Morphological diversity and evolution of anther glands in Mimosoideae (Leguminosae).

Em Mimosoideae (Leguminosae), glândulas em anteras são estruturas de destaque, pois ocorrem em muitos de seus representantes e provêm caracteres com valor taxonômico no clado. O conhecimento a respeito destas glândulas era baseado principalmente em observações de morfologia externa, faltando dados a respeito de sua anatomia, ontogenia, seu funcionamento e seus aspectos evolutivos, o que contribuiria para uma caracterização mais detalhada e uma classificação mais natural dessas estruturas, além da inferência de funções ecológicas mais objetivas. Assim, o objetivo do estudo foi revisitar a estrutura das glândulas nas anteras, investigar sua origem meristemática e o seu desenvolvimento, acompanhar seu funcionamento, inferir funções ecológicas e, então, reconstruir sua evolução em Mimosoideae. Para contemplar os objetivos propostos, o presente estudo investigou aspectos das glândulas em anteras em 25 espécies de Mimosoideae, que representam grupos com divergências mais antigas e recentes no clado. Foram encontrados quatro tipos principais de glândulas: (1) O tipo Piptadenia, caracterizado por uma glândula pedunculada com ápice esférico e que inclui três subtipos: Piptadenia, com ápice composto de células isodiamétricas; Adenanthera, com ápice composto de uma camada de células epidérmicas alongadas que delimitam células isodiamétricas; e Entada, com ápice composto de uma camada de células epidérmicas volumosas que delimitam um lúmen. (2) O tipo Pentaclethra, uma glândula robusta composta de uma camada de células epidérmicas em paliçada e células centrais isodiamétricas, com pedúnculo vascularizado. (3) O tipo Prosopis africana, uma glândula robusta, séssil e vascularizada. (4) Finalmente o tipo Gagnebina, uma glândula acuminada com células alongadas. Apesar da diversidade morfológica, as glândulas de anteras compartilham uma origem meristemática comum. Elas surgem de divisões simultâneas nas primeiras três camadas de células meristemáticas que cobrem a porção apical da antera jovem, sendo classificadas como emergências. Os tipos de glândulas podem apresentar diferenças quanto a: (1) estrutura na pré-antese e antese; (2) longevidade; (3) aspecto e composição química do exsudato e (4) forma de liberação do exsudato. Baseados na caracterização e forma de liberação do exsudato, e no comportamento dos visitantes florais, nossos dados apontam que as glândulas atuam principalmente (a) na produção de substâncias que atuam na adesão do pólen ao corpo do polinizador ou à antera e (b) na produção de fragrância floral atuando como osmóforos. A presença de glândulas em anteras parece ter uma origem única em Mimosoideae, com alguns eventos de perda. O estado ancestral da estrutura das glândulas provavelmente corresponde ao tipo Piptadenia subtipo Adenanthera. Ainda, parece existir uma relação positiva entre ausência de glândulas e a presença de inflorescências de eixo curto, de estames numerosos e de estames unidos. Nossos dados indicam que em Mimosoideae, as glândulas em anteras são caracteres com potencial significado filogenético. A origem meristemática conservada das glândulas além da origem única da presença de glândulas em anteras Mimosoideae favorece a hipótese de homologia dessas estruturas e apoia a monofilia do clado.In Mimosoideae (Leguminosae), anther glands are prominent structures because they occur in many taxa and provide characters with taxonomic value in the clade. The knowledge about the anther glands was mainly based on external morphological observations, lacking data concerning the anatomy, ontogeny, function and evolutionary aspects of the glands, which would contribute to a more detailed characterization and a more natural classification of these structures, as well as the inference of more objective ecological functions. The aims of this study were to revisit the structure of the anther glands, to investigate their meristematic origin, and their developmental stages, their functioning mechanisms, to infer ecological functions, and thus reconstruct the evolutionary history of these glands in Mimosoideae. For this, the present study investigated several aspects of the anther glands in 25 species of Mimosoideae representing groups with early and late divergences in the clade. We found four major types of glands in anther Mimosoideae: (1) Piptadenia type, characterized by a stalked gland with a spherical apex, and that includes three subtypes: Piptadenia, with the apex composed of isodiametrical cells; Adenanthera, with the apex composed a layer of elongated epidermal cells which delimits isodiametric cells; and Entada with the apex composed of a layer of epidermal bulky cells delimiting a lumen. (2) Pentaclethra type,a robust gland comprised of a layer of epidermal and palisade cells isodiametric central cells with vascularized peduncle. (3) The Prosopis africana type, a robust sessile and vascularized gland. (4) Finally, the Gagnebina type, an acuminate gland with elongated cells. Despite the morphological diversity, the anthers glands share a common meristematic origin. They arise from simultaneous divisions in the first three layers of meristematic cells covering the apical portion of the young anther, being classified as emergences. The types of glands may differ as to: (1) structure in the pre-anthesis and anthesis; (2) longevity; (3) aspect and chemical composition of exudate and (4) release form of exudate. Considering the characterization and release form of exudate, and behavior of floral visitors, our data show that the glands act mainly (a) in the production of substances that act on the accession of pollen to the bee body or anther and (b) in production of floral fragrance acting as osmophores. The presence of anther glands appears to have a single origin in Mimosoideae, with some loss events. The ancestral state of the structure of the glands probably corresponds to the type Piptadenia subtype Adenanthera. Still, there seems to be a negative relation between the presence of the glands, the presence of inflorescences with short axis, numerous stamens and united stamens. Our data indicate that in Mimosoideae, the glands in anthers provide characters with potential phylogenetic significance. The conserved meristematic origin of the anther glands beyond the single evolutionary origin of the presence of glands in anthers favor the hypothesis of homology of these structures and support the monophyly of Mimosoideae

