Generalized pollination system: Are floral traits adapted to different pollinators?

Flowers that are pollinated both during the day and at night could exhibit two different groups of pollinators and produce two different sets of attractants and rewards. We explored the patterns of emission of flower scents and production of nectar in the cactus Echinopsis chiloensis ssp. chiloensis, in relation to the patterns of activity of its diurnal and nocturnal pollinators. We measured frequency of flower visitors, analyzed floral scents, measured nectar production and sugar concentration, and performed pollination exclusion experiments. Bees were the main visitors at daytime and hawkmoths at nighttime. Diurnal scents were dominated by several compounds that can attract a wide range of pollinators, whereas nocturnal scents were less diverse and were dominated by (E)-nerolidol, a compound eliciting antennal responses in hawkmoths. Nectar volume and sugar concentration at night were similar to those recorded in hawkmoth-pollinated flowers. Daytime nectar volume was higher than those commonly found in bee-pollinated flowers, but similar to those found in flowers pollinated by several pollinators. Daytime sugar concentration was similar to those recorded in bee-pollinated flowers. Flowers of E. chiloensis ssp. chiloensis seem morphologically adapted to hawkmoth pollination, but diurnal and nocturnal pollinators contribute to similar extents to reproductive success. Additionally, diurnal and nocturnal pollinators showed a synergic effect on the product of fruit set and seed set. The results are discussed in terms of the linkage between floral traits and perception abilities and requirements of pollinators.     

A quantitative review of pollination syndromes: do floral traits predict effective pollinators?

The idea of pollination syndromes has been largely discussed but no formal quantitative evaluation has yet been conducted across angiosperms. We present the first systematic review of pollination syndromes that quantitatively tests whether the most effective pollinators for a species can be inferred from suites of floral traits for 417 plant species. Our results support the syndrome concept, indicating that convergent floral evolution is driven by adaptation to the most effective pollinator group. The predictability of pollination syndromes is greater in pollinator‐dependent species and in plants from tropical regions. Many plant species also have secondary pollinators that generally correspond to the ancestral pollinators documented in evolutionary studies. We discuss the utility and limitations of pollination syndromes and the role of secondary pollinators to understand floral ecology and evolution.     

Phylogenetic analysis of floral integration in Schizanthus (Solanaceae): does pollination truly integrate corolla traits?

To assess whether floral integration patterns result from the action of pollinator selection on functionally related traits, we compared corolla integration patterns in eight Schizanthus species differing in pollination systems and in their degree of pollinator dependence across a molecular phylogeny. Integration patterns differed among species and these differences were not related to their phylogenetic relatedness. When the putative original function of some corolla traits was lost in pollinator-dependent species, the integration among nonfunctional characters and the rest of the corolla traits was disrupted. This pattern was not presented in species adapted for late autonomous selfing, which exhibited higher corolla integration than their pollinator-dependent relatives. These results suggest that corolla integration in pollinator-dependent species was shaped by pollinator-mediated selection. Decoupling of nonfunctional traits in these species may result from a relaxation of correlational selection or from selection acting against a default covariation provided by genetic and developmental connections.     

A survey of floral traits, breeding systems, floral visitors, and pollination systems of the angiosperms of the Juan Fernández Islands (Chile)

