Morphology of floral papillae in Maxillaria Ruiz & Pav. (Orchidaceae)

BACKGROUND AND AIMS: The labellar papillae and trichomes of Maxillaria Ruiz & Pav. show great diversity. Although papillae also occur upon other parts of the flower (e.g. column and anther cap), these have not yet been studied. Labellar trichomes of Maxillaria are useful in taxonomy, but hitherto the taxonomic value of floral papillae has not been assessed. The aim of this paper is to describe the range of floral papillae found in Maxillaria and to determine whether papillae are useful as taxonomic characters. METHODS: Light microscopy, histochemistry, low-vacuum scanning and transmission electron microscopy. KEY RESULTS: A total of 75 taxa were studied. Conical papillae with rounded or pointed tips were the most common. The column and anther cap usually bear conical, obpyriform or villiform papillae, whereas those around the stigmatic surface and at the base of the anther are often larger and swollen. Labellar papillae show greater diversity, and may be conical, obpyriform, villiform, fusiform or clavate. Papillae may also occur on multiseriate trichomes that perhaps function as pseudostamens. Labellar papillae contain protein but most lack lipid. The occurrence of starch, however, is more variable. Many papillae contain pigment or act as osmophores, thereby attracting insects. Rewards such as nectar or a protein-rich, wax-like, lipoidal substance may be secreted by papillae onto the labellar surface. Some papillae may have a protective role in preventing desiccation. Species of diverse vegetative morphology may have identical floral papillae, whereas others of similar vegetative morphology may not. CONCLUSIONS: Generally, floral papillae in Maxillaria have little taxonomic value. Nevertheless, the absence of papillae from members of the M. cucullata alliance, the occurrence of clavate papillae with distended apices in the M. rufescens alliance and the presence of papillose trichomes in some species may yet prove to be useful.     

Nectary structure and nectar secretion in Maxillaria coccinea (Jacq.) L.O. Williams ex Hodge (Orchidaceae)

BACKGROUND AND AIMS: It had previously been assumed that Maxillaria spp. produce no nectar. However, nectar has recently been observed in Maxillaria coccinea (Jacq.) L.O. Williams ex Hodge amongst other species. Furthermore, it is speculated that M. coccinea may be pollinated by hummingbirds. The aim of this paper is to investigate these claims further. METHODS: Light microscopy, histochemistry, scanning and transmission electron microscopy. KEY RESULTS: This is the first detailed account of nectar secretion in Maxillaria Ruiz & Pav. A 'faucet and sink' arrangement occurs in M. coccinea. Here, the nectary is represented by a small protuberance upon the ventral surface of the column and nectar collects in a semi-saccate reservoir formed by the fusion of the labellum and the base of the column-foot. The nectary comprises a single-layered epidermis and three or four layers of small subepidermal cells. Beneath these occur several layers of larger parenchyma cells. Epidermal cells lack ectodesmata and have a thin, permeable, reticulate cuticle with associated swellings that coincide with the middle lamella between adjoining epidermal cells. Nectar is thought to pass both along the apoplast and symplast and eventually through the stretched and distended cuticle. The secretory cells are collenchymatous, nucleated and have numerous pits with plasmodesmata, mitochondria, rough ER and plastids with many plastoglobuli but few lamellae. Subsecretory cells have fewer plastids than secretory cells. Nectary cells also contain large intravacuolar protein bodies. The floral morphology of M. coccinea is considered in relation to ornithophily and its nectary compared with a similar protuberance found in the entomophilous species M. parviflora (Poepp. & Endl.) Garay. CONCLUSIONS: Flowers of M. coccinea produce copious amounts of nectar and, despite the absence of field data, their morphology and the exact configuration of their parts argue strongly in favour of ornithophily.     

