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Mechanisms of long-distance dispersal of seeds by wind.

Long-distance dispersal (LDD) is central to species expansion following climate change, re-colonization of disturbed areas and control of pests. The current paradigm is that the frequency and spatial extent of LDD events are extremely difficult to predict. Here we show that mechanistic models coupling seed release and aerodynamics with turbulent transport processes provide accurate probabilistic descriptions of LDD of seeds by wind. The proposed model reliably predicts the vertical distribution of dispersed seeds of five tree species observed along a 45-m high tower in an eastern US deciduous forest. Simulations show that uplifting above the forest canopy is necessary and sufficient for LDD, hence, they provide the means to define LDD quantitatively rather than arbitrarily. Seed uplifting probability thus sets an upper bound on the probability of long-distance colonization. Uplifted yellow poplar seeds are on average lighter than seeds at the forest floor, but also include the heaviest seeds. Because uplifting probabilities are appreciable (as much as 1 5%), and tree seed crops are commonly massive, some LDD events will establish individuals that can critically affect plant dynamics on large scales.

Computer Simulation↗

Effects of distance and pollen competition on gene flow in the wind-pollinated grass Festuca pratensis Huds.

Pollen dispersal and gene flow in the grass meadow fescue (Festuca pratensis Huds.) were studied using two populations which were homozygous for different allozymes at the Gpi-2 locus. The populations were established in a concentric donor-acceptor field experiment. Gene flow was found mainly to be affected by the distance between the donor and acceptor plants. Analysing 21 132 acceptor plant progenies, gene flow was shown to decrease rapidly with distance to the donor field up to 75 m, and beyond this distance much more slowly. The ability of donor pollen to fertilize acceptor plants depended very much on the density of the acceptor plants. Pairs of acceptor plants produced more compatible pollen locally, and captured significantly less donor pollen than single-plants. Despite the higher seed production of acceptor plants planted in pairs, the absolute number of heterozygous seeds carrying the donor allele was always lower than for single-plants. Wind direction had only a slight effect upon the type of pollen captured. Because of pollen production within the two plant populations being continuous and overlapping, the time when anthesis occurred had little effect on gene flow between the populations. Vigorous and tall acceptor plants with many panicles, high seed yield and high 1000-seed weight were able to capture more donor pollen than shorter plants. The results may be used to assess the risk of gene flow and to develop strategies for monitoring the spread of transgenes from genetically modified grasses.

Gene Frequency↗

Responses of the biting midge Culicoides impunctatus to acetone, CO2 and 1-octen-3-ol in a wind tunnel.

Responses of host-seeking female Culicoides impunctatus (Goetghebuer) (Diptera: Ceratopogonidae) to acetone, carbon dioxide and 1-octen-3-ol were measured in a wind tunnel. Carbon dioxide, presented as a filamentous plume, increased upwind flight in a dose-dependent manner, up to 0.09% concentration. A homogenous CO2 plume elicited similar upwind responses at concentrations up to 0.09%, whereas higher plume concentrations (> 0.1%) induced erratic responses with a suppression of upwind flight. Bovine equivalent concentrations of acetone (1.5 x 10(-6)g/l) and 1-octen-3-ol (1.3 x 10(-8)g/) failed to induce any significant upwind response when tested alone. In the presence of CO2, however, 1-octen-3-ol showed highly significant increases in upwind responses at concentrations of 1.3 x 10(-1) - 10(-8)g/l. Mixtures of CO2+ acetone also enhanced upwind flight at 1.5 x 10(-9)g/l. High tunnel concentrations of both 1-octen-3-ol and acetone inhibited upwind responses. These findings are discussed in relation to host finding by C. impunctatus and known mechanisms by which upwind flight is initiated and arrested at high odour concentrations.

Acetone↗

DNA from bird-dispersed seed and wind-disseminated pollen provides insights into postglacial colonization and population genetic structure of whitebark pine (Pinus albicaulis).

