Comparative morphology of the carpel in the Liliaceae: Colchiceae (Androcymbium)

The pistil of Androcymbium closely resembles that of Colchicum : it is tricarpellate usually, syncarpous and multiovulate, and the carpels of most species have open sutures and bitegmic ovules. The only species with closed carpellary sutures, A. dregei has monotegmic ovules. There are always three dorsal bundles and three compound septal bundles, which latter may bifurcate into simple septal bundles. Six placental bundles (two per carpel) are differentiated, either separately from the compound septal bundles or as lateral branches of them. A statistical evaluation of 47 species (6 genera) of the hemisyncarpous Wurmbaeoideae shows a significant tendency for bitegmic ovules and two simple septal bundles per septum to be associated with open sutures and for monotegmic ovules and no septal bundles to be associated with closed sutures.     

Comparative morphology of the carpel in the Liliaceae: Uvularieae

Three genera of the Uvularieae (Kreysigia, Schelhammera, Uvularia) have tricarpellate, syncarpous pistils. Ventral bundles (presumably the united simple septal and placental bundles of a carpellary wing) may be present in Kreysigia and Schelhammera. In Kreysigia the two presumptive ventral bundles from adjoining carpels are fused basipetally in each septum. The septal bundles of the other two genera are either simple (Schelhammera) or in part compound (united) below and simple (separate) above (Uvularia) , hence fused acropetally. In Uvularia , the dorsal bundle of the carpel and the median bundle of the tepal are uniquely tripartite and probably homologous. No raphides were found in the carpels of these genera.     

Comparative morphology of the carpel in the Liliaceae: Veratreae

The genera of the Veratreae, a tribe of the Melanthioideae, have many features in common: there are usually many ovules, except for Amianthium (with 2 4), arranged in 2 -4 longitudinal placental rows per carpel; all are bitegmic, basipetal, and campylotropous. Of 37 species examined, only 2 have open sutures at the lowermost level of ovular insertion, but 13 species have holes in the centre of the pistil. These holes may represent possible stages in the evolutionary closure of previously open sutures. Most flowers were epigynous, only 11 being hypogynous-perigynous. The tribe as a whole is marked by the presence of 3 composite (heterologous) vascular bundles, composed of joined staminal and tepallary bundles alone and 3 composite bundles, as above, fused to a dorsal bundle. The bundles were united below the locular base in all genera except Schoenocaulon and Toxicoscordion. Two major kinds of central cylinder arrangement occurred at the level of the lowermost ovular insertion: either 6 inverted ventral bundles or 6 simple septal bundles, with normally arranged (or sometimes inverted) xylem and phloem centrifugally located and 6 simple placental bundles, with inverted xylem and phloem, at the centripetal end of the septum. Generally each septal bundle united with its nearest adjoining placental bundle about the mid-locular level.     

Comparative morphology of the carpel in the Liliaceae: Baeometra, Burchardia and Walleria

The pistils in Baeometra, Burchardia and Walleria ate tricarpellate, and their ovules are mostly bitegmic. Baeometra has free styles and deep septal invaginations between the carpels. Its pistil is innervated by three dorsal bundles, three compound septal bundles (each of which may divide into two simple septal bundles above), six placental bundles, and six adjoining auxiliary placental bundles. The pistil of Burchardia resembles that of Baeometra , except that there are six simple septal bundles throughout and no auxiliary placental bundles. In Walleria the wings of adjoining carpels are completely fused (except for rare septal glands); there is a single compound style; additional vascular tissue is present in the central axis of the pistil up to the lowermost ovules; the carpels are fused with the floral cup above the base of the locules; and raphide idioblasts are present. Walleria has six "ventral" bundles, each of which appears to be the fusion product of a placental bundle with a simple septal bundle. Tribal affinities of these genera are discussed.     

Comparative morphology of the carpel in the Liliaceae: Colchiceae (Colchicum)

The pistil of Colchicum is syncarpous, the carpels having open sutures or well-marked commissures and many bitegmic ovules of variable orientation. Although the vascularization of the carpel is also variable, there are usually three dorsal bundles and three alternate, septal bundles at the base of the pistil, with occasionally some placental bundles at that level. More often the placental bundles, differentiating basipetally, appear to establish connections with the septal bundles higher up, at the lowermost ovular insertion level. The septal bundles divide in two more frequently in pistils in which the carpellary suture is open than in those in which it is closed.     

