Functional ecology of ecotypic differentiation in the Californian serpentine sunflower (Helianthus exilis)

* Here, we examined phenotypic differences between locally adapted serpentine and riparian populations of the serpentine sunflower Helianthus exilis from northern California, USA. * Within a common environment, plants from serpentine and riparian sites were grown in regular potting soil or serpentine soil. Physiology, morphology, phenology and fitness-related traits were measured. * Overall, riparian plants grew more rapidly, attained a larger final size, produced larger leaves, and smaller flowering heads. Riparian plants also invested less in root biomass and were more water-use-efficient than the serpentine plants. Serpentine and riparian plants also differed in leaf concentrations of boron, magnesium, sodium and molybdenum. * These ecotypic differences suggest contrasting adaptive strategies to cope with either edaphic stress in serpentine sites or intense above-ground competition at riparian sites. There was a significant population origin x soil type crossing interaction in one fitness trait (average dry weight) that mirrored local adaptation previously documented for these riparian and serpentine ecotypes. However, because all other fitness traits did not exhibit this crossing interaction in our common garden study, it is possible that phenotypic differences underlying local adaptation may be amplified in the field as a result of biotic and abiotic interactions.     

Within- and trans-generational plasticity affects the opportunity for selection in barbed goatgrass (Aegilops triuncialis)

• Premise of the study: Environments are composed of selective agents, and environments may also modify the efficacy of these agents. Environments affect the rate of maximum evolutionary change by influencing variation in relative fitness (i.e., the opportunity for selection, or I). Within- and transgenerational plastic environmental responses may affect I, speeding or slowing processes of local adaptation. • Methods: We determined whether environmental factors affected the opportunity for selection (I) in Aegilops triuncialis (barbed goatgrass) by measuring I as a within- and transgenerational plastic response to two maternal glasshouse environments (serpentine/dry and loam/moist). We also determined whether this species’ two most common genetic lineages (determined by DNA microsatellite length polymorphism) varied in response to glasshouse treatments. • Key Results: Opportunity for selection was less for plants grown in the dry serpentine environment than for plants grown in the moist loam environment. This response varied between genetic lineages. The east lineage exhibited a within-generation response to the dry serpentine environment. For both seed mass and average seed weight in this lineage, the opportunity for selection was lower in dry serpentine than in moist loam. The west lineage had a transgenerational response to the dry serpentine such that the opportunity for selection for seed number and seed mass was lower for plants produced by mothers grown in dry serpentine than for plants produced by mothers in moist loam. • Conclusions: Phenotypic variation in relative fitness is constrained by the dry serpentine environment, which leads to lower evolvability in this environment. Within- and transgenerational effects of the environment may slow local adaptation to serpentine soils.     

Local adaptation, patterns of selection, and gene flow in the Californian serpentine sunflower (Helianthus exilis)

The traditional view of the species as the fundamental unit of evolution has been challenged by observations that in heterogeneous environments, gene flow may be too restricted to overcome the effects of local selection. Whether a species evolves as a cohesive unit depends critically on the dynamic balance between homogenizing gene flow among populations and potentially disruptive local adaptation. To examine this evolutionary balance between "global" gene flow and local selection, we studied northern Californian populations of Helianthus exilis, the serpentine sunflower, within a mosaic of contrasting serpentine and nonserpentine areas that differ considerably in soil chemistry and water availability. Local adaptation to riparian and serpentine habitats was studied in Helianthus exilis along with an analysis of gene flow patterns among populations within these habitats. Local adaptation was assessed in H. exilis during 2002 and 2003 using reciprocal transplant experiments at multiple locations within serpentine and riparian habitats. Effects of competition and germination date on the expression of local adaptation were also examined within the reciprocal transplant experiments. Local adaptation was detected in both years at the local site level and at the level of habitat. The analysis of the transplanted populations indicated that the patterns of selection differed considerably between riparian and serpentine sites. Differential survivorship occurred in serpentine habitats, whereas selection on reproductive output predominated in riparian habitats. Local adaptation was expressed only in the absence of competition. Local adaptation in terms of survivorship was most strongly expressed in treatments with delayed seed germination. Microsatellite markers were used to quantify population genetic parameters and examine the patterns of gene flow among sampled populations. Analysis of molecular markers revealed a system of population patches that freely exchange genes with each other. Strong selection seems to maintain ecotypic variation within this endemic sunflower species, while extensive gene flow among populations prevents local speciation between serpentine and riparian ecotypes.     

