Female fertility per flower and trade-offs between size and number in Claytonia virginica (Portulacaceae)

A consistent and paradoxical feature in flowering plants is the production of many more flowers than appear required for female fertility through fruit and seed production. Many mechanistic hypotheses for this observation share key assumptions about (1) limited resources available for reproduction and (b) greater female fertility benefits from larger flowering-time investment. Here I investigate these assumptions in two populations of Claytonia virginica. I also test predictions from theoretical analyses, comparing patterns of flowering allocation and fertility per flower in 18 populations of C. virginica. Results support the assumption that larger benefits accrue from greater flowering-time investment. The between-population pattern of flowering allocation and fertility per flower is also consistent with theoretical expectation, although not statistically significant. Not supported is the assumption that reproduction occurs under strong resource constraint. Possible reasons for this discrepancy are discussed.     

A new pollination system: brood-site pollination by flower bugs in Macaranga (Euphorbiaceae)

Background and Aims Macaranga: (Euphorbiaceae) is a large genus of dioecious trees with approx.260 species. To date, only one pollination study of the genushas reported brood-site pollination by thrips in M. hullettii.In this study, the pollination system of Macaranga tanariusis reported. Methods: The study was conducted on Okinawa and Amami Islands, Japan.Flower visitors on M. tanarius were collected and their pollenload and behaviour on the flowers examined, as well as inflorescencestructure and reward for the pollinators. Key Results: The most abundant flower visitors found on the male and femaleinflorescences were Orius atratus (Anthocoridae, Hemiptera),followed by Decomioides schneirlai (Miridae, Hemiptera). Pollenload on O. atratus from flowering pistillate inflorescenceswas detected as well as from staminate flowers. Orius atratusand D. schneirlai are likely to use the enclosed chambers formedby floral bracts as breeding sites before and during floweranthesis, and feed on nectar on the adaxial surface of flowerbracts. The extrafloral nectary has a ball-shaped structureand the contained nectar is not exposed; the hemipterans piercethe ball to suck out the nectar. Conclusions: The results indicate that the plant is pollinated by flowerbugs breeding on the inflorescences. This study may be the firstreport of pollination systems in which flower bugs are the mainpollinators. Similarity of pollination systems between M. hullettiiand M. tanarius indicates that the two brood-site pollinationsystems have the same origin. The pollinator species belongsto a predacious group, whose major prey includes thrips. Thepollination system might represent a unique example of evolutionfrom predatory flower visitors feeding on the pollinators (thrips)to the main pollinators.     

Dynamic optimization of fed-batch bioprocesses using flower pollination algorithm

Bioprocess and Biosystems Engineering - There exist several optimization strategies such as sequential quadratic programming (SQP), iterative dynamic programing (IDP), stochastic-based methods such...     

Genetics of flower size and nectar volume in Petunia pollination syndromes

The two related Petunia species, P. axillaris and P. integrifolia, are sympatric at various locations in South America but do not hybridise. Divergent pollinator preferences are believed to be in part responsible for their reproductive isolation. The volume of nectar produced and several components of flower morphology might contribute to pollinator-dependant reproductive isolation. In this study, we aimed to identify the genetic changes underlying the quantitative differences observed between these two Petunia species in flower size and nectar volume. We mapped quantitative trait loci (QTL) responsible for the different phenotypes of P. axillaris and P. integrifolia in an inter-specific backcross population. QTL of small to moderate effect control the differences in flower size and volume of nectar. In addition, we observed strong suppression of meiotic recombination in Petunia, even between closely related species, which precluded a fine resolution of QTL mapping. Thus, our data suggest that flower size and nectar volume are highly polygenic. They are likely to have evolved gradually through pollinator-mediated adaptation or reinforcement, and are not likely to have been primary factors in early steps of pollinator isolation of P. axillaris and P. integrifolia.     

Roles of pollination and short-chain saturated fatty acids in flower senescence

Pollination induced an immediate increase in ethylene production in Dianthus caryophyllus and Petunia hybrida. In Cymbidium, a lag of several hours was observed. In all three species, pollination induced premature flower senescence. Treatment of the stigmatic surface with aminoethoxyvinylglycine prior to pollination effectively blocked the increase in ethylene production and alleviated the detrimental effect of pollination on flower life.In all three tested species, octanoic and decanoic acids, when applied to the stigmatic surface, had no effect on ethylene production and flower life. In isolated Cymbidium lips placed with their cut base in solutions containing these fatty acids, no effects on red colouration, ethylene production, and ethylene forming enzyme activity were observed. In addition, ethylene sensitivity of isolated lips was not affected. The putative regulatory role of short-chain saturated fatty acids in (pollination-induced) flower senescence is discussed.     

