Current and historical factors drive variation of reproductive traits in unisexual mosses in Europe: A case study

Unisexual bryophytes provide excellent models to study the mechanisms that regulate the frequency of sexual versusasexual reproduction in plants, and their ecological and evolutionary implications. Here, we determined sex expression, phenotypic sex ratio, and individual shoot traits in 242 populations of the cosmopolitan moss Pseudoscleropodium purum spanning its whole distributional range. We tested whether niche differentiation, sex-specific differences in shoot size, and biogeographical history explained the spatial variation of reproductive traits. We observed high levels of sex expression and predominantly female-biased populations, although both traits showed high intraspecific variation among populations. Sex expression and sex ratio were partly explained by current macroscale environmental variation, with male shoots being less frequent at the higher end of the environmental gradients defined by the current distribution of the species. Female bias in population sex ratio was significantly lower in areas recolonized after the last glacial maximum (recent populations) than in glacial refugia (long-term persistent populations). We demonstrated that reproductive trait variation in perennial unisexual mosses is partially driven by macroscale and historical environmental variation. Based on our results, we hypothesize that sexual dimorphism in environmental tolerance and vegetative growth contribute to sex ratio bias over time, constraining the chances of sexual reproduction, especially in long-term persistent populations. Further studies combining genetic analyses and population monitoring should improve our understanding of the implications of the intraspecific variation in the frequency of sexual versusasexual reproduction in bryophyte population fitness and eco-evolutionary dynamics.     

Genetic variation in sex allocation in a parasitic wasp: variation in sex pattern within sequences of oviposition

In order to maximize their fitness under Local Mate Competition (LMC), arrhenotokous female wasps have to produce a precise sex ratio when encountering hosts. Recent progress in the theory of hymenopterous parasitoid reproduction suggest that they manage to do it by laying male and female eggs in a particular order and that such reproductive strategies are adaptive. Therefore, the determinism of such sequential patterns would be regulated by genetic control on which natural selection could act. To test this hypothesis, sequences of oviposition were recorded in a set ofTrichogramma brassicae Bezdenko (Hymenoptera; Trichogrammatidae) females and in their daughters by providing themEphestia kuehniella Zeller (Lepidoptera; Pyralidae) eggs. In order to describe accurately sex pattern within these oviposition sequences, I present a joined non-parametric and multivariate statistical method. It is shown thatT. brassicae females do not produce male and female eggs in random sequences. Moreover, the way they organize the sequence of the sexes in their progeny seems to be under a strong genetic control. The evolutionary consequences of such results are discussed.     

Temporal variation in temperature and rainfall differentially affects ectomycorrhizal colonization at two contrasting sites

We examined the roles that seasonal shifts in precipitation and temperature played in the ectomycorrhizal (ECM) colonization of pinyon pine ( Pinus edulis Engelm.) at two contrasting sites in northern Arizona. Pinyons growing in ash and cinder soils experienced much greater water and nutrient stress than pinyons growing nearby in sandy-loam soils. Over a one year period, we obtained monthly measurements of ECM colonization, root zone soil moisture and temperature, and air temperature and precipitation. Four major patterns emerged. Firstly, although climate as measured by ambient temperature and precipitation did not vary between the two sites, soil temperature was significantly higher and soil moisture significantly lower at the cinder site than at the sandy-loam site. Secondly, ECM colonization was significantly higher at the cinder site for 5 of 12 months. Thirdly, although nearly 70% of the variation in ECM colonization of pinyons growing in cinder soil was predicted by a combination of soil moisture and soil temperature, these same variables had little predictive power for pinyons growing in sandy-loam soils. Air temperature and precipitation were also significantly correlated with ECM colonization at the cinder site but not the sandy-loam site. Fourthly, a watering experiment showed that ECM colonization significantly increased with supplemental water at the cinder site, but not at the sandy-loam site. Thus, in two sites that did not differ in plant community or climate, ectomycorrhizas in cinder soils were far more sensitive to changes in moisture and temperature than ectomycorrhizas in sandy-loam soils.     

