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Fetal development in the pig in relation to genetic merit for piglet survival.

The objective of this study was to investigate if litters with different genetic merit for piglet survival differ in late fetal development. In total, 507 fetuses from 46 litters were delivered by Caesarean section at, on average, d 111 of gestation. All litters had known estimated breeding values for piglet survival (EBVps). The obtained range of EBVps of the litters was continuous, and the difference between litters with the lowest and highest EBVps was 16.4%. Analysis of relationships between fetal characteristics and EBVps was performed with litter averages, using linear regression analysis with inclusion of EBVps as a covariate. An increase in EBVps of the litter was associated with decreases in average placental weight (P = 0.01) and within-litter variation in placental weight (P = 0.02), and an increase in average placental efficiency (P = 0.08). Average fetal length decreased with increasing EBVps (P = 0.04), but weights of liver (P = 0.02), adrenals (P = 0.0001), and small intestine (P = 0.01) showed relative increases with increasing EBVps. Average serum cortisol concentrations increased with increasing EBVps (P = 0.0001), but the other blood characteristics (hematocrit, glucose, fructose, albumin, estradiol-17beta) were not related to EBVps. Glycogen concentrations in liver (P = 0.07) and longissimus dorsi muscle (P = 0.04) and total liver glycogen content (P = 0.05) increased with increasing EBVps, whereas heart glycogen concentration decreased with increasing EBVps (P = 0.005). The percentage of carcass fat increased with increasing EBVps (P = 0.05). Relationships of relative liver weight, relative small intestinal weight, and liver and muscle glycogen levels with EBVps were absent after adjustment for differences in cortisol levels between litters. The observed differences in fetal development in relation to EBVps suggest a higher degree of physiological maturity in litters with high EBVps. Differences in fetal cortisol most likely accounted for most of these maturational differences. The results imply that selection for improved piglet survival will lead to slightly smaller piglets that nevertheless have an improved ability to cope with hazards during birth or within the first days of life.

Animals↗

Heritability, reliability of genetic evaluations and response to selection in proportional hazard models.

The purposes of this study were 1) to investigate the heritability, reliability, and selection response for survival traits following a Weibull frailty proportional hazard model; and 2) to examine the relationship between genetic parameters from a Weibull model, a discrete proportional hazard model, and a binary data analysis using a linear model. Both analytical methods and Monte Carlo simulations were used to achieve these aims. Data were simulated using the Weibull frailty model with two different shapes of the Weibull distribution. Breeding values of 100 unrelated sires with 50 to 100 progeny (with different levels of censoring) were generated from a normal distribution and two different sire variances. For analysis of longevity data on the discrete scale, simulated data were transformed to a discrete scale using arbitrary ends of discrete intervals of 400, 800, or 1200 d. For binary data analysis, an individual's longevity was either 0 (when longevity was less than the end of interval) or 1 (when longevity was equal or greater than the end of interval). Three different statistical models were investigated in this study: a Weibull model, a discrete-time model (a proportional hazard model assuming that the survival data are measured on a discrete scale with few classes), and a linear model based upon binary data. An alternative derivation using basic expressions of reliabilities in sire models suggests a simple equation for the heritability on the original scale (effective heritability) that is not dependent on the Weibull parameters. The predictions of reliabilities using the proposed formulae in this study are in very good agreement with reliabilities observed from simulations. In general, the estimates of reliability from either the discrete model or the binary data analysis were close to estimates from the Weibull model for a given number of uncensored records in this simplified case of a balanced design. Although selection response from the binary data analysis depends on the end of interval point, there is a relatively good agreement between selection responses in the Weibull model and the binary data analysis. In general, when the underlying survival data is from a Weibull distribution, it appears that the method of analyzing data does not greatly affect the results in terms of sire ranking or response to selection, at least for the simplified context considered in this study.

Animals↗

Evidence that an imprinted gene on the X chromosome increases ovulation rate in sheep.

