Covariance of floral and vegetative traits in four species of Ranunculaceae: a comparison between specialized and generalized pollination systems

Theory predicts that tighter correlation between floral traits and weaker relationship between floral and vegetative traits more likely occur in specialized flowers than generalized flowers,favoring by precise fit with pollinators.However,traits and trait correlations frequently vary under different environments.Through detecting spatiotemporal variation in phenotypic traits (floral organ size and vegetative size) and trait correlations in four Ranunculaceae species,we examined four predictions.Overall,our results supported these predictions to a certain degree.The mean coefficient of variation (CV) of floral traits in two specialized species (Delphinium kamaonense and Aconitum gymnandrum) was marginally significantly lower than that of another two generalized species (Trollius ranunculoides and Anemone obtusiloba).The two specialized species also showed marginally significantly smaller CV in floral traits than vegetative size across the two species.The absolute mean correlation between floral and vegetative traits,or that between floral traits in species with specialized flowers was not significantly lower,or higher than that in generalized plants,weakly supporting the predictions.Furthermore,we documented a large variation in trait correlations of four species among different seasons and populations.Study of covariance of floral and vegetative traits will benefit from the contrast of results obtained from generalized and specialized pollination systems.     

Strategies for defining traits when calculating economic values for livestock breeding: a review

The objective of the present review was (i) to survey different approaches for choosing the complex of traits for which economic values (EVs) are calculated, (ii) to call attention to the proper definition of traits and (iii) to discuss the manner and extent to which relationships among traits have been considered in the calculation of EVs. For this purpose, papers dealing with the estimation of EVs of traits in livestock were reviewed. The most important reasons for incompatibility of EVs for similar traits estimated in different countries and by different authors were found to be inconsistencies in trait definitions and in assumptions being made about relationships among traits. An important problem identified was how to choose the most appropriate criterion to characterise production or functional ability for a particular class of animals. Accordingly, the review covered the following three topics: (i) which trait(s) would best characterise the growth ability of an animal; (ii) how to define traits expressed repeatedly in subsequent reproductive cycles of breeding females and (iii) how to deal with traits that differ in average value between sexes or among animal groups. Various approaches that have been used to solve these problems were discussed. Furthermore, the manner in which diverse authors chose one or more traits from a group of alternatives for describing a specific biological potential were reviewed and commented on. The consequences of including or excluding relationships among economically important traits when estimating the EV for a specific trait were also examined. An important conclusion of the review is that, for a better comparability and interpretability of estimated EVs in the literature, it is desirable that clear and unique definitions of the traits, complete information on assumptions used in analytical models and details on inter-relationships between traits are documented. Furthermore, the method and the model used for the genetic evaluation of specific traits in a certain breeding organisation are important for the exact definition of traits, for which the economic values will be calculated, and for the inclusion or exclusion of relationships among traits in the calculation of the EVs in livestock breeding.     

Phenotypic variation and fluctuating asymmetry in sexually dimorphic feather ornaments in relation to sex and mating system

Secondary sexual characters have been hypothesized to demonstrate increased phenotypic variation between and within individuals as compared to ordinary morphological traits. We tested whether this was the case by studying phenotypic variation, expressed as the coefficient of variation (CV), and developmental instability, measured as fluctuating asymmetry (FA), in ornamental and non-ornamental traits of 70 bird species with feather ornamentation while controlling for similarity among species due to common descent. Secondary sexual characters differed from ordinary morphological traits by showing large phenotypic CV and FA. This difference can be explained by the different mode of selection operating on each kind of trait: a history of intense directional (ornaments) and stabilizing selection (non-ornaments). Phenotypic variation is reduced in the sex with more intense sexual selection (males), but does not differ among species with different mating systems. The strength of stabilizing selection arising from natural selection is associated with decreased CV (wing CV is smaller than tarsus or tail CVs). We found evidence of FA being reduced in ornamental feathers strongly affected by aerodynamics (tail feathers) compared to other ornaments, but only in females. In conclusion, CV and FA were not related, suggesting mat phenotypic plasticity and developmental instability are independent components of phenotypic variation.     

