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PubMed1995

Identification of genes required for pollen-stigma recognition in Arabidopsis thaliana.

In higher plants, cell-cell recognition reactions taking place following pollination allow the selective restriction of self-pollination and/or interspecific pollination. Many of these systems function by regulating the process of water transfer from the cells found at the stigmatic surface to the individual pollen grain. Interspecific pollination studies on the cruciferous weed Arabidopsis thaliana revealed only a broad specificity of pollen recognition such that pollen from all tested members of the crucifer family were recognized, whereas pollen from almost all other species failed to hydrate. Genetic analysis of A. thaliana has identified three genes that are essential for this recognition process. Recessive mutations in any of these genes result in male sterility due to the production of pollen grains that fail to hydrate when placed on the stigma, but that are capable of hydrating and growing a pollen tube in vitro. Results from mixed pollination experiments suggest that the mutant pollen grains specifically lack a functional pollen-stigma recognition system. All three mutations described also result in a defect in the wax layer normally found on stems and leaves, similar to previously described eceriferum (cer) mutations. Genetic complementation and mapping experiments demonstrated that the newly identified mutants are allelic to the previously identified genes cer1, cer3 and cer6. TEM analysis of the ultrastructure of the pollen coating revealed that all of the mutant pollen grains bear coatings of normal thickness and that tryphine lipid droplets are missing in cer1-147, are reduced in size in cer6-2654 and appear normal in cer3-2186.(ABSTRACT TRUNCATED AT 250 WORDS)

Arabidopsis↗
PubMed2005

Broom and honeybees in Australia: an alien liaison.

Facilitative interactions between non-indigenous species are gaining recognition as a major driver of invasion success. Cytisus scoparius (L.) Link (Fabaceae), or Scotch broom, is a cosmopolitan invasive shrub that lacks the capacity for vegetative reproduction and is a good model to study facilitative interactions. Its success in pioneer environments is determined by constraints on its reproduction. We determined whether pollinators were required for seed set in C. scoparius at Barrington Tops, NSW, Australia, where the species has infested ca. 14,000 ha across the plateau. Field and laboratory experiments showed that C. scoparius is an obligate outcrossing species at Barrington Tops. Monitoring of plants (10.7 h) showed that the flowers of C. scoparius have to be tripped to effect seed set and the only pollinator to do this was the introduced honeybee, Apis mellifera L. Most floral visits by honeybees result in fruit set (84 %) and because fruits have many ovules (10 - 18 per ovary) a single bee on an average foraging day can effect the production of over 6000 seeds. A review of C. scoparius pollination across four continents revealed major differences in pollen quantity, which may explain differences in the efficiencies of honeybees as pollinators of C. scoparius. The incorporation of pollinator management in an integrated approach for the control of C. scoparius is discussed.

Animals↗
PubMed2004

Economic value of tropical forest to coffee production.

Can economic forces be harnessed for biodiversity conservation? The answer hinges on characterizing the value of nature, a tricky business from biophysical, socioeconomic, and ethical perspectives. Although the societal benefits of native ecosystems are clearly immense, they remain largely unquantified for all but a few services. Here, we estimate the value of tropical forest in supplying pollination services to agriculture. We focus on coffee because it is one of the world's most valuable export commodities and is grown in many of the world's most biodiverse regions. Using pollination experiments along replicated distance gradients, we found that forest-based pollinators increased coffee yields by 20% within approximately 1 km of forest. Pollination also improved coffee quality near forest by reducing the frequency of "peaberries" (i.e., small misshapen seeds) by 27%. During 2000-2003, pollination services from two forest fragments (46 and 111 hectares) translated into approximately 60,000 USD per year for one Costa Rican farm. This value is commensurate with expected revenues from competing land uses and far exceeds current conservation incentive payments. Conservation investments in human-dominated landscapes can therefore yield double benefits: for biodiversity and agriculture.

Animals↗
PubMed1993

Origin of floral isolation between ornithophilous and sphingophilous plant species.