Ovulation rate records from 1311 female progeny of 50 Coopworth rams were used to study the inheritance of ovulation rate in a screened high prolificacy sheep flock. Breeding values (BV) for ovulation rate for 33 sires used within the screened flock and ovulation rate deviations for a further 17 sires progeny tested in commercial flocks suggest that a major gene (WOODLANDS: gene) for ovulation rate with a non-Mendelian inheritance pattern is segregating in a family line. Rams assigned as carriers of the putative gene did not produce carrier sons (zero of three), and this coupled with the observation that daughters of carrier rams had ovulation rates of 0. 39 (standard error of difference [SED] = 0.06) higher than contemporaries without a significant increase in the variance of log ovulation rate strongly suggests that the gene is on the X chromosome. The evidence suggests that the gene is also maternally imprinted because ovulation rate data indicate that it is expressed where females inherit a paternal allele but is silenced when inherited on a maternal allele. Maternal granddaughters of carrier rams had mean ovulation rates that were only 0.02 (SED = 0.06) higher than noncarrier ewes from the same flock. Furthermore, carrier dams expressing the gene (paternal allele) had 24 sons, none of which had female offspring that expressed the gene, whereas carrier dams not expressing the gene (maternal allele) had 7 out of 17 sons that had female progeny expressing the gene. There is no evidence of the infertility that occurs in homozygous ewes carrying the X-linked Inverdale gene. Collectively, these results suggest the existence of a novel gene for prolificacy located on the X chromosome that is maternally imprinted. The WOODLANDS: gene was only expressed upon paternal inheritance from carrier males that were the progeny of nonexpressing carrier dams. The gene was not expressed in ewes that received it from either carrier dams (expressing or nonexpressing) or from carrier males that were the progeny of expressing carrier dams.

Animals↗

Maximization of lactation milk production without decreasing persistency.

This study treats each daily estimated breeding value (EBV) of the lactation as a separate trait to modify the lactation curve on a daily basis. Six selection strategies for improving lactation milk without decreasing persistency were compared: 1) index I(R1), subject to the restriction of equal genetic gains at days in milk (DIM) 60 and 280, 2) I(R2), subject to the restriction of zero gain at DIM 60, 3) desired gains index I(d), designed to increase lactation milk without altering the lactation curve, 4) index I(u), comprising lactation EBV and persistency without standardization, 5) index I(w), consisting of lactation EBV (EBV(L)) and persistency with standardization, and 6) conventional selection on EBV(L) and used as a basis for comparison. Of the 6 selection strategies compared, I(R2) yielded the greatest persistency, but achieved the smallest response in EBV(L), suggesting that it is impractical to increase persistency by inhibiting change in the peak yield. Index I(u) showed the same response in lactation milk as conventional selection on EBV(L), but resulted in the same decreased persistency. Although both I(R1) and I(d) achieved constant persistency, the former produced a greater lactation response (669 kg EBV) than the latter (560 kg EBV). Thus, I(R1) is a viable strategy for improving EBV(L) while holding persistency constant. None of the 6 selection strategies excelled in both lactation milk and persistency. Index I(w) appears to be a reasonable choice for improving both traits, although responses would depend on the relative economic importance of the 2 traits. Differential responses between I(u) and I(w) emphasize the need to weight the EBV of different traits by the inverse of their standard deviations in index construction when the EBV vary widely in variance. The general formula developed here provides a useful genetic means of modifying the lactation curve by restricting differential genetic gains among different days of the lactation.

Animal Nutritional Physiological Phenomena↗

Impact of genetic selection on management of boar replacement.

Boars in an artificial insemination centre have been selected for their superior genetic potential, with 'superior' being defined as having traits the customer wants transmitted to his herd. The ability to meet the customers' needs depends on the heritability of the trait, the geneticist's success in devising a selection scheme for the trait in balance with other economically important traits, and the boar's ability to produce sperm that can fertilise oocytes. Genetic evaluation research over the past 20 years has greatly increased the number of traits for which a boar can be selected: currently in the Canadian national program, these include age at 100 kg, backfat at 100 kg, feed efficiency, lean yield and litter size. In the near future, traits that are very likely to be added to this selection list include piglet survival, marbling, loin eye area and structure traits. In Canada, sires are ranked on two estimated breeding value (EBV) indices; one, focused on development of terminal sire lines, is based on the growth and yield traits and another, primarily focused on maternal line development, de-emphasises these traits and incorporates litter size. Boars that are in Canadian AI centres because of their excellent growth traits are typically in the top 5-10% of the national population for terminal sire line index, but they may be only average or substandard for litter size. Conversely, boars selected to be in the top 5-10% for conveying such reproductive traits as litter size may only be in the top 33% for growth traits. The more offspring from a superior boar in either of these indices, the faster the population average for the trait improves. The original sire gets knocked out of the elite group, is culled and replaced by a higher ranked young boar from the now improved general population. Although genetic superiority should govern an AI centre's selection and culling of boars, decision-making in real life is seldom that simple. Selection criteria may be contradictory as above, or a boar with truly superior traits may be excluded because a newly-developed molecular genetics test determines he carries an undesirable gene such as PSS, RN or others being developed. Selection for terminal sire or maternal line traits can ignore important practical factors that affect an AI centre--boars with superior genetics may not produce good semen because skeletal or penile problems prevent ejaculation, or because sperm production is poor due to a genetic flaw, disease, or some other cause. Interestingly, selection pressure for one trait may inadvertently select for a trait that is linked but whose linkage is unrecognised, and such unintentionally selected genes could benefit, harm, or have no effect on production traits. An AI centre serving a variety of customers must select boars in anticipation of their customers' needs (including new, foreign and niche markets). A centre should also review its genetic evaluation results and progeny records, both to critique its own selection success and to try to detect unexpected linkages. Finally, an AI centre needs to predict its own future, selecting not just for production traits for the swine producer, but also for factors that enhance the centre's efficiency including boar conformation and temperament, and sperm quantity, quality and hardiness. Can we select for efficiency? Our colleagues in dairy cattle AI evaluate bull performance--should the swine industry consider evaluation of male fertility traits?

