Floral organogenesis of Chloranthus sessilifolius, with special emphasis on the morphological nature of the androecium of Chloranthus (Chloranthaceae)

 Floral organogenesis of Chloranthus sessilifolius K. F. Wu is described. The inflorescence primordium is dome-like in the beginning and then elongates, and bract primordia initiate almost decussately. Each floral primordium, arising from the axil of a bract, soon becomes a scale-like structure, with three primordia of androecial lobes originating from its abaxial part, and the gynoecial primordium in adaxial position. As the androecial lobes become more distinct, four thecae are already in differentiation, and the gynoecial primordium appears as a shallow disc. The androecial lobes do not extend their length until the thecae approach maturity and the stigma is differentiated. The androecial lobes are united at all the stages of development, and the entire androecium falls off as a unit at the end of anthesis. Based on these results, combined with published evidence from neobotany, palaeobotany and phylogenetic studies, the morphological nature of the androecium of Chloranthus is further discussed. Our studies support the viewpoint that the androecial structure of Chloranthus may have arisen by splitting of a single stamen with 2 marginal thecae. Received May 2, 2001 Accepted December 18, 2001     

Oxypetalum banksii subsp. banksii: a taxon of Asclepiadaceae with an extragynoecial compitum

 The floral morphology of seven Oxypetalum species and, in particular, the spatial relationship between the five stigmatic chambers and two separate ovaries of their flowers with respect to transmission of the pollen tube are studied. In all species, except O. banksii subsp. banksii, floral morphology is similar to that in other Asclepiadeae, and the flowers pollinated with one pollinium develop only one follicle, which means compitum absence. In O. banksii subsp. banksii flowers, the secretory interstaminal tissue lines the inner walls of the stigmatic chambers as in the other species studied, but it also reaches the upper part of the inner surface of the filament tube, where it surrounds the styles, an unprecedented feature for Asclepiadaceae. This tissue secretes nectar and mucilage; the latter acts as transmitting medium for the growth of pollen tubes from pollinia inserted and hydrated in stigmatic chambers (“hyperstigmas”). Mucilage also functions as an extragynoecial compitum: in flowers pollinated with one pollinium both carpels develop into a follicle. Received August 28, 2001; accepted April 9, 2002 Published online: October 14, 2002 Addresses of the authors: Milene Faria Vieira (e-mail: mfvieira@mail.ufv.br), Departamento de Biologia Vegetal, Universidade Federal de Vi?osa, 36571-000, Vi?osa, Minas Gerais, Brazil. George John Shepherd, Departamento de Botanica, Instituto de Biologia, Universidade Estadual de Campinas, C.P. 6109, 13083-970, Campinas, S?o Paulo, Brazil.     

Floral biology and pollinators of three co-occurring Cistus species (Gistaceae)

BOSCH, J., 1992. Floral biology and pollinators of three co-occurring Cistus species (Cistaceae). The pollination of three species of Cistus, Cistus albidus L., Cistus salvifolius L. and Cistus monspeliensis L. is studied. Insect visitors were censused and some were captured to analyse pollen distribution on their bodies. Floral phenology, breeding systems, some aspects of floral biology (pollen and nectar production, pollen-ovule ratios) and floral morphology were also studied for each species. Flowers of C. albidus produce more pollen and nectar than the other two species studied, and attract more insects, especially those with higher nutritional requirements. All three species are highly self-incompatible, but C. salvifolius and C. monspeliensis show some features that hinder xenogamy, and they have lower pollen-ovule ratios than C. albidus. As a result, this latter species apparently receives higher levels of cross-pollination, but at the cost of a higher investment to attract and reward reliable pollinators.     

Understanding plant reproductive diversity

Flowering plants display spectacular floral diversity and a bewildering array of reproductive adaptations that promote mating, particularly outbreeding. A striking feature of this diversity is that related species often differ in pollination and mating systems, and intraspecific variation in sexual traits is not unusual, especially among herbaceous plants. This variation provides opportunities for evolutionary biologists to link micro-evolutionary processes to the macro-evolutionary patterns that are evident within lineages. Here, I provide some personal reflections on recent progress in our understanding of the ecology and evolution of plant reproductive diversity. I begin with a brief historical sketch of the major developments in this field and then focus on three of the most significant evolutionary transitions in the reproductive biology of flowering plants: the pathway from outcrossing to predominant self-fertilization, the origin of separate sexes (females and males) from hermaphroditism and the shift from animal pollination to wind pollination. For each evolutionary transition, I consider what we have discovered and some of the problems that still remain unsolved. I conclude by discussing how new approaches might influence future research in plant reproductive biology.     

The sweet jelly of Combretum lanceolatum flowers (Combretaceae): a cornucopia resource for bird pollinators in the Pantanal, western Brazil

 The pollination biology of the neotropical scandent shrub Combretum lanceolatum was studied in the seasonally-flooded Pantanal region in western Brazil. This plant bears horizontally oriented inflorescences, whose yellowish green flowers begin to expand at dusk and are fully open at dawn. Instead of fluid nectar the flowers produce sweet gelatinous secretion in form of pellets. The glandular complex of the flower is composed of the inner wall of the receptacle and its tubular extension, being equivalent to the nectariferous disk of the nectar-producing species within the genus. The jelly is produced at night, contains mannan and is imbibed by free hexoses. It originates by swelling and disintegration of the inner wall, after contact with the nectar generated concomitantly in the mesophyll. Combretum lanceolatum is unique within the genus in its production of jelly pellets instead of liquid nectar. A new term, the jelly-flower, is proposed for flowers with this kind of reward. The pellet is not replaced once removed by a bird, and thus resembles a fruit in its availability to consumers, another unique feature that distinguishes this species within the genus. The jelly pellets offered by the many flowered branches attract a great diversity of bird visitors (28 species from eight families), which feed on this copious food resource and pollinate the flowers. The most effective pollinators probably are thrushes, tanagers, and orioles. Flocking parakeets and macaws sometimes feed on the petals, thus acting as flower plunderers. Combretum lanceolatum presents a high fruit set under natural conditions, which likely favours its spreading and becoming a weed species. Received July 11, 2000 Accepted November 18, 2000     