Three plant groups in temperate western North America contain closely related ornithophilous and sphingophilous species: the Aquilegia formosa/Aquilegia caerulea group (Ranunculaceae), the Ipomopsis aggregata group (Polemoniaceae), and the Diplacus longiflorus group (Scrophulariaceae). The ornithophilous and sphingophilous species are products of allopatric speciation on the diploid level. Geographical races which are adapted to one class of pollinators in one area where these pollinators are abundant and effective and to another class of pollinators in another geographically isolated area (pollination races) represent a probable intermediate stage in the process of allopatric speciation. Mechanical and ethological isolation (collectively, floral isolation) is a byproduct of the divergence in pollination systems. Selection for reproductive isolation per se has not played any detectable role in the origin of the floral isolation in the three plant groups.

Journal Article↗
PubMed1994

Modes and origins of mechanical and ethological isolation in angiosperms.

Mechanical and ethological isolation between species is widespread in angiosperms with specialized animal-pollinated flowers, being recorded in 29 species groups belonging to 27 genera and 16 families. Mechanical isolation occurs in two forms. (i) The common type, designated the Salvia type, operates when two or more species of flowers are adapted for different groups of pollinators with different body sizes and shapes. (ii) In the Pedicularis type two flower species have the same species of pollinator but pick up pollen from different parts of the pollinator's body. Four forms of ethological isolation are recognized. (i) In the Aquilegia type, which is widespread, ethological isolation is a side effect of mechanical isolation. (ii) The flower-constancy type, as the name suggests, is based on flower-constant foraging behavior. (iii) In the Ophrys type, floral scents attract male bees or wasps and play a role in their mating behavior; different species of flowers, often orchids, have different scents and attract different sets of hymenopteran species. (iv) The monotropy type occurs in plants pollinated by hymenopterans with species-specific or group-specific flower preferences for nutritive purposes (monotropic and oligotropic bees and fig wasps). Three modes of origin of floral isolation are confirmed by evidence: (i) mechanical and ethological isolation arising as a by-product of allopatric speciation, (ii) ethological isolation developing by selection for reproductive isolation per se, and (iii) mechanical isolation arising as a by-product of character displacement. Mode of origin i accounts for the Salvia and Aquilegia types of isolation in nine known species groups and for the Ophrys type in one group. Mode of origin ii accounts for the flower-constancy type of ethological isolation in two species groups. Mode of origin iii explains mechanical isolation in two groups. Sympatric origin of floral isolation by hybrid speciation and by flower constancy has been proposed, but these modes are undocumented and improbable.

Journal Article↗
PubMed2006

Patterns of speciation in the Yucca moths: parallel species radiations within the Tegeticula yuccasella species complex.

The interaction between yuccas and yucca moths has been central to understanding the origin and loss of obligate mutualism and mutualism reversal. Previous systematic research using mtDNA sequence data and characters associated with genitalic morphology revealed that a widespread pollinator species in the genus Tegeticula was in fact a complex of pollinator species that differed in host use and the placement of eggs into yucca flowers. Within this mutualistic clade two nonpollinating "cheater" species evolved. Cheaters feed on yucca seeds but lack the tentacular mouthparts necessary for yucca pollination. Previous work suggested that the species complex formed via a rapid radiation within the last several million years. In this study, we use an expanded mtDNA sequence data set and AFLP markers to examine the phylogenetic relationships among this rapidly diverging clade of moths and compare these relationships to patterns in genitalic morphology. Topologies obtained from analyses of the mtDNA and AFLP data differed significantly. Both data sets, however, corroborated the hypothesis of a rapid species radiation and suggested that there were likely two independent species radiations. Morphological analyses based on oviposition habit produced species groupings more similar to the AFLP topology than the mtDNA topology and suggested the two radiations coincided with differences in oviposition habit. The evolution of cheating was reaffirmed to have evolved twice and the closest pollinating relative for one cheater species was identified by both mtDNA and AFLP markers. For the other cheater species, however, the closest pollinating relative remains ambiguous, and mtDNA, AFLP, and morphological data suggest this cheater species may be diverged based on host use. Much of the divergence in the species complex can be explained by geographic isolation associated with the evolution of two oviposition habits.

Animals↗
PubMed2003

Three-way coexistence in obligate mutualist-exploiter interactions: the potential role of competition.

