Vascular floral anatomy of the east asianHeloniopsis orientalis (Thunb.) C. Tanaka (Liliaceae-Helonieae)

Observations on the vascular floral anatomy, carpel morphology and floral biology ofHeloniopsis orientalis are presented. The lower flowering pedicel has six large bundles which lack an enclosing sclerenchymatous sheath. At mid-pedicel, branch bundles originate via radial divisions from each of these bundles. Subsequently, there is a vascular ring of 12 bundles below the receptacle. The six smaller bundles which are derived from alternate pedicel bundles eventually establish all of the ventral gynoecium supply. The six larger bundles supply the tepals, stamens and dorsal gynoecial vasculature. The simple dorsals do not branch or fuse in their vertical ascent. The ventral and placental supplies are far more complex. Fusion occurs between paired sets of the six smaller pedicel bundles along the septal radii and results in a submarginal laminal ventral network. An independent ventral plexus is formed in each septum and from each plexus two septal axials, of which the innermost has a reversed xylem-phloem disposition, and four placental bundles are derived. Two placental bundles are associated with each septal axial. Basally the septa are fused centrally, but are freed at mid-gymoecial height. The broadly tri-lobed, tri-carpellate gynoecium is depressed terminally where the erect, hollow style with its capitate stigma is attached. Dorsal grooves are present: the fruit is loculicidally dehiscent. There are no septal glands due to complete lateral fusion of the septal wings. Basally each of the six equal tepals has a saccate nectary. The similarity in vascular anatomy and carpel morphology of the AsianHeloniopsis and eastern North American endemic,Helonias bullata, justifies their position in the same tribe. Research and publication supported in part by the M. Graham Netting Research Fund through a grant from the Cordelia Scaife May Charitable Trust, the U. S.—Japan Cooperative Science Program Grant GF-41367, the Japan Society for the Promotion of Science, and Grant-in-Aid No. 934053 from the Ministry of Education, Japan.     

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.     

兰花蕉花的形态解剖学

兰花蕉（Orchidantha chinensis)的子房室顶部闭合后向上延长成延长部,实心,但有花柱沟和隔膜蜜腺管通过,隔膜蜜腺管,可分为中央蜜腺管和三条侧蜜腺管;中央蜜腺管位于三个心皮连接处,自子房室区下部产生,向上于延长部的部顶端终止;三条侧管分别位于两个心皮连接处,于子房室区近中部产生,开口于花柱基部。兰花蕉子房室区与延长部均具6枚雄蕊的维管束系统,即3枚心皮背束的伴束与3枚隔膜束,近轴面1枚事膜向上进入唇瓣的维管束系统,位于唇瓣的中央,致使兰花蕉仅具5枚功能雄蕊,唇瓣具双重结构,本文还讨论了兰花蕉科的系统发育位置。     

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 STUDY OF THE FLORAL VASCULATURE IN SAURURACEAE

The floral vascular systems are compared among all six taxa of Saururaceae, including the two species of Gymnotheca which have not been studied previously. All are zygomorphic (dorsiventrally symmetrical), not radial as sometimes reported, in conformity with dorsiventral symmetry during organogenesis. Apocarpy in the two species of Saururus (with four carpels and six free stamens) is accompanied by a vascular system of four sympodia, each of which supplies a dorsal carpellary bundle, two ventral carpellary bundles, and one or two stamen traces. The level at which the ventral bundles diverge is the major difference in vasculature between the two species. The other four taxa are all syncarpous, and share some degree of stamen adnation and/or connation. The vascular systems also show varying degrees of fusion. The two species of Gymnotheca (with four carpels and six stamens) are very similar to each other; in both, the ventral traces of adjacent carpels fuse to form a placental bundle, which supplies the ovules and then splits into a pair of ventral strands. The flowers of Houttuynia cordata (with only three carpels and three adnate stamens) are sessile. Each flower is vascularized by three sympodia; the median adaxial sympodium is longer than the other two sympodia before it diverges to supply the adaxial organs. Three placental bundles also are formed in Houttuynia, but the three bundles differ in their origin. The median abaxial placental bundle diverges at the same level as the three sympodial bundles of the flower, while the other two lateral placental bundles diverge at a higher level from the median adaxial sympodium. Anemopsis californica, with an inferior ovary of three carpels, sunken in the inflorescence axis, and six stamens adnate to the carpels, has a vascular system very similar to that of Houttuynia cordata. The modular theory of floral evolution is criticized, on the bases of the known behavior of apical meristems and properties of vascular systems. The hypothesis is supported that saururaceous plants may represent a line of angiosperms which diverged very early.     

