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Genotype-environment interactions in pollen competitive ability in an anemophilous tree, Betula pendula Roth.

This study describes genotype-environment interactions in pollen competitive ability expressed as pollen-tube growth rate and seed-siring success in Betula pendula Roth. A factorial crossing design was applied using the same maternal and paternal clones in two different environmental conditions, in a B. pendula seed orchard established in a greenhouse and at an outdoor clone collection. Both single donor and two-donor pollinations were employed. Female inflorescences were collected after a fixed time of germination, pollen-tube lengths were measured for each cross, and paternity of the seeds sired by two-donor pollen mixtures was analyzed using isozyme markers. The pollination site had a significant influence on pollen-tube growth rate and seed-siring success. Significant interactions between pollination site and pollen donor indicated genotype-environment interactions in pollen-tube growth rate and seed-siring success. A highly significant positive correlation between pollen-tube growth rate and seed-siring success was found in the greenhouse but not at the outdoor clone collection. These results suggest that the pollen-tube growth rate can be a predictor of seed-siring success in controlled greenhouse conditions, where differences among maternal plants are mainly of genetic origin, but not in more heterogeneous outdoor conditions. In natural birch stands, environmental maternal effects probably diminish the significance of pollen competition for sexual selection in Betula pendula. At seed orchards, the effects of environmental conditions on pollen competitive ability can have important consequences for the genetic composition of the seed crop.

Journal Article↗

Effect of a gap on gene flow between otherwise adjacent transgenic Brassica napus crops.

Gene flow resulting from cross pollination becomes an issue when transgenic crops are involved and the genetic modification carries a trait of ecological importance. As crop fields are often separated by a barren gap, such as an intervening roadway or unplanted area, I measured cross contamination between two herbicide-resistant transgenic fields (canola, Brassica napus) across a gap of up to 12 m. I focused on pollen exchange from the field border up to 7 m inside each field over two seasons. In the absence of a gap, I found that gene dispersal diminished rapidly with distance, with more than 40% of transgenic progeny found within the first meter from the edge of the adjacent crop. Cross contamination between fields declined more rapidly when there were intervening plants, however. Plants separated from the transgenic source by a gap of 3-4 m, yielded the same level of transgenic progeny as those separated by 1 m of crop. Both insects and wind pollinate canola, and so the explanation for my observations could involve the influence of gaps on wind patterns or on the behaviour of pollinators. The gap effect does not seem to depend only upon the variation in the density of neighbours that surrounds those plants at the crop edge versus those in the crop matrix. On the basis of this study, it is recommended that economic profit would be maximised by removing field borders after flowering rather than by leaving a surrounding gap, which would need to occupy up to threefold as much field surface to achieve the same level of containment.

Analysis of Variance↗

Inbreeding and true seed in tetrasomic potato. IV. Synthetic cultivars.

A true potato seed (TPS) synthetic cultivar from open pollination may broaden the genetic base in potato. A synthetic cultivar can also reduce the effect of inbreeding and maintain the productivity from generation to generation. Selected tetraploid parental genotypes, based on combining-ability tests, were chosen for isolated polycrosses to obtain tetraploid offspring. A series of experiments were undertaken to investigate the development of these TPS synthetic populations in two Peruvian locations: San Ramon, a rainfed, humid, mid-altitude environment, and La Molina, an arid, coastal environment under irrigation. Natural open-pollinated synthetics or control-mixture synthetics, involving two to six parents, had a similar tuber set but plant survival and tuber weight were higher in control synthetics including two or six parents. The results suggest that two-parent open-pollinated TPS synthetics may be a feasible option by selecting the right parents for the base population. A subsequent experiment showed that four-parent control synthetics (i.e. double crosses) were the best for tuber weight and size, followed by the two-parent control synthetics. Synthetic breeding populations could be shared with other breeders, who in cooperation with their local farmers may select promising genotypes for further cultivar release.

Journal Article↗

Cloning and characterization of the gene encoding 3-hydroxy-3-methylglutaryl coenzyme A reductase in melon (Cucumis melo L. reticulatus).

