On the mechanism of floral shifts in speciation: gained pollination efficiency from tongue- to eye-attachment of pollinia in Platanthera (Orchidaceae)

This work explores the mechanism of floral shifts that may result in speciation. The model organisms chosen were the moth-pollinated pollinator-limited orchid species Platanthera bifolia and P. chlorantha . P. bifolia exhibits tongue-attachment of pollinia on pollinators, a character state that has been found previously to be ancestral. The close relative P. chlorantha exhibits the derived state of eye-attachment of pollinia on pollinators. We reasoned that differences between the species in pollination efficiency could give insights into the mechanism of floral shifts and thus cladogenesis. Four populations per species were investigated. In three populations, where the species were growing intermixed and were sharing pollinators, there was significant difference in the pollen export and import efficiency per visit-night (night with pollen export and/or import) between the two species. P. bifolia exported pollinia more efficiently but imported pollen less efficiently than did P. chlorantha . Pollen import was 1.7–4 times faster in P. chlorantha (eye-attachment) than it was in P. bifolia (tongue-attachment). P. chlorantha had a lower risk of interference between pollen import and export. An increase in fitness through greater speed and efficiency of pollen import due to an enlargement of the stigmatic surface and a reduction in the risk of sexual interference may therefore be mechanisms of the floral shift from tongue- to eye-attachment of pollinia on pollinators.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 481–495.     

Characteristics and distribution of intermediates between Platanthera bifolia and P. chlorantha (Orchidaceae) in the Nordic countries

In areas of sympatry Plantanthera bifolia (L.) L. C. Rich. and P. chlorantha (Cust.) Rchb. occasionally produce hybrids. These plants exhibit intermediate floral morphology. The present study presents the distribution, floral morphology, and general characteristics of these intermediates in the Nordic area. The most useful and reliable diagnostic characters for hybrid plants were (1) the distance between viscidia (normally about 1.4 mm), (2) the length of the caudicle of pollinia (about 0.7 mm), and (3) the presence of a distinct pedicel at the connection between caudicle and viscidium. Comparative chemical analyses of the floral fragrances are reported. Material in the principal Nordic herbaria indicated a generally low production of hybrids (0.46%) and that the main area of occurrence is SE Sweden and Alandia, Finland. Intermediates seem more frequent in provinces where P. bifolia is relatively long-spurred. Very few were recorded from Denmark and Norway (hybrids in the latter country here reported for the first time). Aggregations of intermediates displaying patterns of introgressive hybridization were found only on Öland and Gotland. Occurrence of hybrids outside the Nordic area is also discussed and remarks are made on the morphologically similar P. metabifolia Maek. in E Asia. All known Nordic intermediates are listed.     

Nutrient versus pollination limitation in Platanthera bifolia and Dactylorhiza incarnata (Orchidaceae)

Availability of resources and pollination services have been demonstrated to have impact on reproductive success in some orchid species, but to our knowledge no studies have examined the effects of nutrient application and pollination limitation in the same experiment. In this study, factors limiting reproductive success were studied with two terrestrial orchid species in Central Finland during 1996 and 1997. In a field experiment using a factorial design, plants of nectar-producing Platanthera bifolia and nectarless Dactylorhiza incarnata were treated with nutrient application and hand-pollination. Inflorescence size was considered as an indicator of attractiveness to pollinators as well as of the general condition of the plant. In 1996, fertilizer treatment increased relative capsule production in P. bifolia plants with small inflorescences, indicating poorer store of resources in the underground corm and dependence on nutrient availability during capsule maturation, which was not found in large plants . Hand-pollination and large inflorescence size in P. bifolia did not affect capsule initiation, but increased the proportion of mature capsules. P. bifolia may thus be regarded as both resource- and pollination-limited within a year. Pollination success was observed to be the only factor limiting reproduction of the nectarless D. incarnata within a year, because hand-pollination increased capsule production, and there were no significant effects of fertilizer treatment or inflorescence size in 1996. Capsule production in 1996 did not affect the probability of P. bifolia flowering in the following year, while high capsule production decreased the subsequent probability of flowering in D. incarnata . Species with different pollination strategies differed in the use of resources. The nectar-producing P. bifolia had a lower fruit/flower ratio than the nectarless D. incarnata , also after hand-pollination.     