大鹤望兰花部维管束系统的解剖学研究

大鹤望兰梗横切面近三角形,花梗的维管束分散公布在基本组织内。室下区的维管束大致排列三两部分,外方为一到两环维管束组成的外维管束环,中央为分散排列的中央维管束区。前者的维管束进入子房避讳,后者的维管束进入子房的中轴,形成从维管束。至延长部后,胎座维管束逐渐消失。子房壁上的维管束较易识别的有心皮背束、心成背束伴束和隔膜束。３束心皮背束经处长部最终进入花柱。３枚心皮背束伴束最终分别进入一枚１２上轮雄蕊。     

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: 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.     

Complex polyandry in the Magnoliatae: definition, distribution and systematic value

A revised definition of complex polyandry is presented for the Magnoliatae. The differences between primary polyandry and secondary (complex) polyandry are clarified. Fascicled stamens are probably not primitive in Magnoliatae. Complex polyandry finds its origin by an increase of stamens in an oligandrous framework. Multiplications of stamens occur either on primary complex primordia, which follow the regular phyllotactic sequence in the flower, either on a girdling primordium (ringwall). The primary primordia may be fully developed before a secondary proliferation of stamens takes place or provide room for more stamens by a continuing growth process. This happens by the development of a staminal tube (e.g. Malvales) or by the growth of a hypanthium (e.g. Myrtales). A typical floral vasculature is developed concomitantly with complex polyandry. Stamen trunk bundles (stamen fascicle traces) are correlated with preformed complex primordia and the absence of a hypanthium. The inception of a ringwall is responsible for a higher complexity in vasculature and for obscuring the identity of the trunk bundles. The presence of a hypanthium does not permit trunk bundle formation because stamens develop on a limited section of the hypanthial slope. In that case the vasculature is characteristically represented by few large bundles linked with the perianth and the androecium. The distribution of complex polyandry is plotted on a Dahlgrenogram and presented in a table. The systematic relevance of complex polyandry is limited to the lower taxonomic (e.g. familial) levels.     

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．     

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: Neodregeae

The structure of the carpel has been studied in flowers of the Neodregeae ( Dipidax and Neodregea ). Except in D. triquetra , which is syncarpous, the carpels are united below and free above. A dorsal bundle, two or more lateral bundles, and two placental bundles supply each multiovulate carpel. The six placental bundles of the tricarpellate pistil are united by twos in the lower part of the pistil, forming three opposite compound placental bundles in most species of Dipidax and three alternate bundles in D. triquetra and Neodregea : In the latter, an additional septal bundle continues upward as a branch from the compound placental bundle. Sutural openings are usually short and restricted to the top of the locule. All the Neodregeae have monotegmic ovules.     

FLORAL DEVELOPMENT AND VASCULATURE IN HYDROCLEIS NYMPHOIDES (BUTOMACEAE)

The flower of Hydrocleis nymphoides consists of three sepals which arise in spiral succession, three simultaneously arising petals, numerous stamens and staminodia which arise in centrifugal order, and six carpels. A residual apex remains at maturity. The first-formed members of the androecium are stamens and the later-formed members are staminodia which develop below the stamens and which become outwardly displaced during expansion of the receptacle. The androecium is supplied by branching vascular trunk bundles. The carpels are completely open but the ventral margins are slightly conduplicately appressed basally. A single dorsal bundle provides the stigmatic area with vascular tissue, and a network of small placental bundles supplies the numerous laminar ovules. There are no clearly defined ventral bundles. It is suggested that Hydrocleis nymphoides is neither the most primitive nor the most advanced member of the family. A pattern of phylogenetic reduction in the androecium and receptacle is suggested for the entire family.     

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: 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.     

Inflated trichornes in flowers of Bauhinia (Leguminosae: Caesalpinioideae)

TUCKER, S. C., RUGENSTEIN, S. R. & DERSTINE, K. S., 1984. Inflated trichomes in flowers of Bauhinia (Leguminosae: Caesalpinioideae). An unusual kind of bifid multicellular inflated trichome is reported to occur on floral organs of II species of Bauhinia (Leguminosae-Caesalpinioideae), They occur on sepals, petals, stamens, carpels, receptacle, leaves, and stems, but not on all organs in each species. Their development in B. malabarica is described.     

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.     

