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Within-herd heritability estimated with daughter-parent regression for yield and somatic cell score.

Estimates of heritability within herd (h(WH)(2) ) that were generated with daughter-dam regression, daughter-sire regression, and REML were compared, and effects of adjusting lactation records for within-herd heritability on genetic evaluations were evaluated. Holstein records for milk, fat, and protein yields and somatic cell score (SCS) from the USDA national database represented herds in the US Northeast, Southeast, Midwest, and West. Four data subsets (457 to 499 herds) were randomly selected, and a large-herd subset included the 15 largest herds from the West and 10 largest herds from other regions. Subset heritabilities for yield and SCS were estimated assuming a regression model that included fixed covariates for effects of dam yield or SCS, sire predicted transmitting ability (PTA) for yield or SCS, herd-year-season of calving, and age within parity. Dam records and sire PTA were nested within herd as random covariates to generate within-herd heritability estimates that were regressed toward mean h(WH)(2) for the random subset. Heritabilities were estimated with REML using sire models (REML(SIRE)), sire-maternal grandsire models (REML(MGS)), and animal models (REML(ANIM)) for each herd individually in the large-herd subset. Phenotypic variance for each herd was estimated from herd residual variance after adjusting for effects of year-season and age within parity. Deviations from herd-year-season mean were standardized to constant genetic variance across herds, and records were weighted according to estimated error variance to accommodate h(WH)(2) when estimating breeding values. Mean h(WH)(2) tended to be higher with daughter-dam regression (0.35 for milk yield) than with daughter-sire regression (0.24 for milk yield). Heritability estimates varied widely across herds (0.04 to 0.67 for milk yield estimated with daughter-dam regression), and h(WH)(2) deviated from subset means more for large herds than for small herds. Correlation with REML(ANIM) h(WH)(2) was 0.68 for daughter-dam and was 0.45 for daughter-sire h(WH)(2) for milk yield. The correlation between daughter-sire h(WH)(2) and REML(MGS) was greater than the correlation between daughter-dam h(WH)(2) and REML(MGS). Data adjustments had a minimal impact on breeding value bias. Within-herd heritability can be estimated rapidly using regression techniques with moderate accuracy, but adjusting lactation records for h(WH)(2) resulted in only a small improvement in the accuracy of genetic evaluations.

Animals↗

[Study on application of the general selection index].

Several problems associated with selection strategies of breeder rams are discussed using the general selection index. Selection objective is to improve the aggregate breeding value of clean-fleece weight, number of lambs and body weight. Staple length is the assistant trait of selection. Aggregate, restrictive and optimal selection indices are constructed with the information of ram individual and the ram's half-sib sisters and half-sib daughters. The restrictive selection maintains the genetic gain of body weight unchanged. The optimal selection keeps the genetic gain of the clean-fleece weight at 0.4 kg in one generation (20% retention). The selection effects of the three indices are compared. Furthermore, two related problems are discussed. First, if each single trait of the three objective traits is selected to permit the genetic gain of the trait to be as large as possible in a short time, the largest gain of the aggregate breeding value is obtained when the trait is clean-fleece weight. Secondly, when the number of ram's relatives is more than enough, the selection effect of increasing the number is very limited. However, increasing the ram's half-sib daughters is more favorable than increasing the ram's half-sib sisters.

Animals↗

Selection for placental efficiency in swine: genetic parameters and trends.

