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Originally appearing in Volume V12, Page 757 of the 1911 Encyclopedia Britannica.
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CYCADOPHYTA.—A. Cycadales.—Stems tuberous or columnar, not infrequently branched, rarely epiphytic (Peruvian species of Zamia) ; fronds pinnate, bi-pinnate in the Australian genus Bowenia. Dioecious; flowers in the form of cones, except the female flowers of Cycas, which consist of a rosette of leaf-like carpels at the apex of the stem. Seeds albuminous, with one integument; the single embryo, usually bearing two partially fused cotyledons, is attached to a long tangled suspensor. Stems and roots increase in diameter by secondary thickening, the secondary wood being produced by one cambium or developed from successive cambium-rings. The cycads constitute a homogeneous group of a few living members confined to tropical and sub-tropical regions. As a fairly typical and well-known example of the Cycadaceae, a species of the genus Cycas (e.g. C. circinalis, C. revoluta, &c.) is briefly de-scribed. The stout columnar stem may reach a height of 20 metres, and a diameter of half a metre; it remains either unbranched or divides near the summit into several short and thick branches, each branch terminating in a crown of long pinnate leaves. The surface of the stem is covered with rhomboidal areas, which represent the persistent bases of foliage- and scale-leaves. In some species of Cycas there is a well-defined alternation of transverse zones on the stem, consisting of larger areas representing foliage-leaf bases, and similar but smaller areas formed by the bases of scale-leaves (F and S, fig. r). The scale-leaves clothing the terminal bud are linear-lanceolate in form, and of a brown or yellow colour; they are pushed aside as the stem-axis elongates and becomes shrivelled, finally falling off, leaving projecting bases which are eventually cut off at a still lower level. Similarly, the dead fronds fall off ,leaving a ragged petiole, which is afterwards separated from the stem by an absciss-layer a short distance above the base. In some species of Cycas the leaf-bases do not persist as a permanent covering to the stem, but the surface F S F s is covered with a wrinkled bark, as in Cycas siamensis, which has a stem of unusual form (fig. 2). Small tuberous shoots, comparable on a large scale with the bulbils of Lycopodium Selago, are occasionally produced in the axils of some of the persistent leaf-bases; these are characteristic of sickly plants, and serve as a means of vegetative reproduction. In the genus Cycas the female flower is peculiar among cycads in consisting of a terminal crown of separate leaf-like carpels several inches in length ; the apical portion of each carpellary leaf may be broadly triangular in form, and deeply dissected on the margins into narrow woolly appendages like rudimentary pinnae. From the lower part of a carpel are produced several laterally placed ovules, which become bright red or orange on ripening; the bright fleshy seeds, which in some species are as large as a goose's egg, and the tawny spreading carpels produce a pleasing combination of colour in the midst of the long dark-green fronds, which curve grace-fully upwards and outwards from the summit of the columnar stem. In Cycas the stem apex, after producing a cluster of carpellary leaves, continues to elongate and produces more bud-scales, which are afterwards pushed aside as a fresh crown of fronds is developed. The young leaves of Cycas consist of a straight rachis bearing numerous linear pinnae, traversed by a single midrib; the pinnae are circinately coiled like the leaf of a fern (fig. 3). The male flower of Cycas conforms to the type of structure characteristic of the cycads, and consists of a long cone of numerous sporophylls bearing many oval pollen-sacs on their lower faces. The type described serves as a convenient representative of its class. There are eight other living genera, which may be classified as follows: Classification.—A. Cycadeae.—Characterized by (a) the alternation of scale- and foliage-leaves (fig. I) on the branched or unbranched stem; (b) the growth Of the main stem through the female flower; (c) the presence of a prominent single vein in the linear pinnae; (d) the structure of the female flower, which is peculiar in not having the form of a cone, but consists of numerous independent carpels, each of which bears two or more lateral ovules. Represented by a single genus, Cycas. (Tropical Asia, Australia, &c.). B. Zamieae.—The stem does not grow through the female flower; both male and female flowers are in the form of cones. (a) Stangerieae.— Characterized by the fern-like venation of the pinnae, which have a prominent midrib, giving off at a wide angle simple or forked and occasionally anastomosing lateral veins. A single genus, Stangeria, confined to South Africa. (b) Euzamieae.—The pinnae are traversed by several parallel veins. Bowenia, an Australian cycad, is peculiar in having bi-pinnate fronds (fig. 5). The various genera are distinguished from one another by the shape and manner of attachment of the pinnae, the form of the carpellary scales, and to some extent by anatomical characters. Encepha- lartos (South and Tropical Africa).