A survey of the reproductive features of the angiosperm flora of the Juan Fernández Archipelago (Chile) is presented to provide a species-based review of reproduction and pollination, to identify generalizations associated with these systems, to understand the evolution of these features, and to utilize these data to promote conservation. The collection of original data was extensive, based on our own fieldwork, and was combined with data from existing literature. Data recorded include habit, sexual system, flower size, shape, and color, and the hypothesized pollination system of the first colonizers. In addition, the data on compatibility, presence and type of dichogamy, observed floral visitors, presence of floral rewards, and currently known pollination systems are summarized. Pairwise comparisons of different features are tested for statistical association. The flora is typically composed of perennials. The majority of the species have very small or small flowers. Inconspicuous flowers (i.e., a shape character describing flowers with no optical attraction) are widespread, as are dish-shaped flowers. Green is the most frequent flower color, followed by white and yellow. Most species are hermaphroditic, 9% are dioecious, and 9% are monoecious. Some 30% of the species are protandrous and 7% protogynous. Detailed studies of compatibility of about 14% of the flora indicate that 85% of these species are self-compatible (SC). Although most species studied are SC, their level of autogamy is low. Nevertheless, selfing mediated via geitonogamy is the most frequent mechanism of pollen transfer. Outcrossing is mainly achieved through dioecy and self-incompatibility (SI), promoted by dichogamy in the hermaphroditic flowers, and facilitated by wind pollination. About 55% of the species offer nectar rewards, and only 2% offer pollen rewards. Floral visitors are rare to uncommon. Two hummingbird species, one of them endemic, are considered as pollinators for 14 plant species. Flies, moths, and beetles are the native insect visitors to flowers, but they have been documented on only 11 plant species (7%). Even insect visits to these few species were rarely observed. Given the infrequent, irregular, and imprecise nature of native insect association with flowers, there is no certainty that any of the species are truly insect pollinated. Two species of introduced ants and a new endemic bee were recorded as well; however, neither is likely currently important to the pollination of the native flora species. About 9% of the extant flora is currently bird pollinated, and we hypothesize that 47% is wind pollinated. However, we propose that most of the colonizers were ancestrally either insect or wind pollinated. There is association between a number of current floral features and the hypothesized pollination of colonizers. Therefore, to a large extent the flower color, shape, and size of the extant flora may express the pollination syndromes of colonizers rather than representing extant pollination. In addition, the presence of nectar in many species of extant flora does not necessarily indicate biotic pollination. Thus, studies of the reproductive biology on oceanic island plants need to be conducted species by species before broad generalizations can be made, because the observed features can be misleading. Possible changes in the pollination system were assessed by comparison of species for which there are reliable data with the hypothesized pollination of their colonist progenitors. The wind-or bird-pollinated species have retained the pollination system of the colonizers. In other instances, species seem to bear a different pollination system: from ancestral insect systems to current hummingbird-or wind-pollination systems. The lack of alternative means of biotic pollination seems to have led in a number of instances to anemophily—in essence a default pollination system. The lack of strong selection pressure for wind pollination and the relative youth of the archipelago may help explain why the features associated with wind pollination in these species are not so obvious. Because there are many recorded extinctions of vascular plants from islands versus those from continental areas, it is imperative to invest additional effort in protecting the remaining island species. Conservation or restoration programs cannot be effective without a deep and broad understanding of the reproductive biology of the plants. In order to conserve these plants, programs must involve a combination of reproductive and environmental measures. The ultimate fate of some species may depend on preserving the plant-hummingbird relationship, including the web of organisms that affect both plant and pollinator. The populations of introduced animals and weeds must be controlled. Experimentally produced allogamous seeds would enhance diversity in restoration programs. In addition, the preservation of habitat seems to be the central challenge to indirectly protect the unique island species.     

Are changes in N:P ratios in coastal waters the key to increased red tide blooms?

There is mounting evidence of a global increase in nutrient levels of coastal waters through riverine and sewage inputs, and in both the numbers and frequency(as well as the species composition) of red tides. However, it is still not possible to conclude the extent to which the increase in red tides in coastal waters can be attributed to the increase in nutrient levels, since so many other factors are involved. Undoubtedly, a relationship exists between red tides and the N and P load of coastal waters, and many nutrient enrichment experiments have shown that marine phytoplankton blooms are often nutrient limited. What is now becoming clear, however, is that although in classical Liebigian terms minimum amounts can be limiting, nutrient ratios (such as N:P and Si:P) are far more important regulators. This paper reviews evidence collected by the authors from Tolo Harbour, Hong Kong together with data collected in Japanese and North European coastal waters by various authors, which indicates that both long term and relatively short term changes in the N:P ratio are accompanied by increased blooms of non-siliceous phytoplankton groups and, furthermore,that the growth of most red tide causative organisms in Hong Kong coastal water is optimized at a low N:P(atomic) ratio of between 6 and 15. This revised version was published online in August 2006 with corrections to the Cover Date.     

Are there any agricultural effects on the capture rates of male euglossine bees (Apidae: Euglossini)?