FLORAL ONTOGENY OF ILLICIUM FLORIDANUM,WITH EMPHASIS ON STAMEN AND CARPEL DEVELOPMENT

The ontogeny of the flower and fruit of Illicium floridanum Ellis, the Star Anise, was investigated. Each of 5 or 6 bracts in each mixed terminal bud subtends either a vegetative or floral bud. The solitary flowers occur in terminal or axillary positions. Each flower has 3–6 subtending bracteoles arranged in a clockwise helix. The flowers in our material have 24–28 tepals, 30–39 stamens, and usually 13 (rarely 19) uniovulate carpels. Tepals and stamens are initiated in a low-pitched helix; carpels later appear whorled, but arise successively at different levels on the apical flanks. The floral apex is high-convex in outline with a tunica-corpus configuration; it increases in height and width throughout initiation of the floral appendages. Tepals, stamens, and carpels are initiated by one to several periclinal divisions in the subsurface layers low on the apical flanks, augmented by cell divisions in the outer layers of the corpus. The carpel develops as a conduplicate structure with appressed, connivent margins. Procambium development of floral appendages is acropetal and continuous. Bracteoles, tepals, stamens and carpels are each supplied by 1 trace; the carpellary trace splits into a dorsal and an ascending ventral sympodium. The latter bifurcates to form 2 ventral bundles. The ovular bundle diverges from the ventral sympodium. Ovule initiation occurs in a median axillary position to the carpel, an unusual type of ovule initiation. The fruit vasculature is greatly amplified as the receptacle and follicles enlarge. After carpel initiation an apical residuum persists which is not vascularized; a plate meristem develops over its surface to produce a papillate structure.     

Development of the gametophytes, flower, and floral vasculature in Cochliostema odoratissimum (Commelinaceae)

Flowers of Cochliostema odoratissimum are trimerous with three fertile stamens, three unequal antherless staminodes. and three connate carpels. The fertile stamens are on one side of die flower and united by their filaments, forming a compound structure that curves to the flower's right as the flower opens. The thecae arc longitudinally dehiscent, spirally coiled, and enveloped by pctaloid extensions of the filaments of the two lateral stamens contributing to the three-staminate structure. Anther wall development is of the monocotyledonous type. Tapetal raphides are formed and appear to be widespread in CCommelinaceae. Also known from Philydraceae and. perhaps. Haemodoraccae, tapetal raphides and their taxonomic distribution may be of phylogenetic utilitv. Microsporogcnesis is successive, forming both isobilateral and decussate tetrads. Pollen is shed as single binucleatc grains. Each ovary locule contains ten to twelve hemianatropous, crassinucellar. bitegmic ovules on axile placentae. The micropyle is formed by both the inner integument and one side of the outer integument. Megagamctophyte development is of the Polygonum type. The mature megagamctophyte consists of an egg apparatus, fusion nucleus, and three antipodals. the latter showing signs of degeneration. The salient features of the floral vasculature are the same as in the few other commclinaceous species for winch complete data are available. Relative to the floral vasculature in the other species, differences in the vasculature lie primarily in the presence and origin of lateral carpel bundles and in the number of sepal and ovule traces.     

Canrightiopsis,a new Early Cretaceous fossil with Clavatipollenites-type pollen bridge the gap between extinct Canrightia and extant Chloranthaceae

Canrightiopsis with three species (C. intermedia, C. crassitesta, C. dinisii) is described from the Early Cretaceous of Portugal based on small, one-seeded berries. The fruits are derived from bisexual flowers with three stamens borne on one side of the ovary. There are no traces of a perianth. Pollen is of the Clavatipollenites-type, monocolpate, semitectate, reticulate-columellate with heterobrochate reticulum and muri with beaded supratectal ornamentation. The ovary is unilocular with a single pendant, orthotropous and bitegmic ovule. The seed is endotestal. The endotesta consists of one layer of palisade-shaped crystal cells with fibrous infillings. The fruit wall has resin bodies or cavities from presumed ethereal oil cells sometimes seen as stomata-like structures on the fruit surface. A phylogenetic analysis resolves Canrightiopsis as a close relative of extant Chloranthaceae, particularly close to extant Chloranthus and Sarcandra. All three taxa share the one-sided position of the stamens on the ovary. An evolutionary sequence from fossil Canrightia to fossil Canrightiopsis and extant Chloranthus and Sarcandra is suggested by loss of perianth, reduction in number of ovules and stamens and displacement of stamens to one side of the ovary. Canrightiopsis also shares several critical features with extant Ascarina including monoaperturate pollen and beaded supratectal ornamentation of the pollen wall.