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Originally appearing in Volume V12, Page 758 of the 1911 Encyclopedia Britannica.
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GINKGOALES.—This class-designation has been recently proposed to give emphasis to the isolated position of the genus Ginkgo (Salisburia) among the Gymnosperms. Ginkgo biloba, the maiden-hair tree, has usually been placed by botanists in the Taxeae in the neighbourhood of the yew (Taxus), but the proposal by Eichler in 1852 to institute a special family, the Salisburieae, indicated a recognition of the existence of special characteristics which distinguish the genus from other members of the Coniferae. The discovery by the Japanese botanist Hirase of the development of ciliated spermatozoids in the pollen-tube of Ginkgo, in place of the non-motile male cells of typical conifers, served as a cogent argument in favour of separating the genus from the Coniferales and placing it in a class of its own. In 1712 Kaempfer published a drawing of a Japanese tree, which he described under the name Ginkgo; this term was adopted in 1771 by Linnaeus, who spoke of Kaempfer's plant as Ginkgo biloba. In 1797 Smith proposed to use the name Salisburia adiantifolia in preference to the " uncouth " genus Ginkgo and " incorrect " specific term biloba. Both names are still in common use. On account of the resemblance of the leaves to those of some species of Adiantum, the appellation maiden-hair tree has long been given to Ginkgo biloba. Ginkgo is of special interest on account of its isolated position among existing plants, its restricted geographical distribution, and its great antiquity (see PALAEOBOTANY: Mesozoic). This solitary survivor of an ancient stock is almost extinct, but a few old and presumably wild trees are re-corded by travellers in parts of China. Ginkgo is common as a sacred tree in the gardens of temples in the Far East, and often cultivated in North America and Europe. Ginkgo biloba, which may reach a height of over 30 metres, forms a tree of pyramidal shape with a smooth grey bark. The leaves (figs. to and II) have a long, slender petiole terminating in a fan-shaped lamina, which may be entire, divided by a median incision into two wedge-shaped lobes, or subdivided into several narrow segments. The venation is like that of many ferns, e.g. Adiantum; the lowest vein in each half of the lamina follows a course parallel to the edge, and gives off numerous branches, which fork repeatedly as they spread in a palmate manner towards the leaf margin. The foliage-leaves occur either scattered on long shoots of unlimited growth, or at the apex of short shoots (spurs), which may eventually elongate into long shoots. The flowers are dioecious. The male flowers (fig. 12), borne in the axil of scale-leaves, consist of a stalked central axis bearing loosely Flowers. disposed stamens; each stamen consists of a slender filament terminating in a small apical scale, which bears usually two, but not infrequently three or four pollen-sacs (fig. 12, C). The axis of the flower is a shoot bearing leaves in the form of stamens. A mature pollen-grain contains a prothallus of 3 to 5 cells (Fig. 13, Pg) ; the exine extends over two-thirds of the circumference, leaving a thin portion of the wall, which on collapsing pro- duces a longitudinal groove similar to the median depression on the pollen-grain of a cycad. The ordinary type of female flower has the form of a long, naked peduncle bearing a single ovule on either side of the apex (fig. 12), the base of each being enclosed by a small, collar-like rim, the nature of which has been vari- ously interpreted. A young ovule consists of a conical nucellus sur- Py rounded by a single in-FIG. 13.—Ginkgo. Apex of Ovule, and tegument terminating as a Pollen-grain. (After Hirase.) two-le ped micrchamber pollen p, Pollen-tube (proximal end). occupies the apex of the c, Pollen-chamber. nucellus; immediately e, Upward prolongation of megaspore. below this, two or more a, Archegonia. archegonia (fig. 13, a) are Pg, Pollen-grain. developed in the upper Ex, Exine. region of the megaspore, each consisting of a large egg-cell surmounted by two neck-cells and a canal-cell which is cut off shortly before fertilization. After the entrance of the pollen-grain the pollen-chamber becomes roofed over by a blunt protuberance of nucellar tissue. The megaspore (embryo-sac) continues to grow after pollination until the greater part of the nucellus is gradually destroyed; it also gives rise to a vertical outgrowth, which projects from the apex of the megaspore as a short, thick column (fig. 13, e) supporting the remains of the nucellar tissue which forms the roof of the pollen-chamber (fig. 13, c). Surrounding the pitted wall of the ovum there is a definite layer of large cells, no doubt representing a tapetum, which, as in cycads and conifers, plays an important part in nourishing the growing egg-cell. The endosperm detached from a large Ginkgo ovule after fertilization bears a close resemblance to that of a cycad; the apex is occupied by a depression, on the floor of which two small holes mark the position of the archegonia, and the outgrowth from the megaspore apex projects from the centre as a short peg. After pollination the pollen-tube grows into the nucellar tissue, as in cycads, and the pollen-grain itself (fig. 13, Pg) hangs down into the pollen-chamber; two large spirally ciliated spermatozoids are produced, their manner of development agreeing very closely with that of the corresponding cells in Cycas and Zamia. After fertilization the ovum-nucleus divides and cell-formation proceeds rapidly, especially in the lower part of the ovum, in which the cotyledon and axis of the embryo are differentiated; the long, tangled suspensor of the cycadean embryo is not found in Ginkgo. It is often stated that fertilization occurs after the ovules have fallen, but it has been demonstrated by Hirase that this occurs while the ovules are still attached to the tree. The