During 30 days male euglossine bees were bait-sampled at 12 sites, in the central Pacific coast of Colombia (ten days and four sites at each of three adjacent habitats: farmlands, highly disturbed forest and less disturbed forest) and 487 individuals were captured. Most captured individuals belonged to six species, five widely distributed through the American tropics and an endemic species. Two of the frequently captured species presented no different abundances between habitats, while the other four (67.97% of all the captured individuals), all of them big sized or long-tongued, were more frequently captured at the farmlands. A cluster analysis of the data matrix for the 23 captured species and the 12 sampling sites, grouped together the farmland sites, apart from the forest sites. It is proposed that male euglossine bees from generalist, long-tongued or big sized species, forage frequently at the farmlands, where fragrance or nectar resources may be clumped, less diverse, and present an access restricted by deep corollas or by microclimatic conditions of high temperature and low humidity.     

Can phenotypic selection on floral traits explain the presence of enigmatic intermediate individuals in sympatric populations of Platanthera bifolia and P. chlorantha (Orchidaceae)?

Pollinators represent one of the main agents of selection on floral traits. Here, we estimated phenotypic selection on floral morphology and phenology in a sympatric population of two orchid species, Platanthera bifolia and P. chlorantha, including enigmatic individuals with intermediate column morphology (as reflected by the distance between viscidia and caudicle length, two traits involved in assortative mating and reproductive isolation among Platanthera species), but genetically indistinguishable from P. bifolia. Our aim was to clarify whether the occurrence of intermediate phenotypes could be explained by the presence of selective pressures exerted by pollinators. Simple linear and quadratic regressions together with univariate and multivariate analyses were used to evaluate the strength of directional, disruptive and stabilizing selection. We found that selection on phenotypic traits varied between groups and sex functions. Contrary to our hypothesis, selection on the viscidia distance and caudicle length appeared to be consistent in the two P. bifolia groups. Interestingly, the viscidia distance was under significant stabilizing selection through female reproductive success in intermediate individuals. Based on these results, we conclude that, despite a significant selective pressure on some phenotypic traits, the presence of individuals with intermediate phenotype is not due to selection. Stabilizing selection on distance between viscidia in intermediate individuals may suggest that assortative mating play a role in the maintenance of this phenotypic polymorphism.     

A quick screening method to identify β-glucosidase activity in native wine yeast strains: application of Esculin Glycerol Agar (EGA) medium

One hundred and fifty-four yeast strains were isolated from grapes and musts of Uruguayan vineyards and wineries. Only thirty strains showed β-glucosidase activity in Esculin Glycerol Agar (EGA) solid medium. Twenty-one were non-Saccharomyces and nine were Saccharomyces cerevisiae strains. The objective of this study was to evaluate the suitability of Esculin Glycerol Agar (EGA) solid medium for screening β-glucosidase activity in native yeasts strains. Halo sizes measured in the EGA solid medium were correlated to the Glycosyl-Glucose (GG) indexes measured after fermentation of grape musts with each strain. The two S. cerevisiae strains with the best performance were selected for further fermentations on a Muscat Miel grape must, rich in bound monoterpenes. The levels of free linalool, hodiol I and geraniol increased significantly as compared to fermentation with a commercial wine yeast strain. These results show the suitability of this simple and economic medium to identify S. cerevisiae glucosidase producers with a potential impact on real winemaking conditions. On the other hand, great variability was found for the non-Saccharomyces strains, and this would demand further studies for each species. In conclusion, the use of EGA solid medium shows that the screening method is suitable for exploring the glucosidase activity of native strains of S. cerevisiae and shows good correlation with its real impact on free aroma compounds in the final wine.     

Modulating the expression of aquaporin genes in planta: A key to understand their physiological functions?

Aquaporins (AQPs) are believed to act as “cellular plumbers”, allowing plants to rapidly alter their membrane water permeability in response to environmental cues. This study of AQP regulation at both the RNA and protein levels has revealed a large number of possible mechanisms. Currently, modulation of AQP expression in planta is considered the strategy of choice for elucidating the role of AQPs in plant physiology. This review highlights the fact that this strategy is complicated by many factors, such as the incomplete characterization of transport selectivity of the targeted AQP, the fact that AQPs might act as multifunctional channels with multiple physiological roles, and the number of post-translational regulation mechanisms. The classification of AQPs as constitutive or stress-responsive isoforms is also proposed.     

