Morphology and development of anthers and ovules in Croton and Astraea (Euphorbiaceae)

We examined the embryological development of anthers and ovules from Astraea (A. lobata and A. praetervisa) and Croton (C. floribundus, C. fuscescens, C. glandulosus, C. lundianus, C. piptocalyx, C. urucurana and C. triqueter) focusing on features with systematic significance for the group. Some of these features are common in Euphorbiaceae including: a dicotyledonous type of anther wall formation, a secretory tapetum, a mixed origin of the outer ovule integument, an epidermal origin of the inner ovule integument, the occurrence of many archesporial cells inside the ovules and a megagametophyte of the Polygonum‐type. Other features, such as the presence of styloid crystals in the tapetum, an idioblast with a druse in the endothecium, simultaneous and successive microsporogenesis, and a functional micropylar megaspore, have not previously been reported in the family. These characters appear to distinguish Croton and Astraea from other Euphorbiaceae and possibly represents autapomorphies for the tribe Crotoneae.     

Androecium and floral nectaries ofHarungana madagascariensis (Clusiaceae)

The mature flower ofHarungana madagascariensis (Choisy)Poir. has an androecium of five antipetalous fascicles, consisting of four stamens each. The stamen fascicles alternate with five indented nectary scales. A SEM-study of the floral development, as well as a study of the floral anatomy was carried out to understand whether the nectariferous scales represent staminodia or are receptacular in nature and consequently whether or not the androecium ofHarungana, and theClusiaceae in general, is originally diplostemonous. The five petals originate by the splitting of petal-stamen complexes. Next the upper part of each complex differentiates basipetally in four stamens. The stamens remain fascicled and are lifted on a long stalk at maturity. Five carpel primordia are initiated united in a low ringwall. The five nectary scales appear after carpel inception and develop an external morphology reminiscent of anthers. The floral anatomy reveals an independent origin of sepal median traces and common sepal lateral traces, free petal traces, stamen fascicle traces and alternating vascular tissue which supplies the nectaries. The petal-stamen complexes are the result of a retardation in petal inception, linked with the absorption of petal tissue into the stamen primordia. The development of the stamen fascicles is discussed; it is suggested that they are of a secondary nature and do not appear as a reduction from a multistaminate androecium. The external morphology and vascular anatomy of the scales speaks in favour of a staminodial nature. The comparison with some other species of theClusiaceae gives evidence of a diplostemonous ancestry of the androecium.     

Anatomy,ultrastructure, and secretory activity of the floral nectaries in Swietenia macrophylla (Meliaceae)

• Premise of the study: While mahogany (Swietenia macrophylla) is one of the most important forest species in the Amazon region, little is known about its reproductive biology. Knowledge about the nectary structure and dynamics of nectar production of this species represent a key step toward understanding its relationship with pollinators. • Methods: Mahogany tree floral buds and flowers in anthesis were collected, fixed, and processed for study by light and transmission and scanning electron microscopy. The chemical composition of nectar and the nectary pigments was also studied. • Key results: Both staminate and pistillate flowers have nectaries, which contain a papillose epidermis and stomata. The nectariferous tissue is parenchymatous, with the cell cytoplasm primarily containing mitochondria and plastids. Secretory activity initiates at the beginning of anthesis, which occurs at nightfall. Flowers undergoing anthesis become structurally modified, with starch grains in the plastids disappearing. The number of plastoglobuli in the plastids also increases when nectaries change color from pale yellow to intense red. Pistillate and staminate flowers produce meager nectar rewards. • Conclusions: Changes in plastoglobuli number seem to be related to an increase in carotenes and color changes during anthesis. Carotenes can be linked to the protection of the plant against oxidative stress, which results from secretory activities. Nectary color has a limited role as a pollinator attractant. Floral rewards comprise small nectar droplets in both flower types, in addition to a few pollen grains in staminate flowers. These meager rewards are probably adapted to attract small generalist insects.     

Anatomy of the floral nectaries of some neotropical Salacioideae (Celastraceae)

Floral nectaries have contributed to the systematics of different taxonomic groups. Since those of the neotropical genera included in subfamily Salacioideae—Cheiloclinium Miers, Peritassa Miers, Salacia L. and Tontelea Aubl.—have different forms and positions, we explored their anatomy to delimit more precisely the genera of subfamily Salacioideae. Buds and open flowers of six species were treated following the usual techniques in plant anatomy. The obtained data were helpful in characterizing the floral nectary anatomy of the studied species. Furthermore, some features such as form, position and surface of nectaries; form of their epidermal cells; presence and distribution of stomata; occurrence of idioblasts containing druses in the nectariferous parenchyma; and absence of nectary vascularization can contribute to the taxonomy and phylogeny of the Salacioideae studied. In most of the studied species the nectar is probably released by both the stomata and the nectary epidermal surface. In Cheiloclinium cognatum, the structure acknowledged as nectary is actually a vestigial tissue and the functions of attracting and rewarding pollinators has phylogenetically migrated to the stigmatic region. The druses and phenolic substances observed in the nectariferous parenchyma probably help defend flowers against herbivore attacks. The minute size of the nectaries of Salacioideae may explain the absence of vascularization. The floral nectaries of Salacia elliptica are epithelial while those of the other species are mesenchymal.     

