Genetic and phenotypic correlations among feed efficiency,production and selected conformation traits in dairy cows

The difficulties and costs of measuring individual feed intake in dairy cattle are the primary factors limiting the genetic study of feed intake and utilisation, and hence the potential of their subsequent industry-wide applications. However, indirect selection based on heritable, easily measurable, and genetically correlated traits, such as conformation traits, may be an alternative approach to improve feed efficiency. The aim of this study was to estimate genetic and phenotypic correlations among feed intake, production, and feed efficiency traits (particularly residual feed intake; RFI) with routinely recorded conformation traits. A total of 496 repeated records from 260 Holstein dairy cows in different lactations (260, 159 and 77 from first, second and third lactation, respectively) were considered in this study. Individual daily feed intake and monthly BW and body condition scores of these animals were recorded from 5 to 305 days in milk within each lactation from June 2007 to July 2013. Milk yield and composition data of all animals within each lactation were retrieved, and the first lactation conformation traits for primiparous animals were extracted from databases. Individual RFI over 301 days was estimated using linear regression of total 301 days actual energy intake on a total of 301 days estimated traits of metabolic BW, milk production energy requirement, and empty BW change. Pair-wise bivariate animal models were used to estimate genetic and phenotypic parameters among the studied traits. Estimated heritabilities of total intake and production traits ranged from 0.27±0.07 for lactation actual energy intake to 0.45±0.08 for average body condition score over 301 days of the lactation period. RFI showed a moderate heritability estimate (0.20±0.03) and non-significant phenotypic and genetic correlations with lactation 3.5 % fat-corrected milk and average BW over lactation. Among the conformation traits, dairy strength, stature, rear attachment width, chest width and pin width had significant (P<0.05) moderate to strong genetic correlations with RFI. Combinations of these conformation traits could be used as RFI indicators in the dairy genetic improvement programmes to increase the accuracy of the genetic evaluation of feed intake and utilisation included in the index.     

Evolutionary ecology of egg size and number in a seed beetle: genetic trade-off differs between environments

Abstract In many organisms, large offspring have improved fitness over small offspring, and thus their size is under strong selection. However, due to a trade-off between offspring size and number, females producing larger offspring necessarily must produce fewer unless the total amount of reproductive effort is unlimited. Because differential gene expression among environments may affect genetic covariances among traits, it is important to consider environmental effects on the genetic relationships among traits. We compared the genetic relationships among egg size, lifetime fecundity, and female adult body mass (a trait linked to reproductive effort) in the seed beetle, Stator limbatus , between two environments (host-plant species Acacia greggii and Cercidium floridum ). Genetic correlations among these traits were estimated through half-sib analysis, followed with artificial selection on egg size to observe the correlated responses of lifetime fecundity and female body mass. We found that the magnitude of the genetic trade-off between egg size and lifetime fecundity differed between environments–a strong trade-off was estimated when females laid eggs on C. floridum seeds, yet this trade-off was weak when females laid eggs on A. greggii seeds. Also differing between environments was the genetic correlation between egg size and female body mass–these traits were positively genetically correlated for egg size on A. greggii seeds, yet uncorrelated on C. floridum seeds. On A. greggii seeds, the evolution of egg size and traits linked to reproductive effort (such as female body mass) are not independent from each other as commonly assumed in life-history theory.     

Genetic parameters for feed efficiency in Romane rams and responses to single-generation selection

Feeding costs represent one of the highest expenditures in animal production systems. Breeding efficient animals that express their growth potential while eating less is therefore a major objective for breeders. We estimated the genetic parameters for feed intake, feed efficiency traits (residual feed intake (RFI) and feed conversion ratio (FCR)), growth and body composition traits in the Romane meat sheep breed. In these traits, selection responses to single-generation divergent selection on RFI were evaluated. From 2009 to 2016, a total of 951 male lambs were tested for 8 weeks starting from 3 months of age. They were weighed at the beginning and at the end of the testing period. Backfat thickness and muscle depth were recorded at the end of the testing period through ultrasound measurements. Feed intake was continuously recorded over the testing period using the automatic concentrate feeders. The heritability of RFI was estimated at 0.45 ± 0.08, which was higher than the heritability of FCR (0.30 ± 0.08). No significant genetic correlations were observed between RFI and growth traits. A favourable low negative genetic correlation was estimated between RFI and muscle depth (−0.30 ± 0.15), though additional data are needed to confirm these results. The selection of low RFI sires based on their breeding values led to the production of lambs eating significantly less concentrate (3% decrease in the average daily feed intake), but with the same growth as lambs from sires selected based on high RFI breeding values. We concluded that in meat sheep, RFI is a heritable trait that is genetically independent of post-weaning growth and body composition traits. A one-generation divergent selection based on RFI breeding values highlighted that substantial gains in feeding costs can be expected in selection schemes for meat sheep breeds.     