Pseudopollen and food-hair diversity in Polystachya Hook. (Orchidaceae)

Labellar food-hairs in Polystachya Hook. exhibit considerable morphological diversity. The commonest type of trichome is uniseriate, two to four-celled, with a clavate or subclavate terminal cell. This type occurs amongst representatives of most sections examined. Other trichomes are bristle-like with tapering or fusiform terminal cells, whereas representatives of section Polystachya have uniseriate, moniliform trichomes that fragment with the formation of rounded or elliptical component cells. Most contain protein and, while some contain starch, lipid is invariably absent. The presence of particular types of labellar trichomes does not coincide with variations in vegetative morphology. Thus, current taxonomic treatment of the genus indicates that trichome types, with perhaps the sole exception of moniliform, pseudopollen-forming hairs found in section Polystachya only, have limited taxonomic value. However, the remarkable similarity between pseudopollen-forming hairs of Polystachya and those of the Neotropical genus Maxillaria in terms of morphology, cellular dimensions and food content indicates that pseudopollen may have arisen several times and evolved in response to similar pollinator pressures.     

Nectar-secreting floral stomata in Maxillaria anceps Ames & C. Schweinf. (Orchidaceae)

BACKGROUND AND AIMS: Although it was generally assumed that Maxillaria spp. do not produce nectar, in recent years, nectar has been reported for a number of these orchids. Nevertheless, our current understanding of nectary structure and nectar secretion in Maxillaria is based solely on M. coccinea (Jacq.) L.O. Williams ex Hodge, which, since it shows many features characteristic of ornithophilous flowers, is atypical of this largely entomophilous genus. The aim of the present paper is to describe, for the first time, nectar secretion in a presumed entomophilous species of Maxillaria. METHODS: The structure of the nectary of M. anceps Ames & C. Schweinf., nectar composition and the process of nectar secretion were investigated using light microscopy, scanning electron microscopy, transmission electron microscopy, histochemistry, refractometry and high performance liquid chromatography. KEY RESULTS AND CONCLUSIONS: Nectar appears as droplets that are exuded by modified stomata borne upon the labellar callus and collects upon the labellum and at the base of the column-foot. Although such stomata are known to occur in a number of angiosperm families, this is the first time for them to be observed in orchids. The callus consists largely of parenchyma with raphides and is supplied by eight to ten collateral bundles. This tissue, together with the single-layered epidermis, seemingly contains terpenoids. During the bud stage, the callus cells contain an organelle complement consistent with secretory cells whereas by day 4 of anthesis, much of the cell is occupied by a vacuole. The nectar is sucrose-dominant but also contains low concentrations of glucose, fructose, free amino acids and possibly terpenoids. The high sugar concentration (approx. 66 %) is consistent with melittophily and may indicate that, like the majority of Maxillaria spp., M. anceps is visited by stingless bees (Meliponini).     

Pseudopollen in Eria Lindl. section Mycaranthes Rchb.f. (Orchidaceae)

BACKGROUND AND AIMS: Pseudopollen is a whitish, mealy material produced upon the labella of a number of orchid species as labellar hairs either become detached or fragment. Since individual hair cells are rich in protein and starch, it has long been speculated that pseudopollen functions as a reward for visiting insects. Although some 90 years have passed since Beck first described pseudopollen for a small number of Eria spp. currently assigned to section Mycaranthes Rchb.f., we still know little about the character of pseudopollen in this taxon. The use of SEM and histochemistry would re-address this deficit in our knowledge whereas comparison of pseudopollen in Eria (S.E. Asia), Maxillaria (tropical and sub-tropical America), Polystachya (largely tropical Africa and Madagascar) and Dendrobium unicum (Thailand and Laos) would perhaps help us to understand better how this feature may have arisen and evolved on a number of different continents. METHODS: Pseudopollen morphology is described using light microscopy and scanning electron microscopy. Hairs were tested for starch, lipid and protein using IKI, Sudan III and the xanthoproteic test, respectively. KEY RESULTS AND CONCLUSIONS: The labellar hairs of all eight representatives of section Mycaranthes examined are identical. They are unicellular, clavate with a narrow 'stalk' and contain both protein and starch but no detectable lipid droplets. The protein is distributed throughout the cytoplasm and the starch is confined to amyloplasts. The hairs become detached from the labellar surface and bear raised cuticular ridges and flaky deposits that are presumed to be wax. In that they are unicellular and appear to bear wax distally, the labellar hairs are significantly different from those observed for other orchid species. Comparative morphology indicates that they evolved independently in response to pollinator pressures similar to those experienced by other unrelated pseudopollen-forming orchids on other continents.     