Characteristics of floral nectaries and nectar in two species of Crataegus (Rosaceae)

 The structure of floral nectaries of Crataegus coccinea and C. crus-galli was examined using light and scanning electron microscopy. The radial length of the floral nectary, measured from longitudinal sections of flowers, was 30% larger in C. crus-galli than in C. coccinea. For both Crataegus species the glandular tissue thickness was similar – approx. 400 μm. Also, the number of stomata per mm² of nectary surface in C. crus-galli was much higher (by 43%) than for C. coccinea. Stomata were situated in deep hollows. For both taxa the period of nectar secretion was 4 days. The mean quantity of total sugar in nectar per 10 flowers of C. crus-galli and C. coccinea was 3.87 mg and 0.33 mg, respectively. Received August 28, 2002; accepted December 17, 2002 Published online: June 2, 2003     

Micromorphology and ultrastructure of the floral nectaries of Polemonium caeruleum L. (Polemoniaceae)

The Polemoniaceae family forms flowers diverse in the terms of pollination methods and nectar types. The micromorphology of the nectary surface and the tissue structures as well as the ultrastructure of the cells of the floral nectaries in Polemonium caeruleum L. were examined using light, scanning and transmission electron microscopy. A bowl-shaped nectary, detached from the ovary, grows at its base. Its contour shows folds with depressions in the places where the stamens grow, forming five-lobed disc (synapomorphic character). Nectar is secreted through modified anomocytic stomata, which are formed in the epidermis covering the tip and the lateral wall of the projection located between the staminal filaments. The undulate nectary consists of a single-layered epidermis and three to nine layers of parenchymal cells. The cells of the nectary contain a dense cytoplasm, numerous plastids with an osmophilic stroma and starch grains, well-developed endoplasmic reticulum, as well as a large number of mitochondria interacting with the Golgi bodies. The ultrastructure of nectary cells indicates the granulocrine secretion mechanism and diversified transport of nectar.     

Structure,ultrastructure and evolution of floral nectaries in the twinflower tribe Linnaeeae and related taxa (Caprifoliaceae)

Linnaeeae is a small tribe of Caprifoliaceae consisting of six genera and c. 20 species. In Linnaeeae, floral nectaries are located on the corolla‐filament‐tube and nectar is produced from unicellular glandular hairs. We studied 23 taxa using scanning electron microscopy (SEM), light microscopy (LM) and transmission electron microscopy (TEM) and found two distinct nectary morphologies, zonate and gibbous types, and two distinct types of glandular hair, clavate and smooth base types. Plesiomorphic characters associated with the nectary and identified in the tribe include hypocrateriform corollas, dichogamous flowers, zonate nectaries, wet papillate stigmas, vestigial nectary disc and smooth pollen grains. Apomorphic characters include bilabiate corollas, homogamous flowers, bulging nectaries, dry papillate stigmas and echinulate pollen grains. The nectary structure is similar in Vesalea and Linnaea and differs from the rest of the tribe, in accordance with recent phylogenetic results. Nectar secretion is typically granulocrine with subcuticular accumulation of nectar, which we compared with the secretion in multicellular hairs of Adoxa moschatellina. The cuticle on the hair becomes detached from the cell wall and large subcuticular spaces filled with nectar are formed. Nectar is probably released in areas with a thin cuticle. In Zabelia, the smooth basal part of the hair could help to build up the hydrostatic pressure.     

The role of post‐floral persistent perianth in Helleborus viridis subsp. occidentalis (Ranunculaceae)

Species within the genus Helleborus differ in the relative location of their sepals. Previous studies have proved the existence of post‐floral functionality of sepals in H. foetidus, which leads us to consider the possibility of differences in the functionality of the sepals in H. viridis subsp. occidentalis. In this study, we analyzed their influence on the number and weight of seeds through experimental manipulations, which involve a progressive reduction of sepals following fertilization. Our results show that different levels of perianth reduction have no effect on the number and weight of seeds in Helleborus viridis subsp. occidentalis. We propose that differences in the timing of leaf development and changes in the position of floral organs among the different Helleborus species, underlie a distinctive response to the ever‐changing weather conditions of the European winters and springs.     