The objectives of this study were to estimate response to divergent selection for an index of placental efficiency in swine, and to evaluate the effect of placental efficiency on litter size. The selection index (SI) included total born (TB), birth weight (BRWT), and placental weight (PW), and was designed to increase in the high line (H) or decrease in the low line (L) the efficiency of the placental function (PE), defined as the ratio BRWT:PW. (Co)variance components were estimated for direct and maternal additive effects by using an animal model with MTDFREML procedures. Estimated breeding values were calculated by using records on individual BRWT (n = 2,111), PW (n = 2,006), PE (n = 1,677), and SI (n = 1,677). Litter traits were evaluated using records on 193 litters. The model included the fixed effects of contemporary group for all traits, with the addition of sex for individual traits and parity for litter traits. Litter was fitted as an uncorrelated random effect for all traits, and TB was used as a linear and quadratic covariate for BRWT, PW, and PE. Direct heritability estimates from single-trait models were 0.03, 0.25, 0.18, 0.11, and 0.08 for BRWT, PW, PE, SI, and TB, respectively. Estimated breeding values were compared between lines by using a model including generation, line within generation, and replicate within line as the error term. Estimates of genetic divergence were 20.7 +/- 2.7 g, 0.24 +/- 0.03, 0.11 +/- 0.02, and 0.07 +/- 0.02 per generation for PW, PE, SI, and TB, respectively (P < 0.01), but divergence was not significant for BRWT. At Generation 4, direct EBV was higher in L than in H for PW (55.9 +/- 8.7 vs. -24.2 +/- 9.5 g, respectively; P < 0.01) and higher in H than in L for PE (0.58 +/- 0.10 vs. -0.35 +/- 0.09 g, respectively; P < 0.01). However, EBV was not different for BRWT, SI, or TB. These results indicate that PW and PE are susceptible to change by genetic selection; however, the correlated response in TB was an unexpected genetic trend toward a higher TB in L of 0.05 +/- 0.01 piglets per generation (P < 0.01).

Animal Husbandry↗

Impact of paternity errors in cow identification on genetic evaluations and international comparisons.

The impact of paternity identification errors on US genetic evaluations and international comparisons of Holstein dairy bulls for milk, fat, and protein yields was investigated. Sire identification was replaced for 11% of Holstein cows that were sired by AI bulls and had records in the US database for national genetic evaluations; US evaluations were computed based on those modified pedigrees and compared with official national evaluations. Estimated breeding values from the data with introduced paternity errors were biased, especially for later generations. Estimated genetic trends decreased by 11 to 15%. Estimates of standard deviations of sire transmitting ability also decreased by 8 to 9%. International multitrait across-country comparisons of bulls were computed based on national evaluations from the United States, Canada, New Zealand, and The Netherlands. Estimates of genetic correlations between the United States and other countries decreased by 0.04 to 0.06 when US evaluations were based on modified pedigree. The resulting bias toward selection of domestic bulls and the inability to identify truly superior animals that are available internationally could decrease potential selection differentials by 0.07 to 0.09 standard deviation units on the US scale, which corresponds to sire breeding values of approximately 50 kg for milk, 3 kg for fat, and 1.7 kg for protein. Losses for the other countries were lower and ranged from 0.02 to 0.05 standard deviation units, because a correlation of less than unity with the United States decreased the impact of US cow paternity errors on the scales of other countries. Although paternity verification is desirable and technically feasible, commercial implementation would require low testing costs.

Animals↗

Genotype x environment interaction for grazing versus confinement. I. Production traits.

The objective of this study was to investigate the possible existence of a genotype x environment interaction (GxE) for production traits of US Holsteins in grazing versus confinement herds. Grazing herds were defined as those that utilized grazing for at least 6 mo and were enrolled in dairy herd improvement (DHI). Control herds were confinement DHI herds of comparable size in similar regions. The performance of daughters in grazing herds and control herds was examined using linear regression of mature equivalent milk, fat, and protein yield on the November 2000 USDA-DHI predicted transmitting abilities (PTA) of their sires for those traits. Heritabilities and genetic correlations were estimated using restricted maximum likelihood in a bivariate animal model that considered the same trait in different environments as different traits. Product-moment and rank correlations were calculated between sires' estimated breeding values, estimated separately in both environments. For grazing herds, the coefficient of regression of milk, fat and protein on PTA were 0.78, 0.76, and 0.78, respectively. Corresponding coefficients in the control herds were 0.99, 0.96, and 0.98. Estimates of heritability for the traits ranged from 0.2 to 0.25, and differences between grazing and control environments were small. Estimates of the genetic correlations for the traits in both environments were 0.89, 0.88, and 0.91 for milk, fat, and protein, respectively. Within-quartile analyses revealed a lower correlation for milk and protein between the upper and lower grazing quartiles, while the same quartiles for the control herds did not differ from unity. Rank correlation coefficients between sire estimated breeding values from the 2 environments were 0.59, 0.63, and 0.66 for milk, fat, and protein, respectively. The mean rank change for the top 100 sires between the two environments was 27. The regression coefficients indicate that expected daughter differences may be overstated by current sire PTA in grazing herds. Genetic correlations less than unity suggests that there is, at least, some reranking among sires in both environments, while the rank correlations indicate the possibility of sire reranking when evaluations were performed within management system. However, differences are not so large as to justify separate genetic evaluations for each system.