Breeding value prediction for production traits in layer chickens using pedigree or genomic relationships in a reduced animal model

Background

Genomic selection involves breeding value estimation of selection candidates based on high-density SNP genotypes. To quantify the potential benefit of genomic selection, accuracies of estimated breeding values (EBV) obtained with different methods using pedigree or high-density SNP genotypes were evaluated and compared in a commercial layer chicken breeding line.

Methods

The following traits were analyzed: egg production, egg weight, egg color, shell strength, age at sexual maturity, body weight, albumen height, and yolk weight. Predictions appropriate for early or late selection were compared. A total of 2,708 birds were genotyped for 23,356 segregating SNP, including 1,563 females with records. Phenotypes on relatives without genotypes were incorporated in the analysis (in total 13,049 production records).The data were analyzed with a Reduced Animal Model using a relationship matrix based on pedigree data or on marker genotypes and with a Bayesian method using model averaging. Using a validation set that consisted of individuals from the generation following training, these methods were compared by correlating EBV with phenotypes corrected for fixed effects, selecting the top 30 individuals based on EBV and evaluating their mean phenotype, and by regressing phenotypes on EBV.

Results

Using high-density SNP genotypes increased accuracies of EBV up to two-fold for selection at an early age and by up to 88% for selection at a later age. Accuracy increases at an early age can be mostly attributed to improved estimates of parental EBV for shell quality and egg production, while for other egg quality traits it is mostly due to improved estimates of Mendelian sampling effects. A relatively small number of markers was sufficient to explain most of the genetic variation for egg weight and body weight.     

Genetic parameters and selection efficiency using part-records for production traits in strawberries

Summary The efficiency of selection for yield and fruit size using parts of the complete-season record was evaluated using data collected for individual strawberry seedlings (Fragaria x annanasa) in four genetic tests. Part-records were generated for each trait using cumulative weekly sums, and these were compared with complete records to assess the value of extended data collection. In addition, part-records were generated using data from alternate weeks of harvest. Scoring separate sets of seedlings in alternate weeks was assumed to allow a doubling of the unselected population size, halving the selection proportion and increasing selection intensity. Heritabilities estimated for part-records were often larger for mid-season cumulative part-records than for the complete-season records. Seasonal patterns for broad-sense and narrow-sense heritabilities differed, suggesting temporal expression of genes that condition yield and fruit size. Predicted genetic gains were largest using cumulative part-records between 4 and 12 weeks of the 15-week season, depending on trait and breeding objective (breeding value versus clonal value), with 8–9 weeks as a reasonable compromise. Selection using alternate-weeks records was identified as an efficient alternative to selection using total complete-season records. Superiority of the alternate-weeks option depends on the balance between increased selection intensity and decreased individual genetic information, and will differ for specific traits and program designs.     

Genetic parameters for performance,feed efficiency,and carcass traits in Senepol heifers