Fragrance chemotypes of Platanthera (Orchidaceae) - the result of adaptation to pollinating moths?

Floral scent of Platanthera species (Orchidaceae) in S Sweden was collected by head-space adsorption methods in the field and analysed by gas chromatography-mass spectrometry (GC-MS). Variation in scent chemistry of both Platanthera bifolia and P. chlorantha was considerable: different scent chemotypes were found among individuals as well as populations. Mainly linalool, lilac aldehydes and alcohols, geraniol, and methyl benzoate distinguished the chemotypes. Because of a high individual variation, floral scent in Platanthera is not suitable as a taxonomic tool. Scent variation can be the result of differential selection from various pollinators or pollinator groups. However, in a long-spurred race of P. bifolia the scent profile was not clearly different from that of short-spurred races in spite of their different pollinators. The two species form natural hybrids and part of the variation at the individual level may he explained by interspecific introgression. Differences found among populations in different regions may be the result of random genetic drift.     

Variation and post-pollination changes in floral odours released by Platanthera bifolia (Orchidaceae)

Volatile compounds were collected in the field from individual specimens of un-pollinated and hand-pollinated Platanthera bifolia , and investigated by combined gas chromatography-mass spectrometry (GC-MS). In all, 32 compounds were identified of which five were not previously reported in Platanthera. Pollinated flowers wilted significantly faster than unpollinated flowers. Twentyfour hours after pollination the production of volatile compounds had decreased slightly and there was a small change in their composition. Variation in floral scents within and between populations, as well as effects of changes in floral attractiveness on pollination and reproductive success are discussed.     

A multivariate approach to post-pollination changes in the floral scent of Platanthera bifolia (Orchidaceae)

The effect of pollination on floral scent composition and production in the moth-pollinated orchid Platanthera bifolia was studied. A significant decrease in scent production was detected both two and five days after pollination. The proportion of wilted flowers was higher in pollinated compared to control plants after seven days. A principal component analysis (PCA) showed how pollination reduced scent production. All scent compounds were affected by pollination, even though some compounds had larger impact on the overall scent reduction. A classification of samples offered a quantitative test of the changes found in individual plants after pollination. The induced decrease in floral scent production following pollination is suggested to be of adaptive value for both plants and pollinators.     

Volatiles from flowers of Platanthera bifolia (Orchidaceae) attractive to the silver Y moth, Autographa gamma (Lepidoptera: Noctuidae)

We examined the attractiveness of a natural headspace sample of Platanthera bifolia blossoms, synthetic blends and single compounds to the silver Y moth, Autographa gamma, in a flight tunnel. The synthetic blend consisted of previously identified electrophysiologically active compounds from P. bifolia : benzyl benzoate, benzyl salicylate, cinnamyl alcohol, lilac aldehydes, methyl benzoate and methyl salicylate. This blend had a similar attractivity as the natural headspace sample. Subtraction of lilac aldehydes significantly decreased attractiveness of the synthetic blend. When a mixture of lilac aldehydes was tested alone, it showed attractiveness similar to that of the synthetic blend. One or a mixture of lilac aldehydes accounts for the attraction of moths to P. bifolia . All other compounds elicited significantly lower responses. Results are discussed in relation to the pollination biology of P. bifolia .     

Nectar resorption and translocation in Cucurbita pepo L. and Platanthera chlorantha Custer (Rchb.)