紫苏花蜜腺的发育解剖学研究

紫苏花蜜腺位于不均等分裂的花盘裂片上,属于子房基部的盘状蜜腺。3枚小裂片上的蜜腺由分泌表皮和产蜜组织组成,而另一枚大裂片上的指状蜜腺则由分泌表皮、产蜜组织和维管束组成。4枚蜜腺的表皮细胞外均具薄的角质层,仅在指状蜜腺的顶部分布着密集的气孔器。蜜腺来源于花盘表面的2～3层细胞。在蜜腺发育过程中,液泡和淀粉粒呈现有规律的消长变化,这与蜜汁的合成与分泌有关。3枚小裂片蜜腺的原蜜汁来源和泌蜜途径与指状蜜腺不同。     

Morphology and development of floral features recognised by pollinators

The diversity of angiosperm flowers is astounding. The conventional explanation for this diversity is that it represents the great variety of ways in which flowers have adapted to attract an even greater diversity of animal pollinators. Many animal behaviourists are therefore interested in how changes in floral morphology affect pollinator behaviour. The establishment of well-characterised model plant species has greatly furthered our understanding of how floral morphology is generated and varied. Many of these model species are pollinated by animals and attract their pollinators through the production of colour, shape, scent, size and rewards. An understanding of the developmental plasticity of floral morphology, and the constraints upon it, should inform research into animal responses to flowers. The use of genetically characterised model species, and the isogenic and near-isogenic lines available in them, will allow dissection of the different components of floral attraction and reward in natural systems. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: Heikki Hokkanen     

葡萄两性花花蜜腺的发育解剖学研究

葡萄两性花的花蜜腺位于子房基部的花盘上,共5枚,呈椭圆形,与雄蕊相间排列,属于花盘蜜腺,蜜腺由表皮和泌蜜组织组成,缺乏维管束,表皮具薄的角质层,无气孔器,蜜腺原基由子房基部表层细胞恢复分裂能力形成,在蜜腺发育过程中,泌蜜组织的液泡规律 性变化和多糖动态变化均不明显,原蜜汁由子房维管束的韧皮部提供,蜜汁通过表皮细胞排出。     

The systematic significance of floral morphology,nectaries, and nectar concentration in epiphytic cacti of tribes Hylocereeae and Rhipsalideae (Cactaceae)

A long-standing interest in cactus taxonomy has existed since the Linnaean generation, but an appreciation of the reproductive biology of cacti started early in the 1900s. Numerous studies indicate that plant reproductive traits provide valuable systematic information. Despite the extensive reproductive versatility and specializations in breeding systems coupled with the striking floral shapes, the reproductive biology of the Cactaceae has been investigated in approximately 10% of its species. Hence, the systematic value of architectural design and organization of internal floral parts has remained virtually unexplored in the family. This study represents the most extensive survey of flower and nectary morphology in the Cactaceae focusing on tribes Hylocereeae and Rhipsalideae (subfamily Cactoideae). Our objectives were (1) to conduct comparative morphological analyses of flowers and floral nectaries and (2) to compare nectar solute concentration in these two tribes consisting of holo- and semi-epiphytic species. Flower morphology, nectary types, and sugar concentration of nectar have strong taxonomic implications at the tribal, generic and specific levels. Foremost, three types of nectaries were found, namely chamber nectary (with the open and diffuse subtypes), furrow nectary (including the holder nectary subtype), and annular nectary. All Hylocereeae species possess chamber nectaries, in which the nectarial tissue has both trichomes and stomata. The Rhipsalideae are distinguished by two kinds of floral nectaries: furrow and annular, both nectary types with stomata only. The annular nectary type characterizes the genus Rhipsalis. Nectar concentration is another significant taxonomic indicator separating the Hylocereeae and Rhipsalideae and establishing trends linked to nectar sugar concentration and amount of nectar production in relation to flower size. There is an inverse relationship between flower size and amount of nectar production in the smaller Rhipsalideae flowers, in which nectar concentration is more than two-fold higher despite the smaller volume of nectar produced when compared to the large Hylocereeae flowers. Variability of nectary morphology and nectar concentration was also evaluated as potential synapomorphic characters in recent phylogenies of these tribes. In conclusion, our data provide strong evidence of the systematic value of floral nectaries and nectar sugar concentration in the Cactaceae, particularly at different taxonomic levels in the Hylocereeae and Rhipsalideae.     

