Effects of patch size and density on flower visitation and seed set of wild plants: a pan-European approach

1. Habitat fragmentation can affect pollinator and plant population structure in terms of species composition, abundance, area covered and density of flowering plants. This, in turn, may affect pollinator visitation frequency, pollen deposition, seed set and plant fitness. 2. A reduction in the quantity of flower visits can be coupled with a reduction in the quality of pollination service and hence the plants’ overall reproductive success and long-term survival. Understanding the relationship between plant population size and⁄ or isolation and pollination limitation is of fundamental importance for plant conservation. 3. Weexamined flower visitation and seed set of 10 different plant species fromfive European countries to investigate the general effects of plant populations size and density, both within (patch level) and between populations (population level), on seed set and pollination limitation. 4. Wefound evidence that the effects of area and density of flowering plant assemblages were generally more pronounced at the patch level than at the population level. We also found that patch and population level together influenced flower visitation and seed set, and the latter increased with increasing patch area and density, but this effect was only apparent in small populations. 5. Synthesis. By using an extensive pan-European data set on flower visitation and seed set we have identified a general pattern in the interplay between the attractiveness of flowering plant patches for pollinators and density dependence of flower visitation, and also a strong plant species-specific response to habitat fragmentation effects. This can guide efforts to conserve plant–pollinator interactions, ecosystem functioning and plant fitness in fragmented habitats.

Glucosinolate-accumulating S-cells in Arabidopsis leaves and flower stalks undergo programmed cell death at early stages of differentiation

Plant secondary metabolites glucosinolates (GSL) have important functions in plant resistance to herbivores and pathogens. We identified all major GSL that are accumulated in S-cells in Arabidopsis by MALDI-TOF MS, and estimated by LC-MS that the total GSL concentration in these cells is above 130 mM. The precise locations of the S-cells outside phloem bundles in rosette and cauline leaves and in flower stalks were visualised using sulphur mapping by cryo-SEM/EDX. S-cells contain up to 40% of total sulphur in flower stalk tissues. S-cells in emerging flower stalks and developing leaf tissues show typical signs of Programmed Cell Death (PCD) or apoptosis, such as chromatin condensation in the nucleus and blebbing of the membranes. TUNEL staining for DNA double strand breaks confirmed PCD in S-cells in postmeristematic tissues in the flower stalk as well as in the leaf. Our results show that S-cells in postmeristematic tissues proceed to an extreme degree of metabolic specialisation besides PCD. Accumulation and maintenance of a high concentration of GSL in these cells are accompanied by degradation of a number of cell organelles. The substantial changes in the cell composition during S-cell differentiation indicate the importance of this particular GSL-based phloem defence system. The specific anatomy of the S-cells and ability to accumulate specialised secondary metabolites is similar to that of the non-articulated laticifer cells in latex plants and thus indicates a common evolutionary origin.

Integrating drivers influencing the detection of plant pests carried in the international cut flower trade

This paper analyses the cut flower market as an example of an invasion pathway along which species of non-indigenous plant pests can travel to reach new areas. The paper examines the probability of pest detection by assessing information on pest detection and detection effort associated with the import of cut flowers. We test the link between the probability of plant pest arrivals as a precursor to potential invasion, and volume of traded flowers using count data regression models. The analysis is applied to the UK import of specific genera of cut flowers form Kenya between 1996 and 2004. There is a link between pest detection and the Genus of cut flower imported. Hence, pest detection efforts should focus on identifying and targeting those imported plants with a high risk of carrying pest species. For most of the plants studied efforts allocated to inspection have a significant influence on the probabilty of pest detction. However, by better targetting inspection efforts, it is shown that plant inspection effort could be reduced without increasing the risk of pest entry. Similarly, for most of the plants analysed, an increase in volume traded will not necessarily lead to an increase in the number of pests entering the UK. For some species, such as conclude that analysis at the rank of plant Genus is important both to understand the effectiveness of plant pest detection efforts and consequently to manage the risk of introduction of non-indigenous species.