Atypical pseudopollen-forming hairs in Maxillaria (Orchidaceae)

Pseudopollen is found on the labella of several species of Maxillaria and is formed by the fragmentation of uniseriate, multicellular, moniliform trichomes. The resultant component cells are rich in protein. Since flowers of pseudopollen-forming species generally lack nectar, it is probable that pseudopollen gradually replaced nectar as the pollinator reward. However, direct evidence for this hypothesis is lacking. The present paper examines the labellar micromorphology of five Brazilian species of Maxillaria assigned to the M. discolor alliance. The flowers of two of these species, namely M. violaceopunctata and M. villosa , produce both food hairs and a lipoidal, labellar secretion which is rich in aromatic amino acids. Moreover, in the case of M. violaceopunctata , the secretion was found to contain reducing sugars. This may represent an intermediate stage in the transition from nectar-producing to pseudopollen-forming flowers. SEM studies indicate how pseudopollen-forming, moniliform trichomes, such as those found in the Maxillaria grandiflora complex, may have evolved from simple, uniseriate, multicellular hairs.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 143 , 151−158.     

A Rudimentary labellar speculum in Cymbidium lowianum (Rchb.f.) Rchb.f. and Cymbidium devonianum Paxton (Orchidaceae)

BACKGROUND AND AIMS: The labellar 'hairs' of some Cymbidium spp. are said to be thin-walled and to contain 'plasma', oil and sugars and it has long been speculated that they may function as food-hairs. However, the present authors' preliminary studies showed that certain atypical papillae may have a different role and, by reflecting light, function as a speculum. The purpose of the paper is to test this hypothesis. METHODS: Light microscopy, scanning electron microscopy, transmission electron microscopy, histochemistry and ultraviolet photography were used to investigate the structure, food content and light-reflecting properties of these papillae. KEY RESULTS AND CONCLUSIONS: The labellum of Cymbidium lowianum (Rchb.f.) Rchb.f. is densely clothed with obconical to conical papillae with wide bases and pointed tips. However, on either side of the median axis of the lip occur silvery patches comprising papillae with truncated tips and it is thought that these reflect light and thereby attract insect pollinators. Similar patches are also found in Cymbidium devonianum Paxton, and in both species, they are set against a reddish background, which, since bees cannot perceive this colour, probably appears dark to the insect thus enhancing the visual impact of the light-reflecting patches. In Cymbidium tigrinum Parish ex Hook. and Cymbidium mastersii Griff. ex Lindl., however, the labellum is mainly white and no light-reflecting patches were observed. Instead, unlike C. lowianum and C. devonianum, these species are highly fragrant and the attraction of insects probably depends to a greater extent on olfactory cues. In C. lowianum both types of papillae contain protein, starch and lipid bodies but only protein is seemingly present at elevated concentrations. However, lipoidal material also occurs upon the surface of the labellum and it is possible that this may be gathered by insects as reported for C. iridifolium A. Cunn (syn. C. madidum Lindl.). The labellar papillae of C. lowianum, thus, have the potential to function as food-hairs, although direct evidence for this is lacking.     

Comparative account of nectary structure in Hexisea imbricata (Lindl.) Rchb.f. (Orchidaceae)

• Background and Aims Despite the number of orchid speciesthat are thought to be pollinated by hummingbirds, our knowledgeof the nectaries of these orchids is based solely on a singlespecies, Maxillaria coccinea (Jacq.) L.O. Williams ex Hodge.Nevertheless, it is predicted that such nectaries are likelyto be very diverse and the purpose of this paper is to comparethe nectary and the process of nectar secretion in Hexisea imbricata(Lindl.) Rchb.f. with that of Maxillaria coccinea so as to beginto characterize the nectaries of presumed ornithophilous Neotropicalorchids. • Methods Light microscopy, transmission electronmicroscopyand histochemistry were used to examine the histology and chemicalcomposition of nectary tissue and the process of nectar secretionin H. imbricata. • Key Results and Conclusions The nectary of H. imbricatahas a vascular supply, is bound by a single-layered epidermiswith few stomata and comprises two or three layers of subepidermalsecretory cells beneath which lie several layers of palisade-likeparenchymatous cells, some of which contain raphides or mucilage.The secretory cells are collenchymatous and their walls havenumerous pits with associated plasmodesmata. They contain thefull complement of organelles characteristic of secretory cellsas well as intravacuolar protein bodies but some of the secretoryepidermal cells, following secretion, collapse and their anticlinalwalls seem to fold. Nectar secretion is thought to be granulocrineand, following starch depletion, lipid droplets collect withinthe plastids. The nectar accumulates beneath the cuticle whichsubsequently forms swellings. Finally, nectar collects in thesaccate nectary spur formed by the fusion of the margins ofthe labellum and the base of the column-foot. Thus, althoughthe nectary of H. imbricata and M. coccinea have many featuresin common, they nevertheless display a number of important differences.     

