Size-dependent ESS sex allocation in wind-pollinated cosexual plants: fecundity vs. stature effects

To theoretically investigate the single and compound effects of relative fecundity and relative stature of plants on size-dependent sex allocation (SDS) in wind-pollinated cosexual species, we developed a game model and analysed ESS sex allocation of large and small plants having totally or partially different reproductive resources and different pollen and seed dispersal areas in a population. We found that e.g. when both sized plants have large pollen dispersal areas relative to their seed dispersal areas, which plants are male-biased is largely determined by relative fecundity (t) and relative size of seed dispersal area (k) of the large plants to the small plants: If t >k, large plants tend to be more male-biased even if relative size of pollen dispersal area of large to small plants (l) is smaller than k. If t相似文献   

The role of model selection in describing stochastic ecological processes

A great deal of variation in ecological processes can be generated through variation in environmental conditions. Models describing the underlying processes should be able to account for this variation. However, models may not be flexible enough, so that different realizations of a process may be better described by different models. This may lead to uncertainty in model selection.
Here we examine the question of whether two empirical models can provide consistent fits to different realizations of a process affected by environmental variation. We further examine the sensitivity of the model predictions to the amount of data available and the selection of the model. To study this, we simulated pollen dispersal patterns under varying wind conditions and then investigated whether the datasets consistently supported the same model. The role of the model selection and the impact of the spatial range over which the dispersal distances were observed were assessed by comparing model predictions at long dispersal distances.
There was no consistent pattern of one model providing a better fit than the other across simulations. The model providing better fit varied depending on the range of distances over which the dispersal patterns were observed, and on the amount of long-distance dispersal. The model predictions were found to be very sensitive to the selection of the model.
The variation between datasets produced with the same underlying mechanisms cannot be easily described using one model, which also limits our ability to reliably predict the underlying process. Therefore, the amount of information about a model choice provided by an individual field study may be rather limited. If we are to understand processes that are affected by environmental variation then we have to observe the range of possible outcomes of the processes under varying spatio-temporal conditions.     

Evolution of habitat-dependent sex allocation in plants: superficially similar to, but intrinsically different from animals

Because pollen disperses and ovules do not, a basic difference in dispersal abilities of male and female gametes exists in plants. With an analytical model, we show that the combination of such sex-biased dispersal of gametes and variation of habitat quality results in two opposite selective forces acting on the evolution of sex allocation in plants: (i) a plant should overproduce pollen in good patches and overproduce ovules in poor patches in order to equilibrate secondary sex ratios of gametes after pollen dispersal; (ii) a plant should overproduce ovules in good patches and overproduce pollen in poor patches in order to increase the likelihood that its progeny establishes in good patches. Our theoretical results indicate that the evolution of habitat-dependent sex allocation should be favoured in plants, in a direction that depends on the relative dispersal ability of pollen and seeds. We also show that superficially similar predictions obtained for habitat-dependent evolutionarily stable sex allocation in animals actually result from a completely different balance between the two underlying evolutionary forces.     

Sex‐specific selection on plant architecture through “budget” and “direct” effects in experimental populations of the wind‐pollinated herb,Mercurialis annua

Sexual selection may contribute to the evolution of plant sexual dimorphism by favoring architectural traits in males that improve pollen dispersal to mates. In both sexes, larger individuals may be favored by allowing the allocation of more resources to gamete production (a “budget” effect of size). In wind‐pollinated plants, large size may also benefit males by allowing the liberation of pollen from a greater height, fostering its dispersal (a “direct” effect of size). To assess these effects and their implications for trait selection, we measured selection on plant morphology in both males and females of the wind‐pollinated dioecious herb Mercurialis annua in two separate experimental common gardens at contrasting density. In both gardens, selection strongly favored males that disperse their pollen further. Selection for pollen production was observed in the high‐density garden only, and was weak. In addition, male morphologies associated with increased mean pollen dispersal differed between the two gardens, as elongated branches were favored in the high‐density garden, whereas shorter plants with longer inflorescence stalks were favored in the low‐density garden. Larger females were selected in both gardens. Our results point to the importance of both a direct effect of selection on male traits that affect pollen dispersal, and, to a lesser extent, a budget effect of selection on pollen production.     

