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It's our way of making sure we're not accidentally sending you spam. Saturday, July 6, Botany One. Reductions in herkogamy with flower positions may be expected in environments with either low pollinator abundance or low nutrients. The model consists of a sympodial succession of equivalent sympodial units metamers , each of which is orthotropic and determinate in its growth.

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The comparison pollen viability determination among 17 ornamental trees of Prunus. PCR-based for identifying the S-genotypes of Japanese pear cultivars. Identification and partial amino acid sequences of seven S-RNases associated with self-incompatibility of Japanese pear Pyrus pyrifolia Nakai. Lewis D Sexual Incompatibility in Plant. Identification and sequencing of seven new S-RNase genes from Pyrus pyrifolia. Study of Chinese pear S-gene I. Determination of 7 cultivars of the Sgenotype and identification of two new allele of self-incompatibility.

Study on identification of Sgenotypes of Chinese pear cultivars. Characterization and molecular mechanism of gametophytic self-incompatibility in pears. Zhang S L, Hiratsuka S Effects of the stylar S-glycoproteins on the pollen germination and the tube growth in pears Pyrus serotina Rhed. C Details of the short and large bristles of the thorax. D Details of a trichome and conical epidermal cells of the callus. Photographs by C Martel. We recorded unicellular trichomes and conical cells on the adaxial epidermal surface.

Micro-sculpture, as observed by SEM, revealed that they were corrugations of the cuticle, whereas the cell wall was smooth Fig 8. Trichomes presented a prominent nucleus and dense cytoplasm and contained abundant starch grains. In contrast, the epidermis of the abaxial side was relatively flat and did not exhibit trichomes or papillate cells. Starch grains were also frequently observed in almost all the cells of the mesophyll. Raphide-containing idioblasts were scattered throughout the mesophyll.

Neutral red stained the labellum and petals, especially the central region of the flower, corresponding with the white area flanked by the dark red veins see Fig 1.

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However, metabolic activity was recorded over the whole corolla, the latter being responsible for scent emission. A Transverse section showing the adaxial and abaxial epidermis and the mesophyll. B Conical epidermal cells cc on the adaxial epidermis with dense cytoplasm cyt and conspicuous nucleus n.

C Epidermal cells with dense cytoplasm cyt. D raphide-containing idioblast id in the mesophyll. Note the chromocentres c inside the nuclei n. In flower extracts of Te. Thus, the floral odour was predominantly composed of saturated and unsaturated hydrocarbons Table 2. Alkanes and alkenes showed a chain length between 20 and 30 carbons. Numbered peaks as in Table 2. Although the genus Telipogon was suggested to be sexually deceptive more than 50 years ago [ 36 ] and although recurrently asserted since then as being pollinated by sexually aroused male tachinids e.

Thus, our study represents the first confirmed case of a sexually deceptive pollination system in the Oncidiinae by exclusive pre-copulatory behaviour involving tachinid flies. In contrast to pollinator behaviour in other sexually deceptive orchids described to date, the males do not show pseudocopulatory behaviour on Te. We have shown that the pollinia become attached to the legs of the male flies during their attempts to touch and grasp the column and callus of flowers. This behaviour is interpreted as pre-copulatory behaviour, since a similar event has been observed in Eudejeania subalpina males before landing on females Martel pers.

Males of E. In Eucelatoria bryani touching and grasping are the initial steps of pre-mating behaviour. Unfortunately, studies on the mating behaviour of tachinids are almost completely missing and, therefore, further comparisons with other tachinids are not possible. However, males of E. This is a clear indication that males perceived flowers as females, and that Telipogon peruvianus flowers use sexual deception but elicit only the first steps of copulatory behaviour in E.

Although other sexually deceptive orchids have been described in which the removal of pollinia and pollination does not involve pseudocopulation e. Therefore, to the best of our knowledge, this is the first case of pollination in sexually deceptive orchids involving only pre-copulatory behaviour without further copulation attempts. However, we do not rule out that pseudocopulation events occur on flowers of other Telipogon species. We have three assumptions that possibly explain the lack of the development of pseudocopulatory behaviour in E.

These assumptions are not exclusive and can occur together. The first assumption is supported by our results of pollinator behaviour in the presence of female dummies and flowers, in which the flower morphology is not perceived as being similar as the female morphology, since males tried to copulate with the dummies but not with the flowers. The third assumption might not be supported by the comparative results but, as the number of pseudocopulations on female dummies was extremely low, the absence of some odour cues cannot be dismissed. As is already known, macro-structure is important for the stimulation of pseudocopulation in some sexually deceptive orchids [ 57 ] and a combination of odour, visual and tactile cues is used for mating in some Diptera, e.

Drosophila [ 58 ]. Although odour cues play a key role triggering the pseudocopulatory behaviour by pollinators on flowers [ 22 , 59 — 61 ], floral morphology, in the presence of identical odour cues, may influence the frequency and duration of the pseudocopulatory behaviour by pollinators [ 57 ].

