Transfer cells in trichomatous nectary in Adenocalymma magnificum (Bignoniaceae).

Transfer cells are specialized cells that play an important role where there are high energy costs due to facilitation of transmembrane flow of solutes. This study aimed to investigate the ontogenesis, histochemistry and ultrastructure of glandular trichomes focusing on stalk cells and their possible transfer function. Samples of prophylls of axillary buds, calyces of flowers in anthesis, and flower buds in different stages of development were collected, fixed and processed according to the common methods of microscopy. The glandular trichomes are composed of a secretory head with its cells in columnar format. The stalk is formed by two layers of cells, with the upper layer composed of cuboidal cells where the wall starts to thicken at the beginning of the pre-secretory phase. The secretion is heterogeneous, releasing glucose, other carbohydrates, lipids and phenolic compounds, with two types of release - eccrine and granulocrine. These trichomes are functionally termed as nectaries. The stalk cells appear as transfer cells since they have a thicker anticlinal wall with irregular ingrowths. The presence of transfer cells in the nectaries suggests a high specialization because it improves transport capacity of nectar and compensation in the high energy expenditure for its production and release.

Structure and distribution of glandular trichomes in three species of Bignoniaceae / Estrutura e distribuição de tricomas glandulares em três espécies da família Bignoniaceae

Glandular trichomes play a major role in the morphological characterization of the Bignoniaceae. Due to their great diversity of forms and functions, this study aimed to inventory the glandular trichomes present in the aerial vegetative axis of Amphilophium magnoliifolium, Martinella obovata and Stizophyllum riparium, analyze their structure and register the participation of ants in these plants. Fresh samples from the nodal region, petiole and from medium to apical regions of the leaflet blade were fixed and processed according to usual methods in light and scanning electron microscopies. The glandular trichomes found were: peltate, capitate, stipitate, and patelliform/cupular. Peltate trichomes are the most abundant ones and present the most uniform distribution. Patelliform/cupular trichomes occur at specific regions, such as prophylls, leaflet blade and nodal regions. Martinella obovata is the only species that presents capitate and stipitate trichomes, which are widely distributed along the entire aerial vegetative axis. Ants were found in all species, mainly at nodal regions. The occurrence of the capitate-type trichome is reported for the first time to the genus.(AU)

Tricomas glandulares desempenham papel importante na caracterização morfológica de Bignoniaceae e, devido à grande diversidade de formas e funções que esses tricomas apresentam, o objetivo desse estudo foi inventariar os tricomas glandulares presentes no eixo vegetativo aéreo de Amphilophium magnoliifolium, Martinella obovata. e Stizophyllum riparium, bem como analisar sua estrutura e registrar a participação de formigas nessas espécies. Amostras frescas da região mediana à apical das lâmina foliolar, pecíolo e nó foram fixadas e processadas de acordo com os métodos usuais para microscopia de luz e eletrônica de varredura. Os tricomas glandulares encontrados nas espécies foram: peltados, capitados, estipitados e pateliformes/cupuliformes. Dentre esses tipos o mais abundante foi o tricoma peltado, apresentando também uma distribuição mais uniforme que os demais. Os tricomas pateliformes/cupuliformes estavam em regiões mais específicas, como profilos, lâmina foliolar e na região nodal. Martinella obovata foi a única espécie que apresentou os tricomas capitados e estipitados, amplamente distribuídos ao longo de todo eixo vegetativo aéreo. Em todas as espécies foram encontradas formigas, principalmente nas regiões nodais. A ocorrência do tricoma capitado é relatada pela primeira vez para o gênero.(AU)

Structure and distribution of glandular trichomes in three species of Bignoniaceae / Estrutura e distribuição de tricomas glandulares em três espécies da família Bignoniaceae

Glandular trichomes play a major role in the morphological characterization of the Bignoniaceae. Due to their great diversity of forms and functions, this study aimed to inventory the glandular trichomes present in the aerial vegetative axis of Amphilophium magnoliifolium, Martinella obovata and Stizophyllum riparium, analyze their structure and register the participation of ants in these plants. Fresh samples from the nodal region, petiole and from medium to apical regions of the leaflet blade were fixed and processed according to usual methods in light and scanning electron microscopies. The glandular trichomes found were: peltate, capitate, stipitate, and patelliform/cupular. Peltate trichomes are the most abundant ones and present the most uniform distribution. Patelliform/cupular trichomes occur at specific regions, such as prophylls, leaflet blade and nodal regions. Martinella obovata is the only species that presents capitate and stipitate trichomes, which are widely distributed along the entire aerial vegetative axis. Ants were found in all species, mainly at nodal regions. The occurrence of the capitate-type trichome is reported for the first time to the genus.