Secretory structures in vegetative organs of Barbatimão species (Dimorphandra mollis Benth. and Stryphnodendron adstringens (Mart.) Coville - Leguminosae)

Duas espécies de Barbatimão, da família Leguminosae, foram utilizadas como modelo neste trabalho para elucidar as vias ontogenéticas de produção de taninos: Dimorphandra mollis Benth. (Caesalpinioideae) e Stryphnodendron adstringens (Mart.) Coville (Mimosoideae). Apesar de inseridas em grupos distantes de Leguminosae, as duas espécies são grandes produtoras de taninos, geralmente concentrados na casca, o que confere a elas uma grande procura para uso na medicina popular. Partes vegetativas de Indivíduos jovens e adultos foram processadas para observações em microscopia de luz e eletrônica (varredura e transmissão). Dois tipos de estruturas secretoras são responsáveis pela produção e acúmulo de taninos: idioblastos, encontrados já em indivíduos jovens de ambas espécies; e tricomas secretores, encontrados em gemas vegetativas de indivíduos adultos de S. adstringens. O processo de produção de taninos nas duas espécies é semelhante, com alguma diferença nas etapas de acúmulo. Plastídios e RER parecem ser as organelas associadas à produção de taninos nessas espécies. Destaca-se ainda o relato inédito da ocorrência de coléteres de origem protodérmica em D. mollis. Considerando a problemática referente à classificação de estruturas secretoras, pretende-se utilizar os resultados obtidos para reflexão sobre o temaIn this study, two species of Leguminosae, popularly known as Barbatimão, were used as a model to elucidate the ontogenetic process of tannin production: Dimorphandra mollis Benth. (Caesalpinioideae) and Stryphnodendron adstringens (Mart.) Coville (Mimosoideae). Although inserted in non-related groups of Leguminosae, the two species are largely known as tannin producers, a metabolite often concentrated in the bark, which makes them highly sought after for use in folk medicine. Vegetative parts of young and adult plants were processed for observation using light and electron microscopy (scanning and transmission). Two types of secretory structures are responsible for tannin production and accumulation: idioblasts, found even in young plants of both species, and secretory trichomes, observed in vegetative buds of mature plants of S.adstringens. Tannin production is similar in both species, although some differences in the stages of accumulation occur. Plastids and RER organelles appear to be associated with the production of tannins in these species. This study also provides the first report of colleters in D. mollis, which originate from protodermal cells. Considering the controversies regarding the classification and nomenclature of secretory structures, our results can be used to help clarify some issues regarding secretory structures, especially colleter