Little evidence for local adaptation to soils or microclimate in the post-fire recruitment of three Californian shrubs

Background: Seedling recruitment following fire is an infrequent yet critical demographic transition for woody plants in Mediterranean ecosystems.

Aims: Here we examine whether post-fire seedling recruitment of three widespread Californian chaparral shrubs is affected by local adaptation within an edaphically and topographically complex landscape.

Methods: We reciprocally transplanted 6-month-old seedlings of Adenostema fasciculatum, Ceanothus cuneatus and Eriodictyon californicum to serpentine and sandstone soils, and cool northerly and warm southerly slopes.

Results: At the age of 2 years, none of the species manifested higher survival or growth on ‘home’ compared with ‘away’ soils or slopes, indicating an absence of local adaptation with respect to seedling recruitment in these environments. Seedlings of all species manifested lower survival and relative growth on serpentine soils regardless of seedling source, as well as a variety of other destination and source effects.

Conclusions: The ability of these three species to recruit in new environments, such as in restoration settings or in response to shifting climates, is unlikely to be impeded by a need for seeds from sources that closely match their edaphic or topographic destination.     

Clinal resistance structure and pathogen local adaptation in a serpentine flax-flax rust interaction

Because disease resistance is a hallmark signature of pathogen-mediated selection pressure on hosts, studies of resistance structure (the spatial distribution of disease resistance genes among conspecific host populations) can provide valuable insights into the influence of pathogens on host evolution and spatial variation in the magnitude of their effects. To date few studies of wild plant-pathogen interactions have characterized resistance structure by sampling across the host's biogeographic range, and only a handful have paired such investigations with studies of disease levels under natural conditions. I used a greenhouse cross-inoculation experiment to characterize genetic resistance of 16 populations of California dwarf flax (Hesperolinon californicum) to attack by multiple samples of the rust fungus Melampsora lini. I documented a latitudinal cline in resistance structure, manifest across the host's biogeographic range, which mirrored almost identically a cline in infection prevalence documented through field surveys of disease in study populations. These results provide empirical evidence for clinal patterns of antagonistic selection pressure, demonstrate that such patterns can be manifest across broad biogeographic scales, and suggest that rates of disease prevalence in wild plant populations may be tightly linked to the distribution of host resistance genes. Tests for local adaptation of the fungus revealed evidence of the phenomenon (significantly greater infection in sympatric plant-fungal pairings) as well as the potential for substantial bias to be introduced into statistical analyses by spatial patterns of host resistance structure.     

普通小麦与钩刺山羊草杂交F1的农艺性状及细胞遗传学的观察

为有效地利用钩刺山羊草（Ａｅｇｉｌｏｐｓ ｔｒｉｕｎｃｉａｌｉｓ Ｌ．）的抗白粉病基因对小麦（Ｔｒｉｔｉｃｕｍ ａｅｓｔｉｖｕｍ Ｌ．）进行遗传改良,了解两者杂交后杂种Ｆ１的遗传机制是十分必要的。对Ｆ１杂种的研究表明,二价体频率高于理论值,是分别存在于钩刺山羊草Ｃ和Ｕ基因组中的小麦５Ｂ染色体上Ｐｈ基因抑制因子联合作用的结果。以尾状山羊草（Ａｅｇｉｌｏｐｓ ｃａｕｄａｔａ Ｌ．）Ｃ基因组特异重复序列     