Tricomas glandulares desempenham papel importante na caracterização morfológica de Bignoniaceae e, devido à grande diversidade de formas e funções que esses tricomas apresentam, o objetivo desse estudo foi inventariar os tricomas glandulares presentes no eixo vegetativo aéreo de Amphilophium magnoliifolium, Martinella obovata. e Stizophyllum riparium, bem como analisar sua estrutura e registrar a participação de formigas nessas espécies. Amostras frescas da região mediana à apical das lâmina foliolar, pecíolo e nó foram fixadas e processadas de acordo com os métodos usuais para microscopia de luz e eletrônica de varredura. Os tricomas glandulares encontrados nas espécies foram: peltados, capitados, estipitados e pateliformes/cupuliformes. Dentre esses tipos o mais abundante foi o tricoma peltado, apresentando também uma distribuição mais uniforme que os demais. Os tricomas pateliformes/cupuliformes estavam em regiões mais específicas, como profilos, lâmina foliolar e na região nodal. Martinella obovata foi a única espécie que apresentou os tricomas capitados e estipitados, amplamente distribuídos ao longo de todo eixo vegetativo aéreo. Em todas as espécies foram encontradas formigas, principalmente nas regiões nodais. A ocorrência do tricoma capitado é relatada pela primeira vez para o gênero.

Calicinal trichomes of Adenocalymma magnificum (Bignoniaceae) producing lipophilic substances: ultrastructural and functional aspects / Tricomas calicinales de Adenocalymma magnificum (Bignoniaceae) que producen sustancias lipofílicas: aspectos ultraestructurales y funcionales

Plant structures that secrete lipids and phenolic compounds are often associated with the protection and development of organs against desiccation, in addition to the protection they provide against animals, as the capitate trichomes of Adenocalymma magnificum. Understanding the glandular activities that occur in these trichomes has required the study of their ontogeny, structure, ultrastructure and histochemical aspects; the interpretation of their ecological functions or evolutionary history is complicated by the scarcity of reports on calicinal trichomes that are not nectar-secreting. Samples of floral calyx in anthesis and flower buds at different stages of development were fixed and processed according to the methods for light and electron microscopy. The trichomes are randomly distributed throughout the entire inner surface of the calyx and are also visible on the flower buds. These capitate glandular trichomes were composed of a peduncle, having up to nine cells, and a multicellular secretory head with their cells in columnar format and arranged in disc form. The collar cell, which is under the secretory head, divides anticlinally and arranges itself side by side with the mother cell. As they develop, they bend with some of them becoming adpressed to the calyx. Histochemical tests indicate that the secretory head cells produce lipid substances, acidic lipids and phenolic compounds. In the secretory head, the vacuome is dispersed and the cytoplasm possesses a great number of smooth endoplasmic reticulum and leucoplasts, organelles involved in the production of osmiophilic substances. In some regions of the secretory cells, cuticle detachment was observed; however, the accumulation of secretions was not observed. This study describes, for the first time, the origin, development, and secretion process of the calicinal trichomes of Adenocalymma magnificum, showing that production of lipophilic substances is important for this plant, possibly the trichomes may be involved in the plant’s chemical defense against insects, offering protection against herbivores. Rev. Biol. Trop. 63 (2): 537-544. Epub 2015 June 01.

Las estructuras que secretan lípidos y compuestos fenólicos estan frecuentemente asociadas a la protección y desarrollo de órganos que protegen contra la desecación, además de la protección que estas proporcionan contra el ataque de animales. Los tricomas capitados de Adenocalymma magnificum son estructuras de este tipo. Para comprender las actividades glandulares que ocurren en estos tricomas es necesario estudiar su ontogenia, estructura, ultraestructura y aspectos histoquímicos. La interpretación de sus funciones ecológicas o historia evolutiva es compleja, y pocos son los estudios que se enfocan en los tricomas calicinales no secretores de néctar. Las muestras del cáliz floral en antesis y botones florales en diferentes estadios de desarrollo fueron fijadas y procesadas de acuerdo con los métodos para microscopia de luz y electrónicos. Los tricomas de A. magnificum están aleatoriamente distribuidos en toda la extensión de la superficie interna del cáliz, también observados en los botones florales. Estos tricomas son glandulares capitados, compuestos de un pedúnculo largo que posee hasta nueve células y una cabeza secretora multicelular, con células en columnares dispuestas en disco. La célula del collar, que esta sobre las células de la cabeza secretora, se divide anticlinalmente organizándose lado a lado con las células madre. Con el crecimiento de los tricomas, estos mismos se curvan y algunos quedan adpresos al cáliz. Las pruebas histoquímicas indicaron que las células de la cabeza secretora producen sustancias lipídicas, lípidos ácidos y compuestos fenólicos. En la cabeza secretora la vacuoma esta dispersa y posee abundante retículo endoplasmático liso y leucoplastos, los cuales son organelos involucrados en el proceso de producción de sustancias osmiofílicas. En algunas regiones de las células secretoras la distensión de la cutícula fue observada, sin embargo, no fue verificada la acumulación de secreción. Este trabajo describe por primera vez el origen, desarrollo y proceso de secreción de tricomas calicinales de A. magnificum, mostrando que la producción de substancias lipofílicas es importante para esta especie, ofreciendo posiblemente protección contra la herbivoría.