Animals↗

Estimation of genotypic and environmental variation in plants.

The frequency of genotypes with the desired degree of expression of economically important quantitative characters within a hybrid or mutant population is usually very low. Therefore, the early identification and selection of such genotypes involves the analysis of very large populations. Because the breeding values of individuals in a population are masked by environmental, competitional, and ontogenic noises, special quantitative--genetic methods of analysis have to be used in order to eliminate their disturbing effects. The present chapter deals with a new approach to such an analysis by using either a simple background character or a background index obtained as a linear function of two, or more than two, background characters. It is believed that the use of this approach would greatly increase the efficiency of selection and shorten the time needed to produce improved new crop cultivars. As the analyses require the handling of large amounts of measurement data, plant breeders must use computer facilities.

Environment↗

Quantitative traits in chicken associated with DNA fingerprint bands.

In an unselected control line of a selection experiment in chicken the following traits were evaluated: number of eggs, average egg weight, egg mass, age at first egg, body weight at 20 weeks, body weight at 40 weeks, feed consumption and feed efficiency (feed consumption/egg mass). Data and blood samples of 143 females of the same age were available for analysis. The animals were ranked for each trait according to the phenotypic performance adjusted for hatch and laying house effect rather than the breeding value. DNA mixes of the top 5 and the bottom 5 hens were compared to each other by DNA fingerprinting. The most striking differences could be observed with the probe pV47 in the mixes for body weight at 20 weeks and the feed efficiency. In the following the study focused on feed efficiency. Five groups of 10 hens each, around quantiles 0%, 25%, 50%, 75% and 100%, were analyzed with respect to the occurrence of 7 particular DNA fingerprint bands. The association of the groups' mean feed efficiency with the groups' band frequencies was assessed by linear regression. For 2 bands significant regression coefficients (P < 0.005) were found.

Alleles↗

Positive assortative mating with selection restrictions on group coancestry enhances gain while conserving genetic diversity in long-term forest tree breeding.

Selection and mating principles in a closed breeding population (BP) were studied by computer simulation. The BP was advanced, either by random assortment of mates (RAM), or by positive assortative mating (PAM). Selection was done with high precision using clonal testing. Selection considered both genetic gain and gene diversity by "group-merit selection", i.e. selection for breeding value weighted by group coancestry of the selected individuals. A range of weights on group coancestry was applied during selection to vary parent contributions and thereby adjust the balance between gain and diversity. This resulted in a series of scenarios with low to high effective population sizes measured by status effective number. Production populations (PP) were selected only for gain, as a subset of the BP. PAM improved gain in the PP substantially, by increasing the additive variance (i.e. the gain potential) of the BP. This effect was more pronounced under restricted selection when parent contributions to the next generation were more balanced with within-family selection as the extreme, i.e. when a higher status effective number was maintained in the BP. In that case, the additional gain over the BP mean for the clone PP and seed PPs was 32 and 84% higher, respectively, for PAM than for RAM in generation 5. PAM did not reduce gene diversity of the BP but increased inbreeding, and in that way caused a departure from Hardy-Weinberg equilibrium. The effect of inbreeding was eliminated by recombination during the production of seed orchard progeny. Also, for a given level of inbreeding in the seed orchard progeny or in a mixture of genotypes selected for clonal deployment, gain was higher for PAM than for RAM. After including inbreeding depression in the simulation, inbreeding was counteracted by selection, and the enhancement of PAM on production population gain was slightly reduced. In the presence of inbreeding depression the greatest PP gain was achieved at still higher levels of status effective number, i.e. when more gene diversity was conserved in the BP. Thus, the combination of precise selection and PAM resulted in close to maximal short-term PP gain, while conserving maximal gene diversity in the BP.

Computer Simulation↗

Effects of inbreeding on coastal Douglas fir growth and yield in operational plantations: a model-based approach.

