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Stem Development, Medullary Bundles, and Wood Anatomy of Croton Glandulosus Var. Septentrionalis (Euphorbiaceae)

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Anatomy and development of vascular tissues in the annual stems of Croton glandulosus var. septentrionalis are described. In primary stages of growth the stem possesses a eustele of bicollateral bundles; internal phloem is notably more extensive than the external. In addition to a vascular cambium and secondary xylem that form in the usual fashion, additional cambia add cells to the internal phloem portion of the bicollateral bundles, forming well-marked medullary bundles at the perimeter of the pith. At first, the perimedullary cambial strands produce only internal secondary phloem; later, internal secondary xylem is also formed in some stems. When internal secondary xylem is present, the medullary bundles have an inverted orientation, i.e., phloem innermost (towards centre of pith) and xylem outermost (near protoxylem). Cells of the medullary bundles include sieve tube elements, vessel ekments, and fibres. Normal (external) secondary phloem is weakly developed. Normal secondary xylem contains short vessel elements with simple perforation plates and alternate intervascular pits, libriform fibres, narrow heterocellular rays, and lacks axial parenchyma.


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