Rapid response to artificial selection on flower size in Phlox

Quantitative characters are often said to evolve rather slowly, taking many generations to exhibit appreciable differences among populations. We tested this notion experimentally by performing bi-directional selection on corolla diameter of plants from a wild population of Phlox drummondii for three generations. By monitoring flower size, tube length and stigma-anther proximity of flowers, we obtained the direct and indirect responses to selection, and calculated genetic correlations, realized and narrow sense heritabilities using offspring-mother regression. Realized heritability of flower size was high (0.83), whereas genetic correlations among traits were weak or not significant. The per-generation average of the response in corolla diameter was about 5%. We found that P. drummondii has a great capacity to respond rapidly to selection, and this capacity may be in part responsible for the observed high degree of differentiation within the species. We also concluded that rapid evolution of morphological floral traits is possible.     

HERITABILITY AND GENETIC CORRELATION OF COROLLA SHAPE AND SIZE IN ERYSIMUM MEDIOHISPANICUM

Flower shape has evolved in most plants as a consequence of pollinator-mediated selection. Unfortunately, no study has explored the genetic variation of flower shape, despite that this information is crucial to understand its adaptive evolution. Our main goal here is to determine heritability of corolla shape in Erysimum mediohispanicum (Brassicaceae). Also, we explore heritability of other pollinator-selected traits in this plant species, such as plant size, flower display, and corolla size. In addition, we investigate genetic correlations between all these traits. We found significant heritability for one plant-size trait (stalk height), for number of flowers, for all corolla-size traits (corolla diameter, corolla tube length and corolla tube width), and for corolla shape. Consequently, this species retains a high ability to respond to the selection exerted by its pollinators. Genetic correlation was strong between all functionally related traits and between flower number and plant size, weak between corolla size and plant size and no correlation between corolla shape and any other trait. Thus, selection affecting some E. mediohispanicum traits would also indirectly affect other functionally related and unrelated traits. More importantly, the observed genetic correlation seems to be at least partially adaptive because positive correlational selection currently acts on the covariance between some of these traits ( Gómez 2003 ; Gómez et al. 2006 ).     

GENETIC AND ENVIRONMENTAL VARIATION IN LIFE-HISTORY TRAITS OF A MONOCARPIC PERENNIAL: A DECADE-LONG FIELD EXPERIMENT

Directional and stabilizing selection tend to deplete additive genetic variance. On the other hand, genetic variance in traits related to fitness could be retained through polygenic mutation, spatially varying selection, genotype-environment interaction, or antagonistic pleiotropy. Most estimates of genetic variance in fitness-related traits have come from laboratory studies, with few estimates of heritability made under natural conditions, particularly for longer lived organisms. Here I estimated additive genetic variance in life-history characters of a monocarpic herb, Ipomopsis aggregata, that lives for up to a decade. Experimental crosses yielded 229 full-sibships nested within 32 paternal half-sibships. More than 5000 offspring were planted as seeds into natural field sites and were followed in most cases through their entire life cycle. Survival showed substantial additive genetic variance (genetic coefficient of variation ≈ 54%). Small differences at seedling emergence were magnified over time, such that the genetic variability in survival was only detectable by tracking the success of offspring for several years starting from seed. In contrast to survival, reproductive traits such as flower number, seeds per flower, and age at flowering showed little or no genetic variability. Despite relatively high levels of additive genetic variation for some life-history characters, high environmental variance in survival resulted in very low heritabilities (0–9%) for all of these characters. Maternal effects were evident in seed mass and remained strong throughout the lengthy vegetative period. No negative genetic correlations between major components of female fitness were detected. Mean corolla width for a paternal family was, however, negatively correlated with the finite rate of increase based on female fitness. That negative correlation could help to maintain additive genetic variance in the face of strong selection through male function for wide corollas.     