Lip anatomy and its implications for the pollination mechanisms of Bulbophyllum species (Orchidaceae)

BACKGROUND AND AIMS: The lip structure of six Brazilian and one Asiatic species of Bulbophyllum with wind-assisted fly pollination (B. involutum, B. ipanemense and B. weddellii) and non-wind-assisted fly pollination (B. epiphytum, B. glutinosum, B. regnellii and B. rothschildianum) was studied to investigate the presence of secretory tissues related to these pollination mechanisms. METHODS: The lip study was carried out through scanning electron microscopy (lip surface) and light microscopy (anatomical features). KEY RESULTS: In most of the species studied, the osmophores (odour glands) were located in the lobes and in the upper surface of the lip callus. Differences in the lip structure were observed between the two groups (the presence of a nectary and the extent of osmophore surface), depending on the mechanism of pollination. Nectaries were found in the cavity callus in B. ipanemense, B. involutum and B. weddellii, even though their pollinators were presumably attracted by the instinct to oviposit. CONCLUSIONS: These findings corroborate the hypothesis that, because pollination in these species is dependent on an unpredictable external factor (wind), nectar is necessary to keep the insect in the flower for a long period. Despite the occurrence of a liquid-like nectar in the flowers of B. epiphytum, B. glutinosum, B. regnelli and B. rothschildianum, no anatomical evidence for nectaries was found in the lips of these species, although a similar structure may occur in another region of the flowers. This observation agrees with the fact that pollination by lip movement in the latter species requires only gravity, with no additional mechanism being needed to keep the flies in the flower.     

Sexual mimicry in Mormolyca ringens (Lindl.) Schltr. (Orchidaceae: Maxillariinae)

BACKGROUND AND AIMS: Pollination through sexual mimicry, also known as pseudocopulation, has been suggested to occur in some genera of the Neotropical orchid subtribe Maxillariinae. However, it has been demonstrated so far only for Trigonidium obtusum. This study reports and illustrates pollination through sexual mimicry in Mormolyca ringens. METHODS: A total of 70 h were dedicated to the observation of flowers and pollinator behaviour, which was photographically recorded. Flower features involved in pollinator attraction were studied using a stereomicroscope and by SEM analyses. Preliminary observations on the plant breeding system were made by manually self-pollinating flowers. The chemical composition of the fragrance volatiles was determined by GC/MS analysis. KEY RESULTS: The flower features of M. ringens parallel those of other pseudocopulatory flowers. The labellum shape and indument are reminiscent of an insect. Sexually excited drones of Nannotrigona testaceicornis and Scaptotrigona sp. (both in the Apidae: Meliponini) attempt copulation with the labellum and pollinate the flower in the process. In both bee species, the pollinarium is attached to the scutellum. Pollinator behaviour may promote some degree of self-pollination, but preliminary observations indicate that M. ringens flowers are self-incompatible. Flowers are produced all the year round, which ties in with the production of bee males several times a year. The phylogenetic relationships of M. ringens are discussed and a number of morphological and phenological features supporting them are reported. CONCLUSIONS: It is expected that further research could bring to light whether other Maxillariinae species are also pollinated through sexual mimicry. When a definitive and robust phylogeny of this subtribe is available, it should be possible to determine how many times pseudocopulation evolved and its possible evolutionary history.     

