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Effects of forest fragmentation on male and female reproductive success in Cestrum parqui (Solanaceae).

In this paper we evaluate the effects of forest fragmentation on male (pollen removal, pollen load, and pollen tubes) and female reproductive success (fruit- and seed-set) of Cestrum parqui, a self-incompatible, pollination-specialist plant species. We also measure focal individual conspecific density to account for possible density-related effects that could influence the response variables. We calculate an index which incorporates male and female fitness and gives an integrated assessment of overall reproductive success. Forest fragmentation strongly affected the amount of pollen grains on stigmas and number of pollen tubes as well as seed-set, decreasing from continuous forest to small forest fragments, whereas focal individual conspecific density failed to explain any of the variability for the studied variables. Declines in overall reproductive success (i.e. male and female) in small forest fragments are ascribed to decreases in both the quality and quantity of pollination. Self-incompatibility coupled with a specialist pollination system may be particularly important traits determining the negative fragmentation effects observed in C. parqui. Logarithmic regression models described the behaviour of the variables along the fragmentation size gradient, allowing us to detect a threshold below which the effects of fragmentation begin to negatively affect reproductive success in C. parqui. Our results emphasize the importance of evaluating both components of the total plant fitness, as well as including simultaneously several aspects of pollination and reproduction processes when assessing the effects of forest fragmentation on plant reproductive success.

Cestrum↗

Accessory costs of seed production.

Accessory costs of reproduction are those that are necessary to mature a seed, but that do not involve the direct cost of provisioning the seed itself. This study aims to quantify accessory costs in a range of species, and test whether they decrease as a proportion of total reproductive expenditure with increasing seed mass, as might be expected if economies of scale came into play at larger seed sizes. We also test whether accessory costs varied with growth form, pollination mode, and dispersal mode, with the expectation that biotic pollination and dispersal modes should incur greater costs. Reproductive allocation (dry biomass) over one season, was calculated for 14 diclinous angiosperm species. Accessory costs averaged 73% of total reproductive allocation, with the majority spent on packaging and dispersal. Total accessory costs, packaging and dispersal costs, and costs incurred prior to pollination were proportional to direct costs of reproduction in major axis regressions. However, larger seeded species incurred significantly greater costs associated with aborted seeds and fruits, and matured a smaller proportion of ovules. This is consistent with larger seeded species being more selective of the ovules/embryos matured than small-seeded species. Total accessory costs, and proportion of ovules aborted, were also significantly greater for biotically dispersed species, but only due to an association with larger seed masses. Costs associated with abortions were lower for biotically pollinated species, due to a general trend of more ovules per ovary, resulting in greater cost sharing. This study demonstrates that expenditure on items other than seeds accounts for the majority of reproductive allocation in flowering plants. Yet, far more literature exists on seed mass variation than on investment in accessory structures. We found a proportional relationship between accessory costs and seed mass that warrants further investigation within the context of selection on margin returns on investment.

Flowers↗

Experimental investigation of the effect of spatial aggregation on reproductive success in a rewardless orchid.

Plant reproductive success within a patch may depend on plant aggregation through pollinator attraction. For rewardless plants that lack rewards for pollinators, reproductive success may rely strongly on the learning abilities of pollinators. These abilities depend on relative co-flowering rewarding and rewardless plant species spatial distributions. We investigated the effect of aggregation on the reproductive success of a rewardless orchid by setting up 16 arrays in a factorial design with two levels of intraspecific aggregation for both a rewardless orchid and a rewarding co-flowering species. Our results show that increasing aggregation of both species negatively influenced the reproductive success of the rewardless plants. To our knowledge, this is the first experimental study demonstrating negative effects of aggregation on reproductive success of a rewardless species due both to its own spatial aggregation and that of a co-flowering rewarding species. We argue that pollinator learning behaviour is the key driver behind this result.

Analysis of Variance↗

Floral odor variation in two heterostylous species of Primula.