Modelling pollen‐mediated gene flow in rice: risk assessment and management of transgene escape

Fast development and commercialization of genetically modified plants have aroused concerns of transgene escape and its environmental consequences. A model that can effectively predict pollen‐mediated gene flow (PMGF) is essential for assessing and managing risks from transgene escape. A pollen‐trap method was used to measure the wind‐borne pollen dispersal in cultivated rice and common wild rice, and effects of relative humidity, temperature and wind speed on pollen dispersal were estimated. A PMGF model was constructed based on the pollen dispersal pattern in rice, taking outcrossing rates of recipients and cross‐compatibility between rice and its wild relatives into consideration. Published rice gene flow data were used to validate the model. Pollen density decreased in a simple exponential pattern with distances to the rice field. High relative humidity reduced pollen dispersal distances. Model simulation showed an increased PMGF frequency with the increase of pollen source size (the area of a rice field), but this effect levelled off with a large pollen‐source size. Cross‐compatibility is essential when modelling PMGF from rice to its wild relatives. The model fits the data well, including PMGF from rice to its wild relatives. Therefore, it can be used to predict PMGF in rice under diverse conditions (e.g. different outcrossing rates and cross‐compatibilities), facilitating the determination of isolation distances to minimize transgene escape. The PMGF model may be extended to other wind‐pollinated plant species such as wheat and barley.     

Gene Flow,Risk Assessment and the Environmental Release of Transgenic Plants

The release of genetically modified plants is governed by regulations that aim to provide an assessment of potential impact on the environment. One of the most important components of this risk assessment is an evaluation of the probability of gene flow. In this review, we provide an overview of the current literature on gene flow from transgenic plants, providing a framework of issues for those considering the release of a transgenic plant into the environment. For some plants gene flow from transgenic crops is well documented, and this information is discussed in detail in this review. Mechanisms of gene flow vary from plant species to plant species and range from the possibility of asexual propagation, short- or long-distance pollen dispersal mediated by insects or wind and seed dispersal. Volunteer populations of transgenic plants may occur where seed is inadvertently spread during harvest or commercial distribution. If there are wild populations related to the transgenic crop then hybridization and eventually introgression in the wild may occur, as it has for herbicide resistant transgenic oilseed rape (Brassica napus). Tools to measure the amount of gene flow, experimental data measuring the distance of pollen dispersal, and experiments measuring hybridization and seed survivability are discussed in this review. The various methods that have been proposed to prevent gene flow from genetically modified plants are also described. The current “transgenic traits” in the major crops confer resistance to herbicides and certain insects. Such traits could confer a selective advantage (an increase in fitness) in wild plant populations in some circumstances, were gene flow to occur. However, there is ample evidence that gene flow from crops to related wild species occurred before the development of transgenic crops and this should be taken into account in the risk assessment process.     

Should I stay or should I go? Mycorrhizal plants are more likely to invest in long‐distance seed dispersal than non‐mycorrhizal plants

Seed dispersal and mycorrhizal associations are key mutualisms for the functioning and regeneration of plant communities; however, these processes have seldom been explored together. We hypothesised that obligatory mycorrhizal plants will be less likely to have long‐distance dispersal (LDD) syndromes since the probability of finding suitable mycorrhizal partners is likely to decrease with distance to the mother plant. We contrasted the mycorrhizal status and LDD syndromes for 1960 European plant species, using phylogenetically corrected log‐linear models. Contrary to our expectation, having specialised structures for LDD is more frequent in obligate mycorrhizal plants than in non‐mycorrhizal plants, revealing that lack of compatible mutualists does not constrain investment in LDD structures in the European Flora. Ectomycorrhizal plants associated with wind‐dispersing fungi are also more likely to have specialised structures for wind dispersal. Habitat specificity and narrower niche of non‐mycorrhizal plants might explain the smaller investment in specialised structures for seed dispersal.     