Nectar resorption and sugar translocation were studied in Cucurbita pepo (Cucurbitaceae) and Platanthera chlorantha (Orchidaceae) by micro-autoradiography. In both species, nectar was resorbed in pollinated and unpollinated flowers and ovules developing into seeds were found to be the main sugar sink. In C. pepo, the mobility of resorbed sugars in pollinated female flowers was higher than in unpollinated ones; male flowers showed lower mobility of resorbed sugar. In P. chlorantha, radioactivity was detected in pollinated flowers below and above labelled unpollinated ones: the nearer the flower, the stronger the accumulation of label in developing fruits.     

Floral longevity and nectar secretion of Platanthera chlorantha (Custer) Rchb. (Orchidaceae)

Flowering and nectar secretion were studied in Platanthera chlorantha in two years. Nectar was secreted and accumulated in this orchid's spur, originating from part of the labellum. The nectary spur was, on average, 32 mm long. It produced 6.86 micro l nectar in 1999 and 7.84 micro l in 2000. The number of flowers per inflorescence and the volume of nectar secreted per flower were not correlated. Nectar secretion and flower longevity differed depending on pollination and flower position in the inflorescence. Among pairs of pollinated and unpollinated flowers there was no difference in the volume of nectar produced; however, the life span of pollinated flowers was shorter than that of unpollinated ones. Within an inflorescence, the lowest-positioned flowers had the largest nectar production and the longest life compared with flowers positioned higher up.     

Pollination Biology of Hosta sieboldiana (Lodd.) Engler and H. sieboldii (Paxton) J. Ingram (Liliaceae)

Abstract The pollination biology of Hosta sieboldiana and H. sieboldii is investigated comparatively in Central Japan. Both species have homogamous, one-day flowers pollinated by bumblebees. The abdomens of the bees touch the stigma on the extended style when they land on the anthers inside the herkogamous flower, and autogamy is effectively prevented. However, the flowers are fairly self-compatible, and geitonogamy may occur rather frequently because two or more flowers on a scape very often bloom at the same time and many ramets are contiguous. The pollen/ovule ratios suggest that these species are facultative outbreeders. The flower of H. sieboldii seems completely suited to bumblebee pollination. In H. sieboldiana the stigma of the flower, whose style strongly protrudes, is not always touched by bumblebees, but frequent visitation of bumblebees results in pollination of almost all the flowers. Both species have similar pollination systems but seem reproductively isolated by blooming times and habitats. Their common pollinators, however, may sometimes cause introgressive hybridization in contiguous populations.     

Relationships of the purple-fringed orchids Platanthera Psycodes and P. grandiflora

The vegetative and floral differences between two frequently confused species are discussed in terms of function.Platanthera psycodes is a widespread species with flowers adapted to visits by short-tongued lepidoptera, especially skippers of the genusPolites; the pollinaria are attached to the base of the proboscis.Platanthera grandiflora is a geographically more limited species with flowers adapted to visits by longer-tongued lepidoptera; the pollinaria are probably attached to compound eyes. The species overlap geographically and phenologically. Chromosome numbers are 2n = 42 in each species. Cultivated plants of each are partially self-incompatible but produce apparently normal embryos when intercrossed. Mechanical, seasonal, ethological, and distributional differences isolate the species.Habenaria andPlatanthera are considered distinct but closely related genera.Platanthera psycodes is based upon a type at L,P. grandiflora on a type at AMES.Platanthera grandiflora andP. psycodes have affinities with three other eastern North American species characterized by tripartite, fringed, or erose labella. Species pairs within this group, based on column structure, includegrandifloraperamoena andpsycodeslacera. The column ofP. leucophaea is morphologically intermediate in form.     