Floral traits such as odor, color, and morphology are important pollinator attractants. Variation in floral traits may influence floral constancy, the tendency of pollinating insects to visit flowers of only one type. We investigated for the first time variation in odor between floral morphs in heterostylous species. We analyzed inter- and intraspecific odor variation in the "pin" and "thrum" floral morphs of sympatric Primula elatior and P. farinosa (Primulaceae). Floral volatiles were sampled with headspace sorption. Quantitative analysis and chemical identification were performed by gas chromatography coupled to mass spectrometry. The species produced different floral bouquets. P. elatior emitted mostly limonene with small amounts of alpha-pinene, myrcene, and sabinene. P. farinosa produced benzaldehyde, 4-oxoisophorone (2,6,6-trimethyl-2-cyclohexene-1,4-dione), benzyl alcohol, and benzyl acetate. These interspecific differences may play a role in promoting floral constancy and maintaining species integrity. Conversely, no differences were detected between the scents of pin and thrum morphs within each species. Heterostyly relies on pollinators visiting both floral morphs. There may be stabilizing selection against divergences in traits that may cause pollinators to develop floral constancy to only one of the floral morphs.

Animals↗

Postpollination changes in floral odor in Silene latifolia: adaptive mechanisms for seed-predator avoidance?

Floral odor is a key trait for pollinator attraction in many plants, but may also direct antagonists like herbivores to flowers. In this study, we examined how floral scent changes after pollination in Silene latifolia, which has a specialized relationship with the seed predator Hadena bicruris. We found an overall decrease in total scent emission and considerable changes in relative amounts of scent compounds after pollination. Lilac aldehydes A and B as well as veratrole contributed most to the decrease in scent emission. These three compounds are known to be key signals for the attraction of H. bicruris to the flowers. A specific downregulation of these compounds may increase the reproductive success of the plant by reducing seed predation after pollination.

Animals↗

Detection and transcript expression of S-RNase gene associated with self-incompatibility in apricot (Prunus armeniaca L.).

The identity and expression of S-RNase genotypes in the self-compatible (SC) apricot cultivar 'Katy' and the self-incompatible (SI) cultivar 'Xinshiji' were examined. We used allele specific polymerase chain reaction (AS-PCR) and designated the alleles in 'Katy' and 'Xinshiji' as S(8)Sc and S(9)S(10), respectively. The S-RNase gene was expressed in style at the balloon stage in both genotypes. Using real-time fluorescence quantification RT-PCR technology (FQRT-PCR), spatio-temporal expression patterns of S-RNase gene between 'Katy' and 'Xinshiji' were compared. The results revealed that the expression of the S-RNase gene in 'Katy' and 'Xinshiji' were different. The transcript abundance was distinctly diverse at the key stage (i.e., at 24 h after self-pollination) in both genotypes, and was greater in 'Xinshiji' (SI) than 'Katy' (SC). In addition, the abundance of the S-RNase transcript was higher in upper-half of style than in the lower-half of style or in the ovary. In the SI cultivar 'Xinshiji', the expression of S-RNase reminded a relatively high level after cross-pollination, but it dropped continuously after self-pollination and un-pollination.

Alleles↗

Allergen-controlled study of intranasal immunotherapy for ragweed hay fever.

Previous studies of intranasal immunotherapy have not included control groups treated with an irrelevant allergen. In the present double-blind study, we tested the effectiveness of intranasal immunotherapy in 20 patients sensitive to both short ragweed (SRW) and orchard grass (OG). Patients sprayed increasing concentrations of either SRW (n = 11) or OG (n = 9) extract intranasally six times per day for 8 wk before the SRW pollination season. The effects of this treatment were determined by analysis of symptom score diaries and clinical examinations during the SRW pollination season. SRW-treated patients received cumulative AgE doses from 3 to 59 micrograms (mean 21); this mean dose was approximately sevenfold less than that used in a previous study from our laboratory. All patients reported immediate hay fever symptoms after use of the nasal spray. Five patients (four SRW- and one OG-treated) reported episodes of mild epistaxis during treatment; no other unexpected side effects were noted. During the treatment period, more SRW-treated patients showed signs of nasal obstruction and edematous nasal mucosa than OG-treated control patients (p less than 0.03). During the SRW pollination season, the SRW-treated patients reported lower mean weekly symptom scores than the OG-treated control patients, but the difference was not statistically significant. Supplemental antihistamine use was significantly higher (p less than 0.016) in the OG-treated control patients during the SRW pollination period. Subjective assessment of treatment efficacy by patients was similar in both treatment groups. We conclude that intranasal immunotherapy was of only marginal benefit in this study.