—Large cones; the .carpellary scales terminate in a peltate di, tal expansion. Macrozamia (Australia).—Similar to Encephalartos except in the presence of a spinous projection from the swollen distal end of the carpels. Zamia (South America, Florida, &c.).—Stem short and often divided into several columnar branches. Each carpel terminates in a peltate head. Cerato- zamia (Mexico).—Similar in habit to Macrozamia, Young Frond. spinous processes on the apex of the carpels. Microcycas (Cuba).—Like Zamia, except that the ends of the stamens are flat, while the apices of the carpels are peltate. Dioon (Mexico) (fig. 4).—Characterized by the woolly scale-leaves and carpels; the latter terminate in a thick laminar expansion of triangular form, bearing two placental cushions, on which the ovules are situated. Bowenia (Australia).—Bi-pinnate fronds; stem short and tuberous (fig. 5). The stems of cycads are often described as unbranched; it is true that in comparison with conifers, in which the numerous branches, springing from the main stem, give a characteristic form stem to the tree, the tuberous or columnar stem of the Cyca- aad leaf daceae constitutes a striking distinguishing feature. Branching. however, occurs not infrequently: in Cycas the tall stem often produces several candelabra-like arms; in Zamia the main axis may break up near the base into several cylindrical branches; in species of Dioon (fig. 4) lateral branches are occasion-ally produced. The South African Encephalartos frequently produces several branches. Probably the oldest example of this genus in cultivation is in the Botanic Garden of Amsterdam, its age is considered by Professor de Vries to be about two thousand years: although an accurate determination of age is impossible, there is no doubt that many cycads grow very slowly and are remarkable for longevity. The thick armour of petiole-bases enveloping the stem is a characteristic C y c a d e a n feature; in Cycas the alter-nation of scale-leaves and fronds is more clearly shown than in other cycads; in Encephalartos, Dioon, &c., the persistent scale-leaf bases are almost equal in size to those of the foliage-leaves, and there is no , regular alternation of zones such as characterizes some species of Cycas. Another type of stem is illustrated by Stangeria and Zamia, also by a few forms of Cycas (fig. 2), in which the fronds fall off completely, leaving a comparatively smooth stem. The Cyas type of frond, except as regards the presence of a midrib in each pinna, characterizes the cycads generally, except Bowenia and Stangeria. In the monotypic genus Bowenia the large fronds, borne singly on the short and thick stem, are bi-pinnate (fig. 5); the segments, which are broadly ovate or rhomboidal, have several forked spreading veins, and resemble the large pinnules of some species of Adiantum. In Stangeria, also a genus represented by one species (S. paradoxa of South Africa), the long and comparatively broad pinnae, with an entire or irregularly incised margin, are very fern-like, a circumstance which led Kunze to describe the plant in 1835 as a species of the fern L omaria. In rare cases the pinnae of cycads are lobed or branched: in Dioon spinulosum (Central America) the margin of the segments bears numerous spinous pro-Oases; in some species of Encephalartos, e.g. E. horridus, the lamina is deeply lobed; and in a species of the Australian genus Macrozamia, AT. heteromera, the narrow pinnae are dichotomously branched almost to the base (fig. 6), and resemble the frond of some species of the fern Schizaea, or the fossil genus Baiera (Ginkgoales). An interesting species of Cycas, C. Micholitzii, has recently been described by Sir William Thiselton-Dyer from Annam, where it was collected by one of Messrs Sanders & Son's collectors, in which the pinnae instead of being of the usual simple type are From a photograph of a plant in the Peradeniya Gardens, Ceylon, by Professor R. If. Yapp. dichotomously branched as in Macrozamia heteromera. In Cerato- cases, the pollen is conveyed to the ovules by animal agency. zamia the broad petiole-base is characterized by the presence of two lateral spinous processes, suggesting stipular appendages, comparable, on a reduced scale, with the large stipules of the Marattiaceae among Ferns. The vernation varies in different genera; in Cycas the rachis is straight and the pinnae circinately coiled (fig. 3) ; in Encephalartos, Dioon, &c., both rachis and segments are straight; in Zamia the rachis is bent or slightly coiled, bearing straight pinnae. The young leaves arise on the stem-apex as conical protuberances with winged borders, on which the pinnae appear as rounded humps, usually in basipetal order; the scale-leaves in their young condition resemble fronds, but the lamina remains undeveloped. A feature of interest in connexion with the phylogeny of cycads is the presence of long hairs clothing the scale-leaves, and forming a cap on the summit of the stem-apex or attached to the bases of petioles; on some fossil cycadean plants these outgrowths have the form of scales, and are identical in