Odour-mediated nectar foraging in the silver Y moth, Autographa gamma (Lepidoptera: Noctuidae): behavioural and electrophysiological responses to floral volatiles

Naive male and female silver Y moths, Autographa gamma (Lepidoptera: Noctuidae), were attracted in a flight tunnel assay to potted creeping thistle, Cirsium arvense (Asteraceae), butterfly-orchid, Platanthera bifolia (Orchidaceae), soapwort, Saponaria officinalis (Caryophyllaceae), greater knapweed, Centaurea scabiosa (Asteraceae), red clover, Trifolium pratense (Fabaceae), and catnip, Nepeta faasseni (Labiatae), plants with flowers. The most attractive plants were C. arvense , P. bifolia and S. officinalis that elicited 87, 78 and 65% source contacts, respectively. C. scabiosa was less attractive eliciting 43% response. T. pratense and N. faasseni showed the least attraction eliciting 28 and 26% source contacts, respectively. A cotton plant used as control, was not attractive. Floral volatiles from the investigated plant species were collected using headspace sampling technique. Samples were analysed using gas chromatography coupled with electroantennographic detection, and electrophysiologically active compounds were identified by coupled gas chromatography/mass spectrometry. Consistent electrophysiological responses were elicited by twelve compounds from headspace of C. arvense , thirteen compounds from P. bifolia , eleven compounds from S. officinalis , nine from C. scabiosa , ten from T. pratense and two from N. faasseni . Most of the active compounds were specific for one or two species, while benzyl benzoate was present in four and benzaldehyde and benzyl alcohol in three species. Floral scents of C. arvense , P. bifolia and S. officinalis , the most attractive flowers, were dominated by aromatic compounds that were not abundant in the scent of other flowers. To conclude, the results demonstrate the absence of a common denominator of odours present in flowers of different plants visited by A. gamma .     

侧柏和北美香柏(柏科)的传粉机制(英)

观察了侧柏（Phaycladusorientalis（L．）Franco）和北美香柏（ThujaoccidentalisL．）散粉后花粉进入珠孔的过程。在散粉期,这两种植物的胚珠均分泌出传粉滴。当花粉落到传粉滴上后,引起传粉滴表面的形状发生改变或减弱胚珠的继续分泌,使得该传粉滴蒸腾加快,导致其比未授粉的传粉滴明显收缩。观察结果表明：不同植物的花粉导致侧柏传粉滴的收缩速率不尽相同。其中,与侧柏亲缘关系较近的植物花粉引起传粉滴的收缩速率和侧柏自身花粉引起的传粉滴收缩速率相似;反之,收缩速率变慢。侧柏传粉滴的收缩可能主要是由于花粉减弱胚珠分泌的结果。     

Orientation and withdrawal of pollination drops in Cupressaceae s. l. (Coniferales)

Pollination in the Cupressaceae is studied ex situ, focused on orientation and withdrawal of pollination drops. Orientation of pollination drops is a constant feature in most taxa studied and important for pollen capture. Conspecific pollen causes a withdrawal of pollination drops, varying in time among species from 8 to 24 min, but with little variation within species. Pollination drops of each tested Cupressaceae taxon are also withdrawn when pollinated with foreign, but Cupressaceous pollen. However, they remain unchanged and are not withdrawn immediately when pollinated with pollen of other seed plants. The results clearly indicate that the time for the total withdrawal of pollination drops is strongly influenced by the evolutionary distance of the taxa being involved in the pollination process. Among closely related taxa the withdrawal is much more rapid than in distantly related ones. This points to an effective recognition system regulating the withdrawal of pollination drops, probably controlled by the nucellus. This recognition system can be regarded as an important preadaption for the evolution of a self-incompatibility mechanism. The withdrawal of pollination drops is thus not exclusively a physically induced process as suggested in some earlier studies. Pollination drops of several ovules can fuse to form a large common one, perhaps increasing by this way successful pollen capture.     