Uniparentally inherited mitochondrial (mt)DNA and chloroplast (cp)DNA microsatellites (cpSSRs) were used to examine population genetic structure and biogeographic patterns of bird-dispersed seed and wind-disseminated pollen of whitebark pine (Pinus albicaulis Engelm.). Sampling was conducted from 41 populations throughout the range of the species. Analyses provide evidence for an ancestral haplotype and two derived mtDNA haplotypes with distinct regional distributions. An abrupt contact zone between mtDNA haplotypes in the Cascade Range suggests postglacial biogeographic movements. Among three cpSSR loci, 42 haplotypes were detected within 28 cpSSR sample populations that were aggregated into six regions. Analysis of molecular variance (amova) was used to determine the hierarchical genetic structure of cpSSRs. amova and population pairwise comparisons (FST ) of cpSSR, and geographical distribution of mtDNA haplotypes provide insights into historical changes in biogeography. The genetic data suggest that whitebark pine has been intimately tied to climatic change and associated glaciation, which has led to range movements facilitated by seed dispersal by Clark's nutcracker (Nucifraga columbiana Wilson). The two hypotheses proposed to explain the genetic structure are: (i) a northward expansion into Canada and the northern Cascades in the early Holocene; and (ii) historical gene flow between Idaho and the Oregon Cascades when more continuous habitat existed in Central Oregon during the late Pleistocene. Genetic structure and insights gained from historical seed movements provide a basis on which to develop recovery plans for a species that is at risk from multiple threats.

Animals↗

Genetic effects of chronic habitat fragmentation in a wind-pollinated tree.

Habitat fragmentation poses a serious threat to plants through genetic changes associated with increased isolation and reduced population size. However, the longevity of trees, combined with effective seed or pollen dispersal, can enhance their resistance to these effects. The European beech (Fagus sylvatica) dominates forest over large regions of Europe. We demonstrate that habitat fragmentation in this species has led to genetic bottlenecks and the disruption of the species' breeding system, leading to significantly elevated levels of inbreeding, population divergence, and reduced genetic diversity within populations. These results show that, in contrast with the findings of previous studies, forest fragmentation has a negative genetic impact, even in this widespread, wind-pollinated tree. The identification of significant effects of forest fragmentation in beech demonstrates that trees are not at reduced risk from environmental change. This should be accounted for in the management of remaining natural and seminatural forest throughout the world.

Ecology↗

Wind-dispersed pollen mediates postglacial gene flow among refugia.

A long-term genetic legacy of refugial isolation has been postulated and was demonstrated for maternal refugial lineages for numerous plant and animal species. The lineages were assumed to have remained separated from each other for several glacial periods. The conifer Abies alba Miller, silver fir, is an excellent model to test whether pollen-mediated gene flow may eliminate the genetic imprints of Pleistocene refugial isolation. Two DNA markers with contrasting modes of inheritance were applied to 100 populations covering the entire range of silver fir in Europe. The markers exhibited each two highly conserved alleles based on an insertion/deletion of 80 bp in the fourth intron of the mitochondrial nad5 gene and on a synonymous substitution in the chloroplast psbC gene. The geographical distribution of the maternally inherited mitochondrial variation supported the existence of at least two refugia with two recolonizing maternal lineages remaining largely separated throughout the range. The cline of the nad5 allele frequencies was much steeper than the one of the two psbC alleles. The psbC cline was as wide as the whole range of the species. Our results provide striking evidence that even a species with very long generation times and heavy pollen grains was able to establish a highly efficient pollen-mediated gene flow between refugia. Therefore we postulate that an exchange of genetic information between refugia by range-wide paternal introgression is possible in wind-pollinated plant species.

Abies↗

Effects of ambient temperature, age and wind speed on the thermal balance of layer-strain fowls.

