the PVM in the transport and compartmentation of nitrogen reserves in soybeans is discussed.