Mutations in CAX1 produce phenotypes characteristic of plants tolerant to serpentine soils

Plant tolerance of serpentine soils is potentially an excellent model for studying the genetics of adaptive variation in natural populations. A large-scale viability screen of Arabidopsis thaliana mutants on a defined nutrient solution with a low Ca(2+) : Mg(2+) ratio (1 : 24 mol : mol), typical of serpentine soils, yielded survivors with null alleles of the tonoplast calcium-proton antiporter CAX1. cax1 mutants have most of the phenotypes associated with tolerance to serpentine soils, including survival in solutions with a low Ca(2+) : Mg(2+) ratio; requirement for a high concentration of Mg(2+) for maximum growth; reduced leaf tissue concentration of Mg(2+); and poor growth performance on 'normal' levels of Ca(2+) and Mg(2+). A physiological model is proposed to explain how loss-of-function cax1 mutations could produce all these phenotypes characteristic of plants adapted to serpentine soils, why 'normal' plants are unable to survive on serpentine soil, and why serpentine-adapted plants are unable to compete on 'normal' soils.     

Relative growth rate and biomass allocation in 20 Aegilops (Poaceae) species

This paper analyses relationships between relative growth rate ( rgr ), seed mass, biomass allocation, photosynthetic rate and other plant traits as well as habitat factors (rainfall and altitude) in 20 wild species of Aegilops L. and one closely related species of Amblyopyrum (Jaub. & Spach) Eig., which differ in ploidy level (diploid, tetraploid and hexaploid). The plants were grown hydroponically for 20 d in a growth chamber. The relationships between parameters were calculated either using the phylogenetic information (phylogenetically independent contrasts, PIC) or without using the phylogenetic information (trait values of taxa, TIP). The results using the two approaches were very similar, but there were a few exceptions in which the results were different (e.g. rgr vs. seed mass). Specific leaf area ( sla ) was positively correlated with leaf area ratio ( lar ) and negatively correlated with net assimilation rate ( nar ), which together resulted in the absence of a correlation between sla and rgr . Leaf photosynthetic rates (expressed on a mass or area basis) showed no correlation with rgr . rgr was positively correlated with the stem mass ratio and negatively with root mass ratio. Species with a lower d. wt percentage have a higher rgr . Aegilops species from locations with higher annual rainfall invested less biomass in roots and more in shoots (leaves and stems) and had a higher rgr . Diploid species had a lower seed mass and initial mass than the hybrids (tetraploid and hexaploid species), but there was no correlation of rgr with ploidy level. Polyploid species, which have higher seed mass, occur at a higher altitude than diploid species. Our results show that variation in rgr in Aegilops and Amblyopyrum spp. is associated mainly with variation in biomass allocation (proportion of biomass in stems and roots) and d. wt percentage, and not with variation in sla , leaf photosynthetic rates or seed mass.     

Aegilops (Poaceae) in Italy: taxonomy,geographical distribution,ecology, vulnerability and conservation

ABSTRACT

Aegilops L. includes wild species from which, over millennia, man has cultivated forms of Triticum L. Ten species of Aegilops occur in Italy. Three species are allochthonous and eight out of ten species are recorded in the Apulia region. Five out of the ten species have been included in Red Lists. Each taxon is presented and discussed, citing old and new sites of occurrence, by examining specimens from many different herbaria, and describing their ecology and habitats, according to the Directive 92/43 EEC. A new taxonomic key, for the identification of all Aegilops species growing in Italy, is provided. The occurrence of Aegilops caudata L., A. peregrina (Hack. in J. Fraser) Maire & Weiller and A. speltoides Tausch in Italy is doubtful.     