Heritability of phenotypic udder traits to improve resilience to mastitis in Texel ewes

There are no estimates of the heritability of phenotypic udder traits in suckler sheep, which produce meat lambs, and whether these are associated with resilience to mastitis. Mastitis is a common disease which damages the mammary gland and reduces productivity. The aims of this study were to investigate the feasibility of collecting udder phenotypes, their heritability and their association with mastitis in suckler ewes. Udder and teat conformation, teat lesions, intramammary masses (IMM) and litter size were recorded from 10 Texel flocks in Great Britain between 2012 and 2014; 968 records were collected. Pedigree data were obtained from an online pedigree recording system. Univariate quantitative genetic parameters were estimated using animal and sire models. Linear mixed models were used to analyse continuous traits and generalised linear mixed models were used to analyse binary traits. Continuous traits had higher heritabilities than binary with teat placement and teat length heritability (h²) highest at 0.35 (SD 0.04) and 0.42 (SD 0.04), respectively. Udder width, drop and separation heritabilities were lower and varied with udder volume. The heritabilities of IMM and teat lesions (sire model) were 0.18 (SD 0.12) and 0.17 (SD 0.11), respectively. All heritabilities were sufficiently high to be in a selection programme to increase resilience to mastitis in the population of Texel sheep. Further studies are required to investigate genetic relationships between traits and to determine whether udder traits predict IMM, and the potential benefits from including traits in a selection programme to increase resilience to chronic mastitis.     

Egg shell quality in Japanese quail: characteristics,heritabilities and genetic and phenotypic relationships

The objective of the present study was to estimate heritabilities as well as genetic and phenotypic correlations for egg weight, specific gravity, shape index, shell ratio, egg shell strength, egg length, egg width and shell weight in Japanese quail eggs. External egg quality traits were measured on 5864 eggs of 934 female quails from a dam line selected for two generations. Within the Bayesian framework, using Gibbs Sampling algorithm, a multivariate animal model was applied to estimate heritabilities and genetic correlations for external egg quality traits. The heritability estimates for external egg quality traits were moderate to high and ranged from 0.29 to 0.81. The heritability estimates for egg and shell weight of 0.81 and 0.76 were fairly high. The genetic and phenotypic correlations between egg shell strength with specific gravity, shell ratio and shell weight ranging from 0.55 to 0.79 were relatively high. It can be concluded that it is possible to determine egg shell quality using the egg specific gravity values utilizing its high heritability and fairly high positive correlation with most of the egg shell quality traits. As a result, egg specific gravity may be the choice of selection criterion rather than other external egg traits for genetic improvement of egg shell quality in Japanese quails.     

GENOTYPE-BY-ENVIRONMENT INTERACTIONS IN THE DETERMINATION OF THE SIZE OF A SECONDARY SEXUAL CHARACTER IN THE COLLARED FLYCATCHER (FICEDULA ALBICOLLIS)

Although genetic variation in characters closely related to fitness is expected to either become depleted by selection or masked by environmental variation, “good gene” models of sexual selection require moderate to high heritabilities of secondary sexual characters to explain the occurrence of costly female mate preferences. In this study, I investigated whether the estimated heritability of a condition-dependent secondary sexual character (i.e., the white forehead badge) in the collared flycatcher varied depending on environmental conditions experienced during offspring growth. The data were collected over a period of 14 years making it possible to exploit natural variation in natal conditions. In addition, natal conditions were experimentally altered through brood size manipulations. During unfavorable conditions caused by generally poor weather or experimentally enlarged brood size, no significant heritability based on father-sons regressions could be demonstrated (0.19 ? h² ? 0.27). In contrast, sons reared during years with favorable weather or in experimentally reduced broods significantly resembled their fathers (0.44 ? h² ? 0.65). In addition, the heritability estimates declined with increasing maternal age. The strong effect of natal environmental condition on the estimated heritability of forehead badge size suggests that the potential genetic benefit from mate choice vary according to environmental conditions (e.g., the benefit is reduced during unfavorable rearing conditions). Because sons reared during poor conditions have probably experienced a natal environment different from that experienced by their fathers, the low heritability estimates obtained under poor conditions seem to be caused by low additive genetic variation expressed in such environments and/or a low genetic correlation between the expression of the trait in the two different environments (i.e., good vs. bad). Both of these explanations imply the presence of genotype-by-environment interactions. If such interactions frequently affect the expression of secondary sexual characters, this may offer an explanation of the high heritabilites sometimes reported for such traits, despite their exposure to long-term directional selection.     