We have isolated a cDNA for Cm-HMGR, encoding 3-hydroxy-3-methylglutaryl coenzyme A (HMG-CoA) reductase in melon (Cucumis melo L. reticulatus; Genbank Accession No. AB021862). Cm-HMGR encodes a polypeptide of 588 amino acids that contains two transmembrane domains and a catalytic domain. Database searches revealed that Cm-HMGR shows homology to HMG1 (63.7%) and HMG2 (70.3%) of tomato, to HMG1 (77.2%) and HMG2 (69.4%) of Arabidopsis thaliana, and to HMGR of tobacco (72.6%). Functional expression in a HMG-CoA reductase-deficient mutant yeast showed that Cm-HMGR products mediate the synthesis of mevalonate. Northern analysis revealed that the level of Cm-HMGR mRNA in the fruit increased after pollination and markedly decreased at the end of fruit enlargement. During ripening, Cm-HMGR mRNA levels increased markedly in the fruit. In parallel with mRNA expression, Cm-HMGR activity increased after pollination, whereas no Cm-HMGR activity was detectable during fruit ripening. Our results suggest that Cm-HMGR is important during early post-pollination development of the fruit in melon.

Amino Acid Sequence↗

Seasonal changes in pollen-packaging schedules in the protandrous plant Chamerion angustifolium.

The rate at which pollen is presented varies widely both among and within species of flowering plants. Although several studies have tried to explain between-species variation in this trait, few explanations exist for the variation that exists among individuals of the same species. I used protandrous fireweed, Chamerion angustifolium (Onagraceae), to examine seasonal changes in pollinator visitation, population sex ratio, and pollen presentation schedules. Most studies that make predictions about optimal pollen presentation assume conditions are constant throughout the flowering season. However, data presented here show that pollen presentation schedules vary seasonally: in early season flowers, anthers dehisce over a number of days, while late season flowers present all anthers simultaneously. I show that pollinator visitation rates to individual plants are higher but more variable early in the season than late in the season. Furthermore, per-male availability of female-phase flowers increases over the course of the flowering season in this population. I suggest that seasonal differences in the variability of pollinator visitation and the availability of female flowers has played an important role in selection for pollen presentation schedules.

Animals↗

Flowering phenology and compensation for herbivory in Ipomopsis aggregata.

The mechanisms and circumstances that affect a plant's ability to tolerate herbivory are subjects of ongoing interest and investigation. Phenological differences, and the timing of flowering with respect to pollinators and pre-dispersal seed predators, may provide one mechanism underlying variable responses of plants to herbivore damage. The subalpine wildflower, Ipomopsis aggregata, grows across a wide range of elevations and, because phenology varies with elevation, phenological delays associated with elevation may affect the ability of I. aggregata to compensate for or tolerate browsing. Thus, we examined the response of I. aggregata to herbivory across an elevation gradient and addressed the interactions among phenological delays imposed by damage, elevation, pre-dispersal seed predation and pollination, on I. aggregata's compensatory response. Among high and low elevation populations in areas near the Rocky Mountain Biological Laboratory (RMBL) in Gothic, Colorado, we compared the responses of naturally browsed, artificially browsed (clipped), and unbrowsed (control) plants of I. aggregata. We compared responses in the date of initiation of flowering, timing of peak bloom, floral display, nectar production and sugar concentration, oviposition and fruit destruction by the pre-dispersal seed predator Hylemya sp. (Anthomyiidae), fruit production, and aboveground biomass production. Clipping had the greatest effect on reproductive success and clipped plants at high elevation exhibited the lowest tolerance for herbivory. The effects of browsing appear to be mediated by flowering phenology, and both browsing and elevation delayed flowering phenology. Time needed for regrowth delays flowering, and thus affects the overlap with seed predators and pollinators. As a result of delayed flowering, naturally browsed and clipped plants incurred lower rates of seed predation. In the absence of seed predation, plants would exhibit a lower tolerance to herbivory since naturally and artificially browsed plants had fewer fruits destroyed by Hylemya larvae. We provide additional evidence that, for populations near the RMBL, clipping and natural browsing do not have the same effect on I. aggregata plants. This may be due to the selection of larger plants by herbivores. Although under some conditions plants may tolerate browsing, in areas where the growing season is short a phenological delay imposed by damage is likely to significantly reduce plant fitness. Identifying the mechanisms that allow plants to tolerate herbivore damage will help to develop a general framework for understanding the role of tolerance in plant population and community dynamics, as well as plant-herbivore interactions.

Adaptation, Physiological↗

Density-dependent seed set in the Haleakala silversword: evidence for an Allee effect.

Plant species may be subject to Allee effects if individuals experience a reduction in pollination services when populations are small or sparse. I examined temporal variation in reproductive success of the monocarpic Haleakala silversword (Argyroxiphium sandwicense subsp. macrocephalum) over five years, to determine if plants flowering out of synchrony with most of the population (i.e., in low flowering years) exhibited lower percent seed set than synchronously-flowering plants (i.e., those flowering in high flowering years). Through two pollination experiments conducted over multiple years, I also measured pollen limitation and self-incompatibility in this species. The number of flowering plants varied greatly among years, as did reproductive success. Percent seed set was significantly correlated with the number of plants flowering annually, such that plants flowering in high flowering years (1997 and 2001) exhibited significantly higher percent seed set than did plants flowering in low flowering years (1998-2000). In the 3-year pollen limitation study, plants flowering asynchronously were pollen-limited, whereas plants flowering synchronously were not. This species is strongly self-incompatible. Results of this study demonstrate that the Haleakala silversword experiences reduced reproductive success in low flowering years, and suggest that this Allee effect is pollinator-mediated. Allee effects in plants are an understudied yet potentially important force with implications for the population dynamics and conservation of rare species.