Distribution and structural aspects of extrafloral nectaries in leaves of Pyrostegia venusta (Bignoniaceae) / Distribuição e aspectos estruturais de nectários extraflorais em folhas de Pyrostegia venusta (Bignoniaceae)

Pyrostegia venusta (the orange trumpet or commoly called cipَ-de-Sمo-Joمo in Brazil), a medicinal plant that grows with other plants, has an ecological importance due to the presence of nectaries on the leaves. The aim of this work was to study structural and histochemical aspects and the distribution of extrafloral nectaries (ENFs) in P. venusta leaves. Young leaves were collected, fixed and processed by usual techniques, and studied under light microscopy and scanning electron microscopy. Analyses showed that the extrafloral nectaries are dispersed throughout the leaf, with concentrations mainly in the basal third section. Nectaries lie in epidermal depressions and can be found in abaxial and adaxial leaf blades. Their morphology may be characterized macroscopically by achlorophyllous halos and microscopically as structures consisting of a short pedicel, oval multicellular head, covered by a thin cuticular layer with a secretory pole. Reducing sugars, neutral polysaccharides, proteins, starch and phenolic compounds were detected in extrafloral nectaries.(AU)

. Pyrostegia venusta (cipَ-de-Sمo-Joمo) é uma planta medicinal que cresce junto a outras plantas e que apresenta também importância ecolَgica, pela presença de nectلrios na regiمo dos folيolos. O objetivo deste trabalho foi estudar aspectos estruturais, histoquيmicos e da distribuiçمo de nectلrios extraflorais (NEFs) em folhas de P. venusta. Folhas jovens foram coletadas, fixadas e processadas por técnicas usuais e estudadas sob microscopia de luz e eletrônica de varredura. As anلlises mostraram que os NEFs estمo dispersos por toda a folha, concentrando-se preferencialmente no terço basal. Os nectلrios estمo localizados em depressُes epidérmicas e podem ser encontrados nas faces abaxial e adaxial da lâmina foliar. Macroscopicamente, os NEFs podem ser caracterizados como halos aclorofilados e microscopicamente, como estruturas com pedicelo curto, cabeça ovoide pluricelular, recobertos por uma fina camada cuticular e um polo secretor. Nos nectلrios extraflorais foram detectados açْcares redutores, polissacarيdeos neutros, proteيnas, amido e compostos fenَlicos.(AU)

Distribution and structural aspects of extrafloral nectaries in leaves of Pyrostegia venusta (Bignoniaceae) / Distribuição e aspectos estruturais de nectários extraflorais em folhas de Pyrostegia venusta (Bignoniaceae)

Pyrostegia venusta (the orange trumpet or commoly called cipó-de-São-João in Brazil), a medicinal plant that grows with other plants, has an ecological importance due to the presence of nectaries on the leaves. The aim of this work was to study structural and histochemical aspects and the distribution of extrafloral nectaries (ENFs) in P. venusta leaves. Young leaves were collected, fixed and processed by usual techniques, and studied under light microscopy and scanning electron microscopy. Analyses showed that the extrafloral nectaries are dispersed throughout the leaf, with concentrations mainly in the basal third section. Nectaries lie in epidermal depressions and can be found in abaxial and adaxial leaf blades. Their morphology may be characterized macroscopically by achlorophyllous halos and microscopically as structures consisting of a short pedicel, oval multicellular head, covered by a thin cuticular layer with a secretory pole. Reducing sugars, neutral polysaccharides, proteins, starch and phenolic compounds were detected in extrafloral nectaries.