Improving feed efficiency is a key breeding goal in the beef cattle industry. In this study, we estimated the genetic parameters for feed efficiency and carcass traits in Senepol cattle raised in tropical regions. Various indicators of feed efficiency [gain to feed ratio (G:F), feed conversion ratio (FCR), residual weight gain (RG), residual intake and body weight gain (RIG), and residual feed intake (RFI)] as well as growth [final BW, average daily gain (ADG), and DM intake (DMI)], and carcass [rib-eye area (REA), backfat thickness (BF), intramuscular fat score, and carcass conformation score] traits were included in the study. After data editing, records from 1 393 heifers obtained between 2009 and 2018 were used for the analyses. We fitted an animal model that included contemporary group (animals from the same farm that were evaluated in the same test season) as the fixed effect, and a linear effect of animal age at the beginning of the test as a covariate; in addition to random direct additive genetic and residual effects. The (co)variance components were estimated by Bayesian inference in uni- and bivariate analyses. Our results showed that feed efficiency indicators derived from residual variables such as RG, RIG, and RFI can be improved through genetic selection (h² = 0.14 ± 0.06, 0.13 ± 0.06, and 0.20 ± 0.08, respectively). Variables calculated as ratios such as G:F and FCR were more influenced by environmental factors (h² = 0.08 ± 0.05 and 0.09 ± 0.05), and were, therefore, less suitable for use in breeding programs. The traits with the greatest and impact on genetic progress in feed efficiency were ADG, REA, and BF. The traits with the greatest and least impact on growth and carcass traits were RG and RFI, respectively. Selection for feed efficiency will result in distinct overall effects on the growth and carcass traits of Senepol heifers. Direct selection for lower RFI may reduce DMI and increase carcass fatness at the finishing stage, but it might also result in reduced growth and muscle deposition. Residual BW gain is associated with the highest weight gain and zero impact on REA and BF, however, it is linked to higher feed consumption. Thus, the most suitable feed efficiency indicator was RIG, as it promoted the greatest decrease in feed intake concomitant with faster growth, with a similar impact on carcass traits when compared to the other feed efficiency indicators.     

Effect of alfalfa meal in diets of laying quails on performance, egg quality and some serum parameters

This study was conducted to investigate the effects of increasing levels of alfalfa meal in the diet of laying quails on egg production, feed consumption, feed efficiency, egg quality, egg yolk cholesterol and selected serum parameters. In this study, 192, 10-week old quails (Coturnix coturnix japonica) were evenly distributed to four groups with four replicates of 12 quails each. The control group was fed a basal diet containing 0% alfalfa meal and the remaining groups received 3, 6 or 9% alfalfa meal for 12 weeks. Live weight, feed consumption, and egg production were recorded and feed efficiency were calculated. Eggs were examined for interior and exterior quality and egg yolk cholesterol content. At the end of the experiment, blood samples were collected and sera were analysed for serum Ca, inorganic P (P(i)), Mg, triglycerides and total cholesterol. Any level of alfalfa meal had no effect on live weight, egg production, feed consumption, feed efficiency, egg weight, and egg yolk index. Six percent and 9% alfalfa meal increased specific gravity of whole egg and eggshell thickness as well as serum P(i) levels. Nine percent alfalfa meal reduced serum triglycerides, total cholesterol levels and egg yolk cholesterol content. The results of this experiment indicated that addition of 9% alfalfa meal into the laying quail diet may improve eggshell quality and reduced serum triglycerides and serum and egg yolk cholesterol without any adverse effect on performance.     

Can sexual selection drive female life histories? A comparative study on Galliform birds

Sexual selection has been identified as a major evolutionary force shaping male life history traits but its impact on female life history evolution is less clear. Here we examine the impact of sexual selection on three key female traits (body size, egg size and clutch size) in Galliform birds. Using comparative independent contrast analyses and directional discrete analyses, based on published data and a new genera-level supertree phylogeny of Galliform birds, we investigated how sexual selection [quantified as sexual size dimorphism (SSD) and social mating system (MS)] affects these three important female traits. We found that female body mass was strongly and positively correlated with egg size but not with clutch size, and that clutch size decreased as egg size increased. We established that SSD was related to MS, and then used SSD as a proxy of the strength of sexual selection. We found both a positive relationship between SSD and female body mass and egg size and that increases in female body mass and egg size tend to occur following increases in SSD in this bird order. This pattern of female body mass increases lagging behind changes in SSD, established using our directional discrete analysis, suggests that female body mass increases as a response to increases in the level of sexual selection and not simply through a strong genetic relationship with male body mass. This suggests that sexual selection is linked to changes in female life history traits in Galliformes and we discuss how this link may shape patterns of life history variation among species.     