HERITABILITIES OF DOMINANCE-RELATED TRAITS IN MALE BANK VOLES (CLETHRIONOMYS GLAREOLUS)

A number of studies have shown that in several animal species females prefer dominant males as mating partners, but fewer attempts have been made to measure possible indirect benefits of this choice. One reason for this may be that, even though dominance is a widely used concept, the definition of dominance still remains controversial Furthermore, defining and measuring the heritability of social behaviors is problematic because they are not individual traits but, by definition, involve interactions between at least two individuals. In this study we estimated heritabilities and coefficients of additive genetic variances (CV_A) for male traits that are closely associated with dominance and female mating preferences in bank voles (Clethrionomys glareolus). The heritability values were estimated using father-offspring regression. All heritability estimates were relatively high ranging from 0.531 (urine marking) to 0.767 (preputial glands). The CV_A-values indicated high levels of additive genetic variance especially in the characters most closely related to dominance: the weight of preputial glands and urine marking behavior. All phenotypic correlations among the traits measured were significantly positive and the genetic correlations were of similar magnitude as the corresponding phenotypic counterparts. Even though heritabilities may be lower in the natural environment than under controlled laboratory conditions, our results suggest that characters closely related to dominance may be at least partly genetically determined.     

Heritability of female extra-pair paternity rate in song sparrows (Melospiza melodia)

The forces driving the evolution of extra-pair reproduction in socially monogamous animals remain widely debated and unresolved. One key hypothesis is that female extra-pair reproduction evolves through indirect genetic benefits, reflecting increased additive genetic value of extra-pair offspring. Such evolution requires that a female's propensity to produce offspring that are sired by an extra-pair male is heritable. However, additive genetic variance and heritability in female extra-pair paternity (EPP) rate have not been quantified, precluding accurate estimation of the force of indirect selection. Sixteen years of comprehensive paternity and pedigree data from socially monogamous but genetically polygynandrous song sparrows (Melospiza melodia) showed significant additive genetic variance and heritability in the proportion of a female's offspring that was sired by an extra-pair male, constituting major components of the genetic architecture required for extra-pair reproduction to evolve through indirect additive genetic benefits. However, estimated heritabilities were moderately small (0.12 and 0.18 on the observed and underlying latent scales, respectively). The force of selection on extra-pair reproduction through indirect additive genetic benefits may consequently be relatively weak. However, the additive genetic variance and non-zero heritability observed in female EPP rate allow for multiple further genetic mechanisms to drive and constrain mating system evolution.     

THE GENETIC BASIS OF ZEBRA FINCH VOCALIZATIONS

Animal vocalizations play an important role in individual recognition, kin recognition, species recognition, and sexual selection. Despite much work in these fields done on birds virtually nothing is known about the heritability of vocal traits in birds. Here, we study a captive population of more than 800 zebra finches ( Taeniopygia guttata ) with regard to the quantitative genetics of call and song characteristics. We find very high heritabilities in nonlearned female call traits and considerably lower heritabilities in male call and song traits, which are learned from a tutor and hence show much greater environmental variance than innate vocalizations. In both sexes, we found significant heritabilities in several traits such as mean frequency and measures of timbre, which reflect morphological characteristics of the vocal tract. These traits also showed significant genetic correlations with body size, as well as positive genetic correlations between the sexes, supporting a scenario of honest signaling of body size through genetic pleiotropy ("index signal"). In contrast to such morphology-related voice characteristics, classical song features such as repertoire size or song length showed very low heritabilities. Hence, these traits that are often suspected to be sexually selected would hardly respond to current directional selection.     

