Rescuing Robert Brown—The Origins of Angio-Ovuly in Seed Cones of Conifers

Abstract

Robert Brown (1827) recognized the distinction between the two major classes of seed plants on the basis, respectively, of exposed ovules, which receive pollen directly, and enclosed ovules, which do not, at the time of pollination. The two groups, unfortunately, became known as “Gymnosperms” and “Angiosperms”, a distinction not made by Brown. The names are, at best, useful only as identifying labels. As is established here, gymno-ovuly and angio-ovuly would have been better words and should serve as the most consistent distinguishing character among groups of seed plants. Although conifers can hardly be claimed to be members of any lineage that led to the Angiosperms, the gymno-ovulate method of reproduction must have occurred in the direct ancestors of flowering plants. Interpolating development and functional evidence serves to resurrect Robert Brown’s important but neglected observation and correct misstatements implied in generally accepted systematic terminology. The result of this approach is to discover that the conifers in their reproductive diversity show some trends indicative of those characters that define the flowering plants. Although non-homologous these character manifestations provide useful models which can lead to a better understanding of the origin of “Angiosperms” from “Gymnosperms.” In the complex and extended process of reproductive development in both major groups regular transpositions occur, depending on whether “gymno-”and “angio-”are applied to ovules or seeds. Gymno-ovuly and angio-spermy are the most common conditions in conifers but result in intrinsically opposed processes of pollen reception and subsequent seed protection, which are resolved by successive changes in seed-cone structure. The modern conifers illustrate considerable diversity of ovulate and seed cone structure in relation to several biological functions, including traits that anticipate the angio-ovulate condition, in cone inception, ovule enclosure, pollen and embryo development and seed dispersal. From the paleobotanical record we do have a good grasp of the structural changes in the progressive modification of the coniferous seed cone in conifers, starting with the antecedent Carboniferous Cordaitales. These changes have involved the progressive condensation of a lax, branched strobilus and end with the highly condensed uniovulate “cone” as in many Podocarpus species in which the equivalent of a bitegmic ovule exists. The progressive changes can be perceived typologically among existing conifers but need to be understood as the result of biological processes that suggest increased reproductive efficiency, and can be added to our present quite robust understanding of conifer phylogeny.