Asteraceae↗

Experimental evaluation and thermo-physical analysis of thermogenesis in male and female cycad cones.

Thermogenically elevated cone temperatures were measured in two Macrozamia cycad species that differ in their daily heating time. Mathematical models of the cones' thermo-physics were tested for their accuracy in predicting these cone temperatures and for comparison of the energetics of both species and the sexes within species. These models accurately predicted temperatures over approximately 8-h periods with average errors of: 0.46 degrees C for Macrozamia lucida, pollinated by the thrips, Cycadothrips chadwicki, that moves during mid-day concurrent with cone heating; and 0.38 degrees C for Macrozamia machinii, pollinated by the weevil, Tranes sp., that moves after sunset during cone heating. The combination of models and experiments revealed a thermogenic sexual dimorphism in both species. For M. lucida, the estimated female mass specific metabolisms, and their theoretically possible and actual temperature increases due to thermogenic metabolism were only 57, 67, and 76% of males. In addition, female thermogenic metabolisms began and peaked much earlier and lasted significantly longer than males (all differences >1 h), and female metabolic peaks preceded their temperature peaks by 65 vs. 46 min for males. The timing of almost all male cone metabolic peaks was optimized with respect to the diurnal ambient heating cycle so that cone temperatures achieved a maximum temperature gain, whereas most female metabolic peaks occurred much earlier than optimal. In M. machinii, thermogenic sexual dimorphism is much larger since its male peak metabolisms are larger, and its females' peaks are much smaller compared to those of M. lucida. This study provides new information regarding the energetics of cycad cones that is relevant to understanding the interactions of the plant traits with their obligate pollinators' behavior.

Animals↗

Variation in resource limitation of plant reproduction influences natural selection on floral traits of Asclepias syriaca.

The availability of both pollen and resources can influence natural selection on floral traits, but their relative importance in shaping floral evolution is unclear. We experimentally manipulated pollinator and resource (fertilizer and water) availability in the perennial wildflower Asclepias syriaca L. Nine floral traits, one male fitness component (number of pollinia removed), and two female fitness components (number of pollinia inserted and number of fruits initiated) were measured for plants in each of three treatments (unmanipulated control, decreased pollinator access, and resource supplementation). Although decreasing pollinators' access to flowers did result in fewer pollinia inserted and removed, fruit set and phenotypic selection on floral traits via female and male fitness did not differ from the control. In contrast, resource supplementation increased fruit set, and phenotypic selection on seven out of nine floral traits was stronger via female than male fitness, consistent with the prediction that selection via female fitness would be greater when reproduction was less resource-limited. Our results support the hypothesis that abiotic resource availability can influence floral evolution by altering gender-specific selection.

Analysis of Variance↗

Conflict resolution in an ant-plant interaction: Acacia constricta traits reduce ant costs to reproduction.

Many plant species attract ants onto their foliage with food rewards or nesting space. However, ants can interfere with plant reproduction when they visit flowers. This study tests whether Acacia constricta separates visiting ant species temporally or spatially from newly opened inflorescences and pollinators. The diurnal activity patterns of ants and A. constricta pollinators peaked at different times of day, and the activity of pollinators followed the daily dehiscence of A. constricta inflorescences. In addition to being largely temporally separated, ants rarely visited open inflorescences. A floral ant repellent contributes to the spatial separation of ants and inflorescences. In a field experiment, ants of four species were given equal access to inflorescences in different developmental stages. On average, the frequency with which ants made initial, antennal contact with the floral stages did not differ, but ants significantly avoided secondary contact with newly opened inflorescences relative to buds and old inflorescences, and old inflorescences relative to buds. Ants also avoided contact with pollen alone, indicating that pollen is at least one source of the repellent. The results suggest A. constricta has effectively resolved the potential conflict between visiting ants and plant reproduction.

Acacia↗

An objective classification system of air mass types for Szeged, Hungary, with special attention to plant pollen levels.