Genetic and environmental factors affecting reproductive variation in Allium vineale

Traits related to allocation of resources to sexual and asexual reproduction, together with seed production, were scored on Allium vineale plants sampled from five sites in southern Sweden during a period of 4 years. In addition, random amplified polymorphic DNA (RAPD) fingerprinting of the sampled plants allowed the identification of genets. Integration of genetic and phenotypic data from field and greenhouse provided for the analysis of among‐year, among‐site, and among‐genet variance components. These variance components were taken to represent the influences of short‐term environmental changes, persistent site divergence, and within‐site genet differences, respectively. It was shown that differences among sites and among genets explained a large part of the phenotypic variation of allocation traits, whereas among‐year differences had a larger influence on the variation in seed production. Together, the results support the conclusions of a recent model on the evolution of mixed reproductive systems, that predicts a stable balance between sexual and asexual reproduction because of annual fluctuations in fecundity through the two modes.     

Evidence for male-biased effective sex ratio and recent step-by-step colonization in the bivalve Pinctada mazatlanica

This paper presents a comparison of the geographical distribution of genetic variability at mitochondrial and nuclear loci among pearl oyster populations from the tropical American Pacific coast (Pinctada mazatlanica). Surprisingly, both mitochondrial and nuclear gene variability decreased regularly from north to south of the studied area, which, altogether with a significant correlation between genetic and geographical distances for mtDNA, suggests a recent colonization or re-colonization of the southern areas. However, the loss of diversity between north and south was much more important for mitochondrial than for nuclear DNA, and this did not translate into measurable fixation index at nuclear loci (theta = 0.03, n.s.), contrary to the mitochondrial data (theta = 0.18*). Smaller effective size of mtDNA accentuated by a strong male-biased effective sex ratio and step-by-step colonization from northern areas can explain this discrepancy among natural populations of this protandric species.     

Genetic by environmental variation but no local adaptation in oysters (Crassostrea virginica)

Functional trait variation within and across populations can strongly influence population, community, and ecosystem processes, but the relative contributions of genetic vs. environmental factors to this variation are often not clear, potentially complicating conservation and restoration efforts. For example, local adaptation, a particular type of genetic by environmental (G*E) interaction in which the fitness of a population in its own habitat is greater than in other habitats, is often invoked in management practices, even in the absence of supporting evidence. Despite increasing attention to the potential for G*E interactions, few studies have tested multiple populations and environments simultaneously, limiting our understanding of the spatial consistency in patterns of adaptive genetic variation. In addition, few studies explicitly differentiate adaptation in response to predation from other biological and environmental factors. We conducted a reciprocal transplant experiment of first‐generation eastern oyster (Crassostrea virginica) juveniles from six populations across three field sites spanning 1000 km in the southeastern Atlantic Bight in both the presence and absence of predation to test for G*E variation in this economically valuable and ecologically important species. We documented significant G*E variation in survival and growth, yet there was no evidence for local adaptation. Condition varied across oyster cohorts: Offspring of northern populations had better condition than offspring from the center of our region. Oyster populations in the southeastern Atlantic Bight differ in juvenile survival, growth, and condition, yet offspring from local broodstock do not have higher survival or growth than those from farther away. In the absence of population‐specific performance information, oyster restoration and aquaculture may benefit from incorporating multiple populations into their practices.     