Are abdominal prolegs serially homologous with the thoracic legs in panorpidae (Insecta: Mecoptera)? embryological evidence

It is a long standing question whether the abdominal prolegs of holometabolous insect larvae are serially homologous with their thoracic legs. The histology and ultrastructure of proleg embryonic development in the scorpionfly Panorpa magna were studied using light and scanning electron microscopy. During the early embryonic development, paired primary abdominal appendages appeared laterally in line with the thoracic legs. Several hours later, a pair of proleg primordia arose along the midventral line on each of the first eight abdominal segments mesial to the primary abdominal appendages, which then ceased to grow and eventually degenerated into flat vestiges. Histological observation showed that the thoracic legs were obviously connected with lateral thoracic muscle cells, whereas the abdominal prolegs resembled secondary outgrowths. No apparent contact was observed between the lumen of abdominal prolegs and the hemocoel. After dorsal closure, each thoracic segment bore a pair of well‐developed five‐jointed legs, whereas the prolegs were unjointed, fleshy structure. The remnants of the primary abdominal appendages could still be clearly seen in the mature embryo. On the basis of the histological and morphological observation of the embryonic development, we confirm that the abdominal prolegs of Panorpidae lack the characters of the primary appendages; hence they are not serially homologous with the thoracic legs. The reasons why the primary abdominal legs are reduced in scorpionflies are briefly discussed. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

Review: Are there indigenous Saccharomyces in the digestive tract of livestock animal species? Implications for health,nutrition and productivity traits

All livestock animal species harbour complex microbial communities throughout their digestive tract that support vital biochemical processes, thus sustaining health and productivity. In part as a consequence of the strong and ancient alliance between the host and its associated microbes, the gut microbiota is also closely related to productivity traits such as feed efficiency. This phenomenon can help researchers and producers develop new and more effective microbiome-based interventions using probiotics, also known as direct-fed microbials (DFMs), in Animal Science. Here, we focus on one type of such beneficial microorganisms, the yeast Saccharomyces. Saccharomyces is one of the most widely used microorganisms as a DFM in livestock operations. Numerous studies have investigated the effects of dietary supplementation with different species, strains and doses of Saccharomyces (mostly Saccharomyces cerevisiae) on gut microbial ecology, health, nutrition and productivity traits of several livestock species. However, the possible existence of Saccharomyces which are indigenous to the animals’ digestive tract has received little attention and has never been the subject of a review. We for the first time provide a comprehensive review, with the objective of shedding light into the possible existence of indigenous Saccharomyces of the digestive tract of livestock. Saccharomyces cerevisiae is a nomadic yeast able to survive in a broad range of environments including soil, grass and silages. Therefore, it is very likely that cattle and other animals have been in direct contact with this and other types of Saccharomyces throughout their entire existence. However, to date, the majority of animal scientists seem to agree that the presence of Saccharomyces in any section of the gut only reflects dietary contamination; in other words, these are foreign organisms that are only transiently present in the gut. Importantly, this belief (i.e. that Saccharomyces come solely from the diet) is often not well grounded and does not necessarily hold for all the many other groups of microbes in the gut. In addition to summarizing the current body of literature involving Saccharomyces in the digestive tract, we discuss whether the beneficial effects associated with the consumption of Saccharomyces may be related to its foreign origin, though this concept may not necessarily satisfy the theories that have been proposed to explain probiotic efficacy in vivo. This novel review may prove useful for biomedical scientists and others wishing to improve health and productivity using Saccharomyces and other beneficial microorganisms.     

Energy expenditure in Crocidurinae shrews (Insectivora): is metabolism a key component of the insular syndrome?

A cascade of morphological, ecological, demographical and behavioural changes operates within island communities compared to mainland. We tested whether metabolic rates change on islands. Using a closed circuit respirometer, we investigated resting metabolic rate (RMR) of three species of Crocidurinae shrews: Suncus etruscus, Crocidura russula, and C. suaveolens. For the latter, we compared energy expenditure of mainland and island populations. Our measurements agree with those previously reported for others Crocidurinae: the interspecific comparison (ANCOVA) demonstrated an allometric relation between energy requirements and body mass. Energy expenditure also scaled with temperature. Island populations (Corsica and Porquerolles) of C. suaveolens differed in size from mainland (gigantism). A GLM showed a significant relationship between energy expenditure, temperature, body mass and locality. Mass specific RMR allometrically scales body mass, but total RMR does not significantly differ between mainland and island, although island shrews are giant. Our results are consistent with other studies: that demonstrated that the evolution of mammalian metabolism on islands is partially independent of body mass. In relation to the insular syndrome, we discuss how island selective forces (changes in resource availability, decrease in competition and predation pressures) can operate in size and physiological adjustments.     