The pollination mechanism in Trigonidium obtusum Lindl (Orchidaceae: Maxillariinae): sexual mimicry and trap-flowers

The pollination process in Trigonidium obtusum Lindl. (Epidendroideae: Maxillariinae) is documented. The flowers are pollinated by sexually excited drones of Plebeia droryana (Meliponinae). When attempting to copulate either with sepals or petals, these bees slip on the waxy perianth surface and become trapped in the funnel-like flower tube. Bees trying to escape from the flowers may instead access the space between the column and lip, fixing the pollinarium on their scutellum. Pollinarium-bearing bees may pollinate the flowers when repeating the above-mentioned steps, leaving pollinia on the concave stigmatic surface, thus effecting pollination. Recently removed pollinaria are too broad to enter the stigma but they begin to dehydrate and within 40 min of removal are small enough to fit the stigmatic cavity. This mechanism prevents insect-mediated self-pollination and promotes cross-pollination. Preliminary evidence based on experiments with cultivated plants suggests that they are self-compatible but that fruit set is pollinator-dependent. The data obtained are discussed in a phylogenetic context. It is suggested that the pseudocopulatory syndrome in Trigonidium could have evolved from rewardless (food advertising) ancestors. Pseudocopulation in the context of the long flowering period of this orchid species (about 7 months) is understandable since the eusocial Plebeia bees produce fertile individuals several times a year.     

苹毛丽金龟触角嗅感器超微结构

利用扫描电镜和透射电镜对苹毛丽金龟Proagopertha lucidula (Faldermann)雌雄成虫触角嗅感器超微结构特征进行研究。结果表明:嗅感器集中于触角鳃片上,分布在触角鳃片表皮内陷形成的凹腔里,有5种感器,以板状感器为主,以外还有锥状感器、腔状感器、腔锥状感器和毛状感器。板状感器根据盘体形状的不同可分为4种类型,锥状感器根据锥体形状的差异可分为2种类型,腔锥状感器根据形状分为2种类型。嗅感器表皮有微孔和孔道微管。嗅感器内神经元的数目并不一致,1～2个不等。雄虫鳃片嗅感器总数显著多于雌性,是雌虫嗅感器总数1.8倍。其中雄性锥状感器的数目是雌性2倍,雌虫腔锥状感器数量是雄虫的4倍,雄虫板状感器数量是雌虫的2倍,雌雄腔状感器的数目无显著性差异。     

Structure and development of stellate trichomes in Andradea ftoribunda Fr. Allem. (Nyctaginaceae)

LOURO, R. P., MIGUENS, F. C. & MACHADO, R. D., 1992. Structure and development of stellate trichomes in Andradea ftoribunda Fr. Allem. (Nyctaginaceae). Trichomes occur on both faces of young leaves. They are peltate-stellate on the abaxial face, and comprise a stalk and radiating cells with a rudimentary central apex. On the adaxial face the trichomes arc stellate with a large apex comprising one to three cells. In both cases the stalk is formed by three to six cells of which the most distal may contain a tannoid substance. In the adult leaf only the abaxial surface exhibits stellate trichomes, with two to three celled stalks. The central region of radial cells is depressed. On the adaxial side the hairs are shed during maturation of the leaf.     

Adhesion by Paired Pectoral and Pelvic Fins in a Mountain-Stream Catfish, Pseudocheneis sulcatus (Sisoridae)

We describe the adhesive nature of the pectoral and pelvic fins in the catfish Pseudocheneis sulcatus, as examined by scanning electron microscopy. The outer rays of these fins are modified into structures that bear prominent transverse ridges and grooves. The outer epidermal cells of the ridges are thrown into elongated spines. Mucous pores (openings of mucous glands) are frequently present (100m apart) in the epidermis of the ridges and show entangled mucus droplets. In the pectoral fins, they are present towards the contour areas of the outer rays, but they are absent in the pelvic fins. In the latter, mucous pores are present near the base of the ridges (distal to the inner rays). Spines as well as mucous pores are absent in the cells that line the groove between two adjacent ridges. We suggest that in this species adhesion is effected by suction pressure generated by the musculature attached to the grooves and ridges, and that mucus and the spines aid in this process.     