马先蒿属花冠形态的多样性与传粉式样的关系

马先蒿属(Pediculais)是有花植物中花冠形态多样化最为集中的属。该属主要的传粉者是熊蜂属(Bormbus)昆虫;在北美,熊蜂和蜂鸟是马先蒿植物一些种类有效的传粉者;也发现壁蜂(Osmia)为其传粉。不同的传粉机制要求某一特定的取食式样储藏和释放花粉。本文讨论了花冠类型的进化趋势与传粉式样和花粉形态的关系。传粉者的选择压力是决定花冠多样化的重要因素之一;花冠类型与传粉者和传粉行为紧密相关。马先蒿植物和传粉者的相互依赖与其花冠类型、功能和物候互相适应,但花冠类型与花粉形态两者之间似乎没有明显的一一对应关系。通过北美、日本和喜马拉雅不同地理分布马先蒿种类的比较研究表明,具有相同花冠类型的种类有着相同的传粉方式,花冠形态与传粉式样存在紧密的协同进化关系。     

Viability and longevity of pollen from transgenic and nontransgenic tall fescue (Festuca arundinacea) (Poaceae) plants

Pollen is an important vector of gene flow in plants, particularly for outcrossing species like tall fescue. Several aspects of pollination biology were investigated using pollen from transgenic and nontransgenic plants of tall fescue (Festuca arundinacea Schreb.), the most important forage species worldwide of the Festuca genus. To effectively assess in vitro pollen viability in tall fescue, an optimized germination medium (0.8 mol/L sucrose, 1.28 mmol/L boric acid and 1.27 mmol/L calcium nitrate) was developed. Treatment with relatively high temperatures (36° and 40°C) and high doses of UV-B irradiation (900-1500 μW/cm(2)) reduced pollen viability, while relative humidity did not significantly influence pollen viability. Viability of pollen from transgenic progenies (T1 and T2) was similar to that from seed-derived control plants. Pollen from primary transgenics (T0) and primary regenerants (R0) had various levels of viability. Hand pollination using the primary regenerants and transgenics revealed that no seed set could be obtained when pollen viability was lower than 5%. Pollen from transgenic progenies and nontransgenic control plants could survive up to 22 h under controlled conditions in growth chamber. However, under sunny atmospheric conditions, viability of transgenic and nontransgenic pollen reduced to 5% in 30 min, with a complete loss of viability in 90 min. Under cloudy atmospheric conditions, pollen remained viable up to 240 min, with about 5% viability after 150 min. This report is the first on pollen viability and longevity in transgenic forage grasses and could be useful for risk assessment of transgenic plants.     

POLLINATION OF RAFFLESIA (RAFFLESIACEAE)

The genus Rqfflesia includes about 13 species of parasitic flowering plants, among which are the largest known flowers. The flower with subtending scales is the only part of the plant external to the host and is produced solitary on roots (rarely stems) of the genus Tetrastigma (Vitaceae). Field studies were made of the pollination process in R. pricei, a species endemic to the Crocker Range in the Malaysian state of Sabah (northern Borneo). Pollination is mediated by carrion (bluebottle) flies of the genera Lucilla and Chrysomya. Experimental data indicate that both visual and olfactory cues are important in attracting flies to flowers. Flies (mostly female L. papuensis) obtain loads of the viscous liquid pollen matrix by visiting male flowers and entering anther grooves on the central column of the flower, precisely guided by ridges armed with hairs that force the fly into a position in which the pollen is positioned on the dorsal part of the thorax. “Windows” on the inside of the perigone diaphragm apparently help orient their flight inside flowers. Pollen-loaded flies visiting female flowers may enter the infradiscoidal sulcus formed by a broad ring of stigmatic tissue above and the expanded base of the column below. On entering the sulcus the fly is wedged in so tightly that pollen is rubbed off the thorax onto the stigma. Only large flies could be effective in picking up pollen from male flowers and transferring it to female flowers. The pollination syndrome is sapromyophily, in which the flower closely parallels trap flowers of several other plant families, although it is not a trap. The flower provides no reward for pollinators but deceives them by an apparent offering of food and possibly brood place. Rafflesia plants are extremely rare, perhaps in part because of infrequency of pollination, which requires neighboring male and female flowers simultaneously in bloom.     