Post-abscission, pre-dispersal seeds of Digitalis purpurea remain in a developmental state that is not terminated by desiccation ex planta

Seed quality may be compromised if seeds are harvested before natural dispersal (shedding). It has been shown previously that slow or delayed drying can increase potential quality compared with immediate rapid drying. This study set out to investigate whether or not there is a critical moisture content, below which drying terminates maturation events for seeds harvested after mass maturity but before dispersal. Seeds of foxglove (Digitalis purpurea) in the post-abscission pre-dispersal phase were held at between 15 and 95 % RH for 4 or 8 d, with or without re-hydration to 95 % RH for a further 4 d, before drying to equilibrium at 15 % RH. In addition, dry seeds were primed for 48 h at -1 MPa. Subsequent seed longevity was assessed at 60 % RH and 45 degrees C. Rate of germination and longevity were improved by holding seeds at a wide range of humidities after harvest. Longevity was further improved by re-hydration at 95 % RH. Priming improved the longevity of the seeds dried immediately after harvest, but not of those first held at 95 % RH for 8 d prior to drying. Maturation continued ex planta in these post-abscission, pre-dispersal seeds of D. purpurea dried at 15-80 % RH at a rate correlated positively with RH (cf. ageing of mature seeds). Subsequent re-hydration at 95 % RH enabled a further improvement in quality. Priming seeds initially stored air-dry for 3 months also allowed maturation events to resume. However, once individual seeds within the population had reached maximum longevity, priming had a negative impact on their subsequent survival.

The role of flower inclination, depth, and height in the preferences of a pollinating beetle (Coleoptera : Glaphyridae)

Amphicoma ( Glaphyridae) beetles are important pollinators of red bowl-shaped flowers in the Mediterranean. The role of color and shape in flower choice is well studied but the roles of inclination, depth, and height have seldom been investigated. Under field conditions, models were used to experimentally manipulate these three characters and visitation rates of beetles were recorded. Models with red horizontal surfaces were visited significantly more often than models with red vertical surfaces. Shallow flower models were visited significantly more than deeper equivalents. Models below or at the height of natural flower populations elicited significantly more landings than models above the height of flowers. Inclination, depth, and height characteristics are all likely to be important components in the flower preferences exhibited by pollinating beetles.

Nectar resource diversity organises flower-visitor community structure

Communities of nectar-producing plants show high spatio-temporal variation in the patterns of volume and concentration presentation. We illustrate a novel approach for quantifying nectar reward structures in complex communities, demonstrating that nectar resource diversity (defined as the variety of nectar volume-concentration combinations available) may be a fundamental factor organising nectarivore communities. In a series of diverse bee and entomophilous flower communities in Israel, our measure of nectar resource diversity alone explains the majority of variation in bee species richness, while other nectar variables (volume, concentration, energy value, and water content) have little predictive value per se. The new measure of nectar resource diversity is highly correlated with floral species richness and particularly with the species richness of annuals, yet it is additive in its effect on bee diversity. We conclude that relying solely upon measurements of mean nectar volume and mean nectar concentration overlooks a key characteristic of community-level reward structure, nectar resource diversity, so that previous studies may have failed to identify an important determinant of flower-visitor community structure.

The impact of Solanum elaeagnifolium, an invasive plant in the Mediterranean, on the flower visitation and seed set of the native co-flowering species Glaucium flavum

We examined the effect of the invasive Solanum elaeagnifolium (Solanaceae) on flower visitation patterns and seed set of the co-flowering native Glaucium flavum (Papaveraceae). We observed flowering G. flavum plants in invaded and uninvaded sites and found that G. flavum flowers in uninvaded sites received significantly more total visits. In addition, we hand-pollinated flowers on plants of G. flavum with (i) pure conspecific pollen, (ii) pure S. elaeagnifolium pollen and (iii) three different mixtures of the two types of pollen (containing 25, 50 and 75% invasive pollen). As a control, flowers were left unmanipulated or were permanently bagged. Seed set did not differ significantly between flowers receiving pollen mixtures and pure conspecific pollen. However, in the open pollination treatment, seed set was significantly lower than in the 100% conspecific pollen treatment, which suggests pollen limitation. Bagged flowers had very low seed set. G. flavum was generally resilient against the deposition of S. elaeagnifolium pollen.