1. In White Leghorn laying hybrids aged 14 to 399 d the effects were measured of ambient temperature (Ta) and wind speed (WS) on heat production (HP), evaporative heat loss (EHL), nonevaporative heat loss with or without considering heat storage (NEHL1 or NEHL2), conductance (c), total insulation (I) and colonic temperature (Tc) as well as on biological optimum temperature (BOT), thermoneutral temperature (TNT) and threshold temperature for evaporative heat loss (Te). 2. All thermal balance variables were affected by age, Ta and WS. 3. The relationships between Ta and HP or NEHL2 were best expressed in terms of a polynomial function at low WS, and in terms of a linear function at medium and high WS; the relationship between Ta and NEHL1 at all WS studied was best described in terms of linear functions, the slope of which increased with decreasing Ta and rising WS; those between Ta, EHL and c were best described in terms of exponential functions and between Ta and I in terms of quadratic functions, the rate of increase of which was higher at low Ta and medium and high WS. 4. The description of relationships between the age of birds and their HP, EHL, NEHL1 and NEHL2 was performed after logarithmic transformation and yielded a linearly decreasing function with advancing age. 5. Threshold temperatures increased as WS increased from 0.2 to 1.2 m/s; for EHL (Te) by up to 7.5 K, for Tc (BOT) and for HP (TNT) by 10 K. 6. The effects of age, Ta and WS on HP and both forms of NEHL were summarised by multiple regression equations.

Aging↗

A drainage wind flow model.

A method of predetermining drainage wind velocities is derived in the form of an incremental velocity model. The model requires the topographical data and a general description of surface roughness. Using these fixed inputs with the model yields velocity factors. Velocity factors are determined as a function of height above the terrain and elapsed time since the occurrence of the maximum surface temperature. The velocity factors multiplied by a cooling factor give velocities. Time or space mean velocity may be found by summation techniques. By computing velocity factors for several points within an area of concern, one can determine drainage flow patterns and pollutant movements for any given night.

Air Movements↗

New insights into the effect of wind assistance on sprinting performance.

Here, a new mathematical model of sprinting is proposed. The prediction method rests on the construction of an energy balance incorporating a mathematical representation of each of the major terms in the balance. The term expressing the degradation of mechanical energy into thermal energy is formulated to express a dependence on wind speed. The dependence of the drag term on the change in mean body angle relative to the horizontal is taken into account. Whereas the effect of modifying the degradation term is shown to be significant, changes in body lean angle are shown to have little effect. Comparisons of the present predictions with a previous statistical analysis of 100-m track data show good agreement. Sensitivity analyses show which variables have the greatest influence on sprinting performance.

Biomechanical Phenomena↗

Numerical-assisted design of a wind tunnel used in the estimation of volatilization from water, waste or soil.

Numerical simulations of the flow inside two wind tunnel geometries used for determining emission volatilization have been conducted. The flow pattern was examined in a plane close to the emitting surface. A reduced surface, characterizing the low velocity zones in this plane, has been defined. The proposed geometry modification, consisting of a convergent/divergent system, decreases the low velocity regions by a factor ranging between 2 and 3.5. Large vortices present in the conventional configuration are almost eliminated. The new velocity distribution, close to the emitting surface, is more uniform and should therefore generate a more efficient mass transfer to the flowing air.

Air Pollutants↗

Application of a wind model for the El Paso-Juarez airshed.

A time-dependent, 3-dimensional mesoscale model, version 5 (MM5), developed by the Penn State University and the National Center for Atmospheric Research, was applied to study the meteorology over complex terrain of the El Paso-Juarez area. MM5 meteorological output data were compared against experimental data from the Texas Natural Resource Conservation Commission on days of reported high ozone concentrations. Model runs were conducted for a 36-, 12-, 4-km grid arrangement. Results indicate that the dispersion of pollutants by wind plays a significant role on days of low peak ozone concentration.

Air Pollution↗

The wind-frequency allocation method on discharge loading of function zones.

This paper introduces a new allocation method on discharge loading of each function zone in a total emission control region. The wind frequency, the position of each district, and the pollutant's influence area were taken into account in this new method. The concept of "average downwind distance" was brought forward in this paper. The method here is more reasonable than the original method of area distribution, which was proposed by the "A-value" method in regulation of total emissions in China, by means of the simulation of annual average concentration in the total emission control region.

