An overview of secretion in floral bracts of Tillandsioideae (Bromeliaceae), with emphasis on the secretory scales.

Bromeliaceae display many water-use strategies, from leaf impounding tanks to Crassulacean acid metabolism (CAM) photosynthesis and absorbing trichomes. Recent studies show that trichomes in inflorescences of bromeliads can exude viscous secretions, protecting against various stresses, including excessive water loss. In light of this, and considering the knowledge gap regarding inflorescence trichomes in bromeliads, we aimed to investigate the presence, source and chemical nature of inflorescence secretions in species of the Tillandsioideae (Bromeliaceae) and to describe the anatomy of their floral bracts focusing on trichome structure and position. We conducted a prospection of secretory activity and anatomy in floral bracts in 52 species of Tillandsioideae and 1 early divergent Bromeliaceae species. We used histochemical tests to investigate the presence and nature of secretion combined with standard light microscopy methods. Secretion appears in all studied species of tribe Vrieseeae, in Guzmania species, Wallisia cyanea, Tillandsia streptopylla (Tillandsieae) and Catopsis morreniana (Catopsideae). It is absent in Vriesea guttata (Vrieseeae), Racinaea crispa and various Tillandsia species (Tillandsieae). Secretion is produced by peltate trichomes on the adaxial surface of young bracts and comprises hydrophilic and lipophilic substances. Bract anatomy revealed an internal mucilage-secreting tissue with wide distribution within the subtribe Vrieseinae. Our results point to a broad occurrence of secretion associated with bracteal scales in inflorescences of Tillandsioideae. Secretory function is strongly related to trichomes of the adaxial surface, whereas the indumentum of the abaxial side is lacking or likely associated with water absorption; the latter case is especially related to small, xeric plants. Exudates might engage in colleter-like roles, protecting against desiccation, high-radiation and herbivores. Directions for future research are presented.

First Report of Colleters in Araceae: A Case Study in Anthurium andraeanum Reveals Diverse Mucilage Glands Associated with the Developing Shoot.

Araceae comprises a diverse group of plants that grow in various habitats, ranging from submerged aquatics to lithophytes. Thus, aroids are likely to show diverse glands acting in several plant-environment interactions, including colleters that protect young shoots. Based on this premise and the lack of studies regarding secretory structures in Araceae, we employed standard light and electron microscopy methods to test the hypothesis that colleters are present in Anthurium. Our main goals were to identify mucilage glands in A. andraeanum by conducting a detailed anatomical study of their structure, ultrastructure, and secretory activity. We found finger-like colleters in the apex of young leaves, spathes, and unexpanded cataphylls as well as secreting zones at the apex of expanded cataphylls, at the margins of non-fused cataphylls, and throughout the keels in two-keeled cataphylls. The colleters develop precociously and senesce shortly afterwards. Ultrastructural data and histochemistry confirmed the production of a polysaccharide-rich secretion that fills the spaces within the developing shoot. As far we know, this is the first time that colleters have been reported for Araceae. The functional roles of the secretion and the position of finger-like colleters concerning the 'precursor tip' of monocotyledons are discussed. Future research correlating secretory activity in colleters of species from different habitats might reveal a great diversity of mucilage glands with ecological and evolutionary significance to the family.

A unified view of homology.

As it spread through time and into distinct areas of science-from comparative anatomy to evolutionary biology, cladistics, developmental and molecular biology-the homology concept has changed considerably, presenting various meanings. Despite many attempts at developing a comprehensive understanding of the concept, this context-sensitive notion of homology has been a subject of an ongoing debate. Inspired by that and following Kevin de Queiroz and Richard Mayden's view on species concept and delimitation, we presented in this article an attempt to systematize and advance the understanding of the homology problem. Our main goals were: (i) to present a comprehensive checklist of 'concepts of homology'; (ii) to identify which are really concepts with ontological definitions (theoretically rooted in structural correspondence and common ancestry), and which are, in fact, not concepts, but epistemological (empirical and methodological) criteria of homology delimitation; (iii) to provide a synonymy of the concepts and criteria of homology delimitation; (iv) to present a hierarchy of homology concepts within Hennig's hologenetic system; and (v) to endorse the adoption of a unified view of homology by treating homology as a correspondence of spatio-temporal properties (genetic, epigenetic, developmental and positional) at the level of the individual, species or monophyletic group. We found 59 'concepts of homology' in the literature, from which 34 were categorically treated as concepts, 17 as criteria of homology delimitation, Four were excluded from our treatment, and Müller's five concepts were rather treated as approaches to homology. Homology concepts and criteria were synonymized based on structural correspondence, replicability, common ancestry, genetic and epigenetic developmental causes, position and optimization. Regarding the synonymy, we conclusively recognized 21 different concepts of homology, and five empirical and four methodological criteria. Hierarchical ontological aspects of homology were systematized under Hennig's hologenetic system, based on the existence of ontogenetic, tokogenetic and phylogenetic levels of homology. The delimitation of tokogenetic and phylogenetic homologies depends on optimization criteria. The unified view of homology is discussed in the context of the ancestral angiosperm flower.

Extranuptial nectaries in bromeliads: a new record for Pitcairnia burchellii and perspectives for Bromeliaceae.