四川大头茶在不同群落中的遗传分化及适合度成分

研究了四川大头茶在 3个群落中的遗传分化和适合度成分。 12个引物的 RAPD分析表明 ,仅有 10 %左右的遗传多样性存在于种群间。四川大头茶纯林结实率 (31.86 % )显著高于针阔混交林 (2 2 .5 3% )和常绿阔叶林 (2 4 .5 2 % )。每果种子数和每果种子均重各种群间差异均显著 ,每果种子数和每果种子均重分别为常绿阔叶林 (2 9.5 0 ,0 .0 15 4 g) ,四川大头茶纯林 (2 8.39,0 .0 172 g) ,针阔混交林 (2 7.4 4 ,0 .0 195 g) ,表明二者间存在着负耦联关系 (trade off)。而以 R =∑lxbx表示的适合度却以常绿阔叶林最大     

Genetic variation in restriction patterns among mouse amylase gene complexes

The expression of pancreatic amylase in the mouse exhibits pronounced genetic variation. Congenic lines with various amylase complexes on a common C3H/As background have different numbers and forms of isoenzymes. The relative ratio of these isoenzymes may vary, as does the overall production of pancreatic amylase, which in some lines is three- to fourfold higher than in others. DNA from a number of lines was digested with endonucleases and hybridized to an amylase cDNA probe. The restriction patterns from inbred stocks and the corresponding congenic lines are identical, demonstrating that the majority of (if not all) amylase-like DNA sequences is found within the amylase complex. Congenic lines with specific amylase expression, for instance, in enzyme production, show different restriction patterns, whereas three lines with the same amylase phenotype have a uniform pattern. Most of the variation in amylase expression is represented among congenic lines derived from Danish mice. A comparison of such lines with others of remote geographic origin reveals that the restriction patterns of the Danish lines have by far the highest degree of resemblance. This observation seems to exclude major rearrangements within the amylase complex as the cause of the differences in enzyme expression, which instead are likely to be due to variation in regulatory elements associated with the active structural amylase genes in the complex.This work was supported by Grant 11-2290 from the Danish Natural Science Research Council.     

The effects of genetic drift during range expansion on geographical patterns of variation: a computer simulation of the colonization of Australia by Bufo marinus

A computer simulation is performed of allele frequency changes resulting from genetic drift at eleven loci during the probable course of colonization of Australia by populations of the Giant Toad, Bufo marinus. The history of twelve populations for which allele frequency data are available is modelled. Account is taken of the likely pattern of relationship among the populations, the effective size of the populations (as indicated by the observed variance of allele frequencies) and the probable isolation of the populations from each other following their separation. In all simulations, allele frequencies at some loci show a significant association with latitude while others do not. In six of ten simulations, there is a significant association between degree of variation at a locus and the presence of a latitudinal cline of allele frequencies. There are also indications of this kind of association in simulations of a uni-directional range expansion. These results demonstrate that such associations, which were also observed in the data from the actual populations, can result from genetic drift during a range expansion, and therefore cannot be taken as evidence of the action of natural selection.     

Inter‐ and intraspecific variation in mycotoxin tolerance: A study of four Drosophila species

Many mycophagous Drosophila species have adapted to tolerate high concentrations of mycotoxins, an ability not reported in any other eukaryotes. Although an association between mycophagy and mycotoxin tolerance has been established in many Drosophila species, the genetic mechanisms of the tolerance are unknown. This study presents the inter‐ and intraspecific variation in the mycotoxin tolerance trait. We studied the mycotoxin tolerance in four Drosophila species from four separate clades within the immigrans‐tripunctata radiation from two distinct locations. The effect of mycotoxin treatment on 20 isofemale lines per species was studied using seven gross phenotypes: survival to pupation, survival to eclosion, development time to pupation and eclosion, thorax length, fecundity, and longevity. We observed interspecific variation among four species, with D. falleni being the most tolerant, followed by D. recens, D. neotestacea, and D. tripunctata, in that order. The results also revealed geographical variation and intraspecific genetic variation in mycotoxin tolerance. This report provides the foundation for further delineating the genetic mechanisms of the mycotoxin tolerance trait.     

