is filled with tissue as in typical Gymnosperms, and from some of the
superficial cells 3 to 5 archegonia are developed, characterized by
long multicellular necks. The archegonia are separated from one
another, as in Pinus, by some of the prothallus-tissue, and the cells
next the egg-cells (tapetal layer) contribute food-material to their
development. After fertilization, some of the uppermost bracts
below each flower become red and fleshy; the perianth develops into
a woody shell, while the integument remains membranous. In some
species of Ephedra, e.g. E. altissima, the fertilized eggs grow into
tubular proembryos, from the tip of each of which embryos begin to
be developed, but one only comes to maturity. In Ephedra helvetica,
as described by Jaccard, no proembryo or suspensor is formed; but
the most vigorous fertilized egg, after undergoing several divisions,
becomes attached to a tissue, termed the columella, which serves the
purpose of a primary suspensor; the columella appears to be formed
by the lignification of certain cells in the central region of the embryo-sac.
At a later stage some of the cells in the upper (micropylar)
end of the embryo divide and undergo considerable elongation,
serving the purpose of a secondary suspensor. The secondary wood
of Ephedra consists of tracheids, vessels and parenchyma; the
vessels are characterized by their wide lumen and by the large simple
or slightly-bordered pits on their oblique end-walls.
Fig. 17.—Gnetum Gnemon. (After Lotsy.)
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Gnetum.—This genus is represented by several species, most of which are climbing plants, both in tropical America and in warm regions of the Old World. The leaves, which are borne in pairs at the tumid nodes, are oval in form and have a Dicotyledonous type of venation. The male and female inflorescences have the form of simple or paniculate spikes. The spike of an inflorescence bears whorls of flowers at each node in the axils of concrescent bracts accompanied by numerous sterile hairs (paraphyses); in a male inflorescence numerous flowers occur at each node, while in a female inflorescence the number of flowers at each node is much smaller. A male flower consists of a single angular perianth, through the open apex of which the flower-axis projects as a slender column terminating in two anthers. The female flowers, which are more complex in structure, are of two types, complete and incomplete; the latter occur in association with male flowers in a male inflorescence. A complete female flower consists of a nucellus (fig. 17, A, n), surrounded by a single integument (fig. 17, A, i), prolonged upwards as a narrow tube and succeeded by an inner and an outer perianth (fig. 17, A, p′ and p′). The whole flower may be looked upon as an adventitious bud bearing two pairs of leaves; each pair becomes concrescent and forms a perianth, the apex of the shoot being converted into an orthotropous ovule. The incomplete female flowers are characterized by the almost complete suppression of the inner perianth. Several embryo-sacs (megaspores) are present in the nucellus of a young ovule, but one only attains full size, the smaller and partially developed megaspores (fig. 17, B and C, e) being usually found in close association with the surviving and fully-grown megaspore. In Gnetum Gnemon, as described by Lotsy, a mature embryo-sac contains in the upper part a large central vacuole and a peripheral layer of protoplasm, including several nuclei, which take the place of the archegonia of Ephedra; the lower part of the embryo-sac, separated from the upper by a constriction, is full of parenchyma. The upper part of the megaspore may be spoken of as the fertile half (fig. 17, B and C, F) and the lower part, which serves only as food-reservoir for the growing embryo, may be termed the sterile half (fig. 17, B and C, S). (Coulter, Bot. Gazette, xlvi., 1908, regards this tissue as belonging to the nucellus.) At the time of pollination the long tubular integument secretes a drop of fluid at its apex, which holds the pollen-grains, brought by the wind, or possibly to some extent by insect agency, and by evaporation these are drawn on to the top of the nucellus, where partial disorganization of the cells has given rise to an irregular pollen-chamber (fig. 17, A, pc). The pollen-tube, containing two generative and one vegetative nucleus, pierces the wall of the megaspore and then becomes swollen (fig. 17, B and C, pt); finally the two generative nuclei pass out of the tube and fuse with two of the nuclei in the fertile half of the megaspore. As the result of fertilization, the fertilized nuclei of the megaspore become surrounded by a cell-wall, and constitute zygotes, which may attach themselves either to the wall of the megaspore or to the end of a pollen-tube (fig. 17, C, z and z′); they then grow into long tubes or proembryos, which make their way towards the prothallus (C, z′), and eventually embryos are formed from the ends of the proembryo tubes. One embryo only comes to maturity. The embryo of Gnetum forms an out-growth from the hypocotyl, which serves as a feeder and draws nourishment from the prothallus. The fleshy outer portion of the seed is formed from the outer perianth, the woody shell being derived from the inner perianth. The climbing species of Gnetum are characterized by the production of several concentric cylinders of secondary wood and bast, the additional cambium-rings being products of the pericycle, as in Cycas and Macrozamia. The structure of the wood agrees in the main with that of Ephedra.