Pyrostegia venusta (cipó-de-São-João) é uma planta medicinal que cresce junto a outras plantas e que apresenta também importância ecológica, pela presença de nectários na região dos folíolos. O objetivo deste trabalho foi estudar aspectos estruturais, histoquímicos e da distribuição de nectários extraflorais (NEFs) em folhas de P. venusta. Folhas jovens foram coletadas, fixadas e processadas por técnicas usuais e estudadas sob microscopia de luz e eletrônica de varredura. As análises mostraram que os NEFs estão dispersos por toda a folha, concentrando -se preferencialmente no terço basal. Os nectários estão localizados em depressões epidérmicas e podem ser encontrados nas faces abaxial e adaxial da lâmina foliar. Macroscopicamente, os NEFs podem ser caracterizados como halos aclorofilados e microscopicamente, como estruturas com pedicelo curto, cabeça ovoide pluricelular, recobertos por uma fina camada cuticular e um polo secretor. Nos nectários extraflorais foram detectados açúcares redutores, polissacarídeos neutros, proteínas, amido e compostos fenólicos.

Acquisition and diversification of tendrilled leaves in Bignonieae (Bignoniaceae) involved changes in expression patterns of SHOOTMERISTEMLESS (STM), LEAFY/FLORICAULA (LFY/FLO), and PHANTASTICA (PHAN).

Leaves have undergone structural modifications over evolutionary time, and presently exist in many forms. For instance, in Fabaceae and Bignoniaceae, leaf parts can be modified into tendrils. Currently, no data are available on genic control of tendrilled leaf development outside Fabaceae. Here, we conducted a detailed study of three representatives of Bignonieae: Amphilophium buccinatorium, Dolichandra unguis-cati, and Bignonia callistegioides, bearing multifid, trifid, and simple-tendrilled leaves, respectively. We investigated the structure of their petioles, petiolules, leaflets, and tendrils through histological analyses. Additionally, the expression of SHOOTMERISTEMLESS (STM), PHANTASTICA (PHAN), and LEAFY/FLORICAULA (LFY/FLO) during leaf development was analyzed by in situ hybridizations. Tendrils share some anatomical similarities with leaflets, but not with other leaf parts. Transcripts of both STM and LFY/FLO were detected in leaf primordia, associated with regions from which leaflets and tendril branches originate. PHAN expression was found to be polarized in branched tendrils, but not in simple tendrils. In Bignonieae, tendrils are modified leaflets that, as a result of premature completion of development, become bladeless organs. Bignonieae leaves develop differently from those of peas, as both LFY/FLO and STM are expressed in developing leaves of Bignonieae. Moreover, PHAN is probably involved in tendril diversification in Bignonieae, as it has distinct expression patterns in different leaf types.

Trichome structure and evolution in Neotropical lianas.

BACKGROUND AND AIMS: Trichomes are epidermal outgrowths generally associated with protection against herbivores and/or desiccation that are widely distributed from ferns to angiosperms. Patterns of topological variation and morphological evolution of trichomes are still scarce in the literature, preventing valid comparisons across taxa. This study integrates detailed morphoanatomical data and the evolutionary history of the tribe Bignonieae (Bignoniaceae) in order to gain a better understanding of current diversity and evolution of trichome types. METHODS: Two sampling schemes were used to characterize trichome types: (1) macromorphological characterization of all 105 species currently included in Bignonieae; and (2) micromorphological characterization of 16 selected species. Individual trichome morphotypes were coded as binary in each vegetative plant part, and trichome density and size were coded as multistate. Ancestral character state reconstructions were conducted using maximum likelihood (ML) assumptions. KEY RESULTS: Two main functional trichome categories were found: non-glandular and glandular. In glandular trichomes, three morphotypes were recognized: peltate (Pg), stipitate (Sg) and patelliform/cupular (P/Cg) trichomes. Non-glandular trichomes were uniseriate, uni- or multicellular and simple or branched. Pg and P/Cg trichomes were multicellular and non-vascularized with three clearly distinct cell layers. Sg trichomes were multicellular, uniseriate and long-stalked. ML ancestral character state reconstructions suggested that the most recent common ancestor (MRCA) of Bignonieae probably had non-glandular, Pg and P/Cg trichomes, with each trichome type presenting alternative histories of appearance on the different plant parts. For example, the MRCA of Bignonieae probably had non-glandular trichomes on the stems, prophylls, petiole, petiolule and leaflet veins while P/Cg trichomes were restricted to leaflet blades. Sg trichomes were not present in the MRCA of Bignonieae independently of the position of these trichomes. These trichomes had at least eight independent origins in tribe. CONCLUSIONS: The patterns of trichome evolution indicate that most morphotypes are probably homologous in Bignonieae and could be treated under the same name based on its morphological similarity and common evolutionary history, in spite of the plethora of names that have been previously designated in the literature. The trichome descriptions presented here will facilitate comparisons across taxa, allowing inferences on the relationsthips between trichome variants and future studies about their functional properties.