Animals↗

Genetic evaluation of somatic cell score in dairy cattle considering first and later lactations as two different but correlated traits.

A test-day (TD) random regression model (RRM) was described for the genetic evaluation of somatic cell score (SCS) where first and later lactations were considered as two different but correlated traits. A two-step covariance function procedure was used to estimate variance-covariances and associated genetic parameters. Analysis of estimated breeding values (EBV), ranking of top bulls and cows and some computational aspects were used to compare RRM with TD repeatability model (RPM) and lactation average model (LAM). Residuals were analysed to assess the relative fit of TD models. Comparison between RRM and RPM showed that RRM has lower mean squared error and gave better fit to the data. For young bulls and cows, the standard deviation (SD) of EBVs was highest for RRM and lowest for LAM implying efficient utilization of information on SCS, in terms of revealing more genetic variation. A much lower correlation of EBVs ranging from 0.80 to 0.92 and significant re-ranking of top bulls and cows were observed between RRM and LAM. The lower across-lactation correlation between RRM and LAM indicated that LAM is directed to give more weight to first lactation breeding values. The RRM, where SCS in the first and later lactations was considered as two different but correlated traits was able to make effective use of available information on young bulls and cows, and could offer an opportunity to breeders to utilize EBVs for first and later lactations.

Animals↗

Extent and benefits of multi-country progeny testing of young dairy sires.

One of the current trends within the artificial insemination industry is to progeny test young dairy bulls in multiple countries. The objectives of this study were to assess the extent of multi-country progeny testing and to measure the corresponding gains in reliability of international breeding value estimates. Data of Holstein bulls that were born between July 1, 1992, and December 31, 1994, and progeny tested in countries that participate in the International Bull Evaluation Service were used in the present study, because these were the youngest bulls that had completed multi-country progeny testing before the study. Based on August 1999 international sire evaluation data, a total of 562 bulls from 10 countries were multi-country sampled for production traits during this 2.5-yr period, and 233 bulls from seven countries were multi-country sampled for type traits. The United States, Canada, The Netherlands, France, and Germany were most active in multicountry progeny testing, and Germany, New Zealand, Australia, France, and The Netherlands were the most common countries of foreign sampling. Mean reliabilities of international breeding values were calculated within each country. Means for milk yield were 0.89 for single-country sampled bulls with local progeny (i.e., progeny in the home country), 0.71 for single-country sampled bulls with no local progeny, 0.90 for multicountry sampled bulls with local progeny, and 0.78 for multi-country sampled bulls with no local progeny. Mean reliabilities for teat placement for these groups of bulls were 0.80, 0.71, 0.88, and 0.83, respectively, and means for rear udder width were 0.79, 0.60, 0.85, and 0.68, respectively. Gains in reliability in the country of foreign sampling were greatest when foreign progeny were located in countries that had low genetic correlations with the home country.

Animals↗

Genotype x environment interaction for protein yield in Dutch dairy cattle as quantified by different models.