Explosive flower opening in ornithophily: a study of pollination mechanisms in some Central African Loranthaceae

In many ornithophilous Loranthaceae pollination is accompanied by an explosive opening of the flowers, and diverse mechanisms have evolved in different genera to bring this about. These are described for the African genera Erianthemum, Englerina, Tapinanthus, Globimetula, Vanwykia and Plicosepalus. In many genera tensions within the stamens cause the tubular corolla to split along the petal junctions to form window-like fenestrae. The flowers are pollinated mainly by sunbirds which insert their beaks through the fenestrae in search of the abundant nectar. This action causes the tube to split and the stamens to coil inwards explosively. In Globimetula and many species of Tapinanthus pigment is secreted along the edges of the specialized petal segments of the head, the spathulae. Probing along these secretory junctions causes the spathulae to reflex; further probing splits the corolla tube, and allows the stamens to coil inwards explosively. In Globimetula reflexure of the petals exposes the central column of stamens, between which secondary fenestrae are developed. In Plicosepalus curvature of the corolla tube is connected with a more specialized fenestral structure; flower opening is not explosive, and the open flowers continue to be visited regularly by sunbirds. In Vanwykia an early stage in the development of explosive flower-opening is found.     

Phylogenetic evidence for a flower size and number trade-off

The size and number of flowers displayed together on an inflorescence (floral display) influences pollinator attraction and pollen transfer and receipt, and is integral to plant reproductive success and fitness. Life history theory predicts that the evolution of floral display is constrained by trade-offs between the size and number of flowers and inflorescences. Indeed, a trade-off between flower size and flower number is a key assumption of models of inflorescence architecture and the evolution of floral display. Surprisingly, however, empirical evidence for the trade-off is limited. In particular, there is a lack of phylogenetic evidence for a trade-off between flower size and number. Analyses of phylogenetic independent contrasts (PICs) of 251 angiosperm species spanning 63 families yielded a significant negative correlation between flower size and flower number. At smaller phylogenetic scales, analyses of individual genera did not always find evidence of a trade-off, a result consistent with previous studies that have examined the trade-off for a single species or genus. Ours is the first study to support an angiosperm-wide trade-off between flower size and number and supports the theory that life history constraints have influenced the evolution of floral display.     

Size constraints and flower abundance determine the number of interactions in a plant–flower visitor web

The number of interactions with flower visitor species differs considerably among insect pollinated plants. Knowing the causes for this variation is central to the conservation of single species as well as whole plant–flower visitor communities. Species specific constraints on flower visitor numbers are seldom investigated at the community level. In this study we tested whether flower size parameters set constraints on the morphology of the potential nectar feeding visitors and thus determine the number of visitor species. We studied three possible constraints: the depth and width of tubular structures hiding the nectar (nectar holder depth and width) and the size of flower parts that visitors can land on (size of the alighting place). In addition we assess the role of flower abundance on this relationship. We hypothesized that the stronger size constraints and the smaller flower abundance, the smaller the number of visitor species will be. Our study of a Mediterranean plant–flower visitor community revealed that nectar holder depth, nectar holder width and number of flowers explained 71% of the variation in the number of visitor species. The size of the alighting place did not restrict the body length of the visitors and was not related to visitor species number. In a second step of the analyses we calculated for each plant species the potential number of visitors by determining for each insect species of the local visitor pool whether it passed the morphological limits set by the plant. These potential numbers were highly correlated with the observed numbers (r²=0.5, p<0.001). For each plant species we tested whether the observed visitors were a random selection out of these potential visitors by comparing the mean of the observed and expected proboscis length distributions. For most plant species the observed mean was not significantly different from the random means. Our findings shed light on the way plant–flower visitor networks are structured. Knowing the constraints on interaction patterns will be an important prerequisite to formulate realistic null models and understand patterns of resource partitioning as well as coevolutionary processes.     