Hybridization, developmental stability, and functionality of morphological traits in the ground beetle Carabus solieri (Coleoptera, Carabidae)

The assessment of developmental stability in hybrids can provide valuable information in the study of species formation because it allows an evaluation of the degree of incompatibility of genetic systems that control developmental processes. The present study assessed the impact of two hybridization events, assumed to have occurred at different times, on developmental instability in the ground beetle Carabus solieri . Developmental instability was estimated in 678 individuals from 27 populations from the fluctuating asymmetry (FA) levels of four morphological traits: the tibia length of middle and hind legs, which are functional structures, and the length and the proximal width of the hind wings, which are vestigial and thus nonfunctional structures. Significant variations of FA levels between populations were shown only for the wing width. For this trait, FA levels in hybrids were higher than in their parental entities for both hybridization events, indicating a significant divergence of the gene systems controlling development between the parental entities in the two hybridization cases. As expected, wing traits exhibited FA levels at least three times higher than leg trait. Finally, the potential interest of vestigial traits in the particular context of hybridization is discussed.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 151–158.     

Individual differences in developmental precision and fluctuating asymmetry: a model and its implications

In many studies, fluctuating asymmetry (FA) has been used as a measure of individual differences in developmental imprecision. A model of how variation in developmental imprecision is associated with variation in asymmetry is described and applied to important issues about FA. If individual differences in developmental imprecision exist, asymmetry due to developmental error should be leptokurtically distributed. Moreover, the greater the magnitude of individual differences, the greater the leptokurtosis. Asymmetry purportedly due to developmental error in a variety of species is indeed leptokurtically distributed. The level of leptokurtosis suggests that the CV in individual differences in underlying developmental imprecision is generally 20–25, consistent with it being a fitness trait. In addition, data suggest that: (1) the individual differences that underlie the developmental imprecision of different traits are largely shared across traits and not trait-specific; (2) the heritability of these individual differences may average between 35 and 55%, despite small heritabilities of individual trait FAs; and (3) correlations between FA and fitness traits or components suggest high correlations between underlying variation in developmental precision and fitness in many species. Theoretical implications are discussed.     

Revisiting the Holy Grail: using plant functional traits to understand ecological processes

One of ecology's grand challenges is developing general rules to explain and predict highly complex systems. Understanding and predicting ecological processes from species' traits has been considered a ‘Holy Grail’ in ecology. Plant functional traits are increasingly being used to develop mechanistic models that can predict how ecological communities will respond to abiotic and biotic perturbations and how species will affect ecosystem function and services in a rapidly changing world; however, significant challenges remain. In this review, we highlight recent work and outstanding questions in three areas: (i) selecting relevant traits; (ii) describing intraspecific trait variation and incorporating this variation into models; and (iii) scaling trait data to community‐ and ecosystem‐level processes. Over the past decade, there have been significant advances in the characterization of plant strategies based on traits and trait relationships, and the integration of traits into multivariate indices and models of community and ecosystem function. However, the utility of trait‐based approaches in ecology will benefit from efforts that demonstrate how these traits and indices influence organismal, community, and ecosystem processes across vegetation types, which may be achieved through meta‐analysis and enhancement of trait databases. Additionally, intraspecific trait variation and species interactions need to be incorporated into predictive models using tools such as Bayesian hierarchical modelling. Finally, existing models linking traits to community and ecosystem processes need to be empirically tested for their applicability to be realized.     

Effects of Region and Elevation on Adaptation of Leaf Functional Traits of an Invasive Plant <i>Erigeron annuus</i> in China

A key scientific challenge relating to the threat of invasive plants on agriculture at the region level is to understand their adaptation and evolution in functional traits. Leaf functional traits, related to growth and resource utilization, might lead to adaptation of invasive plants to the geographical barriers (region or elevation). In the field experiment, we discussed the effects of region and elevation on leaf functional traits on invasive plant Erigeron annuus in farmland habitats in China. We compared leaf size, coefficient of variation (CV) of leaf traits, and fluctuating asymmetry (FA) of E. annuus from three regions (east vs. center vs. west) and two leaf types (vegetative vs. reproductive leaf), and from nine elevations (980–2100 m) in the west region of China. Our results indicated region and leaf type influenced leaf functional traits, and leaf size was significantly higher and CV of leaf traits and FA in reproductive leaves were significantly lower in the east region than in the west and center regions. Elevation and leaf type affected leaf functional traits, and leaf size was significantly higher and CV of leaf traits in reproductive leaves were significantly lower in moderate elevation. E. annuus has higher leaf size and developmental stability (lower CV and FA) in the eastern region due to the longer adaptation period. Therefore, leaf functional traits play an important role in the adaptation of different longitudes and elevations. It can also facilitate the understanding of the invasiveness and adaptation of leaf traits of invasive plants in the agricultural ecosystem during their spread process in China.     