Effect of Alfalfa Meal in diets of laying quails on performance,egg quality and some serum parameters

This study was conducted to investigate the effects of increasing levels of alfalfa meal in the diet of laying quails on egg production, feed consumption, feed efficiency, egg quality, egg yolk cholesterol and selected serum parameters. In this study, 192, 10-week old quails (Coturnix coturnix japonica) were evenly distributed to four groups with four replicates of 12 quails each. The control group was fed a basal diet containing 0% alfalfa meal and the remaining groups received 3, 6 or 9% alfalfa meal for 12 weeks. Live weight, feed consumption, and egg production were recorded and feed efficiency were calculated. Eggs were examined for interior and exterior quality and egg yolk cholesterol content. At the end of the experiment, blood samples were collected and sera were analysed for serum Ca, inorganic P (P_i), Mg, triglycerides and total cholesterol. Any level of alfalfa meal had no effect on live weight, egg production, feed consumption, feed efficiency, egg weight, and egg yolk index. Six percent and 9% alfalfa meal increased specific gravity of whole egg and eggshell thickness as well as serum P_i levels. Nine percent alfalfa meal reduced serum triglycerides, total cholesterol levels and egg yolk cholesterol content. The results of this experiment indicated that addition of 9% alfalfa meal into the laying quail diet may improve eggshell quality and reduced serum triglycerides and serum and egg yolk cholesterol without any adverse effect on performance.     

Combined effect of divergent selection for breast muscle ultimate pH and dietary amino acids on chicken performance,physical activity and meat quality

Responses to changes in dietary Lys and other essential amino acid (AA) concentrations were evaluated in 480 male and female broilers originating from two lines divergently selected for high (pHu+) or low (pHu−) ultimate pH (pHu) of breast muscle. The two genetic lines were fed with two grower isoenergetic diets differing in both true digestible Lys (control=10.2 g/kg and experimental=7.0 g/kg) and amounts of other essential AA calculated in relation to Lys, which were sufficient for the control diet or in excess for the experimental diet. There were six repetitions per treatment. Birds were weighed individually at days 0, 21, 28 and 43. Feed consumption was recorded per pen and feed conversion was calculated over the growing period. The physical activity and walking ability of broilers were recorded during the whole rearing period. Breast and leg yield, and abdominal fat percentage were measured at 43 days of age, as were pHu, color, drip and cooking loss, Warner–Bratzler shear force, and curing-cooking yield of the breast Pectoralis major and pHu of the thigh Sartorius muscle. Divergent selection greatly affected most breast meat quality traits without significantly changing growth rate or feed efficiency. When subjected to a variation in dietary intake of AA, birds from the two genotypes responded in a similar way in terms of animal’s growth, feed efficiency, body composition and meat quality traits. Although line and diet did not affect physical or feeding activities of the broilers, a significant effect of line-by-diet interaction was observed on gait score. Contrary to the pHu− birds, the walking ability of pHu+ birds was impaired when fed the control diet that favored growth and breast muscle development and limited storage of carbohydrate in muscle.     

Combined analysis of group recorded feed intake and individually recorded body weight and litter size in mink

In the mink industry, feed costs are the largest variable expense and breeding for feed efficient animals is warranted. Implementation of selection for feed efficiency must consider the relationships between feed efficiency and the current selection traits BW and litter size. Often, feed intake (FI) is recorded on a cage with a male and a female and there is sexual dimorphism that needs to be accounted for. Study aims were to (1) model group recorded FI accounting for sexual dimorphism, (2) derive genetic residual feed intake (RFI) as a measure of feed efficiency, (3) examine the relationship between feed efficiency and BW in males (BWM) and females (BWF) and litter size at day 21 after whelping (LS21) in Danish brown mink and (4) investigate direct and correlated response to selection on each trait of interest. Feed intake records from 9574 cages, BW records on 16 782 males and 16 875 females and LS21 records on 6446 yearling females were used for analysis. Genetic parameters for FI, BWM, BWF and LS21 were obtained using a multivariate animal model, yielding sex-specific additive genetic variances for FI and BW to account for sexual dimorphism. The analysis was performed in a Bayesian setting using Gibbs sampling, and genetic RFI was obtained from the conditional distribution of FI given BW using genetic regression coefficients. Responses to single trait selection were defined as the posterior distribution of genetic superiority of the top 10% of animals after conditioning on the genetic trends. The heritabilities ranged from 0.13 for RFI in females and LS21 to 0.59 for BWF. Genetic correlations between BW in both sexes and LS21 and FI in both sexes were unfavorable, and single trait selection on BW in either sex showed increased FI in both sexes and reduced litter size. Due to the definition of RFI and high genetic correlation between BWM and BWF, selection on RFI did not significantly alter BW. In addition, selection on RFI in either sex did not affect LS21. Genetic correlation between sexes for FI and BW was high but significantly lower than unity. The high correlations across sex allowed for selection on standardized averages of animals’ breeding values (BVs) for RFI, FI and BW, which yielded selection responses approximately equal to the responses obtained using the sex-specific BVs. The results illustrate the possibility of selecting against RFI in mink with no negative effects on BW and litter size.     