Floral meristem indeterminacy depends on flower position and is facilitated by acarpellate gynoecium development in Impatiens balsamina

Floral meristems are generally determinate. Termination of their activity varies with species, occurring after carpel or ovule development, depending on the placentation type. In terminal flowering Impatiens balsamina (cv. Dwarf Bush Flowered) some flowers exhibit meristem indeterminacy; they produce organs from the placenta after ovule development. Here we provide a detailed description of gynoecium development in this line and explore the basis of the indeterminate nature of some of its floral meristems. We find that the placenta is sometimes established without complete carpel fusion. Proliferative growth derives from meristematic remnants of the placenta and is more common in the terminal inflorescence. RNA in situ hybridization reveals that IbLFY (Impatiens LFY homologue) is expressed in all meristem states, even in proliferating meristems. Expression of IbAG in axillary flowers is as expected in the meristem, stamens and carpels but absent from the proliferating meristem. We conclude that I. balsamina has cauline placentation. Incomplete suppression of inflorescence identity in flowers of the terminal inflorescence leads to floral meristem proliferation after ovule development in this species.

The regulation of sweet cherry fruit abscission by polar auxin transport

Inconsistency of cropping is an important problem for UK sweet cherry production. Premature fruit abscission in Prunus can reduce yields severely, however, the environmental cues and hormonal signals that trigger abscission have not been identified. Auxin (IAA) is known to delay abscission by reducing the sensitivity of cells in the abscission zone to ethylene, a promoter of abscission. Therefore, the capacity for polar auxin transport (PAT) through sweet cherry pedicels was examined in relation to fruit abscission. Cherry ‘spurs’ (short shoots) with similar leaf areas and different fruit numbers were phloem-girdled to restrict assimilate movement. Abscission from spurs with many fruit (eight or more) occurred within 14 days of girdling, whereas abscission from spurs with few (two) fruit was minimal. The pedicels’ capacity for PAT in spurs with different fruit numbers was determined 1, 3 and 9 days after girdling (DAG). Fruit were analysed for endogenous IAA concentration 3, 5, 7 and 9 DAG. PAT inhibitors 2,3,5-triiodobenzoic acid or 1-N-naphthylphtalamic acid were applied to pedicels of fruit not expected to abscise, i.e. on spurs with few fruit. The effect of these inhibitors on fruit abscission was determined 14 DAG. The proportion of the transported [3H]-IAA was lower from the outset in pedicels from spurs with many fruit. By 9 DAG, symptoms of fruit abscission were apparent and 40% less [3H] -IAA was transported through pedicels on spurs with many fruit. Fruit endogenous IAA concentrations were similar in the two groups of spurs. Application of PAT inhibitors shortly after girdling increased fruit abscission by 30%. The results suggest that although a decline in PAT is not the only cause of fruit abscission, the maintenance of PAT contributes to fruit retention.

Mechanisms and function of flower and inflorescence reversion

Flower and inflorescence reversion involve a switch from floral development back to vegetative development, thus rendering flowering a phase in an ongoing growth pattern rather than a terminal act of the meristem. Although it can be considered an unusual event, reversion raises questions about the nature and function of flowering. It is linked to environmental conditions and is most often a response to conditions opposite to those that induce flowering. Research on molecular genetic mechanisms underlying plant development over the last 15 years has pinpointed some of the key genes involved in the transition to flowering and flower development. Such investigations have also uncovered mutations which reduce floral maintenance or alter the balance between vegetative and floral features of the plant. How this information contributes to an understanding of floral reversion is assessed here. One issue that arises is whether floral commitment (defined as the ability to continue flowering when inductive conditions no longer exist) is a developmental switch affecting the whole plant or is a mechanism which assigns autonomy to individual meristems. A related question is whether floral or vegetative development is the underlying default pathway of the plant. This review begins by considering how studies of flowering in Arabidopsis thaliana have aided understanding of mechanisms of floral maintenance. Arabidopsis has not been found to revert to leaf production in any of the conditions or genetic backgrounds analysed to date. A clear-cut reversion to leaf production has, however, been described in Impatiens balsamina. It is proposed that a single gene controls whether Impatiens reverts or can maintain flowering when inductive conditions are removed, and it is inferred that this gene functions to control the synthesis or transport of a leaf-generated signal. But it is also argued that the susceptibility of Impatiens to reversion is a consequence of the meristem-based mechanisms controlling development of the flower in this species. Thus, in Impatiens, a leaf-derived signal is critical for completion of flowering and can be considered to be the basis of a plant-wide floral commitment that is achieved without accompanying meristem autonomy. The evidence, derived from in vitro and other studies, that similar mechanisms operate in other species is assessed. It is concluded that most species (including Arabidopsis) are less prone to reversion because signals from the leaf are less ephemeral, and the pathways driving flower development have a high level of redundancy that generates meristem autonomy even when leaf-derived signals are weak. This gives stability to the flowering process, even where its initiation is dependent on environmental cues. On this interpretation, Impatiens reversion appears as an anomaly resulting from an unusual combination of leaf signalling and meristem regulation. Nevertheless, it is shown that the ability to revert can serve a function in the life history strategy (perenniality) or reproductive habit (pseudovivipary) of many plants. In these instances reversion has been assimilated into regular plant development and plays a crucial role there.