Genetic Identity of Populus tremuloides Litter Influences Decomposition and Nutrient Release in a Mixed Forest Stand

Recent research has shown that genetic variation can directly impact community and ecosystem level processes. Populus tremuloides (trembling aspen) is an extremely widespread and genetically diverse tree species important to many North American forest ecosystems. Using leaf litter from five genotypes grown in a common garden under two nutrient treatments, we tracked litter decomposition in a natural aspen stand for 1 year. Here we show that aspen leaf litter decomposes and releases carbon, nitrogen, and sulfur in relation to its genetic identity. In a secondary experiment, we show that the genetic diversity of aspen litter mixtures can influence decomposition, however weakly so. Overall, nutrient treatments influenced leaf litter decomposition the most, followed by genetic identity, and then by genetic diversity (if at all in some cases). In this widespread, genetically diverse, and dominant species, genetic variation within a single species is important to ecosystem functioning. The relatively weak effect of genetic diversity on the processes measured here does not preclude its importance to ecosystem functioning, but does suggest that genetic identity and composition are more important than genetic diversity per se.     

Family affiliation,sex ratio and sporophyte frequency in unisexual mosses

Patterns of sex expression and sex ratios are key features of the life histories of organisms. Bryophytes are the only haploid‐dominant land plants. In contrast with seed plants, more than half of bryophyte species are dioecious, with rare sexual expression and sporophyte formation and a commonly female‐biased sex ratio. We asked whether variation in sex expression, sex ratio and sporophyte frequency in ten dioecious pleurocarpous wetland mosses of two different families was best explained by assuming that character states evolved: (1) in ancestors within the respective families or (2) at the species level as a response to recent habitat conditions. Lasso regression shrinkage identified relationships between family membership and sex ratio and sporophyte frequency, whereas environmental conditions were not correlated with any investigated reproductive trait. Sex ratio and sporophyte frequency were correlated with each other. Our results suggest that ancestry is more important than the current environment in explaining reproductive patterns at and above the species level in the studied wetland mosses, and that mechanisms controlling sex ratio and sporophyte frequency are phylogenetically conserved. Obviously, ancestry should be considered in the study of reproductive character state variation in plants. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 163–172.     

Genetic variation of sex allocation in the parasitoid wasp Heterospilus prosopidis

The genetic variation of sex ratio and sex allocation were examined in a series of half-sib analyses on the sex ratio of braconid parasitoid wasp Heterospilus prosopidis populations collected in Hawaii and Arizona. The mean threshold value and the range of the threshold for change in the sex of offspring in response to resource quality (host size) were determined. Estimates of the narrow-sense heritability (h2) of sex ratio at a specific host size ranged from 0.185 to 0.315, and those of the sex changing point (threshold value) ranged from 0.220 to 0.342. The coefficient of variation (CV(A)) of sex ratio was significantly larger than CV(A) of body weight. We discuss factors that maintained the significant additive genetic variation of sex ratio.     

Phantom hitch-hikers mislead estimates of genetic variation in Antarctic mosses

Previous studies of Antarctic mosses employing RAPDs have reported extraordinarily high levels of genetic variation, even within a single clump of moss. This is unexpected given their extreme isolation in Antarctica and lack of sexual reproduction. We offer an alternative explanation: that unusually elevated levels of genetic variability are artefacts from contamination of a number of biota known to be naturally associated with Antarctic mosses. We utilized sequence variation of nrITS and RAPDs to further investigate the effect of naturally occurring contaminants on estimates of genetic variation of mosses. Our results indicate that these ``phantom hitch-hiker' contaminants hinder attempts to accurately and reliably estimate levels of genetic variation by non-specific PCR-based approaches. Furthermore, screening samples via amplification of nrITS failed to identify all contaminated samples, hence we caution against relying solely on ``quick' screening methods and suggest that suspect samples be carefully examined for contamination.     