Phenotypic integration in flowers of neotropical lianas: diversification of form with stasis of underlying patterns.

Phenotypic integration is essential to the understanding of organismal evolution as a whole. In this study, a phylogenetic framework is used to assess phenotypic integration among the floral parts of a group of Neotropical lianas. Flowers consist of plant reproductive organs (carpels and stamens), usually surrounded by attractive whorls (petals and sepals). Thus, flower parts might be involved in different functions and developmental constraints, leading to conflicting selective forces. We found that Bignonieae flowers have very similar patterns of variance/covariance among traits and that such patterns are uncorrelated with the phylogenetic relationships between species. However, in spite of pattern stasis, our results also indicate that diversification of floral morphology in this group has occurred throughout the evolution of magnitudes of correlation among traits. Thus, we suggest that stabilizing selection has played an important role in phenotypic integration, resulting in the long-term stasis of covariance patterns underlying flower diversification during the ca. 50 Myr of evolution of Bignonieae. This is the first report of long-term stasis in the phenotypic integration of angiosperms, suggesting that patterns of floral morphology can be recognizable as specific attributes of distinct botanical families.

Longevity of the Brazilian underground tree Jacaranda decurrens Cham.

Underground trees are a rare clonal growth form. In this survey we describe the branching pattern and estimate the age of the underground tree Jacaranda decurrens Cham. (Bignoniaceae), an endangered species from the Brazilian Cerrado, with a crown diameter of 22 meters. The mean age calculated for the individual was 3,801 years, making it one of the oldest known living Neotropical plants.

Estruturas secretoras em Cipó-d'alho (Mansoa standleyi (Steyerm.) A. H. Gentry, Bignoniaceae): ocorrência e morfologia / Secretory structures in Cipó-d'alho (Mansoa standleyi (Steyerm.) A. H. Gentry, Bignoniaceae): occurrence and morphology

Espécies de Mansoa são denominadas de "cipó-d'alho", por exalarem odor de alho das partes vegetativas e reprodutivas. Contudo, dados sobre morfologia e distribuição das estruturas secretoras presentes em Mansoa são escassos e ausentes para M. standleyi. O presente trabalho objetivou caracterizar a ocorrência e morfologia das estruturas secretoras do eixo vegetativo aéreo de M. standleyi. Para tanto, amostras da lâmina foliolar e de regiões nodais foram fixadas e submetidas às técnicas histológicas e de microscopia eletrônica de varredura. Testes histoquímicos, com os respectivos controles foram aplicados nas estruturas secretoras em fase secretora. Indivíduos de formigas e moscas, que visitavam a espécie foram amostrados, preservados e identificados por entomólogo. As estruturas secretoras do eixo vegetativo aéreo de M. standleyi estão representadas por tricomas glandulares dos tipos pateliformes e peltados. Todos com desenvolvimento assincrônico e presentes nas regiões nodais e lâmina foliolar, principalmente nas partes mais jovens. Nas regiões nodais, os tricomas formam um complexo secretor e, na lâmina foliolar, estão dispersos. As análises histoquímicas revelaram que os tricomas pateliformes são de fato nectários extraflorais e que os tricomas peltados, apresentam uma fração de alcaloides. Os visitantes das glândulas nodais correspondem a formigas Crematogaster (Formicidae) e Ectatomma brunea (Vespoidea, Formicidae) e moscas Oxysarcodexia (Sarcophagidae, subfamília Utitidae (Ulidiidae). Mansoa standleyi possui as estruturas secretoras do eixo vegetativo aéreo semelhantes às citadas pela literatura para Bignoniaceae, sendo esta a primeira vez, que um nectário extrafloral é descrito para a espécie.