Administration, Intranasal↗

A study of the human immune response to Lolium perenne (rye) pollen and its components, Lol p I and Lol p II (Rye I and Rye II). II. Longitudinal variation of antibody levels in relation to symptomatology and pollen exposure and correction of seasonally elevated antibody levels to basal values.

This study used a standardized, dialyzed, Lolium perenne (ryegrass) pollen extract and two of its well-characterized components, Lol p I (Rye I) and Lol p II (Rye II), to characterize the longitudinal variation of both IgE and IgG antibody (Ab) levels, as well as total serum IgE levels, in 20 grass-allergic subjects followed for 13 months. Ab levels declined toward a basal level just before, and increased just after, the grass-pollination season, returning to the same basal level just before the next grass-pollination season. The least complex allergen, Lol II, demonstrated the most uniform pattern of variation in both IgE and IgG Ab levels. Total serum IgE levels demonstrated the least regular pattern of variation. Grass-pollen counts were strongly correlated with symptom-medication scores for these subjects (rs = 0.87). Initial values were correlated with the rise in total IgE and IgE Ab to Lol II across the grass-pollen season. Skin test results were correlated with initial IgE Ab levels for L. perenne pollen extract and Lol II. Finally, a procedure for correcting IgE Ab levels to basal values was proposed and tested. The correction procedure, for each IgE Ab, was based on the average rise during the grass-pollination season (or average decline after the grass-pollination season) observed for all subjects with that IgE Ab.

Adult↗

Pollen waste and unrelated traits in a fig-fig wasp symbiosis: a new behaviour suggesting a host shift.

In a fig-fig wasp symbiosis, we have discovered that male fig pollinators (Alfonsiella fimbriata Waterston) bite into the dehiscent anthers of Ficus natalensis leprieuri Miq., thus scattering the pollen grains throughout the syconium. Female pollinators are the only ones to transfer pollen to conspecific trees, and collect pollen actively from the anthers only. Thus, this male behaviour appears to be antagonistic to the pollination process. We compare different wasp pollinating behaviours between fig species exhibiting dehiscent and non-dehiscent anthers and conclude that this male behaviour is new and not required with spontaneously dehiscent anthers. These findings could suggest a host shift of Alfonsiella fimbriata.

Animals↗

Orchid diversity: an evolutionary consequence of deception?

The Orchidaceae are one of the most species-rich plant families and their floral diversity and pollination biology have long intrigued evolutionary biologists. About one-third of the estimated 18,500 species are thought to be pollinated by deceit. To date, the focus has been on how such pollination evolved, how the different types of deception work, and how it is maintained, but little progress has been made in understanding its evolutionary consequences. To address this issue, we discuss here how deception affects orchid mating systems, the evolution of reproductive isolation, speciation processes and neutral genetic divergence among species. We argue that pollination by deceit is one of the keys to orchid floral and species diversity. A better understanding of its evolutionary consequences could help evolutionary biologists to unravel the reasons for the evolutionary success of orchids.

Journal Article↗

Juniperus oxycedrus: a new allergenic pollen from the Cupressaceae family.