structure with the ramenta (paleae) of the majority of ferns. The male flowers of cycads are constructed on a uniform plan, and in all cases consist of an axis bearing crowded, spirally dis-Flower. posed sporophylls. These are often wedge-shaped and angular; in some cases they consist of a short, thick stalk, terminating in a peltate expansion, or prolonged upwards in the form of a triangular lamina. The sporangia (pollen-sacs), which occur on th% under-side of the stamens, are often arranged in more or less definite groups or sori, interspersed with hairs (paraphyses) ; dehiscence takes place along a line marked out by the occurrence of smaller and thinner-walled cells bounded by larger and thicker-walled elements, which form a fairly prominent cap-like " annulus " near the apex of the sporangium, not unlike the annulus characteristic of the Schizaeaceae among ferns. The sporangial wall, consisting of several layers of cells, encloses a cavity containing numerous oval spores (pollen-grains). In structure a cycadean sporangium recalls those of certain ferns (Marattiaceae, Osmundaceae and Schizaeaceae). but in the development of the spores there are certain peculiarities not met with among the Vascular Cryptogams. With the exception of Cycas, the female flowers are also in the form of cones, bearing numerous carpellary scales. In Cycas revoluta and circinalis each leaf-like carpel may produce several laterally att ed ovules, but in C. Normanbyand the carpel is shorter and the ovules are reduced to two; this latter type brings us nearer to the carpels of Dioon, in which the flower has the form of a cone, and the distal end of the carpels is longer and more leaf-like than in the other genera of the Zamieae, which are characterized by shorter carpels with thick peltate heads bearing two ovules on the morphologically lower surface. The cones of cycads attain in some cases (e.g. Encephalartos) a considerable size, reaching a length of more than a foot. Cases have been recorded (by Thiselton-Dyer in Encephalartos and by Wieland in Zamia) in which the short carpellary cone-scales exhibit a foliaceous form. It is interesting that no monstrous cycadean cone has been described in which ovuliferous and staminate appendages are borne on the same axis: in the Bennettitales (see PALAEOBOTANY: Mesozoic) flowers were produced bearing on the same axis both androecium and gynoecium. The pollen-grains when mature consist of three cells, two small and one large cell; the latter grows into the pollen-tube, as in the Mlcro- Coniferales, and from one of the small cells two large spores ciliated spermatozoids are eventually produced. A and remarkable exception to this rule has recently been recorded by Caldwell, who found that in Microcycas mega- Calocoma the body-cells may be eight or even ten in spores. number and the sperm-cells twice as numerous. One of the most important discoveries made during the latter part of the 19th century was that by Ikeno, a Japanese botanist, who first demonstrated the existence of motile male cells in the genus Cycas. Similar spermatozoids were observed in some species of Zamia by H. J. Webber, and more recent work enables us to assume that all cycads produce ciliated male gametes. Before following the growth of the pollen-grain after pollination, we will briefly describe the structure of a cycadean ovule. An ovule consists of a conical nucellus surrounded by a single integument. At an early stage of development a large cell makes its appearance in the central region of the nucellus; this increases in size and eventually forms three cells; the lowest of these grows vigorously and constitutes the megaspore (embryo-sac) ,which ultimately absorbs the greater part of the nucellus. The megaspore-nucleus divides repeatedly, and cells are produced from the peripheral region inwards, which eventually fill the spore-cavity with a homogeneous tissue (prothallus) ; some of the superficial cells at the micropylar end of the megaspore increase in size and divide by a tangential wall into two, an upper cell which gives rise to the short two-celled neck of the archegonium, and a lower cell which develops into a large egg-cell. Each megaspore may contain 2 to 6 archegonia. During the growth of the ovum nourishment is supplied from the contents of the cells immediately surrounding the egg-cell, as in the development of the ovum of Pinus and other conifers. Meanwhile the tissue in the apical region of the nucellus has been undergoing disorganization, which results in the formation of a pollen-chamber (fig. 7, C) immediately above the mega-spore. Pollination in cycads has always been described as anemophilous, but according to recent observations by Pearson on South African species it seems probable that, at least in some The pollen-grains find their way between the carpophylls, which at the time of pollination are slightly apart owing to the elongation of the internodes of the flower-axis, and pass into the pollen-chamber; the large cell of the pollen-grain grows out into a tube (Pt), which penetrates the nucellar tissue and often branches repeatedly; the pollen-grain itself, with the prothallus-cells, projects freely into the pollen-chamber (fig. 7). The nucleus of the outermost (second small cell (fig. 7, G) divides, and one of the daughter-nuclei passes out of the cell, and may enter the lowest (first) small cell. The outermost cell, by the division of the remaining nucleus, produces two large spermatozoids (fig. 8, a, a). In Microcycas i6 sperm-cells are produced. In the course of division two bodies appear in the cytoplasm, and behave as centrosomes during the karyokinesis; they gradually become threadlike and coil round each daughter nucleus. This thread gives rise to a spiral ciliated band lying in a depression on the body of each spermatozoid; the large spermatozoids eventually escape from the pollen-tube, and are able to perform ciliary movements in the watery liquid which occurs between the thin papery remnant of nucellar tissue and the archegonial necks. Before fertilization a neck-canal cell is formed by the division of the ovum-nucleus. After the body of a spermatozoid has coalesced with the egg-nucleus the latter divides repeatedly and forms a mass of tissue which grows more vigorously in the lower part of the fertilized ovum, and extends upwards towards the apex of the ovum as a peripheral layer of parenchyma surrounding a central space. By further growth this tissue gives rise to a proembryo, which consists, at the micropylar end, of a sac; the tissue at the chalazal end grows into a long and tangled suspensor, terminating in a mass of cells, which is eventually differentiated into a radicle, plumule and two cotyledons. In the ripe seed the integument assumes the form of a fleshy envelope, succeeded internally by a hard woody shell, internal to which is a thin papery membrane—the apical portion of the nucellus—which is easily dissected out as a conical cap covering the apex of the endosperm. A thorough examination of cycadean seeds has recently been made by Miss Stopes, more particularly with a view to a comparison of their vascular supply with that in Palaeozoic gymnospermous seeds (Flora, 1904). The first leaves borne on the seedling axis are often scale-like, and these are followed by two or more larger laminae, which foreshadow the pinnae of the adult frond. The anatomical structure of the vegetative organs of recent cycads is of special interest as affording important evidence of rela- Anatomy. tionship with extinct types, and with other groups of recent plants. Brongniart, who was the first to investigate in detail the anatomy of a cycadean stem, recognized an agreement, as regards the secondary wood, with Dicotyledons and Gymnosperms, rather than with Monocotyledons. He drew attention also to certain structural similarities between Cycas and Ginkgo. The main anatomical features of a cycad stem may be summarized as follows: the centre is occupied by a large parenchymatous pith traversed by numerous secretory canals, and in some genera by cauline vascular bundles (e.g. Encephalartos and Macrozamia). In addition to these cauline strands (confined to the stem and not connected with the leaves), collateral bundles are often met with in the pith, which form the vascular supply of terminal, flowers borne at intervals on the apex of the stem. These latter bundles may be seen in sections of old stems to pursue a more or less horizontal course, passing outwards through the main woody cylinder. This lateral course is due to the more vigorous growth of the axillary branch formed near the base of each flower, which is a terminal structure, and, except in the female flower of Cycas, puts a limit to the apical growth of the stem. The vigorous lateral branch therefore continues the line of the main axis. The pith is encircled by a cylinder of secondary wood, consisting of single or multiple radial rows of tracheids separated by broad medullary rays composed of large parenchymatous cells; the tracheids bear numerous bordered N, Nucellus. G, Generative cell C, Pollen-chamber. (second cell of pollen-tube). a FsG.8: Zamia. Proximal end of Pollen-tube. a, a,Spermatozoids from G of fig. 7; Pg,pollen-grain; c, proximal cell (first cell). (Af ter Webber.) pits on the radial walls. The large medullary rays give to the wood a characteristic parenchymatous or lax appearance, which is in marked contrast to the more compact wood of a conifer. The protoxylem-elements are situated at the extreme inner edge of the secondary wood, and may occur as small groups of narrow, spirally-pitted elements scattered among the parenchyma which abuts on the main mass of wood. Short and reticulately-pitted tracheal cells, similar to tracheids, often occur in the circummedullary region of cycadean stems. In an old stem of Cycas, Encephalartos or Macrozamia the secondary wood consists of several rather unevenly concentric zones, while in some other genera it forms a continuous mass as in conifers and normal dicotyledons. These concentric rings of secondary xylem and phloem (fig. 9) afford a characteristic cycadean feature. After the cambium has been active for some time producing secondary xylem and " • - phloem, the latter consisting of sieve- tubes, phloem-parenchyma and fre- quently thick-walled fibres, a second cambium is developed in the perits cycle ; this produces a second vascular