Air Pollutants↗

Wind-driven surficial oxygen transfer and dinitrogen gas emission from treatment lagoons.

Surficial oxygen transfer plays an important role, when analyzing the complex biochemical and physical processes responsible for ammonia and dinitrogen gas emission in animal waste treatment lagoons. This paper analyzes if currently known nitrogen biochemical pathways can explain the enigmatic dinitrogen gas emissions recently observed from the treatment lagoons, based on the amount of wind-driven oxygen that can be transferred through the air-water interface. The stoichiometric amounts of the maximum dinitrogen gas production potential per unit mass of O(2) transferred were calculated according to three most likely biochemical pathways for ammonia removal in the treatment lagoons-classical nitrification-denitrification, partial nitrification-denitrification, and partial nitrification-Anammox. Partial nitrification-Anammox pathway would produce the largest N(2) emission, followed by partial nitrification-denitrification pathway, then by classical nitrification-denitrification pathway. In order to estimate stoichiometric amount (i.e., maximum) of N(2) emission from these pathways, we assumed that heterotrophic respiration was substantially inhibited due to high levels of free ammonia prevalent in treatment lagoons. Most observed N(2) emission data were below the maximum N(2) emission potentials by the classical nitrification-denitrification pathway. However, one value of observed N(2) emission was much higher than that could be produced by even the partial nitrification-Anammox pathway. This finding suggests yet unknown biological processes and/or non-biological nitrogen processes such as chemodenitrification may also be important in these treatment lagoons.

Adsorption↗

Prediction of clothing thermal insulation and moisture vapour resistance of the clothed body walking in wind.

Clothing thermal insulation and moisture vapour resistance are the two most important parameters in thermal environmental engineering, functional clothing design and end use of clothing ensembles. In this study, clothing thermal insulation and moisture vapour resistance of various types of clothing ensembles were measured using the walking-able sweating manikin, Walter, under various environmental conditions and walking speeds. Based on an extensive experimental investigation and an improved understanding of the effects of body activities and environmental conditions, a simple but effective direct regression model has been established, for predicting the clothing thermal insulation and moisture vapour resistance under wind and walking motion, from those when the manikin was standing in still air. The model has been validated by using experimental data reported in the previous literature. It has shown that the new models have advantages and provide very accurate prediction.

Equipment Design↗

Enhanced transpiration in response to wind effects at the edge of a blue gum (Eucalyptus globulus) plantation.

In Australia, tree planting has been widely promoted to alleviate dryland salinity and one proposed planting configuration is that of strategically placed interception belts. We conducted an experiment to determine the effect of tree position in a belt on transpiration rate. We also assessed how much the effect of tree position can be explained by advection and environmental conditions. Daily transpiration rates were determined by the heat pulse velocity technique for four edge and 12 inner trees in a 7-year-old Tasmanian blue gum (Eucalyptus globulus) plantation in South Australia. Various climatic variables were logged automatically at one edge of the plantation. The relationship between daily sap flow and sapwood area was strongly linear for the edge trees (r2 = 0.97), but only moderately correlated for the inner trees (r2 = 0.46), suggesting an edge effect. For all trees, sap flow normalized to sapwood area (Qs) increased with potential evaporation (PE) initially and then became independent as PE increased further. There was a fairly close correlation between transpiration of the edge and inner trees, implying that water availability was partially responsible for the difference between inner and edge trees. However, the ratio of edge tree to inner tree transpiration differed from unity, indicating differences in canopy conductance, which were estimated by an inverse form of the Penman-Monteith equation. When canopy conductances were less than a critical value, there was a strong linear relationship between Qs of the edge and inner trees. When canopy conductances of the edge trees were greater than the critical value, the slope of the linear relationship was steeper, indicating greater transpiration of the edge trees compared with the inner trees. This was interpreted as evidence for enhancement of transpiration of the edge trees by advection of wind energy.