金钟连翘与东北连翘种间杂交F1代表型性状遗传分析

为了阐明连翘属植物主要观赏性状和抗寒性的遗传特点,以金钟连翘品种‘Lynwood’(Forsythia intermedia‘Lynwood’)和东北连翘(F.mandschurica)杂交获得的F1代群体为研究对象,对杂交群体的花冠口直径、花裂片长度、抗寒性等12个表型性状进行测定,并对这些性状进行相关性分析和混合遗传模型分析。结果表明:金钟连翘与东北连翘杂交群体的表型性状变异丰富,各表型性状均出现大于高亲或小于低亲的超亲个体。除花裂片长度、花裂片宽度、叶片长度外,其他性状的变异度均超过15%,达到中等水平以上。各性状之间存在一定的相关性,其中抗寒性与着花密度呈极显著正相关,与花冠口直径、植株冠幅呈极显著负相关关系。混合遗传模型分析显示,花裂片长度、花裂片宽度、花裂片长宽比、叶片长度、叶片宽度、叶片长宽比、当年生枝条长度、抗寒性由微效多基因控制,花冠口直径和植株冠幅由一对加性—显性主基因控制,着花密度由两对加性—显性主基因控制,株高由两对等加性—显性主基因控制。花冠口直径、冠幅、着花密度和株高的主基因遗传力分别为76.05%、60.3%、72.22%和64.75%。研究结果为定向培育综合性状优良的连翘新品种提供了依据。     

籼型黑米粒形性状的遗传效应及其与矿质元素含量的遗传相关性

采用禾谷类作物种子数量性状的遗传模型,分析了灿型黑米稻品种双列杂交F1和F2种子的粒重,粒长,粒宽和粒长／粒宽等粒形性状的遗传效应及其与米粒中矿质元素Fe,Zn,Mn和P含量的遗传相关性,结果表明：4种粒形性状同时受制于种子直接遗传效应,母体效应和细胞质作用影响,其中种子直接遗传效应比母体效应和细胞质效应的作用更大,且种子直接遗传效应以加性效应占主导,粒重,粒宽和粒长／粒宽的种子直接遗传率较高,杂种早代单粒选择效果较好,粒长的种子直接遗传率和母体遗传率均属中等,较高世代的杂种进行单株选择和单粒选择均有一定效果,4种粒形性状与其米粒中矿质元素Fe,Zn,Mn和P含量表现较强的种子直接加性相关,直接显性相关,细胞质相关,母体加性相关和母体显性相关,在特种稻育种实践中,可以通过粒形性的间接选择,达到改良其矿质元素含量等营养品质性状的目标。     

Predictions of response to selection caused by angling in a wild population of Atlantic salmon (Salmo salar)

1. Recreational angling activities in wild populations of Atlantic salmon may induce a selection pressure towards a reduction in body size and length if the angling season coincides with the return of the largest sea age fish class. 2. Using estimates of heritability for growth traits and estimates of the selection pressure from angling operating on growth, we predicted the response to selection expected to occur in a wild population of Atlantic salmon. 3. The dataset used here comprised individuals from two consecutive generations (parents and offspring) from the River Bidasoa (NW Spain). Offspring were assigned to parents using six highly polymorphic microsatellite loci. Use of restricted maximum likelihood methodology and the animal model allowed us to estimate the heritability for body length and body weight as well as their genetic correlation. 4. Estimated heritabilities (0.32 ± 0.12 for length and 0.32 ± 0.11 for weight) and selection pressure caused by angling were used to obtain predictions of response to selection because of angling. Our results suggested a decline of 1.9 mm in body length and 103.3 g in body weight per generation because of angling pressure. 5. The results derived from this study suggest that the angling season should be annually delayed in order to avoid selective angling of the multi‐year class and further reductions in body weight and length.     

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.     

Heritability and correlation structure of nectar and floral morphology traits in Nicotiana alata

The heritability and genetic basis of nectar traits have been rarely studied in the field, where plants are exposed to environmental factors that could mask underlying genetic effects. Heritabilities and variance components were estimated for nectar and morphological traits of Nicotiana alata , using a partial diallel design. The main experiment was conducted in a Missouri experimental garden using a randomized block design with three plant density treatments, whereas a smaller experiment was conducted near native Brazil habitat to compare the environmental variance in traits between Missouri and Brazil. Significant heritability was detected for nectar volume and energy content, and for corolla tube length. Phenotypic correlations were significant between all traits investigated, whereas significant genetic correlations were only found between nectar volume and energy and between corolla limb width and mouth diameter. There were no significant family-by-density interactions detected in the Missouri field environment. All traits differed significantly between Missouri and Brazil environments, but significant genetic by environment (G × E) interactions between Missouri and Brazil were detected for only one trait. This study shows that nectar traits can be heritable despite considerable environmental variation.     