Morphology and development of floral shoots and organs in certain Zannichelliaeeae

Development of reproductive shoots and associated organs in Vleisia aschersoniana, Althenia filiformis, Lepilaena bilocularis and L. cylindrocarpa (Zannichelliaeeae, sensu Dumortier) has been examined using an epi-illumination technique to provide photographic documentation. Floral shoots share a similar basic developmental pattern. The vegetative shoot is terminated by a unisexual flower, but growth is continued from the axils of leaves immediately below so that fairly regular sympodia develop. Vleisia is most variable in the expression of this pattern. The flowers are simple, consisting of a single stamen or three carpels (one in Vleisia) and show marked similarity in development. Short scale-like appendages, reminiscent of a perianth, develop at the base of the stamen in Althenia and Lepilaena. An outgrowth at the tip of the connective in Lepilaena bilocularis and two at its middle part in Vleisia are initiated at late stages of stamen development. Carpels are subtended by membranous tepal-like appendages that are initiated at the same time as the carpel primordia. Each carpel primordium becomes peltate and develops a bitegmic ovule on the adaxial portion of the carpel wall which in turn overgrows the ovule and ultimately forms a long thin style with either a funnel-shaped (Vleisia, L. cylindrocarpa) , peltate (L. bilocularis) or feathery (Althenia) stigma. Relationships with other Alismatales are discussed.     

芭蕉科花蜜腺形态比较：兼论其与传粉者的关系（英文）

对狭义芭蕉科3个属的代表性种芭蕉(Musa basjoo)、象腿蕉(Ensete glaucum)和地涌金莲(Musella lasiocarpa)的花蜜腺形态进行了比较研究。结果表明它们的蜜腺属于隔膜蜜腺。雌花的蜜腺着生于子房的上部,胚珠的上方;雄花蜜腺占据了整个败育子房的位置。蜜腺结构由许多腔道组成,这些腔道在横切面上呈现出复杂的发散式迷宫状结构。这3种植物花蜜腺的栅栏状表皮细胞、维管束和蜜腺开口方式相似,而从纵切面和横切面上观察其结构存在一些差异。PAS反应显示象腿蕉泌蜜组织中淀粉粒含量高于其他两个种;芭蕉和象腿蕉的蜜腺腔里有许多纤维状物质存在。3种植物的传粉综合征多样化,花序和花的特征(如花序下垂或直立、苞片的颜色、泌蜜量和泌蜜时间等)和传粉样式之间有密切关系。它们的蜜腺结构和传粉者行为之间没有明显的相关性,但是胶质或水质的花蜜对传粉者的取食方式有一定影响。     

芭蕉科花蜜腺形态比较: 兼论其与传粉者的关系

对狭义芭蕉科3个属的代表性种芭蕉(Musa basjoo)、象腿蕉(Ensete glaucum)和地涌金莲(Musella lasiocarpa)的花蜜腺形态进行了比较研究。结果表明它们的蜜腺属于隔膜蜜腺。雌花的蜜腺着生于子房的上部, 胚珠的上方; 雄花蜜腺占据了整个败育子房的位置。蜜腺结构由许多腔道组成, 这些腔道在横切面上呈现出复杂的发散式迷宫状结构。这3种植物花蜜腺的栅栏状表皮细胞、维管束和蜜腺开口方式相似, 而从纵切面和横切面上观察其结构存在一些差异。PAS反应显示象腿蕉泌蜜组织中淀粉粒含量高于其他两个种; 芭蕉和象腿蕉的蜜腺腔里有许多纤维状物质存在。3种植物的传粉综合征多样化, 花序和花的特征(如花序下垂或直立、苞片的颜色、泌蜜量和泌蜜时间等)和传粉样式之间有密切关系。它们的蜜腺结构和传粉者行为之间没有明显的相关性, 但是胶质或水质的花蜜对传粉者的取食方式有一定影响。     

Evolution of perianth and stamen characteristics with respect to floral symmetry in Ranunculales