Fluctuating asymmetry,fecundity and development time in Drosophila: is there an association under optimal and stress conditions?

A number of studies have reported a significant negative association between fluctuating asymmetry (FA) of bilateral morphological traits and individual fitness traits, but almost all of these are unreplicated and based on small sample sizes using single trait estimates of FA. We therefore tested if there was a relationship between the FA of five bilateral traits and fecundity and development time in Drosophila in a multiple replicated experimental design. Stressed treatments were included to increase the variability of traits and to test whether associations among traits were affected by changes in the environment. Significant positive relationships were found between the size of wing characters and mean fecundity for the 5‐day period and this relationship tended to be stronger in the stress treatments. No association was found between FA and mean fecundity for any of the traits measured. Similarly, a significant positive relationship was detected between wing trait size and development time but no association was detected between trait FA and development time. There were no differences between mean fecundity or development time of extreme asymmetry phenotypes compared with modal phenotypes. These results are discussed with reference to suggestions in the literature that FA can be used to estimate individual fitness.     

Phylogenetic conservatism of functional traits in microorganisms

A central question in biology is how biodiversity influences ecosystem functioning. Underlying this is the relationship between organismal phylogeny and the presence of specific functional traits. The relationship is complicated by gene loss and convergent evolution, resulting in the polyphyletic distribution of many traits. In microorganisms, lateral gene transfer can further distort the linkage between phylogeny and the presence of specific functional traits. To identify the phylogenetic conservation of specific traits in microorganisms, we developed a new phylogenetic metric—consenTRAIT—to estimate the clade depth where organisms share a trait. We then analyzed the distribution of 89 functional traits across a broad range of Bacteria and Archaea using genotypic and phenotypic data. A total of 93% of the traits were significantly non-randomly distributed, which suggested that vertical inheritance was generally important for the phylogenetic dispersion of functional traits in microorganisms. Further, traits in microbes were associated with a continuum of trait depths (τ_D), ranging from a few deep to many shallow clades (average τ_D: 0.101–0.0011 rRNA sequence dissimilarity). Next, we demonstrated that the dispersion and the depth of clades that contain a trait is correlated with the trait''s complexity. Specifically, complex traits encoded by many genes like photosynthesis and methanogenesis were found in a few deep clusters, whereas the ability to use simple carbon substrates was highly phylogenetically dispersed. On the basis of these results, we propose a framework for predicting the phylogenetic conservatism of functional traits depending on the complexity of the trait. This framework enables predicting how variation in microbial composition may affect microbially-mediated ecosystem processes as well as linking phylogenetic and trait-based patterns of biogeography.     

Genetic analysis of drought tolerance in maize by molecular markers I. Yield components

Grain yield is a complex trait, strongly influenced by the environment: severe losses can be caused by drought, a stress common in most maize-growing areas, including temperate climatic zones. Accordingly, drought tolerance is one of the main components of yield stability, and its improvement is a major challenge to breeders. The aim of the present work was the identification, in maize genotypes adapted to temperate areas, of genomic segments responsible for the expression of drought tolerance of yield components: ear length, ear weight, kernel weight, kernel number and 50-kernel weight. A linkage analysis between the expression of these traits and molecular markers was performed on a recombinant inbred population of 142 families, obtained by repeated selfing of the F₁ between lines B73 and H99. The population, genotyped at 173 loci (RFLPs, microsatellites and AFLPs), was evaluated in well-watered and water-stressed conditions. A drought tolerance index was calculated as the ratio between the mean value of the trait in the two environments. For the traits measured, a highly positive correlation was found over the two water regimes, and more than 50% of the quantitative trait loci (QTLs) detected were the same in both; moreover, the direction of the allelic contribution was always consistent, the allele increasing the trait value being mostly from line B73. Several QTLs were common to two or more traits. For the tolerance index, however, most of the QTLs were specific for a single component and different from those controlling the basic traits; in addition, a large proportion of the alleles increasing tolerance were provided by line H99. The data suggest that drought tolerance for yield components is largely associated with genetic and physiological factors independent from those determining the traits per se. The implications of these results for developing an efficient strategy of marker-assisted selection for drought tolerance are discussed. Received: 19 October 1998 / Accepted: 28 December 1998     