Species of Mansoa are called "cipó-d'alho" because of the smell of garlic that wafts from their vegetative and reproductive parts. Since data on the morphology and occurrence of their secretory structures are scarce and even absent for M. standleyi, the present work characterizes the distribution and morphology of such structures in the vegetative aerial axis of the latter. To do so, samples of the leaf blade and of nodal regions were fixed and examined using histological and scanning electron microscope techniques. Histochemical tests with appropriate controls were carried out on the secretory structures during the secretory phase. Fly and ant individuals that visit the species were sampled, preserved and identified by an entomologist. The secretory structures of the vegetative aerial axis of M. standleyi are peltate and patelliform glandular trichomes. All develop asynchronously and are present in the nodal regions and leaf blade, mainly in their youngest parts. Trichomes form a secretory complex in the nodal regions while they are scattered in the leaf blade. Histochemical analyses revealed that the cupulate and patelliform trichomes are extrafloral nectaries and that the peltate ones present an alkaloid fraction. The visitors of the nodal glands are ants Crematogaster (Formicidae) and Ectatomma brunea (Vespoidea, Formicidae) and flies Oxysarcodexia (Sarcophagidae, subfamily Utitidae [Ulidiidae]). The secretory structures of the vegetative aerial axis of Mansoa standleyi are similar to those reported for Bignoniaceae. An extrafloral nectary is described for M. standleyi for the first time.

Evolution of disparity between the regular and variant phloem in Bignonieae (Bignoniaceae).

PREMISE OF THE STUDY: The phloem is a plant tissue with a critical role in plant nutrition and signaling. However, little is still known about the evolution of this tissue. In lianas of the Bignoniaceae, two distinct types of phloem coexist: a regular and a variant phloem. The cells associated with these two phloem types are known to be anatomically different; however, it is still unclear what steps were involved in the evolution of such differences. METHODS: Here we studied the anatomical development of the regular and variant phloem in representatives of all 21 genera of Bignonieae and used a phylogenetic framework to investigate the timing of changes associated with the evolution of each phloem type. KEY RESULTS: We found that the variant phloem always appears in a determinate location, between the leaf orthostichies. Furthermore, the variant phloem was mostly occupied by very wide sieve tubes and generally included a higher concentration of fibers, indicating an increase in conduction and mechanical support. On the other hand, the regular phloem included much more parenchyma, more and wider rays, and tiny sieve tubes that resembled terminal sieve tubes from plants with seasonal formation of vascular tissues; these findings suggest reduced conduction and higher storage capacity in the regular phloem. CONCLUSIONS: Overall, differences between the regular and variant phloem increased over time, leading to further specialization in conduction in the variant phloem and an increase in storage specialization in the regular phloem.

Desenvolvimento e produção da carobinha (Jacaranda decurrens Cham. subsp. symmetrifoliolata Farias & Proença) cultivada sob dois arranjos de plantas, com ou sem cobertura de cama-de-frango no solo / Development and yield of carobinha (Jacaranda decurrens Cham. subsp. symmetrifoliolata Farias & Proença) cultivated under two plant arrangements, with or without chicken manure covering on the soil

O objetivo deste trabalho foi avaliar o desenvolvimento e a produção da carobinha (Jacaranda decurrens subsp. symmetrifoliolata) cultivada ex situ sob dois arranjos de plantas, sem ou com cama-de-frango semidecomposta. O trabalho foi desenvolvido sob condições de campo, no Horto de Plantas Medicinais (HPM), da Universidade Federal da Grande Dourados (UFGD) em solo tipo Latossolo Vermelho distroférrico. Estudaram-se os arranjos em fileiras simples e duplas e o uso ou não de cama-de-frango de corte semidecomposta em cobertura, dispostos como fatorial 2 x 2, no delineamento experimental de blocos casualizados, com seis repetições. As maiores alturas máximas das plantas (61,3 cm planta-1) aos 555 dias após o transplante foram daquelas cultivadas sob fileira dupla com cama-de-frango. Os números médios de folhas e de ramificações por planta ao longo do ciclo foram de onze e duas, respectivamente. O diâmetro médio do caule de plantas cultivadas em fileiras simples foi de 8,3 mm, enquanto sob fileiras duplas, foi de 10,8 mm, ambos aos 552 dias de ciclo. As características avaliadas, na colheita, aos 555 dias após o transplante não foram influenciadas pela interação entre arranjos de plantas e o uso da cama-de-frango nem pelos fatores isoladamente. Dessa forma, concluiu-se que o cultivo em fileiras duplas foi favorável à produtividade da carobinha, por possibilitar maior produção por área.