Many mutualisms host "exploiter" species that consume the benefits provided by one or both mutualists without reciprocating. Exploiters have been widely assumed to destabilize mutualisms, yet they are common. We develop models to explore conditions for local coexistence of obligate plant/pollinating seed parasite mutualisms and nonpollinating exploiters. As the larvae of both pollinators and (at a later time) exploiters consume seeds, we examine the importance of intraspecific and (asymmetric) interspecific competition among and between pollinators and exploiters for achieving three-way coexistence. With weak intra- and interspecific competition, exploiters can invade the stable mutualism and coexist with the mutualists (either stably or with oscillations), provided the exploiters' intrinsic birthrate (b(E)) slightly exceeds that of the pollinators. At higher b(E), all three species go locally extinct. When facing strong interspecific competition, exploiters cannot invade and coexist with the mutualists if intraspecific competition in pollinators and exploiters is weak. However, strong intraspecific competition in pollinators and exploiters facilitates exploiter invasion and coexistence and greatly expands the range of b(E) over which stable coexistence occurs. Our results suggest that mutualist/exploiter coexistence may be more easily achieved than previously thought, thus highlighting the need for a better understanding of competition among and between mutualists and exploiters.

Animals↗
PubMed2005

Divergent mating systems and parental conflict as a barrier to hybridization in flowering plants.

Parental conflicts can lead to antagonistic coevolution of the sexes and of parental genomes. Within a population, the resulting antagonistic effects should balance, but crosses between populations can reveal conflict. Parental conflict is less intense in self-pollinating plants than in outcrossers because outcrossing plants are pollinated by multiple pollen donors unrelated to the seed parent, while a self-pollinating plant is primarily pollinated by one individual (itself). Therefore, in crosses between plants with differing mating systems, outcrossing parents are expected to "overpower" selfing parents. We call this the weak inbreeder/strong outbreeder (WISO) hypothesis. Prezygotically, such overpowering can alter pollination success, and we argue that our hypothesis explains a common pattern of unilateral incompatibility, in which pollen from self-incompatible populations fertilizes ovules of self-compatible individuals but the reciprocal cross fails. A postzygotic manifestation of overpowering is aberrant seed development due to parent-of-origin effects such as genomic imprinting. We evaluate evidence for the WISO hypothesis by reviewing published accounts of crosses between plants of different mating systems. Many, but not all, of such reports support our hypothesis. Since parental conflicts can perturb fertilization and development, such conflicts may strengthen reproductive barriers between populations, contributing to speciation.

Flowers↗
PubMed2003

Aechmea pectinata: a hummingbird-dependent bromeliad with inconspicuous flowers from the rainforest in south-eastern Brazil.

The pollination biology of Aechmea pectinata (Bromeliaceae) was studied in a submontane rainforest in south-eastern Brazil. This species has a mainly clumped distribution and its aggregated individuals are likely to be clones. From October to January, during the flowering period, the distal third of its leaves becomes red. The inflorescence produces 1-15 flowers per day over a period of 20-25 d. The flowers are inconspicuous, greenish-white coloured, tubular shaped with a narrow opening, and the stigma is situated just above the anthers. Anthesis begins at 0400 h and flowers last for about 13 h. The highest nectar volume and sugar concentration occur between 0600 and 1000 h, and decrease throughout the day. Aechmea pectinata is self-incompatible and therefore pollinator-dependent. Hummingbirds are its main pollinators (about 90 % of the visits), visiting flowers mainly in the morning. There is a positive correlation between the number of hummingbird visits per inflorescence and the production of nectar, suggesting that the availability of this resource is important in attracting and maintaining visitors. The arrangement of the floral structures favours pollen deposition on the bill of the hummingbirds. Flowers in clumps promote hummingbird territoriality, and a consequence is self-pollination in a broader sense (geitonogamy) as individuals in assemblages are genetically close. However, trap-lining and intruding hummingbirds promote cross-pollination. These observations suggest that successful fruit set of A. pectinata depends on both the spatial distribution of its individuals and the interactions among hummingbirds.

Animal Feed↗
PubMed2003

Floral structure, breeding system and fruit-set in the threatened sub-shrub Tetratheca juncea Smith (Tremandraceae).

