Vascular Connection Between Lateral Roots and Stem in the Ophioglossaceae

Abstract

The vascular connection between lateral roots and stem in the Ophioglossaceae and in two leptosporangiate fern species was examined. Two types of connections were found: “gradual” connections, which resemble leaf traces in ontogeny and morphology, and “abrupt” connections, which resemble the connections between lateral roots and their parent roots. Gradual root-stem connections occur in the genera Ophioglossum and Helminthostachys and in Woodwardia virginica. They are initiated in shoot apices distal to the level where cauline xylem elements mature. They resemble leaf traces in being provascular (procambial) strands that connect the cauline stele with the future vasculature of lateral appendages. As with leaf traces, gradual connections are part of the provascular and, later, protoxylem continuity between stems and lateral appendages. Gradual connections have many features in common with leaf traces, and the term root trace is applicable to them. The order of radial maturation of the primary xylem in gradual connections varies in different parts of the connections. It is endarch near the intersection with the cauline stele and exarch where the connections intersect root steles. Gradual connections resemble the transition regions of certain seed plants where protoxylem is also continuous from stem to root and the order of maturation is found to change continuously from stem to root. Abrupt connections occur in Botrychium and Osmunda cinnamomea. They develop in shoot apices at levels where cauline xylem is mature or maturing. The mature xylem does not dedifferentiate, so provascular and protoxylem continuity of the kind found in root traces does not occur. Also, reorientation of the order of maturation does not occur in abrupt connections. Xylem connectors are found in the region where radially oriented elements of the connections abut the longitudinally oriented cauline elements. Abrupt connections resemble the connection of secondary roots with their parent root systems since xylem connectors and the lack of continuity are also features found in these vascular systems. The resemblance of the vascular pattern of the fern root trace to the transition region of seed plants suggests that the radicle is more closely comparable to the cladogenous roots of pteridophytes than hitherto supposed.