Eucalyptus↗

Vortex wakes generated by robins Erithacus rubecula during free flight in a wind tunnel.

The wakes of two individual robins were measured in digital particle image velocimetry (DPIV) experiments conducted in the Lund wind tunnel. Wake measurements were compared with each other, and with previous studies in the same facility. There was no significant individual variation in any of the measured quantities. Qualitatively, the wake structure and its gradual variation with flight speed were exactly as previously measured for the thrush nightingale. A procedure that accounts for the disparate sources of circulation spread over the complex wake structure nevertheless can account for the vertical momentum flux required to support the weight, and an example calculation is given for estimating drag from the components of horizontal momentum flux (whose net value is zero). The measured circulations of the largest structures in the wake can be predicted quite well by simple models, and expressions are given to predict these and other measurable quantities in future bird flight experiments.

Animals↗

Past and current gene flow in the selfing, wind-dispersed species Mycelis muralis in western Europe.

The distribution of genetic diversity in Mycelis muralis, or wall lettuce, was investigated at a European scale using 12 microsatellite markers to infer historical and contemporary forces from genetic patterns. Mycelis muralis has the potential for long-distance seed dispersal by wind, is mainly self-pollinated, and has patchily distributed populations, some of which may show metapopulation dynamics. A total of 359 individuals were sampled from 17 populations located in three regions, designated southern Europe (Spain and France), the Netherlands, and Sweden. At this within-region scale, contemporary evolutionary forces (selfing and metapopulation dynamics) are responsible for high differentiation between populations (0.34 < F(ST) < 0.60) but, contrary to expectation, levels of within-population diversity, estimated by Nei's unbiased expected heterozygosity (H(E)) (0.24 < H(E) < 0.68) or analyses of molecular variance (50% of the variation found within-populations), were not low. We suggest that the latter results, which are unusual in selfing species, arise from efficient seed dispersal that counteracts population turnover and thus maintains genetic diversity within populations. At the European scale, northern regions showed lower allelic richness (A = 2.38) than populations from southern Europe (A = 3.34). In light of postglacial colonization hypotheses, these results suggest that rare alleles may have been lost during recolonization northwards. Our results further suggest that mutation has contributed to genetic differentiation between southern and northern Europe, and that Sweden may have been colonized by dispersers originating from at least two different refugia.

Analysis of Variance↗

Regional gene flow and population structure of the wind-dispersed plant species Hypochaeris radicata (Asteraceae) in an agricultural landscape.

Using microsatellites, we investigated population structure and gene flow of the short-lived, wind-dispersed plant species Hypochaeris radicata in a fragmented agricultural landscape where more than 99% of the nutrient-poor grasslands have disappeared over the last century. We sampled populations in the few remaining high density populations in conservation areas, as well as individuals that occurred, with lower densities, in linear landscape elements, at two spatial scales. In a re-inventory of the landscape, after 3 years, both extinctions and colonizations of populations were observed. Contrary to expectations, no differences in genetic diversity between high and low density populations were observed. Both types of populations had relatively high levels of diversity. Overall genetic differentiation (theta) was 0.04 and significantly different from zero (P < 0.01). A significant isolation-by-distance pattern was found when all populations were simultaneously analysed (r = 0.24, P = 0.013). Isolation by distance was (marginally) significant at the small scale (r = 0.32, P = 0.06), whereas nonsignificant at the large spatial scale (r = -0.05, P = 0.66). A maximization-of-explained-variance procedure resulted in a threshold distance of 3.5 km above which populations were effectively genetically isolated. An additional partial exclusion Bayesian-based assignment test showed that overall 32.3% of the individuals were assigned to their population of origin, 48% were assigned to another population in the area and 19.7% were not assigned. Together, these results suggest high levels of gene flow. Seed dispersal contributes to the observed gene flow up to several hundred metres, which is higher than previously modelled using aerodynamic models on seed dispersal of H. radicata. We discuss the consequences of these results for an evaluation of the probability of persistence of this species in the fragmented landscape.

Asteraceae↗