Retroelement insertional polymorphisms, diversity and phylogeography within diploid, D-genome Aegilops tauschii (Triticeae, Poaceae) sub-taxa in Iran

Background and Aims: The diploid goat grass Aegilops tauschii (2n = 2x = 14) is nativeto the Middle East and is the D-genome donor to hexaploid breadwheat. The aim of this study was to measure the diversity ofdifferent subspecies and varieties of wild Ae. tauschii collectedacross the major areas where it grows in Iran and to examinepatterns of diversity related to the taxa and geography. Methods: Inter-retroelement amplified polymorphism (IRAP) markers wereused to analyse the biodiversity of DNA from 57 accessions ofAe. tauschii from northern and central Iran, and two hexaploidwheats. Key Results: Eight IRAP primer combinations amplified a total of 171 distinctDNA fragments between 180 and 3200 bp long from the accessions,of which 169 were polymorphic. On average, about eight fragmentswere amplified with each primer combination, with more bandsbeing amplified from accessions from the north-west of the countrythan from other accessions. Conclusions: The IRAP markers showed high levels of genetic diversity. Analysisof all accessions together did not allow the allocation of individualsto taxa based on morphology, but showed a tendency to put accessionsfrom the north-west apart from others regions. It is speculatedthat this could be due to different activity of retroelementsin the different regions. Within the two taxa with most accessions,there was a range of IRAP genotypes that could be correlatedclosely with geographical origin. This supports suggestionsthat the centre of origin of the species is towards the south-eastof the Caspian Sea. IRAP is an appropriate marker system toevaluate genetic diversity and evolutionary relationships withinthe taxa, but it is too variable to define the taxa themselves,where more slowly evolving morphological, DNA sequence or chromosomalmakers may be more appropriate.     

Mitochondrial DNA analysis reveals cryptic large-scale invasion of non-native genotypes of common carp (Cyprinus carpio) in Japan

Wild common carp ( Cyprinus carpio ) are probably suffering from biological invasions of conspecific domesticated strains. However, such invasions may be largely camouflaged by morphological similarities between introduced and native strains. We conducted a large survey of mitochondrial DNA sequences (complete D-loop region) from 11 localities in Japan. From a total of 166 individuals, 28 haplotypes were determined to fit into six divergent clades. One of the six clades included 19 closely related haplotypes with moderate nucleotide differences; however, the remaining five clades each included either a single haplotype or two almost identical haplotypes. Phylogenetic analysis together with the previously published Eurasian haplotypes further demonstrated that the 'monotypic' clades were sisters to various Eurasian lineages, whereas the 19 related haplotypes formed a monophyletic group apart from the whole Eurasian clade. Given their monophyly and genetic diversity, the 19 related haplotypes were thought to originate from the Japanese native strain. Conversely, their phylogenetic affinities to Eurasian lineages and unnaturally low genetic diversities caused the haplotypes of the five monotypic clades to be considered as domesticated strains introduced from Eurasia. These hypotheses were supported by further evidences; i.e. the probable non-native haplotypes were frequently found from Japanese domesticated strains, and the probable native population structure was rescued when the probable non-native haplotypes were excluded from the analyses. This study revealed that almost half or more of the haplotypes in all of the locations studied originated from domesticated strains introduced from Eurasia.     

The invasion route for an insect pest species: the tobacco aphid in the New World

Biological invasions are rapid evolutionary events in which populations are usually subject to a founder event during introduction followed by rapid adaptation to the new environment. Molecular tools and Bayesian approaches have shown their utility in exploring different evolutionary scenarios regarding the invasion routes of introduced species. We examined the situation for the tobacco aphid, Myzus persicae nicotianae, a recently introduced aphid species in Chile. Using seven microsatellite loci and approximate Bayesian computation, we studied populations of the tobacco aphid sampled from several American and European countries, identifying the most likely source populations and tracking the route of introduction to Chile. Our population genetic data are consistent with available historical information, pointing to an introduction route of the tobacco aphid from Europe and/or from other putative populations (e.g. Asia) with subsequent introduction through North America to South America. Evidence of multiple introductions to North America from different genetic pools, with successive loss of genetic diversity from Europe towards North America and a strong bottleneck during the southward introduction to South America, was also found. Additionally, we examined the special case of a widespread multilocus genotype that was found in all American countries examined. This case provides further evidence for the existence of highly successful genotypes or 'superclones' in asexually reproducing organisms.     