Study by transmission and scanning electron microscopy of the morphogenesis of three types of lingual papillae in the albino rat (Rattus rattus)

El‐Bakry, A.M. 2010. Study by transmission and scanning electron microscopy of the morphogenesis of three types of lingual papillae in the albino rat (Rattus rattus).—Acta Zoologica (Stockholm) 91 : 267–278 Tongues were removed from albino rat foetus on days 12 (E12) and 16 (E16) of gestation and from newborns (P0) and from juvenile rats on days 7 (P7), 14 (P14) and 21 (P21) postnatally for investigation by light, scanning, and transmission electron microscopy. Significant changes appeared during the morphogenesis of the papillae. At E12, two rows of rudiments of fungiform papillae were extended bilaterally on the anterior half of the tongue. At E16, the rudiments of fungiform papillae were regularly arranged in a lattice‐like pattern. A rudiment of circumvallate papillae could be recognized. No rudiment of filiform papillae was visible. No evidence of keratinization was recognizable. At P0, rudiments of filiform papillae were visible but had a more rounded appearance, with keratinization. The fungiform and circumvallate papillae were large and their outlines were somewhat irregular as that found in the adult rat. At P7, the filiform papillae were large and slender. The fungiform papillae became large and the shape of circumvallate papillae was almost similar to that observed in the adult. At P14 and P21, the shape and structure of the three types of papillae were irregular as those found in the adult. In conclusion, the rudiments of the fungiform and circumvallate papillae were visible earlier than those of the filiform papillae. The morphogenesis of filiform papillae advanced in a parallel manner with the keratinization of the lingual epithelium, in the period from just before birth to a few weeks after birth.     

Effect of individual height and labellum colour on the pollination of Caladenia (syn. Arachnorchis) behrii (Orchidaceae) in the northern Adelaide region, South Australia

Caladenia behrii, a sexually deceptive orchid, is an endangered terrestrial orchid endemic to the Adelaide Hills, South Australia. We examined pollination success among individuals of different colours and heights in three populations of C. behrii in the northern Adelaide region. The labellum of C. behrii varies from cream to deep maroon. Over five years, the pollination success (pollinia deposited on stigma) of four colour groups (maroon, maroon and cream, hatched, and cream) was assessed for a total of 782 flowers. Labellum colour had no effect on the pollination success of individuals, indicating no frequency-dependent selection. It is likely that pollinators use the sexual pheromone mimic produced by the flowers as the main orientation cue. For three years the height of pollinated C. behrii individuals was recorded, along with the height of the nearest non-pollinated neighbour (distance of labellum from the ground, rather than stem length). Although a range of different heights was pollinated at population level, individual flowers that were taller than their closest neighbours were significantly more likely to be pollinated. Flower head height was not correlated with opening date and was greater for orchids growing among grass-tree leaves. Results imply that pollinators are more successful in finding tall flowers than short ones for visual and/or olfactory reasons; however, the effect of height preference is eliminated at population level.     

纤毛虫念珠异列虫射出胞器的超微结构观察

应用扫描电镜术和透射电镜术显示,纤毛虫念珠异列虫(Anteholosticha monilata)的射出胞器早期发生在细胞质深处,附近有不同类型的囊泡结构。成熟后射出胞器向表膜迁移,结构由不同电子密度片层的体部、结晶状的中心轴杆部和多层膜的帽部组成。受外界刺激时胞器冲破皮层射出,形态呈"蘑菇"状。据上述观察结果推测:该射出胞器具有防御作用,它可能起源于高尔基体活动产生的小泡;在亲缘关系较近的纤毛虫中,其射出胞器可能具有相似的分化特征。     

Ultrastructural studies of the hearts in Arenicola marina L. (Annelida: Polychaeta)

Summary The two hearts in Arenicola are capable of great dilation and contraction. The heart wall consists of myoepithelial cells resting on a basal lamina. On the luminal side of the basal lamina is a layer of collagen fibrils. No true endothelium was observed but occasional haemocytes were observed, subjacent to the collagenous layer. A few chloragogen cells are also found peripherally.The myofibrils are of a non-striated type consisting of thick and thin filaments and scattered Z-bodies. The sarcoplasmic reticulum forms a three-dimensional network. Only peripheral couplings were observed. The myofibrils are in contact with the sarcolemma on the luminal side of the cells, constituting a kind of hemidesmosome. The myoepithelial heart muscle is compared with other muscle types described in invertebrates. Supercontraction is discussed.