Variance components and breeding values for protein yield were estimated with REML without and with correction for heterogeneity of variances. Three different sire models were applied, which all accounted for genotype x environment (G x E) interaction. The first model included a sire x herd-year-season subclass (HYS) interaction. The second model divided all records in four different types of management groups, based on estimated HYS subclass effect. The third model, the reaction norm model, performed a random linear regression on the estimated HYS effect. For comparison, a standard model that did not take G x E interaction into account was also applied. Data consisted of 102,899 305-d first-lactation protein records of Holstein Friesians of 1,000 ofthe largest Dutch dairy herds. All animals calved in 1997, 1998, or 1999. Estimated breeding values (EBV) for 2,150 bulls with at least five daughters were calculated. The interaction model detected an interaction variance of 2.5% of the phenotypic variance. The EBV showed a correlation of 1.00 with those of the standard model without interaction. The model with the division in groups showed correlations between groups ranging from 0.73 to 0.86. The EBV showed correlations from 0.84 to 0.91 with the EBV of the standard model. The reaction norm model calculated EBV that had a correlation of 1.00 with the EBV of the standard model. The reaction norm model was not able to detect significant variance of the slope for the protein data corrected for heterogeneity of variances.

Analysis of Variance↗

A combination of walk-back and optimum contribution selection in fish: a simulation study.

The aim of this paper was to study the performance of a novel fish breeding scheme, which is a combination of walk-back and optimum contribution selection using stochastic simulation. In this walk-back selection scheme, batches of different sizes (50, 100, 1,000, 5,000 and 10,000) with the phenotypically superior fish from one tank with mixed families were genotyped to set up the pedigree. BLUP estimated breeding values were calculated. The optimum contribution selection method was used with the rate of inbreeding (DeltaF) constrained to 0.005 or 0.01 per generation. If the constraint on DeltaF could not be held, a second batch of fish was genotyped etc. Compared with the genotyping of all selection candidates (1000, 5,000 or 10,000), the use of batches saves genotyping costs. The results show that two batches of 50 fish were often necessary. With a batch size of 100, genetic level was 76-92% of the genetic level achieved for schemes with all fish being genotyped and thus candidates for the optimum contribution selection step. More parents were selected for schemes with larger batches, resulting in a higher genetic gain, especially when all selection candidates were genotyped. There was little extra genetic gain in genotyping of 1,000 fish instead of 100 for the larger schemes of 5,000 and 10,000 candidates. The accuracy of breeding values was similar for all batch sizes (approximately 0.30), but higher (approximately 0.5) when all candidates were included. Since only the phenotypically most superior fish were genotyped, BLUP-EBV were biased. Compared with genotyping of all selection candidates, the use of batches saves genotyping costs, while simultaneously maintaining high genetic gains.

Animals↗

Estimation of genetic parameters for milk fat depression in dairy cattle.

The objective of this study was to apply reaction norm models to milk recording data to investigate genetic variation in and environmental sensitivity of susceptibility to milk fat depression (MFD). Data comprised 556,276 test-day records of 80,493 heifers in 1043 herds. Breeding values and genetic variances for fat percentage and fat yield were estimated by applying random regression models to average herd-test-day fat percentage. Genetic and permanent environmental correlations between fat yield expressed in different environments ranged, respectively, from 0.83 to 1.00 and from 0.29 to 1.00. Genetic and permanent environmental correlations between fat percentage expressed in different environments ranged, respectively, from 0.87 to 1.00 and from -0.05 to 0.99. Two traits were defined for MFD. The first trait reflected variation of milk fat percentage of animals within lactation after correction for year-season, herd-test-day, age-at-calving, and stage-of-lactation. This trait had an estimated heritability of about 5% and a genetic correlation between the fifth and 95th percentile of the data of 0.50. The second trait reflected the deviation of an animal's fat percentage on a test-day from its expected fat percentage based on fat percentage on the first test-day. This trait had an estimated heritability of about 4% and a genetic correlation between the fifth and 95th percentile of the data of 0.43. The correlation between estimated breeding values of sires for the 2 MFD traits was -0.3. Our results suggest that genetic variation in susceptibility to MFD is present and that selection for reduced susceptibility to MFD is possible.

