Factors affecting fruit and seed production in Dactylorhixa maculata (Orchidaceae)

It has been argued that some of resources needed for fruit and seed production in terrestrial orchids originate from storage in underground biomass. Resources for female reproductive traits may also originate from current photosynthesis. Orchid mycorrhiza may also influence fruit and seed production. The extent to which current photosynthetic activity and nutrient uptake via mycorrhizal fungi affect fruit and seed production has only rarely been studied experimentally. This paper examines the importance of photosynthesis and mycorrhiza for fruit and seed production, leaf area and plant status in the next season for Dactylorhiza maculata (L.) Soó in two populations. Plants were hand-pollinated in two populations to assure successful fertilization. Photosynthesis in the main leaves and in the green parts of inflorescence was prevented and/or mycorrhizal function inhibited by applying thiabendazole to plants. None of the treatments affected relative capsule production, but all treatments resulted in significant decreases in capsule dry weight and the number of seeds produced. The fact that seed production remained high in spite of the experimental treatments indicates that resources stored in the underground corms were used in fruit development and seed production. Inhibition of photosynthesis decreased leaf area in the year following treatments, while the other treatments had no effect on subsequent growth.     

An experimental study of floral display and fruit set in Chilopsis linearis (Bignoniaceae)

Summary Intact cores from the wet coastal arctic tundra at Barrow, Alaska, were used as microcosms in the measurement of CO₂ fluxes between peat, vegetation, and atmosphere under controlled conditions. Net ecosystem CO₂ uptake was almost twice as high at present summer temperatures (4° C) than at 8°. Lowering the water table from the soil surface to -5 cm also had a pronounced effect in decreasing net ecosystem carbon storage. Warming of the tundra climate could change this ecosystem from a sink for atmospheric CO₂ to a source.     

Pollination efficiency and reproductive patterns in relation to local plant density,population size,and floral display in the rewarding Listera ovata (Orchidaceae)

Pollination efficiency and reproductive success vary strongly among populations of most animal‐pollinated plant species, depending on their size and local density, whereas individual plants within populations experience varying levels of reproductive output as a result of differences in floral display. Although most orchid species have been shown to be severely pollination limited, few studies have investigated the impact of the above‐mentioned factors on pollination success and reproduction, especially in rewarding species. In this study, the impact of population size, local density of flowering plants, and floral display on the rates of pollinia export and fruit production was investigated in 13 natural populations of the rewarding terrestrial orchid Listera ovata. In addition, an emasculation experiment was set up to examine how floral display and local density of flowering plants affected the relative importance of cross‐ vs. geitonogamous pollination in determining fruit set. In the studied populations, pollination efficiency, pollen removal, and fruit set increased with increasing population size until a threshold value of 30–40 flowering plants was reached, above which pollination efficiency and reproductive output decreased again. On average, plants with large floral displays showed higher proportional pollinia removal and fruit set compared with smaller plants. Fruit production was also significantly and positively related to local plant density, whereas emasculation did not affect the relationship between local plant density and fruit set, suggesting that geitonogamous pollination did not affect the outcome of female function. The results of this study are discussed in the light of the flowering mechanism of the species and its generalized pollination system. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157 , 713–721.     

Pollination by deceit and floral mimesis inThelymitra antennifera (Orchidaceae)

Thelymitra antennifera is found to be a general mimic of the search image of other co-blooming yellow or cream-flowered species including members of the following genera:Hibbertia, Goodenia and, to a lesser extent,Helichrysum, Pimelia andStackhousia. The most common pollinators areLasioglossum (Chilalictus) spec. and the introducedSyrphus damaster (hover fly) as well as a solitary wasp,Eurys spec.—T. antennifera acts by deceit, involving opticals (yellow colour and pollen imitation) as well as olfactory (sweet odour) signals. Capsule production is low and density dependent, a situation demonstrated in other species which have evolved pollination syndromes involving deceitful mimicry.     