Ancestral reconstruction of flower morphology and pollination systems in Schizanthus (Solanaceae)

Concerted changes in flower morphology and pollinators provide strong evidence on adaptive evolution. Schizanthus (Solanaceae) has zygomorphic flowers and consists of 12 species of annual or biennial herbs that are distributed mainly in Chile and characterized by bee-, hummingbird-, and moth-pollination syndromes. To infer whether flowers diversified in relation to pollinator shifts, we traced the evolutionary trajectory of flower traits and visitors onto a phylogeny based on sequence data from ITS, waxy, and trnF/ndhJ DNA. Maximum-likelihood ancestral reconstruction of floral traits suggests that ancestral Schizanthus had a bee-pollination syndrome. The hummingbird syndrome evolved in S. grahamii, a high elevation species in the Andes. The moth syndrome evolved in the ancestor of three species that inhabit the Atacama Desert. Results of mapping flower visitors onto the phylogeny show that the shift from bee to hummingbird pollination concurred with a shift in pollinators as predicted by the syndromes. However, the same pattern was not found for the moth syndrome. Visits by moths were observed only in one of the three moth-syndrome species, and at a very low rate. This mismatch suggests either anachronic floral characters or maintenance of rare, imperceptible moth pollination backed up by capacity for autonomous selfing. Overall, results suggest that diversification of flower traits in Schizanthus has occurred in relation to pollinator shifts.     

Pollen movement in a natural population of Arisaema serratum (Araceae), a plant with a pitfall-trap flower pollination system

Arisaema serratum possesses a pitfall-trap flower pollination system. However, little is known about the efficiency and pattern of pollen movement in A. serratum. Thus, the aims of this study are to (1) determine the paternal parents of the seeds and (2) elucidate pollen movement in a natural population. Paternity analysis using microsatellite markers was performed. Seeds were collected from a natural population of A. serratum in 2001 at Horigane, Japan. Small midges became trapped in female spathe tubes during the flowering period. We found that (1) seeds in a fruit were fertilized by multiple sires; (2) seeds sired by a paternal parent were either clumped, exclusively, or randomly distributed on the spadix, depending on the parent; (3) to a great extent, a few males contributed as sires; (4) distance from a female was not a factor in the inequality of reproductive success among males; (5) male reproductive success was not correlated with its size. We conclude that pollen carryover and the trap-flower pollination system are likely to result in multiple paternity and inequality in male success.     

Ecological implications of a flower size/number trade-off in tropical forest trees

BACKGROUND: In angiosperms, flower size commonly scales negatively with number. The ecological consequences of this trade-off for tropical trees remain poorly resolved, despite their potential importance for tropical forest conservation. We investigated the flower size number trade-off and its implications for fecundity in a sample of tree species from the Dipterocarpaceae on Borneo. METHODOLOGY/PRINCIPAL FINDINGS: We combined experimental exclusion of pollinators in 11 species, with direct and indirect estimates of contemporary pollen dispersal in two study species and published estimates of pollen dispersal in a further three species to explore the relationship between flower size, pollinator size and mean pollen dispersal distance. Maximum flower production was two orders of magnitude greater in small-flowered than large-flowered species of Dipterocarpaceae. In contrast, fruit production was unrelated to flower size and did not differ significantly among species. Small-flowered species had both smaller-sized pollinators and lower mean pollination success than large-flowered species. Average pollen dispersal distances were lower and frequency of mating between related individuals was higher in a smaller-flowered species than a larger-flowered confamilial. Our synthesis of pollen dispersal estimates across five species of dipterocarp suggests that pollen dispersal scales positively with flower size. CONCLUSIONS AND THEIR SIGNIFICANCE: Trade-offs embedded in the relationship between flower size and pollination success contribute to a reduction in the variance of fecundity among species. It is therefore plausible that these processes could delay competitive exclusion and contribute to maintenance of species coexistence in this ecologically and economically important family of tropical trees. These results have practical implications for tree species conservation and restoration. Seed collection from small-flowered species may be especially vulnerable to cryptic genetic erosion. Our findings also highlight the potential for differential vulnerability of tropical tree species to the deleterious consequences of forest fragmentation.     

The number of sex-factors per chromosome in Escherichia coli

A substituted sex-factor of Escherichia coli, F'8 gal, was transferred to Proteus mirabilis by conjugation. The DNA of the episome was partially purified from Proteus DNA by preparative equilibrium centrifugation in caesium chloride, and by a bulk method using hydroxyapatite. The buoyant density of the episomal DNA is 1.707, corresponding to a (G+C) content of 47%. By optical renaturation the genetic complexity of the episomal DNA was found to be 76x10(6) daltons. RNA was synthesized in vitro by using the episomal DNA as template. By hybridizing this RNA with DNA extracted from E. coli carrying F'8 gal, it is shown that the number of copies of the episome per replicating chromosome is close to two during exponential growth. The episome makes up about 4.4% of the total DNA of the growing cells. The activities of galactokinase and galactose 1-phosphate uridylyl-transferase in cells with and without episomal and chromosomal gal genes were found to be proportional to the number of gal genes present, when the cells were induced with d-fucose, but not when they were induced with d-galactose.     