Animals↗

Adjusting worm egg counts for faecal moisture in sheep.

The number of eggs from gastrointestinal nematodes per gram of faeces (worm egg count WEC) is commonly used to determine the need for anti-parasite treatments and the breeding value of animals when selecting for worm resistance. Diarrhoea increases faecal moisture and may dilute the number of worm eggs observed. To quantify this effect, egg counts in sheep at pasture were simulated by dosing 15 animals with chromic oxide particles. The simulated WEC diminished as faecal moisture increased. When faeces were dried, simulated WEC per unit dry matter was not influenced by the amount of faecal moisture present prior to drying. The results suggest that adjustment for faecal moisture may provide an improved estimate of FEC. Drying faeces to calculate the WEC per unit dry matter would provide such an adjustment but may not be practical for industry application. In the past, the CSIRO McMaster Laboratory has used an adjustment factor developed by Gordon based on the classification of faecal consistency derived from the morphology of faeces. To examine the utility of an adjustment factor based on faecal consistency score (FCS), the relationships between FCS and simulated WEC and dry matter were examined. Dry matter and simulated WEC exhibited an exponential decline as FCS increased. The relationship between FCS and dry matter was further examined in 368 samples collected over 12 months from sheep at pasture, where it was observed that dry matter showed a linear decline as FCS increased. Adjustment factors based on dry matter were similar to those proposed by Gordon however adjustment factors predicted from simulated WEC diverged from the remainder for FCS>4. As no samples scored FCS 5 in the study of simulated FEC, the adjustment factors based on the larger study that included samples with FCS 5 was therefore considered more robust. Adjustment factors were given by the equation: WEC(estimated)=(WEC(observed)/(34.21-5.15 FCS))x29.06. This equation estimates for samples with FCS>1 the WEC that would be expected if the samples were FCS 1, the faecal consistency score for normal faeces. The impact of adjustment of observed WEC for faecal moisture predicted by FCS on decision points for treatment and on estimated breeding values requires further examination.

Animals↗

Cost-benefit evaluation of artificial insemination for genetic improvement of wool-producing sheep.

The financial costs and benefits associated with the use of artificial insemination (AI) in commercial flocks are evaluated. Benefits are calculated in terms of net present values after summing the discounted value of benefits over 20 years. Two breeding strategies are evaluated. With the first, AI is used to produce flock ewes and wethers. The method is unlikely to be profitable unless high breeding value rams are available for AI programs with fresh semen. With the second, AI is used to produce home-bred rams, which in turn sire flock ewes and wethers. This approach is more likely to be profitable. The cost of AI per lamb weaned from laparoscopic AI programs is about $100. Benefits exceed this cost for rams of very high merit when wool prices are moderate or higher. Flock structure has a significant effect on the benefits. Flocks with low wether retention rates have benefits half that of flocks that retain most wethers to 6 years of age. AI with purchased semen also provides benefits to risk management for owners of commercial flocks who wish to breed their own replacement flock rams.

Animals↗

Association of a lysine-232/alanine polymorphism in a bovine gene encoding acyl-CoA:diacylglycerol acyltransferase (DGAT1) with variation at a quantitative trait locus for milk fat content.