This paper discusses the characteristic air mass types over the Carpathian Basin in relation to plant pollen levels over annual pollination periods. Based on the European Centre for Medium-Range Weather Forecasts dataset, daily sea-level pressure fields analysed at 00 UTC were prepared for each air mass type (cluster) in order to relate sea-level pressure patterns to pollen levels in Szeged, Hungary. The database comprises daily values of 12 meteorological parameters and daily pollen concentrations of 24 species for their pollination periods from 1997 to 2001. Characteristic air mass types were objectively defined via factor analysis and cluster analysis. According to the results, nine air mass types (clusters) were detected for pollination periods of the year corresponding to pollen levels that appear with higher concentration when irradiance is moderate while wind speed is moderate or high. This is the case when an anticyclone prevails in the region west of the Carpathian Basin and when Hungary is under the influence of zonal currents (wind speed is high). The sea level pressure systems associated with low pollen concentrations are mostly similar to those connected to higher pollen concentrations, and arise when wind speed is low or moderate. Low pollen levels occur when an anticyclone prevails in the region west of the Carpathian Basin, as well as when an anticyclone covers the region with Hungary at its centre. Hence, anticyclonic or anticyclonic ridge weather situations seem to be relevant in classifying pollen levels.

Air Movements↗

Pinus pollen in the atmosphere of Vigo and its relationship to meteorological factors.

The Pinus genus has an elevated pollen production and an anemophilous nature. Although considered to be hypoallergenic, numerous cases of allergies caused by Pinus pollen have been cited and different authors believe that its allergenicity should be studied in more depth. In the city of Vigo several patients have tested positive for Pinus pollen extracts in skin tests, some of them being mono-sensitive to such pollens. In order to ascertain the behaviour of Pinus pollen and its correlation to the main meteorological factors, we carried out an aerobiological study in the city of Vigo from 1995 to 1998 by using a Hirst active-impact volumetric sporetrap, model Lanzoni VPPS 2000, placed on the left bank of the Vigo estuary (42 degrees 14'15"N, 8 degrees 43'30"W). Pinus has high quantitative importance in the airborne pollen spectrum of the city. It is one of the best represented taxa constituting 13%-20% of the total annual pollen levels. The quantity of Pinus pollen present in the atmosphere of the city of Vigo throughout a year is 5,751 grains (as the average for the sampled years), with a very long pollination period, from the middle of January until May. The maximum concentration was recorded in 1998 with 1,105 grains/m(3) on 3 March, a much greater value than those for the previous years. At the end of its pollination period there is usually a final increase in Pinus pollen concentrations coinciding with the pollination of Pinus silvestris, which are more abundant in mountainous areas far from the city.

Humans↗

Estragole (4-allylanisole) is the primary compound in volatiles emitted from the male and female cones of Cycas revoluta.

The genus Cycas (Cycadaceae; gymnosperm) have circumstantially been considered to be wind-pollinated. The cones of Cycas revoluta Thunb., however, emit a strong unpleasant odor. The chemical profiles of floral scents often correlate with various pollination modes (pollinators). We collected and analyzed the volatiles emitted from male and female cones of C. revoluta native to Iriomote and Yonaguni Islands, Japan. The analyses indicated that estragole (4-allylanisole) dominated in the volatiles (67.0-92.7%), with small amounts of other benzenoids, e.g., anethole, methyl salicylate, methyl eugenol, and ethyl benzoate. Several fatty acid esters were also detected in the samples from Iriomote Island. The function of estragole in the reproductive biology of C. revoluta is discussed.

Allylbenzene Derivatives↗

Floral scent chemistry of mangrove plants.

The flowers of mangrove plants are pollinated by a variety of pollinators including birds, bats, and insects. This study analyzed the floral scent chemistry of mangroves on Iriomote Island (located near Taiwan) including Bruguiera gymnorrhiza (L.) Lamk. (Rhizophoraceae), Kandelia candel (L.) Druce (Rhizophoraceae), Rhizophora stylosa Griff. (Rhizophoraceae), Sonneratia alba J. Smith (Sonneratiaceae), Nypa fruticans (Thunb.) Wurmb. (Palmae), Lumnitzera racemosa Willd. (Combretaceae), Avicennia marina (Forsk.) Vierh. (Avicenniaceae or Verbenaceae), and Pemphis acidula Forst. (Lythraceae). A total of 61 chemicals (fatty acid derivatives, terpenoids, carotenoid derivatives, benzenoids, nitrogen-containing compounds, 13 unknown chemicals) were detected in the floral scents of the various species. The species displayed a distinct chemical profile ranging from only two chemicals in the floral scent of Kandelia candel to more than 25 chemicals in the floral scent of Nypa fruticans. All of the identified chemicals have been found in the floral scents of other angiosperms. The chemical profile of some species can be correlated with their floral morphology and pollinators.

