Do extrafloral nectaries present a defensive role against herbivores in two species of the family Bignoniaceae in a Neotropical savannas?

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Leaf glands act as nectaries in Diplopterys pubipetala (Malpighiaceae)

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Nectaries and reproductive biology of Croton sarcopetalus (Euphorbiaceae)

Flower morphology, nectary structure, nectar chemical composition, breeding system, floral visitors and pollination were analysed in Croton sarcopetalus, a diclinous-monoecious shrub from Argentina. Male flowers have five receptacular nectaries, with no special vascular bundles, that consist of a uniserial epidermis with stomata subtended by a secretory parenchyma. Female flowers bear two different types of nectaries: inner (IN) and outer (ON) floral nectaries. IN, five in all, are structurally similar to the nectaries of male flowers. The five ON are vascularized, stalked, and composed of secretory, column-shaped epidermal cells without stomata subtended by secretory and ground parenchyma. In addition, ON act as post-floral nectaries secreting nectar during fruit ripening. Extrafloral nectaries (EFN) are located on petioles, stipules and leaf margins. Petiolar EFN are patelliform, stalked and anatomically similar to the ON of the female flower. Nectar sampled from all nectary types is hexose dominant, except for the ON of the female flower at the post-floral stage that is sucrose dominant. The species is self-compatible, but geitonogamous fertilization is rarely possible because male and female flowers are not usually open at the same time in the same individual, i.e. there is temporal dioecism. Flowers are visited by 22 insect species, wasps being the most important group of pollinators. No significant differences were found in fruit and seed set between natural and hand pollinated flowers. This pattern indicates that fruit production in this species is not pollen/pollinator limited and is mediated by a wide array of pollinators. (C) 2001 the Linnean Society of London.

Floral anatomy of Bromeliaceae, with particular reference to the evolution of epigyny and septal nectaries in commelinid monocots

Floral anatomy is described in ten genera of Bromeliaceae, including three members of subfamily Bromelioideae, three Tillandsioideae, and four genera of the polyphyletic subfamily Pitcairnioideae (including Brocchinia, the putatively basal genus of Bromeliaceae). Bromeliaceae are probably unique in the order Poales in possessing septal nectaries and epigynous or semi-epigynous flowers. Evidence presented here from floral ontogeny, vasculature, and the relative positions of nectary and ovules indicates that there could have been one or more reversals to apparent hypogyny in Bromeliaceae, although this hypothesis requires a better-resolved phylogeny. Such evolutionary reversals probably evolved in response to specialist pollinators, and in conjunction with other aspects of floral morphology of Bromeliaceae, such as the petal appendages of some species. The ovary is initiated in an inferior position even in semi-epigynous or hypogynous species. The ovary of all so-called hypogynous Bromeliaceae is actually semi-inferior, because the septal nectary is infralocular; in these species the nectaries have a labyrinthine surface and many vascular bundles. Brocchinia differs from most other fully epigynous species in that each carpel is secretory at the apex and reproductive, rather than secretory, at the base.

Morphological patterns of extrafloral nectaries in woody plant species of the Brazilian cerrado

Extrafloral nectaries are nectar-secreting structures that are especially common among the woody flora of the Brazilian cerrado, a savanna-like vegetation. In this study, we provide morphological and anatomical descriptions of extrafloral nectaries (EFNs) occurring on vegetative and reproductive organs of several plant species from the cerrado, and discuss their function and ecological relevance. We describe the morphology and anatomy of EFNs of 40 species belonging to 15 woody families using scanning electron microscopy and light microscopy. We categorise EFNs following a structural-topographical classification, and characterise the vascularised and complex nectaries, amorphous nectaries and secretory trichomes. Fabaceae, Bignoniaceae, Malpighiaceae and Vochysiaceae were the plant families with the majority of species having EFNs. Ten species possess more than one morphotype of gland structure. Observations and experimental field studies in the cerrado support the anti-herbivore role of EFN-gathering ants in this habitat. Additional morphological studies of EFNs-bearing plants, including other growth forms (e.g. herbs and lianas), are being undertaken and will hopefully cast further light on the ecological relevance of these glands in the cerrado, especially with respect to their attractiveness to multiple visitors.