DGAT1 encodes diacylglycerol O-acyltransferase (EC ), a microsomal enzyme that catalyzes the final step of triglyceride synthesis. It became a functional candidate gene for lactation traits after studies indicated that mice lacking both copies of DGAT1 are completely devoid of milk secretion, most likely because of deficient triglyceride synthesis in the mammary gland. Our mapping studies placed DGAT1 close to the region of a quantitative trait locus (QTL) on bovine chromosome 14 for variation in fat content of milk. Sequencing of DGAT1 from pooled DNA revealed significant frequency shifts at several variable positions between groups of animals with high and low breeding values for milk fat content in different breeds (Holstein-Friesian, Fleckvieh, and Braunvieh). Among the variants was a nonconservative substitution of lysine by alanine (K232A), with the lysine-encoding allele being associated with higher milk fat content. Haplotype analysis indicated the lysine variant to be ancestral. Two animals that were typed heterozygous (Qq) at the QTL based on marker-assisted QTL-genotyping were heterozygous for the K232A substitution, whereas 14 animals that are most likely qq at the QTL were homozygous for the alanine-encoding allele. An independent association study in Fleckvieh animals confirmed the positive effect of the lysine variant on milk fat content. We consider the nonconservative K232A substitution to be directly responsible for the QTL variation, although our genetic studies cannot provide formal proof.

Acyltransferases↗

A method for the dynamic management of genetic variability in dairy cattle.

According to the general approach developed in this paper, dynamic management of genetic variability in selected populations of dairy cattle is carried out for three simultaneous purposes: procreation of young bulls to be further progeny-tested, use of service bulls already selected and approval of recently progeny-tested bulls for use. At each step, the objective is to minimize the average pairwise relationship coefficient in the future population born from programmed matings and the existing population. As a common constraint, the average estimated breeding value of the new population, for a selection goal including many important traits, is set to a desired value. For the procreation of young bulls, breeding costs are additionally constrained. Optimization is fully analytical and directly considers matings. Corresponding algorithms are presented in detail. The efficiency of these procedures was tested on the current Norman population. Comparisons between optimized and real matings, clearly showed that optimization would have saved substantial genetic variability without reducing short-term genetic gains.

Animals↗

Prediction of genetic values of sires for growth traits of crossbred cattle using a multivariate animal model with heterogeneous variances.

The purpose of this study was to evaluate the effect of adjusting for heterogeneous variances across breed groups on prediction of breeding values (PBV) of selected sires and on breed of sire effects. Data on weights at birth (BWT), 200 d (WW), and 365 d (YW) of purebred and crossbred calves from matings of Angus (A), Hereford (H), Polled Hereford, Charolais, Shorthorn, Simmental, Limousin, Maine-Anjou, Chianina, Gelbvieh, Tarentaise, and Salers Bulls to A and H cows were used. Calf performance in H and A dams was treated as a different trait. Models compared included fixed birth year, cow age, and sex classes and crossbreeding effect as a covariate; random direct and maternal genetic and permanent environmental effects were also included, but their variance structure was different. Model I assumed homogeneous variances across breed groups. Model II accounted for heterogeneous variances. Sires were ranked based on PBV from each model, and means of PBV of selected sires were calculated based on Model II. Differences between mean PBV were small for BWT, intermediate for WW, and larger for YW. Differences in PBV of selected sires increased as selection intensity increased, but only for WW and YW. Large differences in mean PBV of selected sires between maternal environments (H vs A) were observed for WW and YW for various sire breeds. Means of PBV of selected sires based on Model II exceeded those based on Model I by 6 to 16 kg of YW for various selection intensities and maternal environments. Estimates of breed of sire effects from Model I or II were similar for BWT and WW, but large differences were found for YW. Results indicate that some additional economic returns may be gained by commercial producers if sires are chosen across breeds based on predicted genetic values computed with models accounting for heterogeneous variances.

Aging↗

Social cues and hormone levels in male Octodon degus (Rodentia): a field test of the Challenge Hypothesis.