Pollination of Pteroglossaspis ruwenzoriensis (Rendle) Rolfe (Orchidaceae) by Beetles in Argentina

Beetle pollination by Euphoria lurida (Scarabaeidae: Cetoninae) is documented for Pteroglossaspis ruwenzoriensis (Orchidaceae: Epidendroideae: Cymbideae: Eulophiinae) in its natural habitat in Central Argentina (South America). Flower features with special emphasis on those related to beetle pollination are given. These include: long rigid and well exposed inflorescences, sturdy inconspicuous and deep flowers with small entrance, emission of a yeast-like fragrance, jelly-like nectar, short column, and a head-attaching pollinarium with a broad saddle-like viscidium. Pollinator behaviour and pattern of flower opening favour cross pollination and probably long distance pollen dispersal. Flowering, which lasts about 3 weeks, peaks in summer past mid-January. In natural conditions about 68 flowers are pollinated for every 100 pollinaria removals.     

Pollinator-mediated competition between two congeners, Limnanthes douglasii subsp. rosea and L. alba (Limnanthaceae)

? Premise of Study: Pollinator visits are essential for reproduction in many plants, yet interspecific movements of pollinators can also lead to competitive interactions between coflowering species. Pollination-mediated reductions in fertility could potentially lead to exclusion of competing plant species, and may generate spatial variation in the associations among coflowering species across a landscape. ? Methods: I documented the potential for heterospecific pollen transfer to cause competitive interactions between two annual grassland species native to California, Limnanthes douglasii subsp. rosea and L. alba, two reproductively incompatible species that have broadly overlapping geographic ranges in the foothills of the Sierra Nevada. I observed pollinator movement in constructed arrays and controlled crosses in the greenhouse and field to investigate the consequences of heterospecific pollen transfer. ? Key Results: Pollinators move readily between species when they are presented together in experimental arrays. In the greenhouse, deposition of heterospecific pollen decreased fertility in both species. The decrease in seeds produced per flower was much more pronounced in L. d. rosea (90.6% reduction) than in L. alba (40.8% reduction). In field experiments, L. d. rosea plants that received pollen from heterospecific neighbors first showed >50% reduction in per-flower fertility. ? Conclusions: Under natural pollination conditions, heterospecific pollen transfer has the ability to decrease the fertility of L. d. rosea when it occurs at low frequency in mixed stands. Accordingly, pollinator-mediated competition may contribute to the locally disjunct distributions of these two species. It may also influence important restoration decisions in vernal pool habitats.     

Wind of change: new insights on the ecology and evolution of pollination and mating in wind-pollinated plants

Background

The rich literature that characterizes the field of pollination biology has focused largely on animal-pollinated plants. At least 10 % of angiosperms are wind pollinated, and this mode of pollination has evolved on multiple occasions among unrelated lineages, and hence this discrepancy in research interest is surprising. Here, the evolution and functional ecology of pollination and mating in wind-pollinated plants are discussed, a theoretical framework for modelling the selection of wind pollination is outlined, and pollen capture and the occurrence of pollen limitation in diverse wind-pollinated herbs are investigated experimentally.

Scope and Conclusions

Wind pollination may commonly evolve to provide reproductive assurance when pollinators are scarce. Evidence is presented that pollen limitation in wind-pollinated plants may not be as common as it is in animal-pollinated species. The studies of pollen capture in wind-pollinated herbs demonstrate that pollen transfer efficiency is not substantially lower than in animal-pollinated plants as is often assumed. These findings challenge the explanation that the evolution of few ovules in wind-pollinated flowers is associated with low pollen loads. Floral and inflorescence architecture is crucial to pollination and mating because of the aerodynamics of wind pollination. Evidence is provided for the importance of plant height, floral position, and stamen and stigma characteristics in promoting effective pollen dispersal and capture. Finally, it is proposed that geitonogamous selfing may alleviate pollen limitation in many wind-pollinated plants with unisexual flowers.Key words: Wind pollination, reproductive assurance, pollen limitation, geitonogamy, sex allocation, inflorescence architecture, mating systems