Effect of canopy closure on pollen dispersal in a wind-pollinated species (Fagus sylvatica L.)

The effect of non-reproductive trees and saplings as a physical barrier to pollen dispersal in wind-pollinated species?? forests has not received enough attention in the literature so far. The neighborhood seedling model was used to fit pollen dispersal models for beech at different stages of gap recolonization and to elucidate the effect of saplings as a physical barrier on pollen dispersal at local scale. Phenological overlap of leaf emergence, and pollen release as well as wind directionality patterns were also examined. As a case study, we used a mixed beech-oak forest that was managed as open woodland until 1974. The ban on entry of cattle has led to the recolonization of empty spaces by seedlings and saplings of beech (Fagus sylvatica L.) and two oak species (Quercus petraea (Matts.) Liebl. and Q. pyrenaica Willd.) and, at last, to canopy closure. The average pollen dispersal distance for the first plants that regenerated in the gaps was almost twice those found for recently installed seedlings and seeds collected in traps, supporting the hypothesis that the understory may act as a physical barrier to pollen dispersal. Although a substantial part of effective pollination directionality is at random, horizontal winds and vertical anabatic winds may explain some of this directionality. At the time of beech pollen release, leaves of beech and sessile oak are fully developed, enhancing pollen interception by the saplings. Explicit models of pollen dispersal for wind-pollinated trees should incorporate the effect of canopy closure caused by growth of saplings and account for leaf phenology of co-occurring species in the forest.     

Pollen clumping and wind dispersal in an invasive angiosperm

Pollen dispersal is a fundamental aspect of plant reproductive biology that maintains connectivity between spatially separated populations. Pollen clumping, a characteristic feature of insect-pollinated plants, is generally assumed to be a detriment to wind pollination because clumps disperse shorter distances than do solitary pollen grains. Yet pollen clumps have been observed in dispersion studies of some widely distributed wind-pollinated species. We used Ambrosia artemisiifolia (common ragweed; Asteraceae), a successful invasive angiosperm, to investigate the effect of clumping on wind dispersal of pollen under natural conditions in a large field. Results of simultaneous measurements of clump size both in pollen shedding from male flowers and airborne pollen being dispersed in the atmosphere are combined with a transport model to show that rather than being detrimental, clumps may actually be advantageous for wind pollination. Initial clumps can pollinate the parent population, while smaller clumps that arise from breakup of larger clumps can cross-pollinate distant populations.     

Long distance seed dispersal by wind: measuring and modelling the tail of the curve

The size and shape of the tail of the seed dispersal curve is important in determining the spatial dynamics of plants, but is difficult to quantify. We devised an experimental protocol to measure long-distance dispersal which involved measuring dispersal by wind from isolated individuals at a range of distances from the source, but maintaining a large and constant sampling intensity at each distance. Seeds were trapped up to 80 m from the plants, the furthest a dispersal curve for an individual plant has been measured for a non-tree species. Standard empirical negative exponential and inverse power models were fitted using likelihood methods. The latter always had a better fit than the former, but in most cases neither described the data well, and strongly under-estimated the tail of the dispersal curve. An alternative model formulation with two kernel components had a much better fit in most cases and described the tail data more accurately. Mechanistic models provide an alternative to direct measurement of dispersal. However, while a previous mechanistic model accurately predicted the modal dispersal distance, it always under-predicted the measured tail. Long-distance dispersal may be caused by rare extremes in horizontal wind speed or turbulence. Therefore, under-estimation of the tail by standard empirical models and mechanistic models may indicate a lack of flexibility to take account of such extremes. Future studies should examine carefully whether the widely used exponential and power models are, in fact, valid, and investigate alternative models. Received: 7 March 1999 / Accepted: 2 April 2000     

Leptokurtic pollen-flow,non-leptokurtic gene-flow in a wind-pollinated herb,Plantago lanceolata L.

