Physiological correlates of the morphology of early vascular plants



RAVEN, J. A., 1984. Physiological correlates of the morphology of early vascular plants. The early evolution of vascular land plants is considered in relation to the physiological problems of life on land. The universal characteristics of vascular plants (xylem, cuticle, stomata, intercellular air spaces, long-distance symplastic transport and alternation of generations) are discussed in terms of the essential properties of a homoiohydric phototroph. Likely precursors of vascular plants, and the physico-chemical and biotic environment in which they occurred, are outlined prior to a discussion of the selective forces acting on the evolution of vascular plants in the Upper Silurian and Lower Devonian. Emphasis is placed on biochemical and structural ‘pre-adaptations’ which may have occurred in the precursors of vascular plants and on which natural selection could have acted with lignified xylem, stomata, etc., as the end-products. Guiding principles in the analysis include the physiology of extant plants, physico-chemical constraints, and compatibility with the fossil record. It is concluded that the likely sequence of acquisition of vascular plant characteristics was: heteromorphic alternation of generations with an erect sporophyte; cuticularization of sporophyte; evolution of xylem; occurrence of intercellular air spaces with pores in the epidermis; stomatal activity of the pores. Endodermis and phloem-type long-distance transport probably originated around stages (3)-(5).