Temporal variation of heterozygosity‐based assortative mating and related benefits in a lesser kestrel population

Heterozygosity as a target of mate choice has received much attention in recent years and there is growing evidence supporting its role in the evolution of mate preferences. In this study we analyse mating patterns in relation to heterozygosity in a lesser kestrel (Falco naumanni) population intensively monitored over six study years (2002–2007). The magnitude of heterozygosity‐based assortative mating varied over time, being particularly patent in the last study years (2006, 2007). We have found evidence that this mating pattern entails both direct and indirect‐genetic benefits. Clutch size increased with female heterozygosity and more heterozygous males raised a higher number of fledglings particularly in those years when the strength of the heterozygosity‐based assortative mating was markedly higher. In the last study year, parent–offspring correlation of heterozygosity was stronger and higher than the expected if individuals would have randomly mated with respect to heterozygosity. Overall, our results offer empirical support to the heterozygous mate hypothesis of sexual selection but suggest that genetic diversity may act as a temporally variable target for mate choice.     

Predicting evolution of floral traits associated with mating system in a natural plant population

Evolution of floral traits requires that they are heritable, that they affect fitness, and that they are not constrained by genetic correlations. These prerequisites have only rarely been examined in natural populations. For Mimulus guttatus, we found by using the Riska-method that corolla width, anther length, ovary length and number of red dots on the corolla were heritable in a natural population. Seed production (maternal fitness) was directly positively affected by corolla width and anther size, and indirectly so by ovary length and number of red dots on the corolla. The siring success (paternal fitness), as estimated from allozyme data, was directly negatively affected by anther-stigma separation, and indirectly so by the corolla length-width ratio. Genetic correlations, estimated with the Lynch-method, were positive between floral size measures. We predict that larger flowers with larger reproductive organs, which generally favour outcrossing, will evolve in this natural population of M. guttatus.     

Early growth invigoration of jack pine seedlings by natural plant growth regulators

 Survival of conifer transplants is often poor on exposed planting sites in the boreal forest. More than one-third of all conifers do not become established. To enhance the competitive ability of jack pine seedlings, seeds were treated with natural plant growth regulators (PGRs; viz., homobrassinolide, salicylic acid, and two polyamines, spermine and spermidine) and growth promotion was studied for 16 days. Homobrassinolide (5 ng l^–1), salicylic acid (100 μg l^–1) and spermine (10 μg l^–1) enhanced elongation growth and elongation rate of whole plant. Homobrassinolide (5 ng l^–1) and salicylic acid (100 μg l^–1) stimulated root elongation by 38% and 10% respectively while spermine (1000 μg l^–1) increased needle growth by 14%. Homobrassinolide stimulated dry weight growth and growth rate. Homobrassinolide recorded over 20% increase in dry matter production, apportioned equally to root and needles, whereas spermine enhanced total dry matter production by almost 10%, mostly by increasing needle weight. Homobrassinolide facilitated nearly 19% increase in growth rate while spermine recorded only a 7% growth promotion. Spermidine inhibited both elongation and dry weight growth at all concentrations. Growth promotion by homobrassinolide, salicylic acid and spermine may be through an acceleration of processes connected to cell elongation, cell division and C allocation and these PGRs showed most promise for the early invigoration and improvement of jack pine seedling growth. Received: 27 August 1997 / Accepted: 8 January 1998     

Age-dependent allozymic variation in a natural population of lizards

An analysis of allozymic variation at 17 loci in a population of the sagebrush lizard (Sceloporus graciosus) in southern Utah yielded an estimate of genic heterozygosity of 0.028. Seven of the loci were variable, but only one, Est-1, was strongly polymorphic. The observation that the frequency of the common genotype (MM) at the Est-1 locus declined monotonically in successive age classes from 0.74 in hatchlings to 0.58 in adults 4 years old or older suggests that allele frequencies at this locus are not independent of selective influences.This research was supported by NSF grant GB-29141 to D.W.T. and NIH grant GM-15769 to R.K.S.     

Multiple mating and sperm displacement in a natural population of Drosophila melanogaster

Summary Mother-offspring data for alcohol dehydrogenase genotypes of a vineyard cellar population of D. melanogaster are best explained by a model that allows 21% of females in the population to mate twice with an 83% level of sperm displacement. A population model with multiple mating and sperm displacement is examined theoretically. A formula for the effective population size is derived under this model. Multiple mating increases the effective population size relative to single mating.     