粳稻品质性状间及其与植株性状和产量性状间的遗传相关

利用朱军等提出的种子性状遗传模型,采用 3×3 NCⅡ正反交设计的亲本和部分组合F2代种子,分析了品质性状糙米率、垩白粒率、垩白面积和AC间及上述品质性状与株高、穗部性状等的遗传相关,以期为粳稻育种后代选择提供指导。结果表明,精米重与糙米率存在极显著的母体加性相关;虽然控制品质性状的主要遗传效应分量与植株性状相应遗传效应分量遗传协方差不显著,但在其他相应遗传效应分量方面存在着复杂的关系。     

Synthesising the trait information of European Chironomidae (Insecta: Diptera): Towards a new database

Chironomidae are among the most conspicuous and ecological diverse group of freshwater invertebrates. They may dominate unimpacted communities in abundance and biomass, accounting for more than 50% of macroinvertebrate species in standing and flowing waters. In deep zones of eutrophic lakes and highly human-impacted streams, they are often the only family of aquatic insects remaining. In bioassessment programmes, Chironomids are often identified at the family and subfamily levels, due to difficulties in the taxonomic identification of larvae resulting from a high intrinsic morphological similarity. This may potentially result in bias as, similarly to Ephemeroptera, Trichoptera or Plecoptera, Chironomidae species, which are replaced along natural and human-impacted gradients due to differences in their ecological requirements. Recently, multiple trait-based approaches have been proposed to complement taxonomic-based assessment of streams and rivers using macroinvertebrates. However, the lack of specific trait information for Chironomidae prevents their use in the functional assessment of communities. Therefore, here, we aimed to: (1) develop a trait database for European Chironomidae genera that can be used in future bioassessment and ecological studies; (2) evaluate, by multivariate analyses, whether our new database provides additional information on Chironomidae compared to the trait information provided in the commonly used European trait database (Tachet et al., 2010); and (3) determine whether the new information on Eltonian traits (proxy to biological traits) translates the most accepted phylogenetic relationships among Chironomidae subfamilies. We gathered information on 744 species and 178 genera, for 37 traits covering 186 trait categories, and found substantial differences between our database and the commonly used European trait database. In addition, available information on traits was not always in agreement with phylogenetic relationships among subfamilies. Orthocladiinae and Chironominae which are considered sister groups in evolutionary terms actually showed confident trait relatedness based on Eltonian traits tree while the remaining relationships between subfamilies are questionable. In addition, different traits can occur in closely related taxa depending on the environmental drivers operating on their habitats. Our study reveals that the usually accepted redundancy within the Chironomidae family and subfamilies must be a product of averaging the information from finer taxonomic resolution added to the substantial lack of information for this insect group.     

Is phenotypic canalization involved in the decline of the endemic Aquilegia thalictrifolia? Rethinking relationships between fluctuating asymmetry and species conservation status

Fluctuating asymmetry (FA), the deviation from the normal symmetrical condition of a morphological trait having specific morphological symmetry, increases in response to environmental and genetic stress, is related to phenotypic plasticity and is considered a tool for monitoring a species conservation status. However, FA–stress relations are dependent on measured traits or species‐specific characteristics such as mating system and habitat. This study investigates the relationships between FA, genetic diversity, population size, density and individual fitness traits (plant height, fruit and seed set), in the endemic Aquilegia thalictrifolia, a mixed breeder that is declining, but maintaining high levels of heterozygosity. Leaf and flower FA and other traits were investigated in 10 populations of A. thalictrifolia, the whole species range. As a result, we found similar patterns of FA in leaves and flowers between populations, indicating a homogenous level of stress between populations that differed for other traits. FA and the other traits were not related, including heterozygosity. Heterozygosity was not related to individual fitness traits with the exception of plant height. In accordance with other studies, we found that the role of FA as a tool for assessing the conservation status of a species or population should be reconsidered. However, we conclude that a low level of FA should not automatically be considered an indicator of good conservation status or low level of stress, because in species that evolved in highly stable environments it may indicate a scarce ability to plastically respond to environmental changes, as a consequence of environmental and genetic canalization. Further investigation of this point is needed.     