The aim of this study was to evaluate the development and yield of "carobinha" (Jacaranda decurrens subsp. symmetrifoliolata) cultivated ex situ under two plant arrangements, with or without semi-decomposed chicken manure. This study was carried out under field conditions at the Medicinal Plant Garden (HPM), Federal University of Grande Dourados (UFGD), in Distroferric Red Latossol. Arrangements with simple and double rows and the use or not of semi-decomposed chicken manure as covering were studied as 2 x 2 factorial arrangement in a randomized block experimental design, with six replicates. The highest maximum heights (61.3 cm plant-1) at 555 days after transplant were obtained by plants cultivated under double rows with chicken manure. The mean numbers of leaves and branches per plant during the cycle were eleven and two, respectively. The mean stem diameter of plants that were cultivated in simple rows was 8.3 mm, while that of plants cultivated in double rows was 10.8 mm, both at 552 days of the cycle. The evaluated characteristics, in the harvest at 555 days after transplant, were not influenced by the interaction between plant arrangements and use of chicken manure or by factors in an isolated way. Thus, cultivation in double rows was favorable to "carobinha" yield for allowing the highest yield per area.

The rise and evolution of the cambial variant in Bignonieae (Bignoniaceae).

Cambial variants represent a form of secondary growth that creates great stem anatomical diversity in lianas. Despite the importance of cambial variants, nothing is known about the developmental mechanisms that may have led to the current diversity seen in these stems. Here, a thorough anatomical analysis of all genera along the phylogeny of Bignonieae (Bignoniaceae) was carried out in order to detect when in their ontogeny and phylogeny there were shifts leading to different stem anatomical patterns. We found that all species depart from a common developmental basis, with a continuous, regularly growing cambium. Initial development is then followed by the modification of four equidistant portions of the cambium that reduce the production of xylem and increase the production of phloem, the former with much larger sieve tubes and an extended lifespan. In most species, the formerly continuous cambium becomes disjunct, with cambial portions within phloem wedges and cambial portions between them. Other anatomical modifications such as the formation of multiples of four phloem wedges, multiple-dissected phloem wedges, and included phloem wedges take place thereafter. The fact that each novel trait raised on the ontogenetic trajectory appeared in subsequently more recent ancestors on the phylogeny suggests a recapitulatory history. This recapitulation is, however, caused by the terminal addition of evolutionary novelties rather than a truly heterochronic process. Truly heterochronic processes were only found in shrubby species, which resemble juveniles of their ancestors, as a result of a decelerated phloem formation by the variant cambia. In addition, the modular evolution of phloem and xylem in Bignonieae seems to indicate that stem anatomical modifications in this group occurred at the level of cambial initials.

Pollination biology of Jacaranda oxyphylla with an emphasis on staminode function.

BACKGROUND AND AIMS: Bignoniaceae is a Neotropical family with >100 genera, only two of which, Jacaranda and Digomphia, have a developed staminode. Jacaranda oxyphylla, whose flowers possess a conspicuous glandular staminode, is a zoophilous cerrado species. Here, the composition of the secretion of the glandular trichome and the influence of the staminode on the pollination biology and reproductive success of J. oxyphylla were studied. METHODS: The floral morphology, pollen viability, stigma receptivity, nectar volume and nectar concentration were studied. Compatibility system experiments were performed and floral visitors were observed and identified. Experiments comparing the effect of staminode presence and absence on pollen removal and pollen deposition efficiency were conducted in open-pollinated flowers. Histochemistry, thin-layer chromatography (TLC) and gas chromatography coupled to flame ionization detection (GC-FID) analyses were performed to determine the main chemical components of the staminode's glandular trichome secretion. KEY RESULTS: Flower anthesis lasted 2 d and, despite the low frequency of flower visitation, pollination seemed to be effected mainly by medium-sized Eulaema nigrita and Bombus morio bees, by the small bee Exomalopsis fulvofasciata and occasionally by hummingbirds. Small bees belonging to the genera Ceratina, Augochlora and Trigona were frequent visitors, collecting pollen. Jacaranda oxyphylla is predominantly allogamous. Staminode removal resulted in fewer pollen grains deposited on stigmas but did not affect total pollen removal. The secretion of capitate glandular trichome occurs continually; the main chemical compounds detected histochemically were phenolic and terpenoid (essential oils and resins). Monoterpene cineole, pentacyclic triterpenes and steroids were identified by TLC and GC-FID. CONCLUSIONS: The staminode of J. oxyphyllla is multifunctional and its importance for female reproductive success was attributed mainly to the secretion produced by capitate glandular trichomes. This secretion is involved in complex chemical interactions with pollinating bees, including the solitary bees Euglossini. These bees are common pollinators of various species of Jacaranda.