Nectar plays important roles in the relationship between plants and other organisms, both within pollination systems and as a defense mechanism. In the latter case, extranuptial nectaries (ENNs) usually attract patrolling arthropods that reduce herbivory. ENNs have been frequently reported within the "xeric clade" of Bromeliaceae, but their occurrence in other groups of bromeliads is largely unexplored, especially considering their position, secretory activity and structure. After observing the presence of ants constantly patrolling the inflorescences of Pitcairnia burchellii Mez, we searched for the presence, secretory activity, and structure of ENNs in this species. We also provide a brief review of the occurrence ENNs in Bromeliaceae. The distribution of nectaries was assessed using ant-exclusion experiments, while structural analysis was performed using standard methods for light and scanning electron microscopy. The presence of sugars in the secretion was assessed by thin-layer chromatography and glucose strip tests. Nectaries in P. burchelli are non-structured glands on the adaxial surface of floral bracts and sepals. Bracts and sepals are distinct spatial units that act over time in the same strategy of floral bud protection. Literature data reveals that ENNs might be more common within Bromeliaceae than previously considered, comprising a homoplastic feature in the family. Future perspectives and evolutionary and taxonomic implications are discussed.

Colleters, Extrafloral Nectaries, and Resin Glands Protect Buds and Young Leaves of Ouratea castaneifolia (DC.) Engl. (Ochnaceae).

Buds usually possess mechanical or chemical protection and may also have secretory structures. We discovered an intricate secretory system in Ouratea castaneifolia (Ochnaceae) related to the protection of buds and young leaves. We studied this system, focusing on the distribution, morphology, histochemistry, and ultrastructure of glands during sprouting. Samples of buds and leaves were processed following the usual procedures for light and electron microscopy. Overlapping bud scales protect dormant buds, and each young leaf is covered with a pair of stipules. Stipules and scales possess a resin gland, while the former also possess an extrafloral nectary. Despite their distinct secretions, these glands are similar and comprise secreting palisade epidermis. Young leaves also possess marginal colleters. All the studied glands shared some structural traits, including palisade secretory epidermis and the absence of stomata. Secretory activity is carried out by epidermal cells. Functionally, the activity of these glands is synchronous with the young and vulnerable stage of vegetative organs. This is the first report of colleters and resin glands for O. castaneifolia. We found evidence that these glands are correlated with protection against herbivores and/or abiotic agents during a developmental stage that precedes the establishment of mechanical defenses.

True nectar or stigmatic secretion? Structural evidence elucidates an old controversy regarding nectaries in Anthurium.

PREMISE: Floral rewards are essential in understanding floral function and evolution of the relationships between flowers and pollinators. Whether sugars are present in stigmatic exudates in Anthurium and whether it has floral nectaries have remained controversial because of the scarcity of structural studies. To solve these questions, we investigated the floral anatomy of A. andraeanum to elucidate whether (1) tepals are secretory organs, (2) tepals possess a structurally recognizable nectary, and (3) tepalar secretion differs from stigmatic secretion. METHODS: Floral structure was assessed through light and electron microscopy of samples of immature, pistillate, and staminate flowers. The dynamics of the starch reserve was investigated using histochemical tests, and the sugar content in the floral exudates was assessed using thin-layer chromatography. RESULTS: Sugar analysis did not detect sucrose, glucose, or fructose in stigmatic secretions, but confirmed their presence in tepalar secretions. Stigmatic secretion was produced by secretory stigmatic papillae; tepalar exudates were produced by nonvascularized nectaries in the apex of tepals. These nectaries were characterized by modified stomata and cells with cytoplasm rich in organelles, and a high content of calcium oxalate crystals. CONCLUSIONS: Our results showed for the first time nectaries on tepals and true nectar secretion for A. andraeanum. Stigmatic secretion appears to be a distinct substance, and its often-reported sugar content seems to be a result of sample contamination. Nectar and stigmatic secretions have been often mistaken in other Anthurium species and deserve a revision for this genus.

More than Scales: Evidence for the Production and Exudation of Mucilage by the Peltate Trichomes of Tillandsia cyanea (Bromeliaceae: Tillandsioideae).

Bromeliad scales have been investigated extensively due to their recognition as a key ecological and evolutionary feature of Bromeliaceae. However, much remains unknown about such trichomes and only recently mucilage exudation was described for them in a species of the subfamily Bromelioideae. The present study aimed to investigate the secretion present in inflorescences of Tillandsia cyanea Linden ex K. Koch (Tillandsioideae) to determine whether the scales of this species also produce and release secretions. Samples of young and mature portions of inflorescences were collected and prepared according to standard methods for light and electron microscopy. Anatomical and ultrastructural results indicate that the secretion is produced by the wing portion of typical peltate trichomes on the adaxial surface of bracts. The secretory activity begins in the early stages of trichome expansion and characteristically occurs in cells exhibiting a porous cuticle and dense cytoplasm with numerous mitochondria and dictyosomes. Histochemical tests confirmed mucilage secretion and revealed proteins in the exudate. These data comprise the first record of mucilage exudation by trichomes within Tillandsioideae and indicate that this capacity may be more relevant to bromeliad biology than previously considered. Functional aspects and colleter-like activity are also discussed.