Secretory structures in vegetative organs of Barbatimão species (Dimorphandra mollis Benth. and Stryphnodendron adstringens (Mart.) Coville - Leguminosae)

Duas espécies de Barbatimão, da família Leguminosae, foram utilizadas como modelo neste trabalho para elucidar as vias ontogenéticas de produção de taninos: Dimorphandra mollis Benth. (Caesalpinioideae) e Stryphnodendron adstringens (Mart.) Coville (Mimosoideae). Apesar de inseridas em grupos distantes de Leguminosae, as duas espécies são grandes produtoras de taninos, geralmente concentrados na casca, o que confere a elas uma grande procura para uso na medicina popular. Partes vegetativas de Indivíduos jovens e adultos foram processadas para observações em microscopia de luz e eletrônica (varredura e transmissão). Dois tipos de estruturas secretoras são responsáveis pela produção e acúmulo de taninos: idioblastos, encontrados já em indivíduos jovens de ambas espécies; e tricomas secretores, encontrados em gemas vegetativas de indivíduos adultos de S. adstringens. O processo de produção de taninos nas duas espécies é semelhante, com alguma diferença nas etapas de acúmulo. Plastídios e RER parecem ser as organelas associadas à produção de taninos nessas espécies. Destaca-se ainda o relato inédito da ocorrência de coléteres de origem protodérmica em D. mollis. Considerando a problemática referente à classificação de estruturas secretoras, pretende-se utilizar os resultados obtidos para reflexão sobre o temaIn this study, two species of Leguminosae, popularly known as Barbatimão, were used as a model to elucidate the ontogenetic process of tannin production: Dimorphandra mollis Benth. (Caesalpinioideae) and Stryphnodendron adstringens (Mart.) Coville (Mimosoideae). Although inserted in non-related groups of Leguminosae, the two species are largely known as tannin producers, a metabolite often concentrated in the bark, which makes them highly sought after for use in folk medicine. Vegetative parts of young and adult plants were processed for observation using light and electron microscopy (scanning and transmission). Two types of secretory structures are responsible for tannin production and accumulation: idioblasts, found even in young plants of both species, and secretory trichomes, observed in vegetative buds of mature plants of S.adstringens. Tannin production is similar in both species, although some differences in the stages of accumulation occur. Plastids and RER organelles appear to be associated with the production of tannins in these species. This study also provides the first report of colleters in D. mollis, which originate from protodermal cells. Considering the controversies regarding the classification and nomenclature of secretory structures, our results can be used to help clarify some issues regarding secretory structures, especially colleter

Beyond pollination: diversity of secretory structures during flower development in different legume lineages

ABSTRACT Floral secretory structures are usually associated with the attraction of pollinators, but may also play an important role in the mechanisms of plant protection. This study aimed to show the diversity of secretory structures present in the developing flowers of 15 legume species belonging to different clades and to associate them with functions other than the pollinator attraction. Buds, flowers and developing axis of inflorescence were processed for surface, histological, and ultrastructural analyses. The species investigated displayed a wide diversity of secretory structures in developing flowers such as phenolic cells and/or tissues, mucilaginous cells, secretory cavities, secretory trichomes and colleters. Each type of secretory structure exhibited variation in morphology and location in the flower and/or axis of inflorescence depending on the species. Special mucilage cells, secretory cavities, secretory trichomes and colleters have great potential for comparative morphological studies due to their diversity of forms or restricted occurrence to certain taxa, contributing to a more robust morphological data base for the new clades emerging in Leguminosae. The scarcity of reports about floral secretory structures of Leguminosae seems to be more related to deficient sampling than to the absence of such structures in the group, which highlights the need for further investigation

Direct Adhesive Pin-Retained Restorations for Severely Worn Dentition Treatment: A 1.5-Year Follow-Up Report