THE ASSOCIATION BETWEEN FLUCTUATING ASYMMETRY,TRAIT VARIABILITY,TRAIT HERITABILITY,AND STRESS: A MULTIPLY REPLICATED EXPERIMENT ON COMBINED STRESSES IN DROSOPHILA MELANOGASTER

A number of hypotheses have been proposed about the association between developmental stability phenotypic variability, heritability, and environmental stress. Stress is often considered to increase both the asymmetry and phenotypic variability of bilateral traits, although this may depend on trait heritability. Empirical studies of such associations often yield inconsistent results. This may reflect the diversity of traits and conditions used or a low repeatability of any associations. To test for repeatable associations between these variables, multiply replicated experiments were undertaken on Drosophila melanogaster using a combination stress at the egg, larval and adult stages of reduced protein, ethanol in the medium, and a cold shock. Both metric and meristic traits were measured and levels of heritable variation for each trait estimated by maximum likelihood and parent-offspring regression over three generations. Trait means were reduced by stress, whereas among-individual variation increased Fluctuating asymmetry (FA) was increased by stress in some cases, but few comparisons were significant. Only one trait orbital bristle, showed consistent increases in FA. Changes in trait means, trait phenotypic variability, and developmental stability as a result of stress were not correlated. Extreme phenotypes tended to have higher levels of FA but only the results for orbital bristles were significant. All traits had low to intermediate heritabilities except orbital bristle, which showed no heritable variation. Only traits with low heritability and high levels of phenotypic variability may show consistent increases in FA under stress. Overall, the independence of phenotypic variability, plasticity, and the developmental stability of traits extend to changes in these measures under stressful conditions.     

Multiple trait multiple interval mapping of quantitative trait loci from inbred line crosses

ABSTRACT: BACKGROUND: Although many experiments have measurements on multiple traits, most studies performed the analysis of mapping of quantitative trait loci (QTL) for each trait separately using single trait analysis. Single trait analysis does not take advantage of possible genetic and environmental correlations between traits. In this paper, we propose a novel statistical method for multiple trait multiple interval mapping (MTMIM) of QTL for inbred line crosses. We also develop a novel score-based method for estimating genome-wide significance level of putative QTL effects suitable for the MTMIM model. The MTMIM method is implemented in the freely available and widely used Windows QTL Cartographer software. RESULTS: Throughout the paper, we provide compelling empirical evidences that: (1) the score-based threshold maintains proper type I error rate and tends to keep false discovery rate within an acceptable level; (2) the MTMIM method can deliver better parameter estimates and power than single trait multiple interval mapping method; (3) an analysis of Drosophila dataset illustrates how the MTMIM method can better extract information from datasets with measurements in multiple traits. CONCLUSIONS: The MTMIM method represents a convenient statistical framework to test hypotheses of pleiotropic QTL versus closely linked nonpleiotropic QTL, QTL by environment interaction, and to estimate the total genotypic variance-covariance matrix between traits and to decompose it in terms of QTL-specific variance-covariance matrices, therefore, providing more details on the genetic architecture of complex traits.     

Variation in fluctuating asymmetry levels across a Robertsonian polymorphic zone of the house mouse

Fluctuating asymmetry (FA) levels were assessed within the Barcelona Robertsonian polymorphic zone of Mus musculus domesticus as a measure of developmental instability (DI). This zone is characterized by populations with a reduced diploid number (2n = 27–39) surrounded by others with standard karyotype (2n = 40). Mice were distributed into four classes according to mean diploid number at each collecting site, and differences in their FA levels were studied. Three different FA indices were calculated on interlandmark distances from two major morphogenetic units of the mandible: the teeth and muscular areas. Three kinds of analysis were performed: trait‐by‐trait, considering the two regions separately and pooling all traits. In univariate analyses few differences were detected between classes. Nevertheless, when characters were pooled, differences in FA values were found between one Robertsonian group and standard mice, and between several Robertsonian classes. Teeth area traits showed higher FA levels in Rb I mice, i.e. those geographically close to the standard populations. However, FA levels decreased in more introgressed Robertsonian groups and showed significant differences with Rb I. Muscular area traits showed lower DI levels in the Robertsonian mice, especially those with an intermediate mean diploid number (Rb II). When all traits were pooled, FA levels increased in Rb I, whilst the other Robertsonian groups showed similar, or even lower, FA values to those of the standard mice. The higher FA levels found in Rb I suggest an increase in the disruption of genetic coadaptation. Furthermore, in Rb I the presence of metacentrics is more recent than in more introgressed populations; we therefore suggest that Rb I have had less time to restore genetic coadaptation. In addition, the teeth region of the mandible seems to be more sensitive to genomic stress than the muscular region. In the light of these results we suggest that hybrids should be separated into groups and that mandible morphogenetic units be differentiated when using FA in hybrid zones to analyse DI.     