Tetratheca juncea Smith (Tremandraceae) has undergone a range contraction of approx. 50 km in the last 100 years and is now listed as a vulnerable sub-shrub restricted to the central and north coast regions of New South Wales, Australia. There are approx. 250 populations in a 110 km north-south distribution and populations are usually small with fewer than 50 plants/clumps. The reproductive ecology of the species was studied to determine why seed-set is reportedly rare. Flowers are bisexual, odourless and nectarless. Flowers are presented dependentally and there are eight stamens recurved around the pistil. Anthers are poricidal, contain viable pollen and basally contain a deep-red tapetal fluid that is slightly oily. Thus flowers are presented for buzz pollinators, although none were observed at flowers during our study. The species was found to be facultatively xenogamous with only one in 50 glasshouse flowers setting seed autogamously, i.e. without pollinator assistance. Field studies revealed fertile fruit in 24 populations but production varied significantly across sites from exceedingly low (0.6 fruits per plant clump) to low (17 fruits per plant clump). Fruit-set ranged from 0 to 65 %, suggesting that pollen vectors exist or that autogamy levels in the field are variable and higher than glasshouse results. Fruit production did not vary with population size, although in three of the five populations in the south-west region more than twice as much fruit was produced as in populations elsewhere. A moderately strong relationship between foliage volume and fruit : flower ratios suggests that bigger plants may be more attractive than smaller plants to pollinators. A review of Tetratheca pollination ecology revealed that several species are poorly fecund and pollinators are rare. The habitat requirements for Tetratheca, a genus of many rare and threatened species, is discussed.

Australia↗
PubMed2005

Stigma development and receptivity in almond (Prunus dulcis).

BACKGROUND AND AIMS: Fertilization is essential in almond production, and pollination can be limiting in production areas. This study investigated stigma receptivity under defined developmental stages to clarify the relationship between stigma morphology, pollen germination, tube growth and fruit set. METHODS: Light and scanning electron microscopy were employed to examine stigma development at seven stages of flower development ranging from buds that were swollen to flowers in which petals were abscising. Flowers at different stages were hand pollinated and pollen germination and tube growth assessed. Artificial pollinations in the field were conducted to determine the effect of flower age on fruit set. KEY RESULTS: Later stages of flower development exhibited greater stigma receptivity, i.e. higher percentages of pollen germination and more extensive tube growth occurred in older (those opened to the flat petal stage or exhibiting petal fall) than younger flowers. Enhanced stigma receptivity was associated with elongation of stigmatic papillae and increased amounts of stigmatic exudate that inundated papillae at later developmental stages. Field pollinations indicated that the stigma was still receptive and nut set was maintained in older flowers. CONCLUSIONS: Stigma receptivity in almond does not become optimal until flowers are past the fully open stage. The stigma is still receptive and fruit set is maintained in flowers even at the stage when petals are abscising. Strategies to enhance pollination and crop yield, including the timing and placement of honey bees, should consider the effectiveness of developmentally advanced flowers.

Flowers↗
PubMed2006

Nectar sugar composition and volumes of 47 species of Gentianales from a southern Ecuadorian montane forest.

BACKGROUND AND AIMS: This study investigates 47 taxonomically related species (Gentianales), all native to a tropical montane forest in southern Ecuador, in terms of nectar chemistry and nectar volumes in relation to pollination biology. METHODS: Nectar volumes of covered (24-h production) and uncovered (standing crop) flowers were measured in the natural habitat. Sucrose, fructose and glucose were quantified in the nectar using high performance liquid chromatography. Flower visitors were observed. KEY RESULTS: Nectar sugar concentration did not differ significantly among the pollination syndromes. Regarding sugar composition, the only significant differences were found in chiropterophilous and myiophilous flowers, which had a significantly lower sugar ratio than sphingophilous flowers. A separation of chiropterophilous and myiophilous flowers from the other pollination syndromes is further substantiated by non-linear multidimensional scaling using the chord-normalized expected species shared index of dissimilarity based on nectar sugar compositions. The matrix test revealed no correlation of observed floral visitors to nectar concentrations; however, a weak significant correlation was found between floral visitors and nectar sugar compositions. The nectar volumes of covered and uncovered flowers are related to, and differ significantly among, pollination syndromes. Matrix tests revealed correlation between floral visitors and nectar volume of covered flowers and, to a lesser extent, of uncovered flowers. CONCLUSIONS: Sucrose is the predominant floral nectar sugar in the order Gentianales, suggesting that nectar sugar composition is a conservative characteristic. However, some degree of an adaptive convergence of floral nectar compositions to principal pollinator type within the constraints set by phylogenetic history is likely. The driving force to visitation appears to be the volume of nectar the visitor can expect to consume.