Reproductive effort and floral photosynthesis in Spiranthes cernua (Orchidaceae)

This study examined the cost of reproduction and photosynthetic characteristics of the reproductive structures of Spiranthes cernua, an agamospermic, terrestrial orchid. Reproduction was frequent: two-thirds of the plants flowered at least 2 yr in a row and one-fourth of the consecutive-year runs were ~3 yr. Neither a significant decrease in leaf area nor a reduced likelihood of flowering was observed following 1 or 2 yr of inflorescence production. While there was a tendency for plants producing >16 flowers to have decreased size the next year, plants with the greatest number of flowers (31+) were the most likely to reproduce. Leaf and reproductive gas exchange were measured in the field. Low but positive rates of net photosynthesis were documented at all stages of inflorescence development. The average rates of photosynthesis for each stage were: leaves, 9.2 mmol CO2/m2s; inflorescence in bud, 3.7 mmol CO2/m2s; inflorescence in flower, 2.5 mmol CO2/m2s and infructescence, 0.2 mmol CO2/m2s. Based on diurnal gas exchange, the contribution of leaves and reproductive structures to seasonal carbon assimilation was 91.6 and 8.4%, respectively. The role of the inflorescence as a source and sink for carbon assimilation may lower the cost of reproduction and support frequent inflorescence production.     

Pollinarium morphology and floral rewards in Brazilian Maxillariinae (Orchidaceae)

BACKGROUND AND AIMS: There is strong support for the monophyly of the orchid subtribe Maxillariinae s.l., yet generic boundaries within it are unsatisfactory and need re-evaluation. In an effort to assemble sets of morphological characters to distinguish major clades within this subtribe, the pollinarium morphology and floral rewards of representative Brazilian species of this subtribe were studied. METHODS: The study was based on fresh material from 60 species and seven genera obtained from cultivated specimens. Variation of pollinarium structure and floral rewards was assessed using a stereomicroscope and by SEM analysis. KEY RESULTS: Four morphological types of pollinaria are described. Type 1 appears to be the most widespread and is characterized by a well-developed tegula. Type 2 lacks a stipe and the pollinia are attached directly to the viscidium. Type 3 also lacks a stipe, and the viscidium is rigid and dark. In Type 4, the stipe consists of the whole median rostelar portion and, so far, is known only from Maxillaria uncata. Nectar, trichomes, wax-like and resin-like secretions are described as flower rewards for different groups of species within the genus Maxillaria. Data on the biomechanics and pollination biology are also discussed and illustrated. In Maxillariinae flowers with arcuate viscidia, the pollinaria are deposited on the scuttellum of their Hymenopteran pollinators. In contrast, some flowers with rounded to rectangular, pad-like viscidia fix their pollinaria on the face of their pollinators. CONCLUSIONS: Pollinarium morphology and floral features related to pollination in Brazilian Maxillariinae are more diverse than previously suggested. It is hoped that the data presented herein, together with other data sources such as vegetative traits and molecular tools, will be helpful in redefining and diagnosing clades within the subtribe Maxillariinae.     

Variation in the floral fragrance of Epidendrum ciliare (Orchidaceae)

Variation in floral fragrance compounds might be a pollination strategy that disrupts associative learning processes and inhibits pollinator's ability to recognize non-rewarding flowers. Floral fragrances of the nectarless, moth-pollinated Epidendrum ciliare (Orchidaceae) were obtained from plants of Puerto Rico and the Virgin Islands. Fragrances were collected by passive diffusion on tenax adsorbent and analyzed using gas chromatography and mass spectrometry (GUMS). The analysis was performed at six different levels: 1) by flower position on the same inflorescence; 2) by flower age on the same plant; 3) by position and age of the flower in the same plant; 4) by different shoots of the same plant; 5) by flowers from different plants; 6) and by plants from different populations. At no level of analyses were fragrances of sample pairs identical. The greatest similarity among fragrances was between flowers of the same plant. The most notable differences were between flowers of different populations. High levels of floral fragrance variation were consistent with the hypothesized deception pollination syndrome, but comparisons with reward-producing flowers remain to be done.     