Longitudinal analyses of correlated response efficiencies of fillet traits in Nile tilapia

Recent studies with Nile tilapia have shown divergent results regarding the possibility of selecting on morphometric measurements to promote indirect genetic gains in fillet yield (FY). The use of indirect selection for fillet traits is important as these traits are only measurable after harvesting. Random regression models are a powerful tool in association studies to identify the best time point to measure and select animals. Random regression models can also be applied in a multiple trait approach to analyze indirect response to selection, which would avoid the need to sacrifice candidate fish. Therefore, the aim of this study was to investigate the genetic relationships between several body measurements, weight and fillet traits throughout the growth period and to evaluate the possibility of indirect selection for fillet traits in Nile tilapia. Data were collected from 2042 fish and was divided into two subsets. The first subset was used to estimate genetic parameters, including the permanent environmental effect for BW and body measurements (8758 records for each body measurement, as each fish was individually weighed and measured a maximum of six times). The second subset (2042 records for each trait) was used to estimate genetic correlations and heritabilities, which enabled the calculation of correlated response efficiencies between body measurements and the fillet traits. Heritability estimates across ages ranged from 0.05 to 0.5 for height, 0.02 to 0.48 for corrected length (CL), 0.05 to 0.68 for width, 0.08 to 0.57 for fillet weight (FW) and 0.12 to 0.42 for FY. All genetic correlation estimates between body measurements and FW were positive and strong (0.64 to 0.98). The estimates of genetic correlation between body measurements and FY were positive (except for CL at some ages), but weak to moderate (−0.08 to 0.68). These estimates resulted in strong and favorable correlated response efficiencies for FW and positive, but moderate for FY. These results indicate the possibility of achieving indirect genetic gains for FW and by selecting for morphometric traits, but low efficiency for FY when compared with direct selection.     

五角枫种群表型多样性

为揭示五角枫种群的表型分化程度、变异模式及地理变异规律,以山西19个种群为研究对象,采用巢式方差分析、相关分析、聚类分析等方法从形态学角度对五角枫种群的叶片、果实、种子等23个表型性状进行了系统分析。结果表明:(1)五角枫23个表型性状中除果柄长、着生痕、种子长/宽以外,其余20个表型性状在种群间和种群内均存在显著和极显著差异。(2)19个种群的平均变异系数为18.07。叶片、果实、种子的平均表型变异系数依次为:果实19.78%叶片18.77%种子10.25%。(3)五角枫种群间表型分化系数(VST)均值为48.82%,种群内变异(51.18%)与种群间变异(48.82%)基本相当。叶片、果实、种子表型分化系数的平均值为:叶片58.08%果实41.24%种子25.87%。(4)五角枫叶片、果实和种子的信息多样性指数(H)为:叶片6.1079果实5.9118种子5.2855;多样性指数平均值(D)分别为:果实0.9967叶片0.9961种子0.9948。(5)主成分分析结果显示:五角枫种群表型多样性基本来源为:叶片贡献率果实贡献率种子贡献率。(6)五角枫表型变异呈现出以经度和纬度变异并存的趋势,少数表型性状与经度和纬度呈现显著或极显著相关。(7)利用欧氏距离对五角枫种群进行UPGMA聚类分析,将五角枫19个种群划分为两大类群。五角枫种群具有较高的表型多样性,种群间和种群内均存在丰富的表型变异,与其遗传特点和分布生境等密切相关,研究结果为今后五角枫种质资源的保护和利用奠定了基础。     

THE FITNESS OF MANIPULATING PHENOTYPES: IMPLICATIONS FOR STUDIES OF FLUCTUATING ASYMMETRY AND MULTIVARIATE SELECTION

Phenotypic manipulation (or phenotypic engineering) that alters trait distributions provides a way to increase the statistical power of detecting relationships between traits and fitness. Manipulations relying on plastic responses, however, assume a specific relationship between the perturbation and the alteration of the traits when multiple traits are involved. We measured several traits, including condition measured as fluctuating asymmetry, in the ladybird beetle Harmonia axyridis under six different diets to examine how altered environments affected multiple traits and their distributions. Although diet affected fluctuating asymmetry, we found no consistent relationship between degree of asymmetry and other phenotypic measures. As expected, individual traits were altered by our treatments. Contrary to expectation, relationships among traits were not constant among diets. Our results suggest that assumptions about the relationship between condition and trait values, especially fluctuating asymmetry, cannot be made. Further, studies that use manipulated phenotypes to statistically determine the form of selection must first demonstrate that the pattern of the phenotypic correlation matrix is not itself altered by the manipulation. If the phenotypic correlation matrix is not constant, then experimental estimates of selection coefficients may not reflect selection that occurs in the wild.