The impact of an insecticide on insect flower visitation and pollination in an agricultural landscape

1 Pesticides are considered a threat to pollinators but little is known about the potential impacts of their widespread use on pollinators. Less still is known about the impacts on pollination, comprising the ecosystem service that pollinators provide to wildflowers and crops. 2 The present study measured flower visitation and pollination in an agricultural landscape, by placing potted flowering plants (Petunia sp.) in vine fields sprayed with a highly toxic insecticide (fenitrothion). During two sampling rounds, insect visitors to the petunias were observed and measures of pollination were recorded by counting and weighing seeds. 3 In the earlier sampling round, a lower species richness of insect visitors was observed in fields that had received an early application of insecticide. No negative impacts were found from later applications. The results obtained suggest a greater potential harm to insect pollinators and flower visitation as a result of insecticide application early in the season. 4 No reduction in pollination was found in fields that received an early insecticide application. Pollination was greater with two insecticide applications between sampling rounds rather than one application. 5 In the present study system, insecticide application had a negative effect on pollinators but a possible positive effect on pollination services. In some cases, it may be that actions for conserving biodiversity will not benefit pollination services to all plants.

Genetic analysis reveals a novel locus in Vicia faba decoupling pigmentation in the flower from that in the extra-floral nectaries

The genetics of the stipule spot pigmentation (SSP) in faba bean (Vicia faba L.) was studied using four inbred lines, of which Disco/2 was zero-tannin (zt2) with colourless stipule spots, ILB938/2 was normal-tannin (ZT2) with colourless stipule spots, and both Aurora/2 and Mélodie/2 were ZT2 with coloured stipule spots. Crosses Mélodie/2 × ILB 938/2, Mélodie/2 × Disco/2, ILB 938/2 × Aurora/2 and ILB 938/2 × Disco/2 (A, B, C and D, respectively) were prepared, along with reciprocals and backcrosses, and advanced through single-seed descent. All F1 hybrid plants had pigmented stipule spots, and in the F2 generation, the segregation ratio fit 3 coloured:1 colourless in crosses A, B and C and 9:7 in cross D. In the F3 generation, the ratio fit 5:3 in crosses A and C and 25:39 in cross D, and in the F4 generation, 9:7 in cross A. SSP was linked to the zero-tannin characteristics (white flower) only in cross B. The results show that coloured stipule spot is dominant to colourless and that colouration is determined by two unlinked complementary recessive genes. We propose the symbols ssp2 for the gene associated with zt2 in Disco/2 and ssp1 for the gene not associated with tannin content in ILB938/2. The novel ssp1 locus was mapped at F5 in cross ‘A’ using Medicago truncatula-derived single-nucleotide polymorphism and was on chromosome 1 of faba bean, in a well-conserved region of M. truncatula chromosome 5 containing some candidate Myb and basic helix–loop–helix transcription factor genes.

Where is the UK's pollinator biodiversity? The importance of urban areas for flower-visiting insects

Insect pollinators provide a crucial ecosystem service, but are under threat. Urban areas could be important for pollinators, though their value relative to other habitats is poorly known. We compared pollinator communities using quantified flower-visitation networks in 36 sites (each 1 km2) in three landscapes: urban, farmland and nature reserves. Overall, flower-visitor abundance and species richness did not differ significantly between the three landscape types. Bee abundance did not differ between landscapes, but bee species richness was higher in urban areas than farmland. Hoverfly abundance was higher in farmland and nature reserves than urban sites, but species richness did not differ significantly. While urban pollinator assemblages were more homogeneous across space than those in farmland or nature reserves, there was no significant difference in the numbers of rarer species between the three landscapes. Network-level specialization was higher in farmland than urban sites. Relative to other habitats, urban visitors foraged from a greater number of plant species (higher generality) but also visited a lower proportion of available plant species (higher specialization), both possibly driven by higher urban plant richness. Urban areas are growing, and improving their value for pollinators should be part of any national strategy to conserve and restore pollinators.