Summary The purpose of this study was to simultaneously measure pollen dispersal distance and actual pollen-mediated gene-flow distance in a wind-pollinated herb, Plantago lanceolata. The pollen dispersal distribution, measured as pollen deposition in a wind tunnel, is leptokurtic, as expected from previous studies of wind-pollinated plants. Gene-flow, measured as seeds produced on rows of male-sterile inflorescences in the wind tunnel, is non-leptokurtic, peaking at an intermediate distance. The difference between the two distributions results from the tendency of the pollen grains to cluster. These pollen clusters are the units of gene dispersal, with clusters of intermediate and large size contributing disproportionately to gene-flow. Since many wind-pollinated species show pollen clustering (see text), the common assumption for wind-pollinated plants that gene-flow is leptokurtic requires re-examination. Gene-flow was also measured in an artifical outdoor population of male-steriles, containing a single pollen source plant in the center of the array. The gene flow distribution is significantly platykurtic, and has the same general properties outdoors, where wind speed and turbulence are uncontrolled, as it does in the wind tunnel. I estimated genetic neighborhood size based on my measure of gene-flow in the outdoor population. The estimate shows that populations of Plantago lanceolata will vary in effective number from a few tens of plants to more than five hundred plants, depending on the density of the population in question. Thus, the measured pollen-mediated gene-flow distribution and population density will interact to produce effective population sizes ranging from those in which there is no random genetic drift to those in which random genetic drift plays an important role in determining gene frequencies within and among populations. Despite the platykurtosis in the distribution, pollen-mediated gene dispersal distances are still quite limited, and considerable within and among-population genetic differentiation is to be expected in this species.     

Increases in air temperature can promote wind-driven dispersal and spread of plants

Long-distance dispersal (LDD) of seeds and pollen shapes the spatial dynamics of plant genotypes, populations and communities. Quantifying LDD is thus important for predicting the future dynamics of plants exposed to environmental changes. However, environmental changes can also alter the behaviour of LDD vectors: for instance, increasing air temperature may enhance atmospheric instability, thereby altering the turbulent airflow that transports seed and pollen. Here, we investigate temperature effects on wind dispersal in a boreal forest using a 10-year time series of micrometeorological measurements and a Lagrangian stochastic model for particle transport. For a wide range of dispersal and life history types, we found positive relations between air temperature and LDD. This translates into a largely consistent positive effect of +3°C warming on predicted LDD frequencies and spread rates of plants. Relative increases in LDD frequency tend to be higher for heavy-seeded plants, whereas absolute increases in LDD and spread rates are higher for light-seeded plants for which wind is often an important dispersal vector. While these predicted increases are not sufficient to compensate forecasted range losses and environmental changes can alter plant spread in various ways, our results generally suggest that warming can promote wind-driven movements of plant genotypes and populations in boreal forests.     

Pollen–plant–climate relationships in sub-Saharan Africa

Aim  To demonstrate that incorporating the bioclimatic range of possible contributor plants leads to improved accuracy in interpreting the palaeoclimatic record of taxonomically complex pollen types.
Location  North Tropical Africa.
Methods  The geographical ranges of selected African plants were extracted from the literature and geo-referenced. These plant ranges were compared with the pollen percentages obtained from a network of surface sediments. Climate-response surfaces were graphed for each pollen taxon and each corresponding plant species.
Results  Several patterns can be identified, including taxa for which the pollen and plant distributions coincide, and others where the range limits diverge. Some pollen types display a reduced climate range compared with that of the corresponding plant species, due to low pollen production and/or dispersal. For other taxa, corresponding to high pollen producers such as pioneer taxa, pollen types display a larger climatic envelope than that of the corresponding plants. The number of species contained in a pollen taxon is an important factor, as the botanical species included in a taxon may have different geographical and climate distributions.
Main conclusions  The comparison between pollen and plant distributions is an essential step towards more precise vegetation and climate reconstructions in Africa, as it identifies taxa that have a high correspondence between pollen and plant distribution patterns. Our method is a useful tool to reassess biome reconstructions in Africa and to characterize accurately the vegetation and climate conditions at a regional scale, from pollen data.     