Excessive occlusal surface wear can result in occlusal disharmony, functional and esthetic impairment. As a therapeutic approach, conventional single crowns have been proposed, but this kind of treatment is complex, highly invasive and expensive. This case report describes the clinical outcomes of an alternative minimally invasive treatment based on direct adhesive-pin retained restorations. A 64-year-old woman with severely worn dentition, eating problems related to missing teeth and generalized tooth hypersensitivity was referred for treatment. Proper treatment planning based on the diagnostic wax-up simulation was used to guide the reconstruction of maxillary anterior teeth with direct composite resin over self-threading dentin pins. As the mandibular remaining teeth were extremely worn, a tooth-supported overdenture was installed. A stabilization splint was also used to protect the restorations. This treatment was a less expensive alternative to full-mouth rehabilitation with positive esthetic and functional outcomes after 1.5 years of follow-up

Health-status outcomes with invasive or conservative care in coronary disease

BACKGROUND In the ISCHEMIA trial, an invasive strategy with angiographic assessment and revascularization did not reduce clinical events among patients with stable ischemic heart disease and moderate or severe ischemia. A secondary objective of the trial was to assess angina-related health status among these patients. METHODS We assessed angina-related symptoms, function, and quality of life with the Seattle Angina Questionnaire (SAQ) at randomization, at months 1.5, 3, and 6, and every 6 months thereafter in participants who had been randomly assigned to an invasive treatment strategy (2295 participants) or a conservative strategy (2322). Mixed-effects cumulative probability models within a Bayesian framework were used to estimate differences between the treatment groups. The primary outcome of this health-status analysis was the SAQ summary score (scores range from 0 to 100, with higher scores indicating better health status). All analyses were performed in the overall population and according to baseline angina frequency. RESULTS At baseline, 35% of patients reported having no angina in the previous month. SAQ summary scores increased in both treatment groups, with increases at 3, 12, and 36 months that were 4.1 points (95% credible interval, 3.2 to 5.0), 4.2 points (95% credible interval, 3.3 to 5.1), and 2.9 points (95% credible interval, 2.2 to 3.7) higher with the invasive strategy than with the conservative strategy. Differences were larger among participants who had more frequent angina at baseline (8.5 vs. 0.1 points at 3 months and 5.3 vs. 1.2 points at 36 months among participants with daily or weekly angina as compared with no angina). CONCLUSIONS In the overall trial population with moderate or severe ischemia, which included 35% of participants without angina at baseline, patients randomly assigned to the invasive strategy had greater improvement in angina-related health status than those assigned to the conservative strategy. The modest mean differences favoring the invasive strategy in the overall group reflected minimal differences among asymptomatic patients and larger differences among patients who had had angina at baseline

Initial invasive or conservative strategy for stable coronary disease

BACKGROUND Among patients with stable coronary disease and moderate or severe ischemia, whether clinical outcomes are better in those who receive an invasive intervention plus medical therapy than in those who receive medical therapy alone is uncertain. METHODS We randomly assigned 5179 patients with moderate or severe ischemia to an initial invasive strategy (angiography and revascularization when feasible) and medical therapy or to an initial conservative strategy of medical therapy alone and angiography if medical therapy failed. The primary outcome was a composite of death from cardiovascular causes, myocardial infarction, or hospitalization for unstable angina, heart failure, or resuscitated cardiac arrest. A key secondary outcome was death from cardiovascular causes or myocardial infarction. RESULTS Over a median of 3.2 years, 318 primary outcome events occurred in the invasive-strategy group and 352 occurred in the conservative-strategy group. At 6 months, the cumulative event rate was 5.3% in the invasive-strategy group and 3.4% in the conservative-strategy group (difference, 1.9 percentage points; 95% confidence interval [CI], 0.8 to 3.0); at 5 years, the cumulative event rate was 16.4% and 18.2%, respectively (difference, 121.8 percentage points; 95% CI, 124.7 to 1.0). Results were similar with respect to the key secondary outcome. The incidence of the primary outcome was sensitive to the definition of myocardial infarction; a secondary analysis yielded more procedural myocardial infarctions of uncertain clinical importance. There were 145 deaths in the invasive-strategy group and 144 deaths in the conservative-strategy group (hazard ratio, 1.05; 95% CI, 0.83 to 1.32). CONCLUSIONS Among patients with stable coronary disease and moderate or severe ischemia, we did not find evidence that an initial invasive strategy, as compared with an initial conservative strategy, reduced the risk of ischemic cardiovascular events or death from any cause over a median of 3.2 years. The trial findings were sensitive to the definition of myocardial infarction that was used