The occurrence of an extended bundle sheath system (paraveinal mesophyll) in the legumes

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Abstract
The tissue previously described as paraveinal mesophyll in soybean leaves is shown to have the characters of bundle sheath rather than mesophyll cells and is renamed "extended bundle sheath" (EBS) tissue. Its presence was surveyed by leaf clearing in 66 species of legumes of all three subfamilies. A complete extended bundle sheath system similar to that previously described in soybean was identified in 21 of the species. This system is a paradermally oriented tissue, one cell deep, between the spongy and palisade mesophylls, consisting of extended bundle sheath cells, which join each other across the interveinal space either directly or via bridging cells of somewhat similar shape and size. A newly recognized, attenuated extended bundle sheath system, in which bundle sheath cells extend but do not form a continuum except in very narrow interveinal spaces, is described; it was found in 32 species. Extended bundle sheath tissue was absent from 13 of the species. The presence or form of extended bundle sheath tissue does not follow traditional taxonomic divisions. Extended bundle sheath systems were also found in 3 of 5 nonlegume species.