Animals↗
PubMed2002

When looks can kill: the evolution of sexually dimorphic floral display and the extinction of dioecious plants.

Dioecious plants (with separate male and female individuals) more often have drab, inconspicuous flowers than related bisexual plants. Models indicate, however, that similar conditions favour the evolution of showy floral displays in dioecious and bisexual plants. One difference, however, is that dioecious plants may evolve floral displays that are sexually dimorphic. We show that males are more likely to evolve showy flowers than females in animal-pollinated plants, especially when pollinators are abundant. We demonstrate that this dimorphism places showy dioecious plants at a much higher risk of extinction during years of low pollinator abundance because pollinators may fail to visit female flowers. The higher extinction risk of showy dioecious plants provides an explanation for the fact that dioecious plants that do persist tend to have inconspicuous flowers and are more often wind pollinated. It may also help explain why dioecious plants are less species-rich than related bisexual plants.

Biological Evolution↗
PubMed2002

Dynamic 1-aminocyclopropane-1-carboxylate-synthase and -oxidase transcript accumulation patterns during pollen tube growth in tobacco styles.

In flowering plants, pollination of the stigma sets off a cascade of responses in the distal flower organs. Ethylene and its biosynthetic precursor 1-aminocyclopropane-1-carboxylate (ACC) play an important role in regulating these responses. Because exogenous application of ethylene or ACC does not invoke the full postpollination syndrome, the pollination signal probably consists of a more complex set of stimuli. We set out to study how and when the pollination signal moves through the style of tobacco (Nicotiana tabacum) by analyzing the expression patterns of pistil-expressed ACC-synthase and -oxidase genes. Results from this analysis showed that pollination induces high ACC-oxidase transcript levels in all cells of the transmitting tissue. ACC-synthase mRNA accumulated only in a subset of transmitting tract cells and to lower levels as compared with ACC-oxidase. More significantly, we found that although ACC-oxidase transcripts accumulate to uniform high levels, the ACC-synthase transcripts accumulate in a wave-like pattern in which the peak coincides with the front of the ingrowing pollen tube tips. This wave of ACC-synthase expression can also be induced by incongruous pollination and (partially) by wounding. This indicates that wounding-like features of pollen tube invasion might be part of the stimuli evoking the postpollination response and that these stimuli are interpreted differently by the regulatory mechanisms of the ACC-synthase and -oxidase genes.

Amino Acid Oxidoreductases↗
PubMed1995

Low Water Potential Disrupts Carbohydrate Metabolism in Maize (Zea mays L.) Ovaries.

Water deficit during pollination increases the frequency of kernel abortion in maize (Zea mays L.). Much of the kernel loss is attributable to lack of current photosynthate, but a large number of kernels fail to develop on water-deficient plants even when assimilate supply is increased. We examined the possibility that assimilate utilization by developing ovaries might be impaired at low water potential ([Psi]w). Plants were grown in the greenhouse in 20-L pots containing 22 kg of amended soil. Water was withheld on the first day silks emerged, and plants were hand-pollinated 4 d later when leaf [Psi]w decreased to approximately - 1.8 MPa and silk [Psi]w was approximately -1.0 MPa. Plants were rehydrated 2 d after pollination. The brief water deficit inhibited ovary growth (dry matter accumulation) and decreased kernel number per ear by 60%, compared to controls. Inhibition of ovary growth was associated with a decrease in the level of reducing sugars, depletion of starch, a 75-fold increase in sucrose concentration (dry weight basis), and inhibition of acid invertase (EC 3.2.1.26) activity. These results indicate that water deficits during pollination disrupt carbohydrate metabolism in maize ovaries. They suggest that acid invertase activity is important for establishing and maintaining reproductive sink strength during pollination and early kernel development.