Use of a novel Förster resonance energy transfer method to identify locations of site-bound metal ions in the U2-U6 snRNA complex

U2 and U6 snRNAs pair to form a phylogenetically conserved complex at the catalytic core of the spliceosome. Interactions with divalent metal ions, particularly Mg(II), at specific sites are essential for its folding and catalytic activity. We used a novel Förster resonance energy transfer (FRET) method between site-bound luminescent lanthanide ions and a covalently attached fluorescent dye, combined with supporting stoichiometric and mutational studies, to determine locations of site-bound Tb(III) within the human U2–U6 complex. At pH 7.2, we detected three metal-ion-binding sites in: (1) the consensus ACACAGA sequence, which forms the internal loop between helices I and III; (2) the four-way junction, which contains the conserved AGC triad; and (3) the internal loop of the U6 intra-molecular stem loop (ISL). Binding at each of these sites is supported by previous phosphorothioate substitution studies and, in the case of the ISL site, by NMR. Binding of Tb(III) at the four-way junction and the ISL sites was found to be pH-dependent, with no ion binding observed below pH 6 and 7, respectively. This pH dependence of metal ion binding suggests that the local environment may play a role in the binding of metal ions, which may impact on splicing activity.     

Broad geographic covariation between floral traits and the mating system in Camissoniopsis cheiranthifolia (Onagraceae): multiple stable mixed mating systems across the species' range?

Background and Aims

Plants vary widely in the extent to which seeds are produced via self-fertilization vs. outcrossing, and evolutionary change in the mating system is thought to be accompanied by genetic differentiation in a syndrome of floral traits. We quantified the pattern of variation and covariation in floral traits and the proportion of seeds outcrossed (t) to better understand the evolutionary processes involved in mating system differentiation among and within populations of the short-lived Pacific coastal dune endemic Camissoniopsis cheiranthifolia across its geographic range in western North America.

Methods

We quantified corolla width and herkogamy, two traits expected to influence the mating system, for 48 populations sampled in the field and for a sub-sample of 29 populations grown from seed in a glasshouse. We also measured several other floral traits for 9–19 populations, estimated t for 16 populations using seven allozyme polymorphisms, and measured the strength of self-incompatibility for nine populations.

Key Results

Floral morphology and self-incompatibility varied widely but non-randomly, such that populations could be assigned to three phenotypically and geographically divergent groups. Populations spanned the full range of outcrossing (t = 0·001–0·992), which covaried with corolla width, herkogamy and floral life span. Outcrossing also correlated with floral morphology within two populations that exhibited exceptional floral variation.

Conclusions

Populations of C. cheiranthifolia seem to have differentiated into three modal mating systems: (1) predominant outcrossing associated with self-incompatibility and large flowers; (2) moderate selfing associated with large but self-compatible flowers; and (3) higher but not complete selfing associated with small, autogamous, self-compatible flowers. The transition to complete selfing has not occurred even though the species appears to possess the required genetic capacity. We hypothesize that outcrossing populations in this species have evolved to different stable states of mixed mating.     

Rhinanthus plants found in calcareous fens on Gotland (Sweden): Are they related to Rhinanthus osiliensis from Saaremaa (Estonia)?

Rhinanthus osiliensis is a narrow endemic growing in calcareous spring fens on the island Saaremaa, Estonia. Plants very similar to R. osiliensis have been found in Sweden on the island of Gotland. Fifteen microsatellite markers were used to characterize genetic variation within R. osiliensis and the unknown taxon from Gotland. Results were compared with common Rhinanthus minor and Rhinanthus angstifolius. Our results showed that genetic diversity was remarkably lower in the Rhinanthus sp. from Gotland compared to Estonian endemic R. osiliensis. The microsatellite loci we used distinguished four studied species. In total, 46 species-specific alleles were found. The Neighbor-Joining dendrogram based on Nei's genetic distances, the Bayesian clustering analysis and the principle coordinate analysis showed that R. osiliensis and Rhinanthus sp. from Gotland are genetically differentiated and could not be the same species. In addition, endemic R. osiliensis was closer to widespread R. angustifolius. Further genetic comparison between unknown Rhinanthus sp. and local R. angustifolius and R. minor from Gotland is necessary for determine the species status and checking the possible alternative hypothesis about the origin of the unclear taxon.