The control of flower initiation by gibberellin in Helianthus annuus L. (sunflower), a non-photoperiodic plant

The involvement of gibberellins in the control of flowering of sunflower was studied by direct application of GA₃ to the apex of the plants, analysis of the endogenous levels of gibberellin-like substances at different plant ages, and indirectly by the application of paclobutrazol, an inhibitor of gibberellin synthesis. GA₃ speeded-up flower initiation and floral apex development. The time of GA₃ application was more critical than the amount of GA₃ applied. The endogenous levels of gibberellin-like compounds increased significantly by day 15 after sowing. The application of paclobutrazol markedly delayed floral initiation and this effect was also depedent on plant age. Both GA₃ and paclobutrazol had their greatest effects between 10 and 20 days after sowing suggesting that an increase in gibberellins in that time period plays a role in floral initiation.     

Reproductive biology and pollinators in two enantiostylous Qualea species (Vochysiaceae) in the Brazilian Cerrado

• Enantiostyly is a floral polymorphism in which two floral forms in the same species differ in deflection of the stigma to right or left position. In monomorphic enantiostylous plants, flowers of the two morphs occur within the same individual, usually in the same proportion. In self‐compatible species the function of monomorphic enantiostyly is proposed to increase outcrossing rates and offer a reproductive advantage under pollination limitation. Enantiostylous species are usually self‐compatible and show heteranthery, with poricide anthers and pollen as pollinator reward; however, there are families, such as Vochysiaceae, that have different characteristics.
• We analysed the reproductive system and pollination biology of Qualea parviflora and Q. multiflora, two enantiostylous species from the Brazilian Cerrado that have specific morphological and physiological traits. For this, we characterized flower traits, performed hand pollinations and studied floral visitors.
• We found no differences between morphs in the proportion of flowers, nectar produced or its concentration, pollen quantity and fruit set. Both species were self‐incompatible and quite generalist regarding floral visitors.
• Enantiostyly in self‐incompatible plants seems to confer a reproductive advantage by reducing self‐interference resulting from stigma clogging. This novel result helps to expand our knowledge on this complex floral polymorphism and opens new avenues for future research on this topic.

     

Current trends and future directions in flower development research

Flowers, the reproductive structures of the approximately 400 000 extant species of flowering plants, exist in a tremendous range of forms and sizes, mainly due to developmental differences involving the number, arrangement, size and form of the floral organs of which they consist. However, this tremendous diversity is underpinned by a surprisingly robust basic floral structure in which a central group of carpels forms on an axis of determinate growth, almost invariably surrounded by two successive zones containing stamens and perianth organs, respectively. Over the last 25 years, remarkable progress has been achieved in describing the molecular mechanisms that control almost all aspects of flower development, from the phase change that initiates flowering to the final production of fruits and seeds. However, this work has been performed almost exclusively in a small number of eudicot model species, chief among which is Arabidopsis thaliana. Studies of flower development must now be extended to a much wider phylogenetic range of flowering plants and, indeed, to their closest living relatives, the gymnosperms. Studies of further, more wide-ranging models should provide insights that, for various reasons, cannot be obtained by studying the major existing models alone. The use of further models should also help to explain how the first flowering plants evolved from an unknown, although presumably gymnosperm-like ancestor, and rapidly diversified to become the largest major plant group and to dominate the terrestrial flora. The benefits for society of a thorough understanding of flower development are self-evident, as human life depends to a large extent on flowering plants and on the fruits and seeds they produce. In this preface to the Special Issue, we introduce eleven articles on flower development, representing work in both established and further models, including gymnosperms. We also present some of our own views on current trends and future directions of the flower development field.     

SPATIAL FLOWER PARAMETERS AND INSECT SPATIAL VISION

The present article reviews recent and older literature on the spatial parameters that flowers display, as well as on the capacities of anthophilous insects to perceive and use these parameters for optimizing their foraging success. Although co-evolution of plants and pollinators has frequently been discussed with respect to floral colours and insect colour vision, it has rarely been assessed with respect to insect spatial vision and spatial floral cues, such as shape, pattern, size, contrast, symmetry, spatial frequency, contour density and orientation of contours. This review is an attempt to fill this gap. From experimental findings and observations on both flowers and insects, we arrive at the conclusion that all of the spatial and spatio-temporal parameters that flowers offer are relevant to the foraging task and are tuned to the insect's visual capacities and visually guided behaviour. We try, in addition, to indicate that temporal cues are closely related to spatial cues, and must therefore be included when flower–pollinator interactions are examined. We include results that show that colour vision and spatial vision have diverged over the course of evolution, particularly regarding the processing of spatio-temporal information, but that colour vision plays a role in the processing of spatial cues that are independent of temporal parameters. By presenting this review we hope to contribute to closer collaboration among scientists working in the vast fields of botany, ecology, evolution, ethology and sensory physiology.