Social interactions are important factors determining and regulating individual behaviors. Testosterone has been related to agonistic interactions, while glucocorticoids have been related to social stress, especially during interactions of dominance. We compared testosterone and cortisol concentrations in male degus (Octodon degus, Rodentia) under laboratory conditions without male social interactions, with data from wild males in nature. Under natural conditions, males should present higher levels of testosterone during the breeding season due to social interactions (Challenge Hypothesis). Alternatively, intense social instability could act as a stressing environment, raising glucocorticoids, which inhibit testosterone concentrations. Our results show a significant increase in agonistic interactions between males during the breeding season, and disappearance of non-agonistic male interactions during this period. Hormone levels in breeding season show nonsignificant differences between laboratory groups, but testosterone concentrations in field males were significantly higher than in laboratory males. Testosterone levels were similar among pre-breeding and breeding periods, but in field animals the concentration was approximately 30% higher than in laboratory degus. In field animals, we found two different mating strategies: resident males, with territorial behavior, and transient males, displayed an opportunistic approach to females. Finally, cortisol presents a similar pattern in both laboratory and field animals; pre-breeding values of cortisol are higher than during the breeding season. This suggests that social interactions in O. degus activate a rise in testosterone, supporting the Challenge Hypothesis, and could be considered as partial support of the Social Stress Hypothesis.

Animals↗

Deleterious mutations, apparent stabilizing selection and the maintenance of quantitative variation.

Apparent stabilizing selection on a quantitative trait that is not causally connected to fitness can result from the pleiotropic effects of unconditionally deleterious mutations, because as N. Barton noted, "...individuals with extreme values of the trait will tend to carry more deleterious alleles...." We use a simple model to investigate the dependence of this apparent selection on the genomic deleterious mutation rate, U; the equilibrium distribution of K, the number of deleterious mutations per genome; and the parameters describing directional selection against deleterious mutations. Unlike previous analyses, we allow for epistatic selection against deleterious alleles. For various selection functions and realistic parameter values, the distribution of K, the distribution of breeding values for a pleiotropically affected trait, and the apparent stabilizing selection function are all nearly Gaussian. The additive genetic variance for the quantitative trait is kQa2, where k is the average number of deleterious mutations per genome, Q is the proportion of deleterious mutations that affect the trait, and a2 is the variance of pleiotropic effects for individual mutations that do affect the trait. In contrast, when the trait is measured in units of its additive standard deviation, the apparent fitness function is essentially independent of Q and a2; and beta, the intensity of selection, measured as the ratio of additive genetic variance to the "variance" of the fitness curve, is very close to s = U/k, the selection coefficient against individual deleterious mutations at equilibrium. Therefore, this model predicts appreciable apparent stabilizing selection if s exceeds about 0.03, which is consistent with various data. However, the model also predicts that beta must equal Vm/VG, the ratio of new additive variance for the trait introduced each generation by mutation to the standing additive variance. Most, although not all, estimates of this ratio imply apparent stabilizing selection weaker than generally observed. A qualitative argument suggests that even when direct selection is responsible for most of the selection observed on a character, it may be essentially irrelevant to the maintenance of variation for the character by mutation-selection balance. Simple experiments can indicate the fraction of observed stabilizing selection attributable to the pleiotropic effects of deleterious mutations.

Alleles↗

Evolution in a changing environment: a case study with great tit fledging mass.

Heritable phenotypic traits under significant and consistent directional selection often fail to show the expected evolutionary response. A potential explanation for this contradiction is that because environmental conditions change constantly, environmental change can mask an evolutionary response to selection. We combined an "animal model" analysis with 36 years of data from a long-term study of great tits (Parus major) to explore selection on and evolution of a morphological trait: body mass at fledging. We found significant heritability of this trait, but despite consistent positive directional selection on both the phenotypic and the additive genetic component of body mass, the population mean phenotypic value declined rather than increased over time. However, the mean breeding value for body mass at fledging increased over time, presumably in response to selection. We show that the divergence between the response to selection observed at the levels of genotype and phenotype can be explained by a change in environmental conditions over time, that is, related both to increased spring temperature before breeding and elevated population density. Our results support the suggestion that measuring phenotypes may not always give a reliable impression of evolutionary trajectories and that understanding patterns of phenotypic evolution in nature requires an understanding of how the environment has itself changed.

Animals↗