zone, which is in turn followed by a third cambium, and so on, until several hollow cylinders are developed. It has been recently shown that several cambium-zones may remain in a state 0 0 0 0 a of activity, so that the formation of a new cambium does not necessarily (After \Vorsdell.) xylem and phloem are developed ppd, Periderm in leaf-bases. internally to some of the vascular lt, Leaf-traces in cortex. rings; these are characterized by an ph, Phloem. inverse orientation of the tissues, x, Xylem. the xylem being centrifugal and the m, Medullary bundles. phloem centripetal in its development. c, Cortical bundles. The broad cortical region, which con- tains many secretory canals, is tra- versed by numerous vascular bundles (fig. 9, c) some of which pursue a more or less vertical course, and by frequent anastomoses with one another form a loose reticulum of vascular strands; others are leaf- traces on their way from the stele of the stem to the leaves. Most of these cortical bundles are collateral in structure, but in some the xylem and phloem are concentrically arranged; the secondary origin of these bundles from procambium-strands was described by Mettenius in his classical paper of 186o. During the increase in thickness of a cycadean stem successive layers of cork-tissue are formed by phello- gens in the persistent bases of leaves (fig. 9, pd), which increase in size to adapt themselves to the growth of the vascular zones. The leaf- traces of cycads are remarkable both on account of their course and their anatomy. In a transverse section of a stem (fig. 9) one sees some vascular bundles following a horizontal or slightly oblique course in the cortex, stretch- ing for a longer or shorter distance in a direction con- centric with the woody cylinder. From each leaf- base two main bundles spread right and left through the cortex of the stem (fig. 9, it), and as they curve gradually towards the vascular ring they present the appearance of two rather flat ogee curves, usually spoken of as the leaf-trace girdles (fig. 9, it). The distal ends of these girdles give off several branches, which traverse the petiole and rachis as numerous collateral bundles. The complicated girdle-like course is characteristic of the leaf-traces of most recent cycads, but in some cases, e.g. in Zamia floi'idana, the traces are described by Wieland in his recent monograph on American fossil cycads (Carnegie Institu- tion Publications, 1906) as possessing a more direct course similar to that in Mesozoic genera. A leaf-trace, as it passes through the cortex, has a collateral structure, the protoxylem being situated at the inner edge,of the xylem; when it reaches the leaf-base the position of the spiral tracheids is gradually altered, and the endarch arrangement (protoxylem internal) gives place to a mesarch structure (protoxylem more or less central and not on the edge of the xylem strand). In a bundle examined in the basal portion of a leaf the bulk of the xylem is found to be centrifugal in position, but internally to the protoxylem there is a group of centripetal tracheids; higher up in the petiole the xylem is mainly centripetal, the centrifugal wood being represented by a small arc of tracheids external to the protoxylem and separated from it by a few parenchymatous elements. Finally, in the pinnae of the frond the centrifugal xylem may disappear, the protoxylem being now exarch in position and abutting on the phloem. Similarly in the sporophylls of some cycads the bundles are endarch near the base and mesarch near the distal end of the stamen or carpel. The vascular system of cycadean seedlings presents some features worthy of note; centripetal xylem occurs in the cotyledonary bundles associated with transfusion-tracheids. The bundles from the cotyledons pursue a direct course to the stele of the main axis, and do not assume the girdle-form characteristic of the adult plant. This is of interest from the point of view of the comparison of recent cycads with extinct species (Bennettites), in which the leaf-traces follow a much more direct course than in modern cycads. The mesarch structure of the leaf-bundles is met with in a less pronounced form in the flower ped- uncles of some cycads. This fact is of importance as showing that the FIG.II.—Ginkgoadiantoides. type of vascular structure, which Fossil (Eocene) leaf from the characterized the stems of many Island of Mull. Palaeozoic genera, has not entirely disappeared from the stems of modern cycarls; but the mesarch bundle is now confined to the leaves and peduncles. The roots of some cycads resemble the stems in producing several cambium- Roots. rings; they possess 2 to 8 protoxylem-groups, and are characterized by a broad pericyclic zone. A common phenomenon in cycads is the production of roots which grow upwards (apogeotropic), and appear as coralline branched.structures above the level of the ground; some of the cortical cells of these roots are hypertrophied, and contain numerous filaments of blue-green Algae (Nostocaceae), which live as endoparasites in the cell-cavities.
End of Article: CYCADOPHYTA
CYBELE, or CYBEBE (Gr. Ku/3X, Ku,(3i7sn)

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