BACKGROUND AND AIMS: Floral symmetry presents two main states in angiosperms, namely polysymmetry and monosymmetry. Monosymmetry is thought to have evolved several times independently from polysymmetry, possibly in co-adaptation with specialized pollinators. Monosymmetry commonly refers to the perianth, even though associated androecium modifications have been reported. The evolution of perianth symmetry is examined with respect to traits of flower architecture in the Ranunculales, the sister group to all other eudicots, which present a large diversity of floral forms. METHODS: Characters considered were perianth merism, calyx, corolla and androecium symmetry, number of stamens and spurs. Character evolution was optimized on a composite phylogenetic tree of Ranunculales using maximum parsimony. KEY RESULTS: The ancestral state for merism could not be inferred because the basalmost Eupteleaceae lack a perianth and have a variable number of stamens. The Papaveraceae are dimerous, and the five other families share a common trimerous ancestor. Shifts from trimery to dimery (or reverse) are observed. Pentamery evolved in Ranunculaceae. Ranunculales except Eupteleaceae, present a polysymmetric ancestral state. Monosymmetry evolved once within Papaveraceae, Ranunculaceae and Menispermaceae (female flowers only). Oligandry is the ancestral state for all Ranunculales, and polyandry evolved several times independently, in Papaveraceae, Menispermaceae, Berberidaceae and Ranunculaceae, with two reversions to oligandry in the latter. The ancestral state for androecium symmetry is ambiguous for the Ranunculales, while polysymmetry evolved immediately after the divergence of Eupteleaceae. A disymmetric androecium evolved in Papaveraceae. The ancestral state for spurs is none. Multiple spurs evolved in Papaveraceae, Berberidaceae and Ranunculaceae, and single spurs occur in Papaveraceae and Ranunculaceae. CONCLUSIONS: The evolution of symmetry appears disconnected from changes in merism and stamen number, although monosymmetry never evolved in the context of an open ground plan. In bisexual species, monosymmetry evolved coincidently with single spurs, allowing us to propose an evolutionary scenario for Papaveraceae.     

Comparative floral development in Lardizabalaceae (Ranunculales)

Lardizabalaceae, one of seven families of Ranunculales, represent a monophyletic group. The family has functionally unisexual flowers with the organs in trimerous whorls, petaloid sepals and sometimes nectariferous petals. Among Ranunculales, Lardizabalaceae share several floral characters and climbing habit with Menispermaceae, but molecular analyses indicate that Circaeasteraceae and Lardizabalaceae form a strongly supported clade. Morphological and ontogenetic studies of flowers have proved to be a good complement to molecular data in clarifying relationships. Floral organogenesis has been studied in very few species of the family. This study investigates the comparative floral development of three species from three genera (Decaisnea, Akebia and Holboellia) of Lardizabalaceae using scanning electron microscopy. Flowers have a whorled phyllotaxis. Within each whorl, the organs are initiated either simultaneously or in a rapid spiral sequence. In Akebia, six sepals are initiated, but one to three sepals of the second whorl do not further develop. The presence of three sepals in Akebia is thus a developmentally secondary simplification. The petals (if present) are retarded in early developmental stages; stamens and petals are different in shape from the beginning of development. The retarded petals may not be derived from staminodes in Lardizabalaceae. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 166 , 171–184.     

Patterns and development of floral asymmetry in Senna (Leguminosae, Cassiinae)

The buzz-pollinated genus Senna (Leguminosae) is outstanding for including species with monosymmetric flowers and species with diverse asymmetric, enantiomorphic (enantiostylous) flowers. To recognize patterns of homology, we dissected the floral symmetry character complex and explored corolla morphology in 60 Senna species and studied floral development of four enantiomorphic species. The asymmetry morph of a flower is correlated with the direction of spiral calyx aestivation. We recognized five patterns of floral asymmetry, resulting from different combinations of six structural elements: deflection of the carpel, deflection of the median abaxial stamen, deflection or modification in size of one lateral abaxial stamen, and modification in shape and size of one or both lower petals. Prominent corolla asymmetry begins in the earl-stage bud (unequal development of lower petals). Androecium asymmetry begins either in the midstage bud (unequal development of thecae in median abaxial stamen; twisting of androecium) or at anthesis (stamen deflection). Gynoecium asymmetry begins in early bud (primordium off the median plane, ventral slit laterally oriented) or midstage to late bud (carpel deflection). In enantiostylous flowers, pronouncedly concave and robust petals of both monosymmetric and asymmetric corollas likely function to ricochet and direct pollen flow during buzz pollination. Occurrence of particular combinations of structural elements of floral symmetry in the subclades is shown.     

3种单子叶蜜源植物花蜜腺的发育解剖学研究

通过解剖镜、扫描电镜观察和石蜡制片等方法对韭菜、萱草和鸢尾的花蜜腺进行了系统研究。结果表明,它们都属于子房蜜腺,其中韭菜为典型的隔膜蜜腺,萱草为非典型的隔膜蜜腺,而鸢尾为心皮边缘蜜腺。其结构都由分泌表皮和产蜜组织构成,萱草蜜腺中含有维管束。三者在开花前后,蜜腺组织中液泡大小都发生有规律变化,蛋白质含量也都发生有规律变化,而淀粉仅在鸢尾蜜腺中有少量积累。由于三者花蜜腺的结构存在一定差异,其蜜汁泌出的途径也不相同。