Cytonuclear disequilibrium and genetic drift in a natural population of ponderosa pine

We measured the cytonuclear disequilibrium between 11 nuclear allozyme loci and both mitochondrial and chloroplast DNA haplotypes in a natural population of ponderosa pine (Pinus ponderosa, Laws). Three allozyme loci showed significant associations with mtDNA variation, while two other loci showed significant association with cpDNA. However, the absolute number of individuals involved in any of the associations was small, such that in none of the nuclear-organellar combinations was the difference between observed and expected numbers >11 individuals. Patterns of association were not consistent across loci or organellar genomes, suggesting that they are not the result of mating patterns, which would act uniformly on all loci. This pattern of disequilibria is consistent with the action of genetic drift and with existing knowledge of the structure of this population and thus does not imply the action of other evolutionary processes. The overall magnitude (normalized disequilibrium) of associations was greater for maternally inherited mtDNA than for paternally inherited cpDNA, though this difference was neither large nor significant. Such significant disequilibria involving the paternally inherited organelle indicate that not only are there a limited number of seed parents, but the effective number of pollen parents is also limited.     

Temporal genetic variation in a population of the pocket gopher,Geomys bursarius

Genetic variation at eight polymorphic loci was examined in a population of a subterranean rodent, the plains pocket gopher (Geomys bursarius), sampled over a 10-yr period. Two loci exhibited relatively minor changes in gene frequencies, while the remaining loci demonstrated major shifts in predominant alleles, loss of minor alleles, and addition of alleles due to migration. Significant deviations from Hardy-Weinberg expectations, concomitant to heterozygote deficiencies, were observed for several loci. Temporal heterogeneity, as measured by F_ST, was high and comparable to that exhibited by local populations sampled over relatively short periods of time. The high degree of temporal genetic variation is consistent with observations that fossorial rodents occur in locally isolated populations with small effective population sizes that are subject to genetic drift, bottlenecking, and inbreeding.     

Host mating system and the spread of a disease-resistant allele in a population

The model presented here modifies a susceptible-infected (SI) host–pathogen model to determine the influence of mating system on the outcome of a host–pathogen interaction. Both deterministic and stochastic (individual-based) versions of the model were used. This model considers the potential consequences of varying mating systems on the rate of spread of both the pathogen and resistance alleles within the population. We assumed that a single allele for disease resistance was sufficient to confer complete resistance in an individual, and that both homozygote and heterozygote resistant individuals had the same mean birth and death rates. When disease invaded a population with only an initial small fraction of resistant genes, inbreeding (selfing) tended to increase the probability that the disease would soon be eliminated from a small population rather than become endemic, while outcrossing greatly increased the probability that the population would become extinct due to the disease.     

Host mating system and the prevalence of disease in a plant population

A modified susceptible-infected-recovered (SIR) host-pathogen model is used to determine the influence of plant mating system on the outcome of a host-pathogen interaction. Unlike previous models describing how interactions between mating system and pathogen infection affect individual fitness, this model considers the potential consequences of varying mating systems on the prevalence of resistance alleles and disease within the population. If a single allele for disease resistance is sufficient to confer complete resistance in an individual and if both homozygote and heterozygote resistant individuals have the same mean birth and death rates, then, for any parameter set, the selfing rate does not affect the proportions of resistant, susceptible or infected individuals at equilibrium. If homozygote and heterozygote individual birth rates differ, however, the mating system can make a difference in these proportions. In that case, depending on other parameters, increased selfing can either increase or decrease the rate of infection in the population. Results from this model also predict higher frequencies of resistance alleles in predominantly selfing compared to predominantly outcrossing populations for most model conditions. In populations that have higher selfing rates, the resistance alleles are concentrated in homozygotes, whereas in more outcrossing populations, there are more resistant heterozygotes.     

Socio-genetic structure and mating system of a wild boar population

Wild boars Sus scrofa have a social organization based on female groups that can include several generations of adults and offspring, and are thus likely matrilineal. However, little is known about the degree of relatedness between animals living in such groups or occupying the same core area of spatial activity. Also, polygynous male mating combined with matrilineal female groups can have strong influences on the genetic structure of populations. We used microsatellite genotyping combined with behavioral data to investigate the fine-scale population genetic structure and the mating system of wild boars in a multi-year study at Châteauvillain-Arc-en-Barrois (France). According to spatial genetic autocorrelation, females in spatial proximity were significantly inter-related. However, we found that numerous males contributed to the next generation, even within the same social group. Based on our genetic data and behavioral observations, wild boars in this population appear to have a low level of polygyny associated with matrilineal female groups, and infrequent multiple paternity. Mortality due to hunting may facilitate the breakup of what historically has been a more predominantly polygynous mating system, and likely accelerates the turnover of adults within the matrilineal groups.     