Genetic variation and local adaptation at a cheatgrass (Bromus tectorum) invasion edge in western Nevada

Cheatgrass ( Bromus tectorum ) is an invasive weed in western North America found primarily growing at elevations less than 2200 m. We asked whether cheatgrass is capable of becoming adapted to a marginal habitat, by investigating a population at a high elevation invasion edge. We used a combination of methods, including reciprocal field transplants, controlled environment studies and molecular analysis. High levels of SSR gene diversity (0.50 vs. 0.43) and comparable variation in phenotypic traits were observed at both the invasion edge and a low elevation, high-density population. Three heterozygotes were observed in the edge population, which is unusual in this predominantly self-pollinating plant. Plants from high elevations germinated more slowly in a growth chamber and had slower seedling growth rates. Survivorship was low at the edge (13%), compared with the low elevation site (55%), but surviving plants were of similar size and had equivalent reproductive output. Seed size positively affected survival and plant performance in the field and this trait was inherited. Emergence timing affected survival at the low elevation site and germination timing was also inherited. Local adaptation was seen in the low, rather than in the high elevation site, because of differential survival. While there was no evidence for local adaptation to the high elevation site observed in the field, family level and genotype-level differences in traits that affected field performance, high genetic diversity at the invasion edge, and evidence of outcrossing in this highly selfing species indicates that the potential for adaptation to a marginal habitat exists within this population.     

Potential limits to the benefits of admixture during biological invasion

Species introductions often bring together genetically divergent source populations, resulting in genetic admixture. This geographic reshuffling of diversity has the potential to generate favourable new genetic combinations, facilitating the establishment and invasive spread of introduced populations. Observational support for the superior performance of admixed introductions has been mixed, however, and the broad importance of admixture to invasion questioned. Under most underlying mechanisms, admixture's benefits should be expected to increase with greater divergence among and lower genetic diversity within source populations, though these effects have not been quantified in invaders. We experimentally crossed source populations differing in divergence in the invasive plant Centaurea solstitialis. Crosses resulted in many positive (heterotic) interactions, but fitness benefits declined and were ultimately negative at high source divergence, with patterns suggesting cytonuclear epistasis. We explored the literature to assess whether such negative epistatic interactions might be impeding admixture at high source population divergence. Admixed introductions reported for plants came from sources with a wide range of genetic variation, but were disproportionately absent where there was high genetic divergence among native populations. We conclude that while admixture is common in species introductions and often happens under conditions expected to be beneficial to invaders, these conditions may be constrained by predictable negative genetic interactions, potentially explaining conflicting evidence for admixture's benefits to invasion.     

QTL mapping for a trade-off between leaf and bud production in a recombinant inbred population of Microseris douglasii and M. bigelovii (Asteraceae, Lactuceae): a potential preadaptation for the colonization of serpentine soils