Journal Article↗
PubMed1989

Abscisic Acid Levels during Early Seed Development in Sechium edule Sw.

The time-course growth of single tissues in pollinated and unpollinated ovules of Sechium edule Sw. is described in relation to the endogenous levels of abscisic acid. Quantitation of abscisic acid (ABA) in the minute amounts of material obtained after ovule dissection has been performed by using a highly specific and sensitive solid-phase radioimmunoassay based on a monoclonal antibody raised against free (S)-ABA. While the absolute amount of ABA rises in both types of ovules, only in unpollinated ones does this leads to an increase in the hormone concentration. Infact in pollinated ovules the rapid growth following pollination prevents, through a dilution effect, the increase in ABA concentration. Growth patterns and endogenous ABA levels are similar for integuments and nucellus tissues either in pollinated or unpollinated ovules. It is suggested that the growth inhibition induced by the increase in ABA concentration after anthesis could be counteracted by the pollination triggered fast ovule growth.

Journal Article↗
PubMed1992

Endosperm cell division in maize kernels cultured at three levels of water potential.

The influence of osmoticum treatments on early kernel development of maize (Zea mays L.) was studied using an in vitro culture method. Kernels with subtending cob sections were placed in culture at 5 days after pollination. Sucrose (0.29, 0.44, or 0.58 molar) and sorbitol (0, 0.15, or 0.29 molar) were used to obtain six media with water potentials of -1.1, -1.6, or -2.0 megapascals. Kernel water potential declined in correspondence with the water potential of the medium; however, fresh weight growth was not significantly inhibited from 5 to 12 days after pollination. In stress treatments with media water potentials of -1.6 or -2.0 megapascals, endosperm tissue accumulated water and solutes from 10 and 12 days after pollination at a rate similar to or greater than that of the control (-1.1 megapascals). In contrast, endosperm cell division was inhibited in all treatments relative to control. At 10 days after pollination, endosperm sucrose concentration was greater in two of the -2.0 megapascal treatments with 0.44 or 0.58 molar media sucrose compared to control kernels cultured in 0.29 molar sucrose at -1.1 megapascals. Significant increases in abscisic acid content per gram of fresh weight were detected in two -2.0 megapascal treatments (0.29 molar sucrose plus 0.29 molar sorbitol and 0.58 molar sucrose) at 10 days after pollination. We conclude that in cultured maize kernels, endosperm cell division was more responsive than fresh weight accumulation to low water potential treatments. Data were consistent with mechanisms involving abscisic acid or lowered tissue water potential, or an interaction of the two factors.

Journal Article↗
PubMed2003

Convergence and coevolution in a mutualism: evidence from a molecular phylogeny of Ficus.

The interaction between Ficus (Moraceae) and their pollinating wasps (Chalcidoidea: Agaonidae; more than 700 species-specific couples) is one of the most specialized mutualisms found in nature. Both partners of this interaction show extensive variation in their respective biology. Here we investigate Ficus life-history trait evolution and fig/fig wasp coadaptation in the context of a well-resolved molecular phylogeny. Mapping out variations in Ficus life-history traits on an independently derived phylogeny constructed from ribosomal DNA sequences (external and internal transcribed spacer) reveals several parallel transitions in Ficus growth habit and breeding system. Convergent trait evolution might explain the discrepancies between morphological analyses and our molecular reconstruction of the genus. Morphological characters probably correlate with growth habit and breeding system and could therefore be subject to convergent evolution. Furthermore, we reconstruct the evolution of Ficus inflorescence characters that are considered adaptations to the pollinators. Our phylogeny reveals convergences in ostiole shape, stigma morphology, and stamen:ovule ratio. Statistical tests taking into account the phylogenetic relationship of the species show that transitions in ostiole shape are correlated with variation in wasp pollinator head shape, and evolutionary changes in stigma morphology and stamen:ovule ratio correlate with changes in the pollination behavior of the associated wasp. These correlations provide evidence for reciprocal adaptations of morphological characters between these mutualistic partners that have interacted over a long evolutionary time. In light of previous ecological studies on mutualism, we discuss the adaptive significance of these correlations and what they can tell us about the coevolutionary process occurring between figs and their pollinators.

Adaptation, Biological↗