Seasonal variation in the mating system of a selfing annual with large floral displays

Background and Aims Flowering plants display considerable variation in mating system, specifically the relative frequency of cross- and self-fertilization. The majority of estimates of outcrossing rate do not account for temporal variation, particularly during the flowering season. Here, we investigated seasonal variation in mating and fertility in Incarvillea sinensis (Bignoniaceae), an annual with showy, insect-pollinated, ‘one-day’ flowers capable of delayed selfing. We examined the influence of several biotic and abiotic environmental factors on day-to-day variation in fruit set, seed set and patterns of mating.Methods We recorded daily flower number and pollinator abundance in nine 3 × 3-m patches in a population at Mu Us Sand land, Inner Mongolia, China. From marked flowers we collected data on daily fruit and seed set and estimated outcrossing rate and biparental inbreeding using six microsatellite loci and 172 open-pollinated families throughout the flowering period.Key Results Flower density increased significantly over most of the 50-d flowering season, but was associated with a decline in levels of pollinator service by bees, particularly on windy days. Fruit and seed set declined over time, especially during the latter third of the flowering period. Multilocus estimates of outcrossing rate were obtained using two methods (the programs MLTR and BORICE) and both indicated high selfing rates of ∼80 %. There was evidence for a significant increase in levels of selfing as the flowering season progressed and pollinator visitation declined. Biparental inbreeding also declined significantly as the flowering season progressed.Conclusions Temporal variation in outcrossing rates may be a common feature of the mating biology of annual, insect-pollinated plants of harsh environments but our study is the first to examine seasonal mating-system dynamics in this context. Despite having large flowers and showy floral displays, I. sinensis attracted relatively few pollinators. Delayed selfing by corolla dragging largely explains the occurrence of mixed mating in I. sinensis, and this mode of self-fertilization probably functions to promote reproductive assurance when pollinator service is limited by windy environmental conditions and competition from co-occurring flowering plants.     

Selection at work in self-incompatible Arabidopsis lyrata: mating patterns in a natural population

Identification and characterization of the self-incompatibility genes in Brassicaceae species now allow typing of self-incompatibility haplotypes in natural populations. In this study we sampled and mapped all 88 individuals in a small population of Arabidopsis lyrata from Iceland. The self-incompatibility haplotypes at the SRK gene were typed for all the plants and some of their progeny and used to investigate the realized mating patterns in the population. The observed frequencies of haplotypes were found to change considerably from the parent generation to the offspring generation around their deterministic equilibria as determined from the known dominance relations among haplotypes. We provide direct evidence that the incompatibility system discriminates against matings among adjacent individuals. Multiple paternity is very common, causing mate availability among progeny of a single mother to be much larger than expected for single paternity.     

Temporal variation in nutrient uptake capacity by intact roots of mature loblolly pine

Nutrient uptake is generally thought to exhibit a simple seasonal pattern, but few studies have measured temporal variation of nutrient uptake capacity in mature trees. We measured net uptake capacity of K, NH⁺₄, NO₃^–, Mg and Ca across a range of solution concentrations by roots of mature loblolly pine at Calhoun Experimental Forest in October 2001, July 2001, and April 2002. Uptake capacity was generally lowest in July; rates in October were similar to those in April. Across a range of concentrations, antecedent nutrient solution concentrations affected the temporal patterns in uptake in July but not in October or April. In July, uptake of NH⁺₄, Mg and Ca was positively correlated with concentration when roots were exposed to successively lower concentrations, but negatively correlated with concentration when exposed to successively higher concentrations. In contrast, uptake in October was constant across the range of concentrations, while uptake increased with concentration in April. As in studies of other species, we found greater uptake of NH⁺₄ than NO₃^–. Temporal patterns of uptake capacity are difficult to predict, and our results indicate that experimental conditions, such as experiment duration, antecedent root conditions and nutrient solution concentration, affect measured rates of nutrient uptake.     

Temporal variation and host association in the Campylobacter population in a longitudinal ruminant farm study

Campylobacter jejuni and C. coli were quantified and typed, using multilocus sequence typing (MLST), from fecal samples collected from a mixed cattle and sheep farm during summer. Cattle had a significantly higher prevalence than sheep (21.9% [74/338] and 14.0% [30/214], respectively), but both decreased over time. There were no differences in the average Campylobacter concentrations shed by cattle (600 CFU g(-1)) and sheep (820 CFU g(-1)), although sheep did show a significant temporal reduction in the number of Campylobacter organisms shed in their feces. A total of 21 different sequence types (STs) (97.7% C. jejuni, 2.3% C. coli) were isolated from cattle, and 9 different STs were isolated from sheep (40.6% C. jejuni, 59.4% C. coli). The Campylobacter population in cattle was relatively stable, and the frequencies of genotypes isolated showed little temporal variation. However, the composition of subtypes isolated from sheep did show significant temporal differences. The cattle and sheep consistently showed significant differences in their carriage of Campylobacter species, STs, and CCs despite the fact that both were exposed to the same farming environment. This work has highlighted the patterns of a Campylobacter population on a ruminant farm by identifying the existence of both temporal and between-host variations.     