BACKGROUND: Cupressaceae allergy is a worldwide pollinosis caused by several species. Some species in limited geographic areas pollinate in fall and winter. Juniperus oxycedrus matches these features. OBJECTIVE: We sought to define the immunochemical, allergologic, and environmental aspects of J. oxycedrus pollen. METHODS: Pollen extract from J. oxycedrus was prepared and characterized by biochemical analysis and human specific IgE binding by means of ELISA and immunoblotting. A 3-year phenological study was conducted to define the pollinating period of J. oxycedrus. Forty consecutive patients allergic to cypress were recruited in two areas and divided into two groups according to their exposure to J. oxycedrus pollen. Clinical evaluation, skin prick tests, and specific IgE determination with J. oxycedrus, J. ashei, and Cupressus arizonica extracts were carried out on both groups. RESULTS: J. oxycedrus pollen extract was obtained, and it showed specific IgE binding and wide cross-reactivity with other Cupressaceae species. The extract caused a positive skin test response in all the patients tested, with about 80% of them having detectable specific IgE. Symptoms related to J. oxycedrus pollen exposure were recorded in 72% of the directly exposed patients and occasionally in 9% of the nonexposed patients. In the Mediterranean coastal area considered, J. oxycedrus was the first Cupressaceae species that started to pollinate at the beginning of November and ended in the first part of December. CONCLUSIONS: J. oxycedrus represents a newly characterized pollen species of the Cupressaceae family that cross-reacts with other members of the same family. Subjects with cypress allergy have in vivo and in vitro positive test responses for J. oxycedrus and can show symptoms when exposed to its pollen. Finally, the most important feature of J. oxycedrus is its early pollinating period in southern Europe (Italy), causing a further extension of the cypress pollen season in areas where other Cupressaceae species are present.

Allergens↗

Hybrid seed production and the challenge of propagating male-sterile plants.

The introduction of hybrid crop varieties has enabled spectacular increases in productivity owing to hybrid vigor and increased uniformity. To produce hybrid seeds, a pollination control system is required to prevent unwanted self-pollination. In crop species with hermaphrodite flowers, this can be a major challenge. Over the past decade, new pollination control systems have been developed with the aid of genetic engineering, mainly based on the generation of nuclear-encoded male sterility. The successful application of these systems for large-scale hybrid seed production depends on whether the male-sterile female parent line can be multiplied efficiently and economically. In spite of its relevance, the propagation of the male-sterile line has often been overlooked in the development of pollination control systems.

Crops, Agricultural↗

Non-mutualistic yucca moths and their evolutionary consequences.

Interspecific mutualisms are regarded as having evolved from antagonistic or commensalistic interactions, with most mutualisms remaining facultative but some having coevolved into obligate reciprocal dependency. Underlying mutualism is an intrinsic conflict between the parties, in that each is under selection for increased exploitation of the other. Theoretical models suggest that this conflict is a source of evolutionary instability, and that evolution of 'cheating' by one party may lead to reciprocal extinction. Here we present phylogenic evidence for reversal of an obligate mutualism: within the yucca moth complex, distinct cheater species derived from obligate pollinators inflict a heavy cost on their yucca hosts by laying their eggs but not pollinating the yucca. Phylogenetic data show the cheaters to have existed for a long time. Coexisting pollinators and cheaters are not sister taxa, supporting predictions that evolution of cheating within a single pollinator is evolutionarily unstable. Several lines of evidence support a hypothesis that host shifts preceded the reversal of obligate mutualism. Host or partner shifts is a mechanism that can provide a route of evolutionary escape among obligate mutualists in general.

Animals↗

The value of bees to the coffee harvest.

The self-pollinating African shrub Coffea arabica, a pillar of tropical agriculture, was considered to gain nothing from insect pollinators. But I show here that naturalized, non-native honeybees can augment pollination and boost crop yields by over 50%. These findings, together with world coffee-harvest statistics and results from field studies of organically shade-grown coffee, indicate that coffee plants would benefit from being grown in habitats that are suitable for sustaining valuable pollinators.

Animals↗

The Great Wall of China: a physical barrier to gene flow?

One population from each of six plant species along both sides of the Juyong-guan Great Wall, together with one population from each of five species along both sides of a path on a mountain top near Juyong-guan, were selected to study the effect of the Great Wall as a barrier on genetic differentiation between two subpopulations using RAPD markers. Significant genetic differentiation was found between the subpopulations on both sides of the Great Wall. A wind-pollinated woody species, Ulmus pumila, showed less genetic differentiation than four insect-pollinated species: Prunus armeniaca, Ziziphus jujuba, Vitex negundo, and Heteropappus hispidus. Cleistogenes caespitosa, a wind-pollinated perennial herb, displayed more genetic differentiation between subpopulations than the insect-pollinated species because of its propagation strategy. Although AMOVA analysis showed that subpopulations divided by a mountain path had diverged genetically, the variance component between the subpopulations on both sides of the Great Wall was significantly larger than that between the subpopulations at the control site. Therefore, it is reasonable to deduce that the Juyong-guan Great Wall has served as a physical barrier to gene flow between subpopulations separated for more than 600 years.