Comparative morphology of the carpel in the Liliaceae: Wurmbaeae

The two genera of Buxbaum's tribe Wurmbaeae, Anguillaria and Wurmbea , have multiovulate carpels. There are deep septal indentations between the carpels of Anguillaria , but the wings of adjoining carpels are fused to solid septa in most species of Wurmbea. In Anguillaria the carpels have open sutures or prominent commissural markings; in Wurmbea the carpels generally lack these characteristics, and some species have a vascularized, columella-like axis in the centre of the pistil. In both genera there are a dorsal bundle, lateral bundles, and two placental bundles in each carpel. At the inner edge of the septum there are one or two septal bundles in Anguillaria and one or none in Wurmbea. The ovules are monotegmic, the integument and funiculus being partly fused in Anguillaria and mostly fused in Wurmbea. An obturator is present in Anguillaria but absent from most species of Wurmbea.     

Comparative morphology of the carpel in the Liliaceae: Helonieae

Most Helonieae have only slight septal indentations between the three carpels: in Xerophyllum deep septal clefts extend centripetally and completely enclosed, narrow septal pockets occur in Metanarthecium . Other unique generic features are found: tepallary-staminal nectarial glands in Heloniopsis , zygomorphy in Chionographis , and dioecism in Chamaelirium . The carpels are biovulate in Chionographis; there are two to several ovules per carpel in Xerophyllum; 8–12 ovules occur in the carpel of Chamaelirium; and numerous bitegmic ovules are borne in many longitudinal rows on enlarged placentae in Helonias, Heloniopsis, Metanarthecium , and Ypsilandra . Except for Metanarthecium , this last-named group of genera displays a near ring composed of 'accessory' placental bundles and a compound septal bundle (with normally oriented xylem and phloem) in cross-section at the inner edge of each septum. Ventral bundles occur in the other four genera.     

Comparative morphology of the carpel in the Liliaceae: Glorioseae

The pistils of the Glorioseae (Gloriosa, Littonia, Sandersonia) are generally tricarpellate and alike. Virtually all have closed sutures at flowering; they have many ovules, some of which are barely bitegmic, with inner integuments often nearly fused with nucellar remnants; and there is usually but one compound septal bundle in the inner edge of a septum. In two species of Littonia , the compound septal bundle divided to form two simple septal bundles; but in many other plants it remained undivided, and in some it died out, still undivided, below the locular apex. Most of the placental and septal bundles are vascularized in large part by three alternate (compound septal) bundles at the base of the locules and sometimes by branches from the lateral bundles. Three large (compound) placental bundles are formed just below the lowermost ovular insertion, and each then divides in two to furnish ovular branches along their ascent. Occasional auxiliary placental bundles lie between the septal bundle and the placental bundles in the septum (Gloriosa, Sandersonia).     

Comparative morphology of the carpel in the Liliaceae: Hewardieae, Petrosavieae, and Tricyrteae

The young pistils in the melanthioid tribes, Hewardieae, Petrosavieae and Tricyrteae, are uniformly tricarpellate and syncarpous. They lack raphide idioblasts. All are multiovulate, with bitegmic ovules. The Petrosavieae are marked by the presence of septal glands and incomplete syncarpy. Tepals and stamens adhere to the ovary in the Hewardieae and the Petrosavieae but not in the Tricyrteae. Two vascular bundles occur in the stamens of the Hewartlieae and Tricyrtis latifolia. Ventral bundles in the upper part of the ovary of the Hewardieae are continuous with compound septal bundles and placental bundles in the lower part. Putative ventral bundles occur in the alternate position in the Tricyrteae and putative placental bundles in the opposite. position in the Petrosavieae. The dichtomously branched stigma in each carpel of the Tricyrteae is supplied by a bifurcated dorsal bundle.     