Genetic variation for postzygotic reproductive isolation between Caenorhabditis briggsae and Caenorhabditis sp. 9

The process of speciation is key to the origins of biodiversity, and yet the Caenorhabditis nematode model system has contributed little to this topic. Genetic studies of speciation in the genus are now feasible, owing to crosses between the recently discovered Caenorhabditis sp. 9 and the well-known C. briggsae producing fertile F(1) hybrid females. We dissected patterns of postzygotic reproductive isolation between these species by crossing eight isogenic strains of C. briggsae reciprocally with six strains of C. sp. 9. We determined that overall patterns of reproductive isolation are robust across these genetic backgrounds. However, we also quantified significant heritable variation within each species for interspecific hybrid incompatibilities for total adult progeny, egg-to-adult viability, and the percentage of male progeny. This demonstrates that intraspecific variation for interspecific hybrid incompatibility occurs despite extensive, albeit incomplete, reproductive isolation. Therefore, this emerging general phenomenon of variable reproductive isolation is not restricted to highly interfertile, early-stage incipient species, but also applies to species in the latest stages of the speciation process. Furthermore, we confirm Haldane's rule and demonstrate strongly asymmetric parent-of-origin effects (Darwin's corollary) that consistently manifest more extremely when hermaphroditic C. briggsae serves as maternal parent. These findings highlight Caenorhabditis as an emerging system for understanding the genetics of general patterns of reproductive isolation.     

Genetic and phenotypic variation among Galaxias maculatus populations reflects contrasting landscape effects between northern and southern Patagonia

1. Understanding the influence of landscape characteristics on genetic and phenotypic intraspecific variability can yield insights into how evolutionary processes work as well as provide essential information for the conservation of biodiversity. 2. Our aim was to compare the genetic structure and phenotypic variation among Galaxias maculatus populations inhabiting two Atlantic Ocean river basins in Patagonia in relation to historical and contemporary landscape characteristics associated with latitude. 3. Population genetic analysis (based on eight microsatellite loci, 505 individuals) indicates that genetic structure is more pronounced in the Río Negro basin (RN) at 39–41°S (10 localities) than in the southernmost Santa Cruz River basin (SCR) at 49–50°S (seven localities). Spatial autocorrelation analysis showed different effects of geographic distance in shaping population differentiation patterns in the two basins. 4. While in northern Patagonia, fragmented populations could have survived the severity of Quaternary climate cycles, at higher latitudes G. maculatus populations were probably extirpated by extensive ice sheets. Extant populations in the upper reaches of the southernmost basin probably originated from refugia close to the Atlantic Ocean. 5. Based on otolith microchemistry, we have documented facultative diadromy for the first time for the species in the SCR basin, while no evidence of migration to the sea was found in individuals from the RN basin. Vertebral number increased with latitude, and within the SCR basin, the higher counts are possibly associated with a migratory life style.     

Spatial and temporal variation in prey color patterns for background matching across a continuous heterogeneous environment

In heterogeneous habitats, camouflage via background matching can be challenging because visual characteristics can vary dramatically across small spatial scales. Additionally, temporal variation in signaling functions of coloration can affect crypsis, especially when animals use coloration seasonally for intraspecific signaling (e.g., mate selection). We currently have a poor understanding of how wild prey optimize background matching within continuously heterogeneous habitats, and whether this is affected by requirements of intraspecific signaling across biological seasons. Here, we quantified color patterns of a wild population of shore skink (Oligosoma smithi), a variably colored lizard endemic to New Zealand, to (a) investigate whether background matching varies across a vegetation gradient; (b) assess potential signaling functions of color; and (c) to determine whether there is a trade‐off between requirements for crypsis and intraspecific signaling in coloration across seasons. Although all pattern types occurred throughout the vegetation gradient, we found evidence for background matching in skinks across the vegetation gradient, where dorsal brightness and pattern complexity corresponded with the proportion of vegetation cover. There was also a significant disparity between ventral color (saturation) of juveniles and adults, and also between sexes, suggestive of sex recognition. However, there was little indication that color was condition‐dependent in adults. Despite some evidence for a potential role in signaling, crypsis did not greatly differ across seasons. Our study suggests that selection favors a mix of generalist and specialist background matching strategies across continuously heterogeneous habitats.