Estimation of additive genetic variance in commercial layer poultry and simulated populations under selection

Changes in genetic parameters over generations for a selected commercial population and simulated populations of poultry with different sizes were studied. The traits analyzed from the commercial population were rate of lay, age at first egg, egg weight, deformation, and body weight. In the simulated population, a trait measured on both sexes and a sex-limited trait, measured only on one sex, each with a heritability of 0.1 and 0.5, were analyzed. In the commercial and simulated populations, males and females were selected on the basis of family selection indexes and data was available only after many generations of selection. Parameters for each generation were estimated by fitting an animal model using derivative free maximum likelihood (DFREML) with different data structures. In structure 1, data included the given (base) generation for which the parameters were to be estimated, and all subsequent generations. In structure 2, only data on birds in the given generation and their progeny were included. In both structures, parents of base-generation birds were assumed unrelated and pedigrees traced back to these parents. With commercial data using structure 1, estimates of _a ² and h² decreased by 14 to 37% across five generations. With structure 2, no trends were observed, though estimates were lower than for structure 1. For simulated data, with a heritability of 0.1, both structures yielded apparently unbiased estimates of the observed additive genetic variances in the (selected) base generation, no matter how many generations of data were utilized, for both sex-limited and normal traits. However, with a heritability of 0.5 the estimated additive genetic variance for both types of trait decreased with a decrease in the number of generations used in the estimation. Estimates based on the first two generations underestimated, while estimates based on five generations of data overestimated, the observed genetic variances in the defined base. The combinations of conditions that lead to varying degrees of bias remain undefined.     

Trait association of a genetic marker near the IGF-I gene in egg-laying chickens

The insulin-like growth factor I (IGF-I) gene was screened for genetic variants associated with trait means and trait correlations. Analysis of an unselected randomly mated White Leghorn population revealed a PstI restriction fragment length polymorphism (RFLP) in the 5' region of the gene which segregated at a frequency of 0.83 for the PstI(+) allele (presence of a PstI restriction site). A comparison of the three genotypic classes revealed that the PstI(-/-) genotype was associated with a significantly lower egg weight measured in three different time periods, while the PstI(+/-) genotype was significantly associated with a higher eggshell weight estimated from the egg weight and egg specific gravity. For eggshell weight, the effect was age dependent and significant only for the last two periods of egg laying. No genotype associations were found for body weight, feed consumption, and egg laying rates. Significant dominance effects of the IGF-I genotype were observed for two of the egg weight measurements and three of the eggshell weight estimates. Partial correlation analyses in the two most frequent genotypic classes, PstI(+/+) and PstI(+/-), revealed the presence of a regulatory loop between feed consumption, body weight, egg weight, and the rate of egg laying. Several aspects of this regulatory loop were different between the two genotypic classes. In particular, for the PstI(+/+) genotype, feed consumption was positively associated with egg weight, while there was no significant association for the PstI(+/-) genotype. Further, the degree of association of body weight with egg weight decreased with age in the genotypic class PstI(+/-), while it was constant for the PstI(+/+) genotype. The results indicated that the marker in the IGF-I gene was not only associated with changes in some trait means, but also with changes in the stability of the coordination between feed intake, body weight, and egg production traits.     