STUDIES ON THE FLORAL ANATOMY OF THE CUNONIACEAE

Twenty-two genera representing sixty-two species of Cunoniaceae and Davidsonia were examined with respect to floral anatomy. Sepals are vascularized by three traces with the lateral traces of adjacent sepals united. Pancheria is unique for the family with species in which the sepals are vascularized by a single, undivided bundle. Petals, when present, and stamens, are uniformly one-trace structures. A general tendency exists within the family for the principal floral bundles to unite in various ways, with fusions evident between calyx, corolla, and androecial vascular supplies. Carpel number ranges from two to five and the gynoecium is generally surrounded by a prominent disc. Gynoecia of Ceratopetalum and Pullea are “half-inferior.” The number of ovules per carpel locule ranges from one to numerous. Ventral carpel sutures range from open to completely sealed at the level of placentation. Carpels of the apocarpous genus Spiraeanthemum (incl. Acsmithia) are vascularized by a dorsal bundle and either three or four bundles constituting the ovular and wing vasculation in the ventral position, a condition unlike other members of the family. Ovules are supplied by the median ventral bundle. More advanced bicarpellate gynoecia within the family are predominately vascularized by a dorsal and two ventral bundles although a variable number of additional lateral wall traces may be present. A major trend exists toward fusion of the ventral bundles of adjacent carpels in the ovary of both bicarpellate and multicarpellate plants. At the base of the styles the fused ventral strands separate and extend along with the dorsal carpellary bundles into styles of adjacent carpels. In Pullea the ventral bundles terminate within the ovules. The united ventral carpellary bundles in Aphanopetalum, Gillbeea, and Aistopetalum lie in the plane of the septa separating adjacent carpels. Ovules are vascularized by traces originating from the vascular cylinder at the base of the gynoecium or by traces branching from the ventral bundles. Ovular traces in each carpel are united, or remain as discrete bundles, prior to entering the placenta. Tannin and druses are common throughout all floral parts. Although floral anatomy generally supports the position of Cunoniaceae near Saxifragaceae and Davidsoniaceae, the evolutionary relationship of the Cunoniaceae to the Dilleniaceae is uncertain.     

Comparative morphology of the carpel in the Liliaceae: Iphigenieae

The pistil in the flowers of the Iphigenieae (Camptorrhiza, Iphigenia, Omithoglossum) is usually tricarpellate. The carpels are coherent generally, with closed sutures and seemingly bitegmic ovules. Camptorrhiza differs from the others in having a single compound style. The pistils of most species of these genera have a common vascular structure: three dorsal bundles which run into the style(s), a number of lateral bundles, six placental bundles, and up to three compound septal bundles. The latter nine bundles usually differentiate from a central vascular plexus above the base of the locules. There may be fewer than three septal bundles in some specieS. When present, the septal bundles usually die out in the ovuliferous region, but in some cases they persist to the apex of the locules.     

Morphology ofCoptis japonica and its meaning in phylogeny

In the genusCoptis, some interesting features are found which are considered important phylogenetically. The median bundles of petiole and petiolule, and the midrib of lamina are double. They seem to represent a transitional situation between a dichotomy and a single median bundle found in usual angiospermous venation. The double bundle is either derived from 2 independent leaf trace bundles or formed by dichotomy of a leaf trace bundle, and it does not seem so important whether the number of trace bundles is even or odd. The nodal structure is trilacunar or pentalacunar with 3, 4, 6 or 8 trace bundles. The upper part of the carpel does not produce ovules and is open from the initiation of the carpel. It is suggested that the carpel becomes open secondarily concomitant with the reduction of ovules. This shows that the closure of the carpel is not perfectly established.     

THE ANATOMY OF THE PISTILLATE INFLORESCENCE AND FLOWER OF CASUARINA VERTICILLATA LAMARCK (CASUARINACEAE)

The pistillate inflorescence of Casuarina verticillata is described as consisting of a primary axis bearing whorls of bracts with a cymule in the axil of each bract of the more central whorls. Each cymule consists of an atepallate, two-carpellate, syncarpous floret and two, lateral, once-lobed bracteoles. A “peripheral intercalary” meristem, in which divisions are primarily periclinal, forms a meshwork beneath the bracts from early development and moves the connate bracts centrifugally around the cymules and extends and binds the bracts, and to some extent the bracteoles, of the fertile part of the inflorescence together. Each bract receives a single trace; each cymule receives two traces. Each bundle extension of a cymule trace supplies: 1) a branch which joins its counterpart to become the anterior common carpellary bundle; 2) a second branch which joins its counterpart to become the posterior common carpellary bundle; and 3) a central branch which supplies a lateral bracteole. Within each floret, each common carpellary bundle provides a dorsal carpellary bundle, two ventral carpellary bundles (fertile anterior carpel) or one common ventral bundle (sterile posterior carpel). The ventral bundle-supplies join and form a single placental bundle which lies in the gynoecial septum, and which, in turn, supplies the two ovules in the anterior carpel. Whether the inflorescence is a simple racemose or a condensed cymose type cannot be determined from this species alone. The function of the sclerenchymatous, enclosing bracteoles and connate bracts is discussed.     