The release of genetically modified grasses. Part 2: the influence of wind direction on pollen dispersal

 In part 1 an experiment was described for determining the extent of pollen dispersal from a Lolium source. The results were used to test Bateman’s pollen dispersal equations, which were found to be not particularly useful for describing variation in pollen deposition with distance. An improvement is suggested here which takes the influence of wind direction into account. For 11 of the 12 datasets the new equations fit significantly better than the original ones. Mean wind directions were used to produce 15 data subsets for testing Bateman’s equations for dispersal downwind of a pollen source. These equations fitted only 4 of the data subsets, all of which were collected from traps facing towards the pollen source. The usefulness of the model equations in estimating the importance of turbulence is brought into question. It is shown that models incorporating only distance and wind direction do not accurately describe pollen deposition. The amount of pollen deposited does not always decrease smoothly with increasing distance from the source. The variation in the amount of pollen deposited is probably influenced by several factors, including wind speed and turbulence. Received: 5 June 1996 / Accepted: 11 October 1996     

A mathematical and statistical framework for modelling dispersal

Mechanistic and phenomenological dispersal modelling of organisms has long been an area of intensive research. Recently, there has been an increased interest in intermediate models between the two. Intermediate models include major mechanisms that affect dispersal, in addition to the dispersal curve of a phenomenological model. Here we review and describe the mathematical and statistical framework for phenomenological dispersal modelling. In the mathematical development we describe modelling of dispersal in two dimensions from a point source, and in one dimension from a line or area source. In the statistical development we describe applicable observation distributions, and the procedures of model fitting, comparison, checking, and prediction. The procedures are also demonstrated using data from dispersal experiments. The data are hierarchically structured, and hence, we fit hierarchical models. The Bayesian modelling approach is applied, which allows us to show the uncertainty in the parameter estimates and in predictions. Finally, we show how to account for the effect of wind speed on the estimates of the dispersal parameters. This serves as an example of how to strengthen the coupling in the modelling between the phenomenon observed in an experiment and the underlying process – something that should be striven for in the statistical modelling of dispersal.     

To succeed globally, disperse locally: effects of local pollen and seed dispersal on tetraploid establishment

Newly formed tetraploid plants in sympatry with their diploid progenitors should face significant obstacles to persistence and population establishment because of low-fitness triploids formed by cross-ploidy pollinations. Prior models have found restrictive conditions for a minority tetraploid subpopulation to persist. A stochastic spatial model, parameterized using snow buttercups (Ranunculus adoneus), was used to examine the influence of limited seed and pollen dispersal distances on the success of minority tetraploids and the interaction of these factors with different rates of self-pollination and tetraploid advantage. Higher rates of self-pollination and increased tetraploid advantage increase the probability of tetraploid persistence. Limiting the dispersal of seeds and pollen further increases the positive impact of any given level of self-pollination and tetraploid advantage. Taxa with short-distance seed and pollen dispersal should face much less stringent barriers to sympatric polyploid speciation than taxa with long-distance dispersal patterns. With short-distance seed and pollen dispersal, polyploid speciation should be possible in the absence of ecological differentiation or recurrent polyploid formation through unreduced gametes.     