Trade-offs between flower number and investment to a flower in selfing and outcrossing varieties of Impatiens hypophylla (Balsaminaceae)

Floral resource allocation was compared on a whole-plant basis between two varieties of Impatiens hypophylla that differ in flower size. There were significant negative correlations between flower number and investments to a flower at both the within-population and between-variety levels. In individual flowers, var. hypophylla with larger flowers invested significantly more resources to male and pollinator-attractive functions, whereas investments to female function did not differ between the varieties. In experimental populations placed in the field, pollinators preferred the larger flowers of var. hypophylla even within the same habitat of var. microhypophylla, which has smaller flowers. There was a significant lack of observed heterozygosity only in var. microhypophylla. Thus, the outcrossing variety had more attractive but fewer flowers, while the selfing variety had less attractive but more abundant flowers.     

The number of generations in freshwater monogeneans per year

It seems that the monogenean Ancyrocephalus paradoxus inhabiting the zander Stizostedion lucioperca (L.) has three generations per year in the Kurish Bay of the Baltic Sea. Diplozoids of cyprinids in the same bay have only two generations. There is evidence that temperature is the chief factor determining the number of generations in Monogenea; but there are also data showing that an increase in abundance of some Monogenea takes place at low temperatures and during congregation of fishes. There is another opinion that sex hormones offish influence the development of Monogenea. However, the appearance of the third generation during winter and early spring is not consistent with this.     

Ovule Fertilization and Seed Number per Pod Determination in Oil Seed Rape (Brassica napus)

Ovule fertilization in relation to seed number determinationin spring rapeseed, Brassica napus var. Maris Haplana, was investigated.Failure of fertilization was found to be the major factor limitingthe number of seeds per pod in both the basally and apicallylocated pods on the terminal inflorescence. Sufficient pollengermination on a stigma did not guarantee full seed set andeven when pollen tubes were present at the micropyle regions,ovules were not penetrated. Some seeds aborted: in some, triplefusion did not take place while other seeds were small and deemedat disadvantage to compete for available nutrients. Brassica napus, fertilization, ovules, seeds     

Influence of flower functionality and pollination system on the pollen size-pistil length relationship

Twenty-five biotically pollinated plants of the Chaco Serrano Forest (Córdoba, Argentina) were studied in order to analyze whether ‘flower functionality’ is related to the relationship between pollen size and pistil length. Because flower functionality may act on the respective mean values of pollen size and pistil length rather than on intraspecific variation in these traits, we expected (1) a high positive correlation between pollen size and pistil length in a set of sympatric species, independent of their degree of pollination specialization or generalization; and (2) no interspecific correlation between the coefficients of variation (CVs) of those traits. On the other hand, on the assumption that pollinators are influencing the variation in floral traits (e.g. in pistil length) we expected lower mean phenotypic variation of pollen size and pistil length in pollination-specialist plants than in pollination-generalist ones. A positive correlation between pollen size and pistil length was found for the set of species, but not between the CVs of these traits. This trend was maintained when pollination-specialist plants were analyzed separately, but no statistical significance was obtained for the correlation in pollination-generalist plants. Contrary to our expectations, pollination-specialist plants did not show less mean intraspecific variation in floral traits than pollination-generalist plants. Therefore, the relationship between pollen size and pistil length among species suggests that the pollination system may be of less importance as a selective force than flower functionality.     

Fitting of a binomial distribution to the number of chiasmata per bivalent

Statistical tests on the distribution of the number of chiasmata per chromosome, collected from literatures, showed that they can be approximated by binomial distributions with one obligatory chiasma, i.e., B(N-1, p). N is proportional to the average number of chiasmata, while p is nearly constant for the species tested.     

Comments on flower evolution and beetle pollination in the generaAnnona andRollinia (Annonaceae)

Dynastid scarab beetle pollination appears basic within the genusAnnona. Those species ofAnnona which are more morphologically derived, as well as allRollinia spp. possess reduced floral chambers and attract small beetles likeNitidulidae orStaphylinidae. Pollination of the primitive species ofAnnona byDynastinae would imply that the genus had not evolved before the Tertiary. The fossil record is in congruence with this hypothesis. Once again it is stressed that the cantharophilous syndrome, as it is found in theAnnonaceae, Magnoliaceae, Eupomatiaceae andCalycanthaceae, with beetles being exclusive pollinators, is a secondary and derived condition and obviously different from the expected basic entomophily of the original angiosperms.