Interpopulational variation of the mating system in the peacock blenny Salaria pavo

The ecology and mating system of two populations of the peacock blenny Salaria pavo that have been reported to differ in sexual behaviour were studied. In the Gulf of Trieste, a northern Adriatic Sea population lives in rocky shores, whilst in the Ria Formosa, an Atlantic population inhabits a coastal lagoon with sandy barrier islands. In the Gulf of Trieste, nest availability was found to be higher and nest aggregation lower than in Ria Formosa. Nesting males courted females more and in higher proportion, and inversely, females directed less courtship displays towards nest holders in the Gulf of Trieste than in Ria Formosa. The relative frequency of small female-like males that reproduce by parasitically fertilizing eggs in the larger males’ nests was lower in the Gulf of Trieste population. This variation in the sexual behaviour of nesting males and females is likely to reflect a plastic behavioural response to the varying ecological conditions. The difference between populations in the relative frequency of parasitic males suggests that the male alternative reproductive tactics are condition-dependent.     

Carbon sequestration in boreal jack pine stands following harvesting

A large area of boreal jack pine (Pinus banksiana Lamb.) forest in Canada is recovering from clear‐cut harvesting, and the carbon (C) balance of these regenerating forests remains uncertain. Net ecosystem CO₂ exchange was measured using the eddy‐covariance technique at four jack pine sites representing different stages of stand development: three postharvest sites (HJP02, HJP94, and HJP75) and one preharvest site (OJP). The four sites, located in the southern Canadian boreal forest, Saskatchewan, Canada, are typical of low productivity jack pine stands and were 2, 10, 29, and 90 years old in 2004, respectively. Mean annual net ecosystem production (NEP) for 2004 and 2005 was ?137±11, 19±16, 73±28, and 22±30 g C m^?2 yr^?1 at HJP02, HJP94, HJP75 and OJP, respectively, showing the postharvest jack pine stands to be moderate C sources immediately after harvesting, weak sinks at 10 years, moderate C sinks at 30 years, then weak C sinks at 90 years. Mean annual gross ecosystem photosynthesis (GEP) for the 2 years was 96±10, 347±20, 576±34, and 583±35 g C m^?2 yr^?1 at HJP02, HJP94, HJP75, and OJP, respectively. The ratio of annual ecosystem respiration (R) to annual GEP was 2.51±0.15, 0.95±0.04, 0.87±0.03, and 0.96±0.03. Seasonally, NEP peaked in May or June at all four sites but GEP and R were highest in July. R at a reference soil temperature of 10 °C, ecosystem quantum yield and photosynthetic capacity were lowest for the 2‐year‐old stand. R was most sensitive to soil temperature for the 90‐year‐old stand. The primary source of variability in NEP over the course of succession of the jack pine ecosystem following harvesting was stand age due to the changes in leaf area index. Intersite variability in GEP and R was an order of magnitude greater than interannual variability at OJP. For both young and old stands, GEP had greater interannual variability than R and played a more important role than R in interannual variation in NEP. Based on year‐round flux measurements from 2000 to 2005, the 10‐year stand had larger interannual variability in GEP and R than the 90‐year stand. Interannual variability in NEP was driven primarily by early‐growing‐season temperature and growing‐season length. Photosynthesis played a dominant role in the rapid rise in NEP early in stand development. Late in stand development, however, the subtle decrease in NEP resulted primarily from increasing respiration.     

Inbreeding depression and heterosis in a subdivided population: influence of the mating system

We investigate the joint effects of gene flow and selfing on the level of inbreeding depression, heterosis and genetic load in a subdivided population at equilibrium. Low gene flow reduces inbreeding depression and substantially increases heterosis. However, in highly self-fertilizing populations, inbreeding depression is independent of the amount of gene flow. When migration occurs via pollen, consanguinity of the reproductive system could have a negative influence on subpopulation persistence, in contrast to the case of isolated populations. However, with only seed migration, genetic load and heterosis depend mildly on the mating system. From an evolutionary point of view, we reach two main conclusions: first, outcrossing is selected for if gene flow is low; second, intermediate levels of gene flow could promote mixed mating systems, especially when migration occurs through pollen.