Welwitschia (Tumboa).—This is by far the most remarkable member of the Gnetales, both as regards habit and the form of its flowers. In a supplement to the systematic work of Engler and Prantl the well-known name Welwitschia, instituted by Hooker in 1864 in honour of Welwitsch, the discoverer of the plant, is superseded by that of Tumboa, originally suggested by Welwitsch. The genus is confined to certain localities in Damaraland and adjoining territory on the west coast of tropical South Africa. A well-grown plant projects less than a foot above the surface of the ground; the stem, which may have a circumference of more than 12 ft., terminates in a depressed crown resembling a circular table with a median groove across the centre and prominent broad ridges concentric with the margin. The thick tuberous stem becomes rapidly narrower, and passes gradually downwards into a tap-root. A pair of small strap-shaped leaves succeed the two cotyledons of the seedling, and persist as the only leaves during the life of the plant; they retain the power of growth in their basal portion, which is sunk in a narrow groove near the edge of the crown, and the tough lamina, 6 ft. in length, becomes split into narrow strap-shaped or thong-like strips which trail on the ground. Numerous circular pits occur on the concentric ridges of the depressed and wrinkled crown, marking the position of former inflorescences borne in the leaf-axil at different stages in the growth of the plant. An inflorescence has the form of a dichotomously-branched cyme bearing small erect cones; those containing the female flowers attain the size of a fir-cone, and are scarlet in colour. Each cone consists of an axis, on which numerous broad and thin bracts are arranged in regular rows; in the axil of each bract occurs a single flower; a male flower is enclosed by two opposite pairs of leaves, forming a perianth surrounding a central sterile ovule encircled by a ring of stamens united below, but free distally as short filaments, each of which terminates in a trilocular anther. The integument of the sterile ovule is prolonged above the nucellus as a spirally-twisted tube expanded at its apex into a flat stigma-like organ. A complete and functional female flower consists of a single ovule with two integuments, the inner of which is prolonged into a narrow tubular micropyle, like that in the flower of Gnetum. The megaspore of Welwitschia is filled with a prothallus-tissue before fertilization, and some of the prothallus-cells function as egg-cells; these grow upwards as long tubes into the apical region of the nucellus, where they come into contact with the pollen-tubes. After the egg-cells have been fertilized by the non-motile male cells they grow into tubular proembryos, producing terminal embryos. The stem is traversed by numerous collateral bundles, which have a limited growth, and are constantly replaced by new bundles developed from strands of secondary meristem. One of the best-known anatomical characteristics of the genus is the occurrence of numerous spindle-shaped or branched fibres with enormously-thickened walls studded with crystals of calcium oxalate. Additional information has been published by Professor Pearson of Cape Town based on material collected in Damaraland in 1904 and 1906–1907. In 1906 he gave an account of the early stages of development of the male and female organs and, among other interesting statements in regard to the general biology of Welwitschia, he expressed the opinion that, as Hooker suspected, the ovules are pollinated by insect-agency. In a later paper Pearson considerably extended our knowledge of the reproduction and gametophyte of this genus.
Authorities.—General: Bentham and Hooker, Genera Plantarum (London, 1862–1883); Engler and Prantl, Die natürlichen Pflanzenfamilien (Leipzig, 1889 and 1897); Strasburger, Die Coniferen und Gnetaceen (Jena, 1872); Die Angiospermen und die Gymnospermen (Jena, 1879); Histologische Beiträge, iv. (Jena, 1892); Coulter and Chamberlain, Morphology of Spermatophytes (New York, 1901); Rendle, The Classification of Flowering Plants, vol. i. (Cambridge, 1904); “The Origin of Gymnosperms” (A discussion at the Linnean Society; New Phytologist, vol. v., 1906). Cycadales: Mettenius, “Beiträge zur Anatomie der Cycadeen,” Abh. k. sächs.