A macro- and micromorphological survey of floral and extrafloral nectaries in the epiphytic cactus Rhipsalis teres (Cactoideae: Rhipsalideae)

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Morphology, development and homologies of the perianth and floral nectaries in Croton and Astraea (Euphorbiaceae-Malpighiales)

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

EXTRAFLORAL NECTARIES IN THE TROPICAL TREE GUAREA-MACROPHYLLA (MELIACEAE)

This paper describes the anatomy and morphology of the complex nectary systems of the tropical tree Guarea macrophylla Vahl (Meliaceae) and presents the first record of extrafloral nectaries occurring on reproductive organs (fruits) of a member of the order Sapindales. The extrafloral nectaries of G. macrophylla occur on petioles, petiolules, the abaxial surface of all leaflets, leaf buds, and over the surface of fruits. All extrafloral nectaries are distinctly raised above the surface. Foraging ants collect extrafloral nectar on Guarea trees both day and night. We suggest that the presence of extrafloral nectaries might be a useful taxonomic character for the identification of Guarea species.

Distribution of extrafloral nectaries in different vegetation types of Amazonian Brazil

Surveys were carried out in terra firme' forest, successional forest, buritirana' (palm vegetation) and shrub canga' (savanna). Extrafloral nectaries (EFNs) were present in 30 plant species belonging to 22 genera and 14 families. Nectary species represented 17.6-53.3% of the species samples in different areas, with local abundances varying from 19.1-50.0%. The percentage of species with EFNs was greater in the flora of the shrub canga than in the terra firme and successional forests. Nectary plants were more abundant in the shrub canga and successional forest. The high abundance of EFNs may be the result of intense foraging activity by ants on plants, leading to the formation of facultative mutualism. -from Authors

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. © 2012 Springer-Verlag Wien.

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. © 2013 Perspectives in Plant Ecology, Evolution and Systematics.

Anatomy of extrafloral nectaries in Fabaceae from dry-seasonal forest in Brazil

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Morfologia de nectários em Leguminosae senso lato em áreas de caatinga no Brasil

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

The floral nectary of Hymenaea stigonocarpa (Fabaceae, caesalpinioideae): Structural aspects during floral development

Background and Aims Considering that few studies on nectary anatomy and ultrastructure are available for chiropterophilous flowers and the importance of Hymenaea stigonocarpa in natural 'cerrado' communities, the present study sought to analyse the structure and cellular modifications that take place within its nectaries during the different stages of floral development, with special emphasis on plastid dynamics.Methods For the structural and ultrastructural studies the nectary was processed as per usual techniques and studied under light, scanning and transmission electron microscopy. Histochemical tests were employed to identify the main metabolites on nectary tissue and secretion samples.Key Results The floral nectary consists of the inner epidermis of the hypanthium and vascularized parenchyma. Some evidence indicates that the nectar release occurs via the stomata. The high populations of mitochondria, and their juxtaposition with amyloplasts, seem to be related to energy needs for starch hydrolysis. Among the alterations observed during the secretory phase, the reduction in the plastid stromatic density and starch grain size are highlighted. When the secretory stage begins, the plastid envelope disappears and a new membrane is formed, enclosing this region and giving rise to new vacuoles. After the secretory stage, cellular structures named 'extrastomatic bodies' were observed and seem to be related to the nectar resorption.Conclusions Starch hydrolysis contributes to nectar formation, in addition to the photosynthates derived directly from the phloem. In these nectaries, the secretion is an energy-requiring process. During the secretion stage, some plastids show starch grain hydrolysis and membrane rupture, and it was observed that the region previously occupied by this organelle continued to be reasonably well defined, and gave rise to new vacuoles. The extrastomatic bodies appear to be related to the resorption of uncollected nectar.

Intercellular pectic protuberances in Hymenaea stigonocarpa (Fabaceae, Caesalpinioideae): Occurrence and functional aspects

A study of the anatomy and ultrastructural aspects of leaf mesophyll and floral nectaries of Hymenaea stigonocarpa Mart. ex Hayne revealed the presence of intercellular pectic protuberances (IPPs) linking adjacent cells in both the leaf palisade cells and the secretory parenchyma of the floral nectary. Samples of the middle third of the leaf blade and of floral nectaries in anthesis were collected, fixed, and processed using standard procedures for light, transmission, and scanning electron microscopies. The IPPs of palisade cells of the mesophyll and the secretory parenchyma cells of the floral nectary take the form of scalae or strands, respectively. No evidence of the specific synthesis of these structures was observed, and they are apparently formed by the separation of adjacent cells due to cell expansion, when intercellular spaces develop. The IPPs observed in H. stigonocarpa increase cellular contact and probably act in apoplastic transport.