Journal Article↗

Strong conservation of floral scent composition in two allopatric yuccas.

Floral scent has been suggested to play a key role in the obligate pollination mutualism between yuccas and yucca moths. We analyzed floral fragrance compounds of Yucca elata with headspace collection followed by gas chromatography and mass spectrometry, and compared the odor blend with the recently characterized blend of the allopatric Yucca filamentosa. A principal component analysis based on 20 scent compounds revealed that the floral odor bouquets of Y. elata and Y. filamentosa are virtually identical. Although the two plants belong to the same section of capsular-fruited yuccas, they rely on different species of Tegeticula moths for pollination and probably have been allopatric for several million years. Yet, their floral odor blends are very similar, which may indicate that strong selection by obligate pollinators counteracts drift or divergence in this trait.

Animals↗

Flower scent of floral oil-producing Lysimachia punctata as attractant for the oil-bee Macropis fulvipes.

Most flowers offer nectar and/or pollen as a reward for pollinators. However, some plants are known to produce mostly fatty oil in the flowers, instead of nectar. This oil is exclusively collected by specialized oil-bees, the pollinators of the oil-plants. Little is known about chemical communication in this pollination system, especially how the bees find their hosts. We collected the floral and vegetative scent emitted by oil-producing Lysimachia punctata by dynamic headspace, and identified the compounds by gas chromatography coupled to mass spectrometry. Thirty-six compounds were detected in the scent samples, several of which were flower-specific. Pentane extracts of flowers and floral oil were tested on Macropis fulvipes in a biotest. Flower and oil extracts attracted the bees, and some of the compounds identified are seldom found in the floral scent of other plants; these may have been responsible for the attraction of the bees.

Animals↗

Microarray analysis reveals similarities and variations in genetic programs controlling pollination/fertilization and stress responses in rice (Oryza sativa L.).

Previously, we identified 253 cDNAs that are regulated by pollination/fertilization in rice by using a 10K cDNA microarray. In addition, many of them also appeared to be involved in drought and wounding responses. To investigate this relationship, we obtained their expression profiles after dehydration and wounding treatments in this study. Venn diagram analysis indicated that 53.8% (136/253) and 21% (57/253) of the pollination/fertilization-related genes are indeed regulated by dehydration and wounding, respectively, and nearly half of the genes expressed preferentially in unpollinated pistils (UP) are responsive to dehydration. These results indicated that an extensive gene set is shared among these responses, suggesting that the genetic programs regulating them are likely related. Among them, the genetic network of water stress control may be a key player in pollination and fertilization. Additionally, 39.5% (100/253) cDNAs that are related to pollination/fertilization appear not to be regulated by the stress treatments (dehydration and wounding), suggesting that the existence of additional genetic networks are involved in pollination/fertilization. Furthermore, comparative analysis of the expression profiles of the 253 cDNAs under 18 different conditions (various tissues, treatments and developmental status) revealed that the genetic networks regulating photosynthesis, starch metabolisms, GA- and defense-responses are involved in pollination and fertilization. Taken together, these results provided some clues to elucidate the molecular mechanisms of pollination and fertilization in rice.

Cluster Analysis↗

Crop-to-crop gene flow using farm scale sites of oilseed rape (Brassica napus) in the UK.

From 2000-2003 a range of Farm Scale Evaluation (FSE) trials were established in the UK to assess the effect of the release and management of herbicide tolerant (HT) crops on the abundance and diversity of farmland wildlife compared with their conventionally managed non-GM-equivalents. The objective of this research project was to investigate gene flow within the winter (WOSR) and spring oilseed rape (SOSR) FSE trials and to develop a statistical model for the prediction of cross-pollination frequency that can be used to evaluate current separation distance guidelines. Seed samples were collected from the non-GM half of the trial sites and were tested for evidence of cross-pollination from the GM HT halves using a quantitative PCR assay specific to the HT (bar) gene. Rates of cross-pollination were found to decrease rapidly with increasing distance from the GM source. The quantitative data were subjected to statistical analysis and a two-step model was found to provide the best fit for the data. Significant differences were found between the results for WOSR, SOSR and varietal association (VA) crops. The model predicted that the %GM content (including upper 95% confidence limits) of a sample taken at a distance of 50 m away from the GM source would be 0.04% (0.84%) for WOSR, 0.02% (0.39%) for SOSR, 0.77% (21.72%) for WOSR VA and 0.37% (5.18%) for SOSR VA. The data and models presented here are discussed in the context of necessary separation distances to meet various possible thresholds for adventitious presence of GM in OSR.

Agriculture↗