FLORAL ANATOMY IN THE SAXIFRAGACEAE SENSU LATO. II. SAXIFRAGOIDEAE AND ITEOIDEAE

The floral anatomy and morphology of 26 species from the Saxifragoideae and three from the Iteoideae are described and compared. The flowers of the Saxifragoideae are predominantly actinomorphic, partially epigynous and/or perigynous, and pentamerous, with two carpels which bear numerous ovules. There is usually some degree of independence between carpels, and the normally separate styles possess both a canal and transmitting tissue. Generally, staminodia are absent and nectariferous tissue, which is not vascularized, is present. The subfamily is characterized by large multicellular trichomes with globular, often glandular, heads. Placentation may be parietal, axile, or transitional between the two; parietal appears to be a derived condition in the subfamily. The vascular cylinder in the pedicel generally consists of several to many discrete bundles from which diverge ten compound traces at the base of the receptacle, leaving an inner cylinder of vascular strands that coalesce at a higher level into either as many ventral bundles as carpels or twice that number. In the former case, each ventral bundle consists of one-half of the vascular supply to each adjacent carpel and separates into individual ventral strands in the distal half of the ovary. The ventral bundles provide vascular traces to the ovules and, along with the dorsals, extend up the style to the stigma. Each trace diverging in a sepal plane typically supplies one or more carpel-wall bundles, a median sepal bundle, and a stamen bundle. Each petal-plane trace usually provides one or more carpel-wall bundles, a lateral trace to each adjacent sepal, a petal bundle and, in flowers with ten stamens, a stamen bundle. Dorsal carpel bundles are usually recognizable and may originate from traces in either perianth plane. While the position of Ribes remains problematical, its floral structure does not easily exclude it from the Saxifragoideae. Floral structure in the Iteoideae is remarkably similar to that in the Saxifragoideae, the main differences being a lesser degree of independence between carpels, generally narrower placentae with somewhat fewer ovules, and the presence of only unicellular, acutely pointed epidermal hairs as opposed to the relatively complex, multicellular trichomes prevalent in the Saxifragoideae.     

兰花蕉花部维管束系统的解剖学研究

兰花蕉花梗的维管束分散排列．子房基部的维管束排成两部分,外方为一轮大维管束环,中央为分散排列的小维管束区。前者的纸管束进入子房壁,后者进入子房的中轴,形成股座纸管束;及至延长都以后,股座维管束逐渐消失．子房壁上的维管束较易识别的有心皮背束、心皮背束伴束和隔膜束．三束心皮背束经延长部最终进入花柱和柱头．心皮背束指心皮背束务与其紧靠的大维管束,三枚心皮背束伴束最终分别进入三枚外轮雄蓝．三枚隔膜束中远轴面的两枚分别进入两校内轮雄蕊,而近轴面的一枚伴随着第六枚雄蓝的缺失最后进入唇瓣中央．子房壁其余的维管束进入延长部后,先向外分出一轮纸管束进入花幕,余下的中央部分排成一轮心形的线管来环．该环远轴面的维管束分为两半分别进入两枚侧生花瓣;近轴面即心形凹陷一侧初为两轮即外轮大的维管束与内轮小的维管束,后排成一轮并与近轴面的隔膜束一同进入唇瓣．兰花蕉的唇瓣既为花瓣成员,又含一枚缺失的雄蓝维管束,与姜目已报道的只来自退化雄蕊的竹芋科的兜状结构和美人蕉科、姜科、闭鞘姜科的唇瓣有明显区别．在旅人蕉科尚未有研究资料的情况下,作者根据已有资料,对姜目雄蕊维管束系统来源和结构进行比较,初步认为在姜目的系统演化上,兰花蕉科与芭蕉料更近．     