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Therefore, a combination of both odour and morphological cues is highly likely to play a role in the absence of pseudocopulatory behaviour of male pollinators. Manipulative experiments are needed in Telipogon flowers to identify the importance of odour and morphological cues in this mimicry and to understand the way that pollinators perceive both the model and mimic. In the sexually deceptive orchids studied so far, the flowers attract males for pollination by mimicking visual cues and sex pheromones of the females of their pollinators [ 22 , 59 — 63 ].

The yellow corolla might also mimic the perianth of flowers in which females wait for males. Although several plant species with a yellow corolla occur in the study area, we have only seen males chasing other tachinids and females of E. Furthermore, both Te. Although plants of Te. Therefore, we can reasonably hypothesize that Te.

If so, this would represent a new combined pollination mechanism of rendez-vous attraction and sexual deception in which the mimic imitates two models, the ligulate flowers of the host D. Thus, Telipogon has probably developed a system of multifarious floral mimicry during its evolution. This may explain why some Telipogon species, and specifically Te. A Inflorescences of a shrub of Dendrophorbium longilinguae Asteraceae. B Groups of several capitules hosting a Eudejeania aff.

Photographs by C Martel left and M Ayasse right. Telipogon peruvianus flowers seem to show an imperfect morphological mimicry as the flower parts and diverse bristle sizes differ between flowers and E. Nevertheless, this imperfect mimicry might still be enough to attract and cheat their male pollinators successfully. Therefore, we cannot exclude that the morphological structures of Te. This is supported by the finding that only a dummy triggered pseudocopulatory behaviour by males, and not flowers.

However, further experimental work is needed to evaluate whether males perceive tactilely similarly flowers and females. Telipogon flowers present some characteristics of the typical insectiform floral structures present in sexually deceptive orchids such as the presence of hairs and pronounced structures and the dull colours at the flower centre [ 24 ]; however, Telipogon has other characteristics that are not usual in sexually deceptive orchids such as petals as large as the labellum, a slight dimorphism between the petal and labellum and relatively large flowers.

Other sexually deceptive species, such as many Ophrys and Chiloglottis species, are clearly insectiform and show only a physical mimicry to the females of their pollinators [ 12 , 15 , 57 , 59 , 63 ]. Furthermore, the presence of only the olfactory stimuli is enough to trigger the mating behaviour of pollinators in the absence of tactile stimuli as shown in Chiloglottis and Drakaea [ 22 , 61 ]. Visual stimuli have been suggested as being the most important cues in attracting the pollinators of Telipogon , with olfactory stimuli not playing any role.

However, contrary to previous reports e. We have identified petals and labella as being the source of floral scent production and emission, since they display a conical epidermis, starch deposits, cells with large nuclei, dense cytoplasm and prominent chromocentres and grooves on the epidermis surface [ 50 , 66 — 68 ].

In several orchids e. Chiloglottis , Drakaea and Ophrys , odour is the most important cue for attracting pollinators and for stimulating copulatory behaviour on the labellum [ 11 , 12 , 22 , 60 — 62 , 69 ]. Alkanes and alkenes are important semiochemicals and often play a role as pheromones in insects [ 70 — 72 ]. Alkanes and alkenes are common in Ophrys , especially in Andrena -pollinated species, and are known for being responsible for triggering copulation attempts in bee males [ 32 , 60 , 69 , 73 — 75 ].

Alkenes also occur in dipteran pheromones and some fly species even use alkenes as major sex pheromone components such as tricosene and pentacosene in Drosophila , Musca and Fannia [ 58 , 71 , 76 — 79 ]. Although alkanes are rather common substances in floral bouquets; alkenes have been reported to be rare, but when present they are usually associated with specialized pollination systems involving males such as sexual deception [ 75 ].

Therefore, the occurrence of both alkanes and alkenes in flowers of Te. However, without further behavioural experiments see [ 80 ] , the role of the floral scents in Telipogon remains speculative. As in many other sexually deceptive orchids, ethological isolation barriers [ 2 ] play a primary role in the highly specific relationship between plants and their pollinators. Usually, only males of one pollinator species are attracted by most of the Ophrys species [ 30 ], Chiloglottis [ 29 ] and Drakaea [ 22 ].

Three Eudejeania species were recorded as being attracted to Te. This might be related to ethological isolation barriers, possibly originating from the composition of the floral odour blend i. Furthermore, the morphology of the flowers also plays a role as a morphological isolation barrier [ 2 ] and allows only males of one pollinator species to remove and transfer a pollinarium.

In order to do so, the stipe size should fit to the length of the fly leg and the viscidium diameter should accord with the femur diameter. Plants of Te. Attracted males usually leave one inflorescence after visiting one flower, thus preventing autogamy and allogamy. Therefore, self-pollination is reduced and pollen flow is encouraged [ 5 , 11 ]. Van der Pijl and Dodson [ 33 ] have noted that Telipogon pollination is successful, as seed pods are often found.