开花式样对传粉者行为及花粉散布的影响

理解植物花的特征可以从单花特征和群体特征两个层次入手。开花式样是植物的花在群体上的特征体现, 通过在开花数目、开花类型以及花的排列上的变化, 不同的开花式样对传粉者具有不同的吸引力, 影响昆虫在植株上的活动, 使花粉运动的方向发生相应变化, 从而影响着植物最终的交配结果。此外开花式样随环境改变也会发生一些变化。本文介绍了开花式样研究的进展, 对开花数目、开花类型以及花的排列等3个方面的已有研究进行了分别阐述, 并提出开花式样研究应更多地考虑影响传粉的各种因素。     

Effects of floral display size and biparental inbreeding on outcrossing rates in Delphinium barbeyi (Ranunculaceae)

Floral display size represents a tradeoff between the benefits of increased pollinator visitation and the quantity of pollen received vs. the costs of increased self-pollination and reduced pollination quality. Plants with large floral displays often are more attractive to pollinators, but pollinators visit more flowers per plant. Intraplant foraging movements should increase self-pollination through geitonogamy, lowering outcrossing rates in large plants. Local genetic structure should also increase inbreeding and decrease outcrossing estimates, if pollinators move between neighboring, related plants. These predictions were tested in a population of larkspurs (Delphinium barbeyi) in Colorado. Allozymes were used to estimate outcrossing rates of plants varying in display size. Floral displays varied widely (2-1400 flowers; 1-26 inflorescences per plant), and outcrossing rate decreased significantly with increasing display size. Large, multistalked plants self over twice as frequently as single-stalked plants (46 vs. 21%). Local population structure is significant, and biparental inbreeding depresses outcrossing in plants surrounded by genetically similar neighbors. Protandry, coupled with stereotypical bottom-up pollinator foraging, reduces self-fertilization by autogamy or geitonogamy within inflorescences. Selfing is predominantly (>60%) by geitonogamy between inflorescences in large plants. Geitonogamy may be a significant cost to plants with large floral displays if inbreeding depression and/or pollen and ovule discounting results. If so, floral display size, particularly inflorescence number, may be under contrasting selection for pollination quantity vs. quality.     

Observations of the floral biology ofPrasophyllum odoratum (Orchidaceae,Spiranthoideae)

Prasophyllum odoratum is a vernal, nectariferous, terrestrial orchid that flowers profusely six to eight months following cyclical fires that disrupt sclerophyll woodlands. The morphology of the column and pollinarium is indicative of taxa placed within the subfam.Spiranthoideae. The orientation of the pollinaria to the stigma appears to prevent mechanical self-pollination. Both cross- and self-pollination appear to be effected by insects that forage within the brightly-colored, scented, non-resupinate flowers. Ants and drosophilid flies remove nectar, but do not appear to transport pollinaria between flowers. The primary pollinators are polytrophic flies in the fam.Syrphidae and opportunistic male bees in the genusLeioproctus (Colletidae). Approximately 52% of the flowers on a raceme set seed. The comparatively short floral tube ofP. odoratum reflects the dependence of this species on short-tongued insects to effect successful dispersal of pollinaria.     

Zingerone in the floral synomone of Bulbophyllum baileyi (Orchidaceae) attracts Bactrocera fruit flies during pollination