Pollen and seed flow under different predominant winds in wind-pollinated and wind-dispersed species <Emphasis Type="Italic">Engelhardia roxburghiana</Emphasis>

In most plants, the contributions of pollen and seed flow to their genetic structures are generally difficult to disentangle. For typical wind-pollinated and wind-dispersed species Engelhardia roxburghiana in a 20-ha natural forest plot in lower subtropic China, because the prevailing wind directions change during its pollen release and seed dispersal seasons, we could compare its genetic structures in different directions, which could result primarily from pollen or seed flow. Furthermore, because the plot has undergone from an open to a closed canopy stage historically, we also examined forest canopy effects on gene flow in different generations and different directions. Using 522 E. roxburghiana individuals mapped in the plot, our results revealed that greater pollen flow led to biased gene flow in the pollen dispersal-predominant direction (pollen direction), while greater seed flow generated less spatial genetic structure in the seed dispersal-predominant direction (seed direction). The results predicted from generalized additive models indicated that canopy closure enhanced resistance to gene flow from the old generation to the new generation. Analyses by landscape genetic models for the new generation revealed that gene flow associated with pollen direction was more strongly affected by canopy than with seed direction. Our study is new by proposing an alternative way to separate effects of the pollen and seed flow on spatial variation patterns in E. roxburghiana. To our knowledge, our study is also the first attempt to use landscape genetic models to represent canopy effects for different dispersal vectors in spatial scales only up to a few hundred meters.     

Analysis of genetic stability of plants regenerated from suspension cultures and protoplasts of meadow fescue (Festuca pratensis Huds.)

A cytological and molecular analysis was performed to assess the genetic uniformity and true-to-type character of plants regenerated from 20 week-old embryogenic suspension cultures of meadow fescue (Festuca pratensis Huds.), and compared to protoplastderived plants obtained from the same cell suspension. Cytological variation was not observed in a representative sample of plants regenerated directly from the embryogenic suspensions and from protoplasts isolated therefrom. Similarly, no restriction fragment length polymorphisms (RFLPs) were detected in the mitochondrial, plastid and nuclear genomes in the plants analyzed. Randomly amplified polymorphic DNA markers (RAPDs) have been used to characterise molecularly a set of mature meadow fescue plants regenerated from these in vitro cultures. RAPD markers using 18 different short oligonucleotide primers of arbitrary nucleotide sequence in combination with polymerase chain reaction (PCR) allowed the detection of pre-existing polymorphisms in the donor genotypes, but failed to reveal newly generated variation in the protoplast-derived plants compared to their equivalent suspensionculture regenerated materials.The genetic stability of meadow fescue plants regenerated from suspension cultures and protoplasts isolated therefrom and its implications on gene transfer technology for this species are discussed.Abbreviations PCR polymerase chain reaction - RAPD random amplified polymorphic DNA - RFLP restriction fragment length polymorphism.     

Pollen dispersal is driven by pollinator response to plant disease and plant spatial aggregation

Most plant species are pollinated by animals, mainly insects, who adjust their foraging behaviour to the spatial distribution of rewards. Any changes in rewards of individual plants could then affect pollen dispersal at the level of plant patches or populations. Such change in floral rewards often results from infection by plant pathogens, for example by anther smuts (i.e. no pollen and reduced nectar in diseased flowers). Here, we tested the hypothesis that the infection of plant populations by anther smuts affects the pattern of pollen dispersal.We investigated the patterns of pollen dispersal in experimental arrays of potted plants differing in the presence of diseased plants and the degree of plant spatial aggregation. We tracked pollen dispersal using a fluorescent dye powder as a pollen analogue, while we simultaneously observed pollinator foraging behaviour.We found that the dispersal of the pollen analogue increased in the presence of diseased plants in experimental arrays, but this effect was strongly dependant on plant spatial aggregation. The parallel observations of pollinator behaviour suggest that this pattern resulted from pollinator discrimination against diseased plants and increased movement in arrays with intermingled diseased plants, provided that plant clusters were close to each other.Our study indicates that pollinators respond to diseased plants in a similar way as to healthy plants with low rewards. Consequently, diseased plants should be treated not only as a potential source of infection but also as a factor influencing pollen dispersal in plant populations.