Estudo anatômico das espécies de cerrado Anemopaegma arvense (Vell.) Stellf. ex de Souza (catuaba), Zeyheria montana Mart. (bolsa-de-pastor) e Jacaranda decurrens Chamisso (caroba) - Bignoniaceae / Anatomical study of species from savanna-like formation, Anemopaegma arvense (Vell.) Stellf. ex de Souza (catuaba), Zeyheria montana Mart. (bolsa-de-pastor) and Jacaranda decurrens Chamisso (caroba) - Bignoniaceae

Foi realizado um estudo anatômico de três espécies medicinais da família Bignoniaceae, típicas de cerrado, em risco de extinção, a saber: Anemopaegma arvense, Jacaranda decurrens e Zeyheria montana, provenientes de Botucatu. Para isto, realizaram-se cortes transversais de caule, limbo e pecíolo e paradérmicos de limbo. Os cortes, corados diferencialmente com azul de alcian e fucsina básica, foram objeto de estudo e descrição anatômica. As três espécies mostram adaptações anatômicas a ambiente xerofítico, representadas por mesofilo foliar compactado, cutícula espessa (Anemopaegma e Jacaranda) e abundante aquisição de tricomas tectores (Zeyheria).

The anatomy of three medicinal species belonging to the Bignoniaceae family, from savanna-like formation and in risk of extinction, was studied. They were: Anemopaegma arvense, Jacaranda decurrens and Zeyheria montana, from Botucatu. Hand-made transversal sections of stem, foliar leaf and petiole and paradermal sections of foliar leaf, differentially stained with alcian blue and basic fuchsin, were studied and described anatomically. The three species show anatomical adaptations to xerophytic environment, expressed by compaction of foliar mesophyll, thickened cuticle (Anemopaegma and Jacaranda) and acquisition of abundant non-glandular trichomes (Zeyheria).

Ovary peltate trichomes of Zeyheria montana (Bignoniaceae): developmental ultrastructure and secretion in relation to function.

BACKGROUND AND AIMS: Nectar production in the Bignoniaceae species lacking a nectariferous functional disc is ascribed to trichomatic glands around the ovary base and/or on the inner corolla wall. Nevertheless, knowledge about the secretion and function of these glands is very incomplete. The purpose of this paper is to study, from a developmental viewpoint, the ultrastructure, histochemistry and secretory process of the peltate trichomes on the ovary of Zeyheria montana, a species in the Bignoniaceae which has a rudimentary disc. METHODS: Samples of the gynoecium at various developmental stages were fixed and processed for light and electron microscopy. Histochemistry and cytochemistry tests were performed to examine the chemical composition of exudates. Thin layer chromatography was used to determine the presence of alkaloids and terpenes in gynoecium and fruit extracts, and in fresh nectar stored in the nectar chamber. KEY RESULTS: Peltate trichomes at different developmental stages appear side by side from floral budding up to pre-dispersal fruit. Large plastids with an extensive internal membrane system consisting of tubules filled with lipophilic material, abundant smooth endoplasmic reticulum, few Golgi bodies, lipophilic deposits in the smooth endoplasmic reticulum and mitochondria, and scattered cytoplasmic oil droplets are the main characteristics of mature head cells. The secretion which accumulates in the subcuticular space stains positively for hydrophilic and lipophilic substances, with lipids prevailing for fully peltate trichomes. Histochemistry and thin layer chromatography detected terpenes and alkaloids. Fehling's test to detect of sugars in the secretion was negative. CONCLUSIONS: The continuous presence and activity of peltate trichomes on the ovary of Z. montana from early budding through to flowering and fruiting set, and its main chemical components, alkaloids and terpenes, suggest that they serve a protective function and are not related to the floral nectar source or to improving nectar quality.

Secretory trichomes, a substitutive floral nectar source in Lundia A. DC. (Bignoniaceae), a genus lacking a functional disc.

This is the first report of corolla-borne secretory trichomes that substitute in role for a non-functional disc in a species of the neotropical genus Lundia A. DC. (Bignoniaceae). The floral biology and flowering phenology of Lundia cordata were investigated at two remnants of tropical rainforest in northeastern Brazil. This species is a typically omithophilous liana, with reddish, tubular, scentless flowers. The flowers are resupinate, protandrous and last for 2 d. There is a vestigial non-functional perigynous disc and nectar is secreted by glandular trichomes distributed along the internal surface of the corolla. The nectar is stored at the base of the corolla tube, thus showing secondary nectar presentation. The nectariferous trichomes are multi-cellular, uniseriate, with a basal foot cell rooting in the epidermis, one neck cell, and a glandular head with 13 cells on average. Three species of hummingbirds (Amazilia fimbriata, Eupetomena macroura and Phaethornis pretrei) serve as pollinators. Phaethornis ruber, Xylocopa bees, wasps and diurnal moths are considered nectar thieves.