title: Labellar anatomy and secretion in Bulbophyllum Thouars (Orchidaceae: Bulbophyllinae) sect. Racemosae Benth. & Hook. f.
abstract: OBJECTIVE
  Floral secretions are common in Bulbophyllum Thouars, and the labella of a number of Asian species are said to produce secretions rich in lipids that act as food rewards for insect pollinators. Although some of these reports are based on simple histochemical tests, a much greater number are anecdotal and, hitherto, neither the ultrastructure of the labellum nor the secretory process has been investigated in detail. Furthermore, sophisticated histochemical approaches have generally not been applied. Here, both the labellar structure and the secretory process are investigated for four species of Asian Bulbophyllum sect. Racemosae Benth. & Hook. f., namely Bulbophyllum careyanum (Hook.) Spreng., B. morphologorum Kraenzl., B. orientale Seidenf. and B. wangkaense Seidenf., and compared with those of unequivocal lipid-secreting orchids.
  
  
  METHODS
  Labellar, secretory tissue was investigated using light microscopy, scanning electron microscopy, transmission electron microscopy and histochemistry.
  
  
  RESULTS
  The adaxial median longitudinal groove of the labellum contained secretory tissue comprising palisade-like epidermal cells, similar to those of certain lipid-secreting Oncidiinae Benth. However, these cells and their secretions gave positive results mainly for protein and mucilage, and their organelle complement was consistent with that of cells involved in protein and mucilage synthesis. Sub-cuticular accumulation of secretion resulted in cuticular distension and blistering. The sub-epidermal layer of isodiametric parenchyma contained starch and, like the epidermal cells, ultrastructure consistent with mucilage synthesis. Lipids were mainly confined to the cuticle, and hardly any intracellular lipid droplets were observed.
  
  
  CONCLUSIONS
  It is proposed that mucilage is produced by dictyosomes present in the palisade-like epidermal cells. Mucilage precursors may also be produced by these same organelles in sub-epidermal cells and are thought to pass along the symplast via plasmodesmata into the adjoining palisade-like secretory cells, which contain abundant arrays of rough endoplasmic reticulum. Here, they become chemically modified and form a protein-rich, mucilaginous secretion that, following vesicle-mediated transport across the cytoplasm, traverses the cell wall and accumulates in blisters formed from the distended cuticle. Rupture of these blisters releases the secretion onto the labellar surface. However, in certain species, there is some evidence that the secretion may traverse the cuticle via cuticular pores, and micro-channels may permit the passage of fragrance. Hydrolysis of sub-epidermal starch probably generates the carbohydrate and, together with mitochondria, much of the energy required for the secretory process. This anatomical organization resembles that found in certain lipid-secreting, Neotropical species of Bulbophyllum and Oncidiinae, but since the chemical composition of their secretions is different, and these taxa occur on a separate continent and have different insect pollinators, parallelism of floral anatomy is likely.
authors.affiliation: School of Earth and Ocean Sciences, Cardiff University, Main Building, Park Place, Cardiff CF10 3AT, UK kevinldavies@btinternet.com.
authors.forename: Kevin L
authors.initials: KL
authors.lastname: Davies
authors.affiliation: University of Warsaw, Faculty of Biology, Botanic Garden Al. Ujazdowskie 4, 00-478 Warsaw, Poland.
authors.forename: Malgorzata
authors.initials: M
authors.lastname: Stpiczyńska
mesh_terms: D000818:Animals; D002473:Cell Wall / Q000648:ultrastructure; D035264:Flowers / Q000033:anatomy & histology* / Q000648:ultrastructure; D007313:Insecta / Q000502:physiology*; D050356:Lipid Metabolism; D008855:Microscopy, Electron, Scanning; D046529:Microscopy, Electron, Transmission; D029595:Orchidaceae / Q000033:anatomy & histology* / Q000648:ultrastructure; D054817:Pollination; D013045:Species Specificity; D008055:Lipids
chemical_list: D008055:Lipids
keywords: Anatomy; Bulbophyllinae; Bulbophyllum; Orchidaceae; histochemistry; lipid; micromorphology; mucilage; orchid; protein; secretion; sect. Racemosae; ultrastructure