THE PALM GYNOECIUM

The morphology, anatomy, and histology of the gynoecia at or close to anthesis are described for 20 genera of palms selected to represent different taxonomic alliances and to include major gynoecial types within the family. Palms may have 1–10 carpels, but most have three. Fifteen genera, including 14 coryphoid palms and the monotypic Nypa fruticans, are apocarpous and the remainder, approximately 190, are syncarpous. Fusion of carpels in some gynoecia begins in the base, in others in the styles. Pseudomonomerous pistils occur in several different alliances: the ovarian parts of two carpels are reduced but three usually equal and functional styles and stigmas are present. The carpel is often follicular in shape with the ventral suture open or, more frequently, partially or completely closed. The carpel may be stipitate or sessile and usually has a conduplicate laminar part. Most carpels are spirally and laterally inserted on the receptacle, but the carpel in some unicarpellate genera (e.g., Thrinax) appears terminal. Stipes, ovarian parts, styles, and stigmas vary in structure and development. Septal nectaries which differ in size, in the presence or absence of specialized canals, and in position, characterize all genera of some groups but only some genera of others. Diverse vascular configurations in the bases of gynoecia vary according to the extent of the floral axis, the development of carpellary stipes, and the connation of the carpels and their adnation to the tip of the floral axis. Four types of carpellary vascular systems are present in the genera described: (1) most palm carpels have three major traces consisting of a dorsal bundle and two ventral bundles, and they may also have up to four pairs of lateral bundles or occasionally more; (2) in certain cocosoid palms no ventral bundles can be distinguished, but a dorsal bundle, many parallel lateral bundles, and a row of immature ventral strands vascularize each carpel; (3) carpels of Phytelephas have a dorsal bundle, two pairs of major lateral bundles and about four pairs of shorter lateral bundles, with no identifiable ventral bundles; (4) carpels of Nypa have many dichotomously branched bundles but none that are recognizable as dorsal, ventral, or lateral strands. Additional peripheral bundles or systems may be present in each of the above types. Ovules are supplied by 1–15 bundles. These are derived either from the carpellary stele; from ventral bundles only; from ventral, lateral, and dorsal bundles; or from a combination of these origins. Certain areas of the gynoecia or certain parts of dorsal carpellary walls in some genera are much less mature at anthesis than surrounding tissues. Implications for floral biology and relationships within the palms and of palms to other groups are discussed.     

PISTIL DEVELOPMENT AND PARIETAL PLACENTATION IN THE PSEUDOMONOMEROUS OVARY OF ZELKOVA SERRATA (ULMACEAE)

Development of female flowers in Zelkova serrata was observed using epi-illuminated microscopy and scanning electron microscopy, with particular attention given to placentation. After the inception of staminodial primordia, the floral apex becomes flat, and the first and subsequently the second carpel primordia appear at opposite comers of the pistil primordium. Inside each carpel primordium a fossette forms. Through differential growth this depression becomes clear and the carpel wall encircles one side of the future placental region. The placental region is detectable even in early stages, but clear signs of ovule inception appear late when the placental region is elevated onto one side of the ovary wall by intercalary growth. Although the relative size of the two carpels varies among flowers, the placental position always appears to be the border between the two carpels and the floral apex. This suggests that the placentation of Zelkova is parietal. The ovule position in tricarpellate ovaries also suggests an evolutionary derivation from ovaries with parietal placentation. Parietal placentation appears to be the original condition in Urticales.     

THE MORPHOLOGY OF THE MYRISTICACEAE. I. FLOWERS OF MYRISTICA FRAGRANS AND M. MALABARICA

Floral histology and vascular anatomy of Myristica fragrans Van Houtt. and M. malabarica Lam. have been investigated from serial sections and specimens cleared in chloro-lacto-phenol. The flowers are unisexual. The androecium is considered to consist of a whorl of laterally concrescent anthers. The bisporangiate anthers are attached by a ridge of tissue to the terminal part of the androphore. In many cases the number of vascular bundles in the androphore is half the number of anthers. The gynoecium consists of a monocarpellate pistil with basal placentation and a single anatropous ovule. Of the many vascular bundles that enter the base of the carpel, two, because of their position and because they provide vascular traces to the ovule, are designated as ventral bundles. Additional ovular traces are provided by the carpel wall vascular system. These additional traces originate at the top of the locule and descend to the ovule. The similarity between the androecia of these two species and the androecium of the ćnellaceae is noted.     

COMPARATIVE MORPHOLOGY OF THE CARPEL IN THE ROSACEAE VII. POMOIDEAE: CHAENOMELES,CYDONIA, DOCYNIA

The multi-ovulate pomoids, Chaenomeles, Cydonia, and Docynia, all have closed sutures and extensive fusion between carpel and floral cup and between ovular and wing bundles. Although the ovules in Docynia are generally apotropic and few in number (4–7), the ovules in the other two genera are pleurotropic and numerous (15–48). A statistical treatment of the whole tribe of Pomoideae shows that in carpels with open sutures ovular and wing bundles definitely tend to be separate while in those with closed sutures these bundles tend to be fused. To a lesser degree carpels with open sutures also tend to have bitegmic ovules, separate carpels, and a lesser extent of fusion between carpel and floral cup, while carpels with closed sutures tend to have monotegmic ovules, united carpels, and a greater extent of fusion between carpel and floral cup.