PATERNAL CARE: DIRECT AND INDIRECT GENETIC EFFECTS OF FATHERS ON OFFSPRING PERFORMANCE

Knowledge of how genetic effects arising from parental care influence the evolution of offspring traits comes almost exclusively from studies of maternal care. However, males provide care in some taxa, and often this care differs from females in quality or quantity. If variation in paternal care is genetically based then, like maternal care and maternal effects, paternal effects may have important consequences for the evolution of offspring traits via indirect genetic effects (IGEs). IGEs and direct–indirect genetic covariances associated with parental care can contribute substantially to total heritability and influence predictions about how traits respond to selection. It is unknown, however, if the magnitude and sign of parental effects arising from fathers are the same as those arising from mothers. We used a reciprocal cross‐fostering experiment to quantify environmental and genetic effects of paternal care on offspring performance in the burying beetle, Nicrophorus vespilloides. We found that IGEs were substantial and direct–indirect genetic covariances were negative. Combined, these patterns led to low total heritabilities for offspring performance traits. Thus, under paternal care, offspring performance traits are unlikely to evolve in response to selection, and variation in these traits will be maintained in the population despite potentially strong selection on these traits. These patterns are similar to those generated by maternal care, indicating that the genetic effects of care on offspring performance are independent of the caregiver's sex.     

QUANTITATIVE GENETICS OF SEXUAL SIZE DIMORPHISM IN THE COLLARED FLYCATCHER,FICEDULA ALBICOLLIS

Quantitative genetic theory predicts that evolution of sexual size dimorphism (SSD) will be a slow process if the genetic correlation in size between the sexes is close to unity, and the heritability of size is similar in both sexes. However, there are very few reliable estimates of genetic correlations and sex-specific heritabilities from natural populations, the reasons for this being that (1) offspring have often been sexed retrospectively, and hence, selection acting differently with respect to body size in the two sexes between measuring and sex identification can bias estimates of SSD; and (2) in many taxa, parents may be incorrectly assigned to offspring either because of assignment errors or because of extrapair paternity. We used molecular sex and paternity identification to overcome these problems and estimated sex-specific heritabilities and the genetic correlation in body size between the two sexes in the collared flycatcher, Ficedula albicollis. After exclusion of the illegitimate offspring, the genetic correlation in body size between the sexes was 1.00 (SE = 0.22), implying a severe constraint on the evolution of SSD in this species. Furthermore, sex-specific heritability estimates were very similar, indicating that neither sex will be able to evolve faster than the other. By using estimated genetic parameters, together with empirically derived estimates of sex-specific selection gradients, we further demonstrated that the predicted selection response in female tarsus length is displaced about 200% in the opposite direction from that to be expected if there were no genetic correlation between the sexes. The correspondence between the biochemically estimated rate of extrapair paternity (about 15 % of the young) and that estimated from the “heritability method” (11%) was good. However, the estimated rate of extrapair paternity with the heritability method after exclusion of the illegitimate young was 22%, adding to increasing evidence that factors other than extrapair paternity (e.g., maternal effects) may be resposible for the commonly observed higher mother-offspring than father-offspring resemblance.     

Intralocus sexual conflict and the genetic architecture of sexually dimorphic traits in Prochyliza xanthostoma (Diptera: Piophilidae)

Because homologous traits of males and females are likely to have a common genetic basis, sex-specific selection (often resulting from sexual selection on one sex) may generate an evolutionary tug-of-war known as intralocus sexual conflict, which will constrain the adaptive divergence of the sexes. Theory suggests that intralocus sexual conflict can be mitigated through reduction of the intersexual genetic correlation (rMF), predicting negative covariation between rMF and sexual dimorphism. In addition, recent work showed that selection should favor reduced expression of alleles inherited from the opposite-sex parent (intersexual inheritance) in traits subject to intralocus sexual conflict. For traits under sexual selection in males, this should be manifested either in reduced maternal heritability or, when conflict is severe, in reduced heritability through the opposite-sex parent in offspring of both sexes. However, because we do not know how far these hypothesized evolutionary responses can actually proceed, the importance of intralocus sexual conflict as a long-term constraint on adaptive evolution remains unclear. In this study, we investigated the genetic architecture of sexual and nonsexual morphological traits in Prochyliza xanthostoma. The lowest rMF and greatest dimorphism were exhibited by two sexual traits (head length and antenna length) and, among all traits, the degree of sexual dimorphism was correlated negatively with rMF. Moreover, sexual traits exhibited reduced maternal heritabilities, and the most strongly dimorphic sexual trait (antenna length) was heritable only through the same-sex parent in offspring of both sexes. Our results support theory and suggest that intralocus sexual conflict can be resolved substantially by genomic adaptation. Further work is required to identify the proximate mechanisms underlying these patterns.     