Furnished cages for laying hens: study of the effects of group size and litter provision on laying location, zootechnical performance and egg quality

The furnished cage is a new housing system for layers. A current trend in furnished cage design is to increase group size and replace the litter box with a mat provided with litter. An experiment was set up to determine the effects of group size and litter provision on laying performance and egg quality of beak-trimmed ISA Brown hens housed in large furnished cages with more than 12 hens. Six treatments, each of 18 furnished cages (768 cm(2)/hen including nest and litter area) were compared in a 3 × 2 experimental trial: three group sizes (S20 (20 hens per cage), S40 (40) and S60 (60)), with or without feed as litter distributed on the mat of the litter area. The provision of facilities per hen was equal in all treatments. Mortality, laying rate, mean egg weight, feed intake and feed conversion ratio were unaffected by group size over the 53-week laying period, and performance exceeded the ISA production standards. The overall percentage of eggs laid in the nest exceeded 95% except that it was slightly lower in group S20 (92.0% ± 6.4% v. S40: 96.0% ± 3.3% and S60: 96.2% ± 2.7%) leading to a higher proportion of dirty eggs (S20: 1.6% ± 2.2%, S40: 1.4% ± 1.5%, S60: 1.0% ± 1.0%). At 66 to 70 weeks, eggs laid outside the nest had a slightly higher count of mesophilic bacteria on the eggshell (5.0 log CFU/egg ± 0.4) than those laid in the nest (4.8 log CFU/egg ± 0.5) but no difference in contamination was observed between group sizes. Litter provision had no effect on mortality, egg weight or egg quality traits except for a higher proportion of broken eggs in cages with litter (5.3% ± 6.2% v. 4.6% ± 5.7%). Providing hens with feed for litter was associated with a higher laying rate (97.3% ± 3.2% v. 94.8% ± 4.4% at 23 weeks) and an apparent improvement in feed efficiency at the beginning of the laying period (feed conversion ratio based on feed consumption at the trough: 2.18 ± 0.06 with litter v. 2.28 ± 0.09 without litter at 25 weeks). The results of this study showed that a high level of productivity and good egg quality could be obtained in large furnished cages. Further research is needed to assess the impact on hens' welfare and performance of using more economically competitive substrates than feed for litter.     

The evolution of egg shape in birds: selection during the incubation period

A recent broad comparative study suggested that factors during egg formation – in particular ‘flight efficiency’, which explained only 4% of the interspecific variation – are the main forces of selection on the evolution of egg shape in birds. As an alternative, we tested whether selection during the incubation period might also influence egg shape in two taxa with a wide range of egg shapes, the alcids (Alcidae) and the penguins (Spheniscidae). To do this, we analysed data from 30 species of these two distantly related but ecologically similar bird families with egg shapes ranging from nearly spherical to the most pyriform eggs found in birds. The shape of pyriform eggs, in particular, has previously proven difficult to quantify. Using three egg‐shape indices – pointedness, polar‐asymmetry and elongation – that accurately describe the shapes of all birds’ eggs, we examined the effects of egg size, chick developmental mode, clutch size and incubation site on egg shape. Linear models that include only these factors explained 70–85% of the variation in these egg‐shape indices, with incubation site consistently explaining > 60% of the variation in shape. The five species of alcids and penguins that produce the most pyriform eggs all incubate in an upright posture on flat or sloping substrates, whereas species that incubate in a cup nest have more spherical eggs. We suggest that breeding sites and incubation posture influence the ability of parents to manipulate egg position, and thus selection acting during incubation may influence egg‐shape variation across birds as a whole.     

Estimation of genetic parameters for partial egg production periods by means of random regression models

We estimated genetic parameters for egg production in different periods by means of random regression models, aiming at selection based on partial egg production from a generation of layers. The production was evaluated for each individual by recording the number of eggs produced from 20 to 70 weeks of age, with partial records taken every three weeks for a total of 17 periods. The covariance functions were estimated with a random regression model by the restricted maximum likelihood method. A model composed of third-order polynomials for the additive effect, ninth-order polynomials for the permanent environment, and a residual variance structure with five distinct classes, was found to be most suitable for adjusting the egg production data for laying hens. The heritability estimates varied from 0.04 to 0.14. The genetic correlations were all positive, varying from 0.10 to 0.99. Selection applied in partial egg production periods will result in greater genetic profit for the adjacent periods. However, as the distance in time between periods increases, selection becomes less efficient. Selection based on the second period (23 to 25 weeks of age), where greater heritability was estimated, would note benefit the final egg-laying cycle periods.     

Inheritance of fertility in broiler chickens

Background

The fertility of a chicken''s egg is a trait which depends on both the hen that lays the egg and on her mate. It is also known that fertility of an individual changes over the laying period.

Methods

Longitudinal models including both random genetic and permanent environmental effects of both the female and her male mate were used to model the proportion of fertile eggs in a pedigree broiler population over the ages 29-54 weeks.