ripe seed, which grows as large as a rather small plum, is enclosed by a thick, fleshy envelope covering a hard woody shell with two or rarely three longitudinal keels. A papery remnant of nucellus lines the inner face of the woody shell, and, as in cycadean seeds, the apical portion is readily separated as a cap covering the summit of the endosperm. The morphology of the female flowers has been variously interpreted by botanists; the peduncle bearing the ovules has been described as homologous with the petiole of a foliage-leaf and as a shoot-structure, the collar-like envelope at the base of the ovules being referred to as a second integument or arillus, or as the representative of a carpel. The evidence afforded by normal and abnormal flowers appears to be in favour of the following interpretation: The peduncle is a shoot bearing two or more carpels. Each ovule is enclosed at the base by an envelope or collar homologous with the lamina of a leaf ; the fleshy and hard coats of the nucellus constitute a single integument. The stalk of an ovule, considerably reduced in normal flowers and much larger in some abnormal flowers, is homologous with a leaf-stalk, with which it agrees in the structure and number of vascular bundles. The facts on which this description is based are derived partly from anatomical evidence, and in part from an account given by a Japanese botanist, Fujii, of several abnormal female flowers; in some cases the collar at the base of an ovule, often described as an arillus, is found to pass gradually into the lamina of a leaf bearing marginal ovules (fig. I B). The occurrence of more than two ovules on one peduncle is by no means rare; aparticularly striking example is described by Fujii, in which an unusually thick peduncle bearing several stalked ovules terminates in a scaly bud (fig. 14, A, b). The frequent occurrence of more than two pollen-sacs and the equally common occurrence of additional ovules have been regarded by some authors as evidence in favour of the view that ancestral types normally possessed a greater number of these organs than are usually found in the recent species. This view receives support from fossil evidence. Close to the apex of a shoot the vascular bundles of a leaf make their Anatomy. appearance as double strands, and the leaf-traces in the upper part of a shoot have the form of distinct bundles, which in the older part of the shoot form a continuous ring. Each double leaf-trace passes through four internodes before becoming a part of the stele; the double nature of the trace is a characteristic feature. Secretory sacs occur abundantly in the leaf-lamina, where they appear as short lines between the veins; they are abundant also in the cortex and pith of the shoot, in the fleshy integument of the ovule, and elsewhere. The secondary wood. of the shoot and root conforms in the main to the coniferous type; in the short shoots the greater breadth FIG. 14.—Ginkgo. Abnormal female of the medullary rays in Flowers. A, Peduncle; b, scaly bud; the more internal part of B, leaf bearing marginal ovule. (After the xylem recalls the Fujii.) cycadean type. The secondary phloem contains numerous thick-walled fibres, parenchymatous cells, and large sieve-tubes with plates on the radial walls; swollen parenchymatous cells containing crystals are commonly met with in the cortex, pith and medullary-ray tissues. The wood consists of tracheids, with circular bordered pits on their radial walls, and in the late summer wood pits are unusually abundant on the tangential walls. A point of anatomical interest is the occurrence in the vascular bundles of the cotyledons, scale-leaves, and elsewhere of a few centripetally developed tracheids, which give to the xylem-strands a mesarch structure such as characterizes the foliar bundles of cycads. The root is diarch in structure, but additional protoxylem-strands may be present at the base of the main root; the pericycle consists of several layers of cells. This is not the place to discuss in detail the past history of Ginkgo (see PAL AEOBOTANY : Mesozoic). Among Palaeozoic genera there are some which bear a close resemblance to the recent type in Geological the form of the leaves; and petrified Palaeozoic seeds, history. almost identical with those of the maidenhair tree, have been described from French and English localities. During the Triassic and Jurassic periods the genus Baiera--no doubt a representative of the Ginkgoales—was widely spread throughout Europe and in other regions; Ginkgo itself occurs abundantly in Mesozoic and Tertiary rocks, and was a common plant in the Arctic regions as elsewhere during the Jurassic and Lower Cretaceous periods. Some unusually perfect Ginkgo leaves have been found in the Eocene leaf-beds between the lava-flows exposed in the cliffs of Mull (fig. II). From an evolutionary point of view, it is of interest to note the occurrence of filicinean and cycadean characters in the maidenhair tree. The leaves at once invite a comparison with ferns; the numerous long hairs which form a delicate woolly covering on young leaves recall the hairs of certain ferns, but agree more closely with the long filamentous hairs of recent cycads. The spermatozoids constitute the most striking link with both cycads and ferns. The structure of the seed, the presence of two neck-cells in the archegonia, the late development of the embryo, the partially-fused cotyledons and certain anatomical characters, are features common to Ginkgo and the cycads. 'The maidenhair tree is one of the most interesting survivals from the past; it represents a type which, in the Palaeozoic era, may have been merged into the extinct class Cordaitales. Through the succeeding ages the Ginkgoales were represented by numerous forms, which gradually became more restricted in their distribution and fewer in number during the Cretaceous and Tertiary periods, terminating at the present day in one solitary survivor.
End of Article: GINKGOALES

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