Our observations in P 1 have confirmed that the reproductive success of Te. Pollination events in sexually deceptive orchids are maximized by their long-lived flowers [ 10 ] and the behaviour of males competing for females [ 27 ], as occurs in Te. Differences in the pollination success between the two Telipogon patches are explained by pollinator occurrence and not because of actual ineffective attraction.

The limited presence of Eudejeania flies in P 2 might be related to differences in vegetation structure and, in particular, the low presence of nectar host plants compared with P 1 Martel pers. Our findings are the first conclusive report of pollination by sexual deception in the genus Telipogon , the subtribe Oncidiinae, involving male tachinids. The next step is to identify the stimuli visual, olfactory or both that play a function in the attraction of male tachinids to flowers of Te.

Further chemical analyses and electrophysiological and behavioural tests are presently in progress. We are deeply grateful to Hermenegildo Martel for field assistance and William Nauray for providing information on Telipogon species from Southern Peru. Two anonymous reviewers provided helpful comments and suggestions. Conceptualization: CM MA.

Formal analysis: CM MA. Funding acquisition: CM MA. Project administration: CM MA. Writing — original draft: CM. Browse Subject Areas? Click through the PLOS taxonomy to find articles in your field. Abstract Several neotropical orchid genera have been proposed as being sexually deceptive; however, this has been carefully tested in only a few cases.

Introduction Animal-pollinated plants have evolved various floral signals in order to attract their pollinators, which are usually rewarded with pollen and nectar while visiting and pollinating flowers [ 1 — 3 ]; however, certain plant species deceive their pollinators by using attractive floral signals that mimic signals that play a role in food-seeking behaviour and the reproductive biology of their cheated pollinators [ 4 — 6 ].

Download: PPT. Flower-visiting behaviour and frequency of pollinators All the observations were carried out in the two previously mentioned patches P 1 and P 2 with, in total, h of observations h in P 1 , 60 h in P 2. Comparison of male behaviour at females and flowers To identify whether pollinator behaviour on a flower was similar to that shown in the presence of female visual and tactile stimuli, an odourless female dummy Soxhlet extracted pollinator female was pinned on a leaf.

Morphology and comparative anatomy Floral diameter was measured by means of a digital calliper, the length of the labellum plus the petal being measured. Scent-producing organs and light microscopic analyses We used neutral red staining and light microscopy to identify potential areas of scent emission on the labellum and petals of flowers. Flower scent collection and chemical analyses In preliminary investigations Headspace samples of flowers, using filters with adsorbents for methodological details see [ 51 ] , were collected.

Pollination process, pollinator behaviour and pollinator frequency Only four fly species, all male tachinids and never females, were attracted to flowers of Te. Tachinid collected attracted by Telipogon peruvianus flowers. Fig 3. Eudejeania aff. Fig 4. Frequency of pollinator visits to Telipogon peruvianus flowers.

Comparison of male behaviour at females and flowers The pattern of behavioural responses of E. Fig 5. Behavioural responses of Eudejeania aff. Table 1. Comparative anatomy Flowers of Te.

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Fig 6. Fig 7. SEM micrographs of Telipogon peruvianus flower and Eudejeania aff. Light-microscopic analyses of scent-producing organs in flowers We recorded unicellular trichomes and conical cells on the adaxial epidermal surface. Fig 8. Anatomical details of Telipogon peruvianus labellum. Chemical analysis In flower extracts of Te. Fig 9. Gas chromatogram of the labellum extract of Telipogon peruvianus. Table 2. Discussion Although the genus Telipogon was suggested to be sexually deceptive more than 50 years ago [ 36 ] and although recurrently asserted since then as being pollinated by sexually aroused male tachinids e.

Concluding remarks and future prospects Our findings are the first conclusive report of pollination by sexual deception in the genus Telipogon , the subtribe Oncidiinae, involving male tachinids.

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Supporting Information. S1 Video. A visit of a male of Eudejeania aff. Acknowledgments We are deeply grateful to Hermenegildo Martel for field assistance and William Nauray for providing information on Telipogon species from Southern Peru. References 1. Faegri K, van der Pijl L. The principles of pollination ecology. New York: Pergamon Press; View Article Google Scholar 2. Grant V. Modes and origins on mechanical and ethological isolation in angiosperms.

Generalization in pollination systems, and why it matters. View Article Google Scholar 4. Cozzolino S, Widmer A. Orchid diversity: an evolutionary consequence of deception? Trends Ecol Evol. Schiestl FP. On the success of a swindle: pollination by deception in orchids. Mechanisms and evolution of deceptive pollination in orchids. Biol Rev.

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Floral mimicry enhances pollen export: the evolution of pollination by sexual deceit outside of the Orchidaceae. Am Nat.