大豆粒形性状的遗传效应分析

采用双子叶植物种子数量性状的遗传模型,分析了大豆品种双列杂交F1和F2种子的粒重、粒宽、粒厚和粒长／粒宽、粒长／粒厚、粒宽／粒厚粒形性状的遗传效应。结果表明：7种粒形性状同时受制于种子直接遗传效应,而且还不同程度的受制于母体和细胞质效应。其中,百粒重、粒长、粒长／粒宽、粒长／粒厚和粒宽／粒厚的遗传以细胞质效应为主;粒宽和粒厚以母体遗传效应为主。粒重、粒长和粒长／粒宽、粒宽／粒厚的种子直接遗传率和细胞质遗传率均属中等,对其4个性状选择可以在较高世代单株和单粒选择均有效果。粒宽和粒厚母体遗传率数值较大,对其性状应以母体单株为单位早代选择,以增加粒宽和粒厚。P2和P7可作为增加百粒重、粒长／粒宽、粒长／粒厚和粒宽／粒厚的理想亲本;P1、P4和P6分别是提高粒长、粒厚和粒宽的理想亲本。     

The quantitative genetics of sexually selected traits,preferred traits and preference: a review and analysis of the data

The maintenance of variation in sexually selected traits is a puzzle that has received increasing attention in the past several decades. Traits that are related to fitness, such as life‐history or sexually selected traits, are expected to have low additive genetic variance (and hence, heritability) due to the rapid fixation of advantageous alleles. However, previous analyses have suggested that the heritabilities of sexually selected traits are on average higher than nonsexually selected traits. We show that the heritabilities of sexually selected traits are not significantly different from those of nonsexually selected traits overall or when separated into the three trait categories: behavioural, morphological and physiological. In contrast with previous findings, the heritability of preference is quite low (h² = 0.25 ± 0.06) and is in the same range as life‐history traits. We distinguish preferred traits as a category of sexually selected traits and find that the heritability of the former is not significantly different than sexually selected traits overall (0.48 ± 0.04 vs. 0.46 ± 0.03). We test the hypothesis that the heritability of sexually selected traits is negatively correlated with the strength of sexual selection. As predicted, there is a significant negative correlation between the heritabilities of sexually selected traits and the strength of selection. This suggests that heritabilities do indeed decrease as sexual selection increases but sexual selection is not strong enough to cause heritabilities of sexually selected traits to deviate from the same type of nonsexually selected traits.     

Comparing Evolvability and Variability of Quantitative Traits

There are two distinct reasons for making comparisons of genetic variation for quantitative characters. The first is to compare evolvabilities, or ability to respond to selection, and the second is to make inferences about the forces that maintain genetic variability. Measures of variation that are standardized by the trait mean, such as the additive genetic coefficient of variation, are appropriate for both purposes. Variation has usually been compared as narrow sense heritabilities, but this is almost always an inappropriate comparative measure of evolvability and variability. Coefficients of variation were calculated from 842 estimates of trait means, variances and heritabilities in the literature. Traits closely related to fitness have higher additive genetic and nongenetic variability by the coefficient of variation criterion than characters under weak selection. This is the reverse of the accepted conclusion based on comparisons of heritability. The low heritability of fitness components is best explained by their high residual variation. The high additive genetic and residual variability of fitness traits might be explained by the great number of genetic and environmental events they are affected by, or by a lack of stabilizing selection to reduce their phenotypic variance. Over one-third of the quantitative genetics papers reviewed did not report trait means or variances. Researchers should always report these statistics, so that measures of variation appropriate to a variety of situations may be calculated.