Many orchids use specific volatile chemical(s) in floral fragrance to attract potential pollinators to locate and visit flowers in the tropical rain forests. Male fruit flies (Bactrocera species) are attracted to the ‘fruity odour’ of Bulbophyllum baileyi. The floral chemical component responsible for this fruit fly attraction is zingerone, which also acts as a floral chemical reward during pollination. Zingerone (5 mg), absorbed into a filter and released via a portable-battery operated fan between 0900 and 1100 h, attracted 5–7 males of Bactrocera indonesiae in an orchard located in Kedah, Malaysia. In Penang island, the fruit fly species attracted are Bactrocera dorsalis (methyl eugenol-sensitive) and Bactrocera albistragata (raspberry ketone-sensitive). An attracted male fruit fly probes and feeds on floral petal or sepal before climbing on to the small see-saw lip (labellum). While on the lip and due to shifting of the fly's weight, the fly is toppled head first into the column cavity and simultaneously touches and consequently gets stuck to the sticky viscidium of pollinarium that still remained in the anther. It spends 23–46 min trying to free itself from being suspended from the floral column. Removal of pollinarium (pollinia, stipe and viscidium) is a long process. A male B. dorsalis that fed on a B. baileyi flower was found to sequester zingerol (a reduced form of zingerone) in the body, suggesting its role as sex pheromone to attract a female during courtship behaviour.     

Patterns of pollinator-generated fruit set inOrchis morio (Orchidaceae)

Reproductive success of a deceptive species,Orchis morio, was investigated in relation to floral display, vertical position of fruits and dispersion pattern. A caging experiment confirmed the necessity of the presence of insect vectors for fruit development. In two Czech populations studied in 1997, insect-pollinated plants had on average 27.3 and 38% fruit set, compared to 96.7% for flowers pollinated by hand. Floral display (number of flowers per spike) was expected to positively affect the probability of pollinator visit. However, fruit set (number of fruits per spike) did not increase linearly with increasing floral display, but rose sigmoidally and approached an asymptotic value. Floral display enhanced reproductive success (fruit/flower ratio) up to a certain number of flowers (11–14), above which the relationship started to decrease and only increased the probability of production of at least one fruit. We also studied the effect of flower position within an inflorescence on the probability of being pollinated and we counted the number of seeds per capsule. A rapid decline in fruit set from the bottom to the top of the inflorescence observed probably reflects the behaviour of pollinators. The number of seeds per one capsule varied from 550 to 12 270. In addition, individuals growing in large clumps or isolated had lower reproductive success than those growing in small clumps. This result implies the existence of an intraspecific competition for pollinators and on the other hand inability of isolated individuals to attract a large number of pollinators.     

Insect pollination and floral mechanisms in South African species ofSatyrium (Orchidaceae)

The morphologically diverse flowers in the genusSatyrium reflect adaptations to a wide range of pollinators. Several recently discovered pollination systems inSatyrium are described and illustrated here; these include pollination by solitary bees, carrion flies, butterflies and moths. Two basic types of floral mechanism are recognised inSatyrium: (1) Species pollinated by lepidoptera and birds have long floral spurs and plate-like viscidia seated in lateral rostellum notches; these viscidia become attached to the proboscis or bill of the pollinator. (2) Species pollinated by flies and bees have relatively short floral spurs and globose viscidia seated in terminal rostellum notches; these viscidia become attached to the face, thorax or eyes of the pollinator.     

Effect of pollination success on floral longevity in the orchid Calypso bulbosa (Orchidaceae)

The lifespan of an individual flower is often affected by pollination success. Species differ regarding whether male function (pollen removal), female function (pollen deposition), or both trigger floral senescence. We studied senescence in the singleflowered, deceptive orchid Calypso bulbosa by manipulating the degree of male and female reproductive success. We found that deposition of any amount of pollen resulted in dramatic changes in shape and color within 4 d, whereas unmanipulated flowers and those that had had pollinia removed remained unchanged for 8-11 d after treatment. Selection may favor the reproductive function that is less easily satisfied as the trigger for senescence, because a flower that senesces after accomplishment of this function is likely to have already succeeded at the more easily satisfied one. Deceptive (i.e., rewardless) flowers are more likely to satisfy male than female function since the latter requires that a pollinator be fooled twice, first to pick up pollen and second to deposit it. A survey of naturally pollinated Calypso showed that male function, pollinium removal, was more likely to occur than female function, deposition (95% vs. 66% of visited flowers); thus floral senescence in Calypso is triggered by achievement of the function less likely to succeed. Studies of senescence triggers in species in which female function is more likely to be achieved than male are necessary to further test this hypothesis.     

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.     

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.