Biological Evolution↗

To succeed globally, disperse locally: effects of local pollen and seed dispersal on tetraploid establishment.

Newly formed tetraploid plants in sympatry with their diploid progenitors should face significant obstacles to persistence and population establishment because of low-fitness triploids formed by cross-ploidy pollinations. Prior models have found restrictive conditions for a minority tetraploid subpopulation to persist. A stochastic spatial model, parameterized using snow buttercups (Ranunculus adoneus), was used to examine the influence of limited seed and pollen dispersal distances on the success of minority tetraploids and the interaction of these factors with different rates of self-pollination and tetraploid advantage. Higher rates of self-pollination and increased tetraploid advantage increase the probability of tetraploid persistence. Limiting the dispersal of seeds and pollen further increases the positive impact of any given level of self-pollination and tetraploid advantage. Taxa with short-distance seed and pollen dispersal should face much less stringent barriers to sympatric polyploid speciation than taxa with long-distance dispersal patterns. With short-distance seed and pollen dispersal, polyploid speciation should be possible in the absence of ecological differentiation or recurrent polyploid formation through unreduced gametes.

Crosses, Genetic↗

Fertilization in Arabidopsis thaliana wild type: developmental stages and time course.

We describe some previously uncharacterised stages of fertilization in Arabidopsis thaliana and provide for the first time a precise time course of the fertilization process. We hand-pollinated wild type pistils with wild type pollen (Columbia ecotype), fixed them at various times after pollination, and analysed 600 embryo sacs using Confocal Laser Scanning Microscopy. Degeneration of one of the synergid cells starts at 5 Hours After Pollination (HAP). Polarity of the egg changes rapidly after this synergid degeneration. Karyogamy is then detected by the presence of two nucleoli of different diameters in both the egg and central cell nuclei, 7-8 HAP. Within the next hour, first nuclear division takes place in the fertilized central cell and two nucleoli can then be seen transiently in each nucleus produced. In a second set of experiments, we hand-pollinated wild type pistils with pollen from a transgenic promLAT52::EGFP line that expresses EGFP in its pollen vegetative cell. Release of the pollen tube contents into the synergid cell could be detected in living material. We show that the timing of synergid degeneration and pollen tube release correlate well, suggesting that either the synergid cell degenerates at the time of pollen tube discharge or very shortly before it. These observations and protocols constitute an important basis for the further phenotypic analysis of mutants affected in fertilization.

Arabidopsis↗

Origin of a complex key innovation in an obligate insect-plant mutualism.

Evolutionary key innovations give organisms access to new ecological resources and cause rapid, sometimes spectacular adaptive radiation. The well known obligate pollination mutualism between yuccas and yucca moths is a major model system for studies of coevolution, and it relies on the key innovation in the moths of complex tentacles used for pollen collecting and active pollination. These structures lack apparent homology in other insects, making them a rare example of a novel limb. We performed anatomical and behavioral studies to determine their origin and found evidence of a remarkably simple mechanism. Morphological analyses of the tentacles and adjacent mouthparts in pollinators and closely related taxa showed that the tentacle appears abruptly in female pollinating yucca moths. Several morphological synapomorphies between the galeae, which constitute the characteristic lepidopteran proboscis, and the tentacle suggest that the tentacle evolved quickly through expression of the genetic template for the galea at an apical growth bud on the first segment of the maxillary palp. Behavioral data indicate that tentacle and proboscis movements are controlled by a shared hydraulic extension mechanism, thus no new mechanism was needed for tentacle function. Known developmental paths from other insects can explain the origin of this sex-specific key innovation in a few steps.

Animals↗