Results

Both the male and the female contribute to variation in fertility. Estimates of heritability of weekly records were typically 7% for female and 10% for male contributions to fertility. Repeatability estimates ranged from 24 to 33%, respectively. The estimated genetic variance remained almost constant for both sexes over the laying period and the genetic correlations between different ages were close to 1.0. The permanent environment components increased substantially towards the end of the analyzed period, and correlations between permanent environment effects at different ages declined with increasing age difference The heritability of mean fertility over the whole laying period was estimated at 13% for females and 17% for males. A small positive correlation between genetic effects for male and female fertility was found.

Conclusion

Opportunities to improve fertility in broiler stocks by selection on both sexes exist and should have an impact throughout the laying period.     

Impaired flight ability--a cost of reproduction in female blue tits

When prey are attacked by predators, escape ability has an obvious influence on the probability of survival. Laboratory studieshave suggested that flight performance of female birds mightbe affected by egg production. This is the first study of changesin take-off ability, and thus potentially in predation risk,during reproduction in wild birds. We trapped individual maleand female blue tits repeatedly during the breeding season.Females were 14% heavier and flew 20% slower (probably as aconsequence of a lower ratio of flight muscle to body mass)during the egg-laying period than after the eggs had hatched.However, flight muscle size did not change to compensate for changes in body mass over this period. In contrast, males showedno changes in either body mass, muscle size, or flight abilityover the same period. Furthermore, the impairment of flightin females increased with the proportion of the clutch thathad been laid, an effect that was independent of body mass and muscle size. This indicates that egg production causes additional physiological changes in the female body that produce impairedlocomotor performance. We suggest that courtship feeding offemale blue tits by their mates might reduce predation riskduring the period when female take-off ability is impairedby reducing the time females have to spend foraging and thusreducing the time they are exposed to increased predation.     

Realized correlated responses to artificial selection on pre-adult life-history traits in a butterfly

We use artificial selection experiments targeted on egg size, development time or pupal mass within a single butterfly population followed by a common-garden experiment to explore the interactions among these life-history traits. Relationships were predicted to be negative between egg size and development time, but to be positive between development time and body size and between egg size and body size. Correlated responses to selection were in part inconsistent with these predictions. Although there was evidence for a positive genetic correlation between egg and body size, there was no support for genetic correlations between larval development time and either egg size or pupal mass. Phenotypic correlations among the three target traits of selection gave comparable results for the relationships between egg mass and development time (no association) as well as between egg mass and pupal mass (positive association), but not for the relation between development time and pupal mass (negative phenotypic correlation). In summary, correlated responses to selection as well as phenotypic correlations were rather unpredictable. The impact of variation in acquisition and allocation of energy as well as of the benign conditions used deserve further investigation.     

Repeatability and heritability of reproductive traits in free-ranging snakes

The underlying genetic basis of life-history traits in free-ranging animals is critical to the effects of selection on such traits, but logistical constraints mean that such data are rarely available. Our long-term ecological studies on free-ranging oviparous snakes (keelbacks, Tropidonophis mairii (Gray, 1841), Colubridae) on an Australian floodplain provide the first such data for any tropical reptile. All size-corrected reproductive traits (egg mass, clutch size, clutch mass and post-partum maternal mass) were moderately repeatable between pairs of clutches produced by 69 female snakes after intervals of 49-1152 days, perhaps because maternal body condition was similar between clutches. Parent-offspring regression of reproductive traits of 59 pairs of mothers and daughters revealed high heritability for egg mass (h2= 0.73, SE=0.24), whereas heritability for the other three traits was low (< 0.37). The estimated heritability of egg mass may be inflated by maternal effects such as differential allocation of yolk steroids to different-sized eggs. High heritability of egg size may be maintained (rather than eroded by stabilizing selection) because selection acts on a trait (hatchling size) that is determined by the interaction between egg size and incubation substrate rather than by egg size alone. Variation in clutch size was mainly because of environmental factors (h2=0.04), indicating that one component of the trade-off between egg size and clutch size is under much tighter genetic control than the other. Thus, the phenotypic trade-off between egg size and egg number in keelback snakes occurs because each female snake must allocate a finite amount of energy into eggs of a genetically determined size.