The different response to growth on serpentine soil is a major autecological difference between the annual asteracean species Microseris douglasii and M. bigelovii, with nearly non-overlapping distribution ranges in California. Early flowering and seed set is regarded as a crucial character contributing to escape drought and thus is strongly correlated with survival and reproductive success on serpentine as naturally toxic soil. M. bigelovii (strain C94) from non-serpentine soil produces more leaves at the expense of bud production in the first growing phase than M. douglasii (B14) from serpentine soil. A QTL mapping study for this trade-off and for other growth-related traits was performed after six generations of inbreeding (F7) from a single interspecific hybrid between B14 and C94 on plants that were grown on serpentine and alternatively on normal potting soil. The trade-off is mainly correlated with markers on one map region on linkage group 03a (lg03a) with major phenotypic effects (phenotypic variance explained [PVE] = 18.8 - 31.7 %). Plants with the M. douglasii allele in QTL-B1 (QTL-NL1) produce more buds but fewer leaves in the first 119 days on both soil types. Three modifier QTL could be mapped for bud and leaf production. In one modifier (QTL-B2 = QTL-NL4) the M. douglasii allele is again associated with more buds but fewer leaves. QTL mapped for bud set in the F6 co-localize with QTL-B1 (major QTL) and QTL-B3. Two additional QTL for leaf length and red coloration of leaves could be mapped to one map region on lg03a. Co-localization of the two QTL loci with major phenotypic effects on bud and leaf production strongly suggests that a major genetic locus controls the trade-off between the two adaptive traits. The importance of mutational changes in major genes for the adaptation to stressful environments is discussed.     

Genomewide analysis of admixture and adaptation in the Africanized honeybee

Genetic exchange by hybridization or admixture can make an important contribution to evolution, and introgression of favourable alleles can facilitate adaptation to new environments. A small number of honeybees (Apis mellifera) with African ancestry were introduced to Brazil ~60 years ago, which dispersed and hybridized with existing managed populations of European origin, quickly spreading across much of the Americas in an example of a massive biological invasion. Here, we analyse whole‐genome sequences of 32 Africanized honeybees sampled from throughout Brazil to study the effect of this process on genome diversity. By comparison with ancestral populations from Europe and Africa, we infer that these samples have 84% African ancestry, with the remainder from western European populations. However, this proportion varies across the genome and we identify signals of positive selection in regions with high European ancestry proportions. These observations are largely driven by one large gene‐rich 1.4‐Mbp segment on chromosome 11 where European haplotypes are present at a significantly elevated frequency and likely confer an adaptive advantage in the Africanized honeybee population. This region has previously been implicated in reproductive traits and foraging behaviour in worker bees. Finally, by analysing the distribution of ancestry tract lengths in the context of the known time of the admixture event, we are able to infer an average generation time of 2.0 years. Our analysis highlights the processes by which populations of mixed genetic ancestry form and adapt to new environments.     

互花米草黄酮含量分析及其生态学意义

互花米草原产于北美洲大西洋沿岸和墨西哥海湾,经引种和其他原因,已扩散到包括中国在内的多个国家和地区,并对被入侵地生态系统的结构和功能均造成了显著影响。有关该物种的入侵机制和控制策略是当前入侵生态学研究领域的热点。互花米草生物质材料中富含黄酮类物质,与该物种向海扩张的能力可能存在一定联系。鉴于黄酮广泛的应用价值,能否借助黄酮成分的开发带动生物质的收割,以达到有效控制该物种的目的,也是当前研究者所关注的问题。以芦丁为标准参照物,NaNO2-Al（ NO3）3法比较温室互花米草不同器官中总黄酮含量的差异,分析盐城滩涂互花米草越冬芽黄酮含量与所在地理位置之间的关系,采用L9（34）正交试验对干储互花米草叶总黄酮的提取工艺进行优化。互花米草叶片中黄酮类物质远高于根、根状茎、茎、叶鞘、颖稃及种子。盐城滩涂上互花米草越冬芽的黄酮含量随着由陆向海方向基本呈升高趋势。互花米草总黄酮最佳提取工艺为料液比1∶30,乙醇浓度70％,水浴温度80℃,水浴时间2h。互花米草植物材料中,尤其在叶部,黄酮含量十分丰富。该黄酮物质与互花米草在海岸带盐沼生态系统中的适应和向海扩张也存在正相关关系,可作为探索互花米草入侵机制的一个方向。利用本研究中所得优化提取工艺可从互花米草干储生物质中获得稳定的黄酮来源。配合黄酮经济成分的开发而推动生物质材料的收割利用,将成为互花米草生态控制的一项潜在策略。