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Resin-secreting, sunken, glandular trichomes have been described from the vegetative organs of Maxillaria and certain related taxa. However, until now, these hairs have not been recorded for Maxillaria flowers, nor have they been examined at the ultrastructural level for any part of the plant. Here for the first time, these hairs are described using LM, SEM and TEM, and their possible functions discussed.
Floral epidermal cells of most species of Bulbophyllum Thouars studied to date produce both lipid-rich foodrewards and fragrance. Since fragrances largely consist of terpenoids and have an affinity for lipophilic stains, the simultaneous presence of lipid-rich food-rewards frustrates identification of fragrance-secreting cells by conventional histochemistry. Furthermore, since both lipid-rich food-rewards and fragrances are probably synthesized by a similar complement of organelles, interpretation of TEM images can prove difficult. All members of section Racemosae Benth. & Hook. f. investigated to date, however, are unusual in their secretion of a predominantly proteinaceous food-reward, and lipids are seemingly absent. This might enable their use as models for the identification and characterization of fragrance-secreting tissues and organelles. Three members of sect. Racemosae were chosen, namely Bulbophyllum dissitiflorum Seidenf., B. lilacinum Ridl. and B. tricorne Seidenf. & Smitinand. All produced food-rewards. Of these, one (B. dissitiflorum) lacked fragrance and was used as a control, whereas the remaining two species produced fragrance. Having established that the food-reward was mainly proteinaceous in each case, and did not test positively for lipid, we undertook further histochemical investigations, as well as light microscopy, SEM and TEM. Specialized palisade-like epidermal cells of all species contained protein bodies and rough endoplasmic reticulum consistent with the production and secretion of a protein-rich food-reward. Cuticular pores were also present. In fragrant species, these cells also contained abundant smooth endoplasmic reticulum, oil droplets and many, well-developed, spherical plastids with numerous plastoglobuli, similar to those found in the osmophores (fragrance-producing structures) of other orchids. Indeterminate, osmiophilic cytoplasmic inclusions were also present. By contrast, the non-fragrant species lacked oil droplets and other osmiophilic inclusions and the plastids were scant, poorly developed, often elongate or irregular in shape and contained few plastoglobuli. Smooth endoplasmic reticulum was also less frequent. Since food-rewards tested negatively for lipid, it is probable that any oil droplets present were involved in fragrance production, especially since they were absent from the non-fragrant species. Thus, the unusual absence of lipids from the food-rewards of sect. Racemosae provided a rare opportunity, permitting, for the first time, the unraveling of these two secretory processes (food-reward and fragrance) in Bulbophyllum and clearly demonstrating the plasticity of these cells and their dual role in secretion.
Ornithophily occurs in a great number of orchid species but despite this, researchers have largely neglected to investigate their nectaries. The aim of this study is to describe the nectary structure of Symphyglossum sanguineum, a species presumed to be pollinated by hummingbirds. The nectary is located at the free margins of auricles, which form a channel for the passage of nectar. The nectary, which consists of a single-layered epidermis and 2-3 layers of subepidermal cells, is supplied by collateral, vascular bundles. The nectary cells of S. sanguineum, like those of other ornithophilous orchids, have thick cellulose cell walls. A remarkable feature of these nectary cells is the dissolution of the middle lamella and the subsequent separation of epidermal cells. It is possible that this latter process facilitates the flow of the nectar to the nectary surface. The cuticle covering the nectary epidermis has micro-channels, but unlike the other species of ornithophilous orchids studied to date, it neither becomes disrupted nor detached from the epidermal cells. Abundant mitochondria, lipid droplets and smooth endoplasmic reticulum (SER) with an osmiophilic material are present in the cytoplasm of nectary cells. Some plastids with few lamellae contain numerous vesicles and osmiophillic globules whereas others accumulate starch. SER lamellae are often closely associated with plastids and the contents of the former organelles closely resemble osmiophillic globules. Secretory vesicles are common, especially near the outer, tangential wall indicating that granulocrine secretion possibly occurs in S. sanguineum.
Most orchids do not produce floral food-rewards. Instead, they attract pollinators by mimicry or deceit. When present, the most common floral food-reward is nectar. To date, nectary structure has been described for only two species of Maxillaria sensu lato, namely Maxillariella anceps and Ornithidium coccineum (formerly Maxillaria anceps and M. coccinea, respectively). Here, we describe that of a third species, Ornithidium sophronitis (formerly Maxillaria sophronitis). This species possesses floral characters concomitant with ornithophily. A ‘faucet and sink’ arrangement is present, with nectar secreted by a protuberance on the ventral surface of the column, collecting between column and tepal bases. The nectary of O. sophronitis shares many features with that of O. coccineum. It has a single-layered epidermis and 3- 5 layers of small, subepidermal, collenchymatous, secretory cells. Beneath these occur 2-3 layers of larger, subsecretory, parenchymatous cells supplied by phloem. Nectary cell vacuoles contain osmiophilic material and proteinaceous intravacuolar bodies. Moreover, distension of the nectary cuticle occurs as nectar accumulates between it and the secretory epidermis. Subsecretory cells, however, have thinner walls and contain flocculent, intravacuolar precipitates that may be related to the presence of flavonoids. Since the floral and nectary structure of O. sophronitis is very similar to that of closely related Ornithidium coccineum, it may have evolved in like manner in response to similar pollinator pressures.
To date, the structure of the cuniculus nectary has not been studied in detail. Furthermore, the secretory mechanism of such nectaries has not been investigated. The present paper describes, for the first time, the structural organization and ultrastructure of the cuniculus nectary in the moth-pollinated orchid Brassavola flagellaris Barb. Rodr. This tubular structure is situated between the perianth tube and ovary and, in its possession of thick, cellulose cell walls, resembles the nectary of ornithophilous taxa. The presence of large secretory vesicles that fuse with the plasmalemma indicate that granulocrine nectar secretion occurs in this species. The lumen of the cuniculus is lined with unicellular hairs. However, the cuticle overlying the whole epidermal surface lining the lumen (both glabrous and pubescent regions) was coated with nectar residues and became distended and cracked, indicating that this entire tissue is probably involved in nectar secretion.
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