In advanced generation seed orchards, tradeoffs exist between genetic gain obtained by selecting the best related individuals for seed orchard populations, and potential losses due to subsequent inbreeding between these individuals. Although inbreeding depression for growth rate is strong in most forest tree species at the individual tree level, the effect of a small proportion of inbreds in seed lots on final stand yield may be less important. The effects of inbreeding on wood production of mature stands cannot be assessed empirically in the short term, thus such effects were simulated for coastal Douglas fir [ Pseudotsuga menziesii var. menziesii (Mirb.) Franco] using an individual-tree growth and yield model TASS (Tree and Stand Simulator). The simulations were based on seed set, nursery culling rates, and 10-year-old field test performance for trees resulting from crosses between unrelated individuals and for inbred trees produced through mating between half-sibs, full-sibs, parents and offspring and self-pollination. Results indicate that inclusion of a small proportion of related clones in seed orchards will have relatively low impacts on stand yields due to low probability of related individuals mating, lower probability of producing acceptable seedlings from related matings than from unrelated matings, and a greater probability of competition-induced mortality for slower growing inbred individuals than for outcrossed trees. Thus, competition reduces the losses expected due to inbreeding depression at harvest, particularly on better sites with higher planting densities and longer rotations. Slightly higher breeding values for related clones than unrelated clones would offset or exceed the effects of inbreeding resulting from related matings. Concerns regarding the maintenance of genetic diversity are more likely to limit inclusion of related clones in orchards than inbreeding depression for final stand yield.

Analysis of Variance↗

Production of doubled haploids in durum wheat (Triticum turgidum L.) through isolated microspore culture.

The objective of this work was to produce doubled haploid plants from durum wheat through the induction of androgenesis. A microspore culture technique was developed and used to produce fertile doubled haploid plants of agronomic interest. Five cultivars, one selected line, plus a collection of 20 F(1) crosses between different genotypes of high breeding value were used. Studies on several factors such as pre-treatments and media components were carried out in order to develop a protocol to regenerate green haploid plantlets. Anthers were pre-treated in 0.7 M mannitol. Microspores, from anther maceration, were plated on a C(17) induction culture medium with ovary co-culture. The optimum regeneration medium J25-8 was used. From 35 microspore isolations, 407 green plantlets were obtained. With this technique mature embryos were obtained. Green plants were regenerated from all genotypes used and approximately 67% of them were spontaneously doubled haploids. Some haploids and a very few polyploids plants were obtained. From the 407 plants, 275 were completely fertile and gave enough seeds to be assayed in the field. This protocol could be used complementary to or instead of the intergeneric crossing with maize as an economically feasible method to obtain doubled haploids from most durum wheat genotypes.

Benzyl Compounds↗

Detection of QTL for milk production on Chromosomes 1 and 6 of Holstein cattle.

Seventy to 75 sons of each of six Holstein sires were assayed for genotypes at a number of microsatellite loci spanning Chromosomes (Chrs) 1 and 6. The number of informative loci varied from three to eight on each chromosome in different sire families. Linkage order and map distance for microsatellite loci were estimated using CRI-MAP. Estimates of QTL effect and location were made by using a least squares interval mapping approach based on daughter yield deviations of sons for 305-d milk, fat, protein yield, and fat and protein percentage. Thresholds for statistical significance of QTL effects were determined from interval mapping of 10,000 random permutations of the data across the bull sire families and within each sire family separately. Across-sire analyses indicated a significant QTL for fat and protein yield, and fat percentage on Chr 1, and QTL effects on milk yield and protein percentage that might represent one or two QTL on Chr 6. Analyses within each sire family indicated significant QTL effects in five sire families, with one sire possibly being heterozygous for two QTLs. Statistically significant estimates of QTL effects on breeding value ranged from 340 to 640 kg of milk, from 15.6 to 28.4 kg of fat, and 14.4 to 17.6 kg of protein.

Alleles↗

Body and egg iron content in shaver and New Hampshire chickens.

1. A comparative study on the iron content in internal organs (blood, liver, spleen, kidneys and heart), carcass and feathers, as well as on the whole iron content has been carried out at different ages (4, 8, 13, 18 and 24 weeks), on two chicken breeds (shaver and New Hampshire). 2. The iron content in internal organs in shaver was higher than in New Hampshire. 3. A tendency of the iron content in the carcass to decrease with age has been observed for both breeds; values were higher for laying shaver hens. 4. The egg-producing strain, shaver, showed a greater iron status with a stable body iron content from 13 weeks on (53-55 ppm), permitting a constant laying frequency (90%) during an 18-month period, with only a small reduction of egg iron content from 33.8 to 31 ppm. 5. The whole body iron decreased progressively with age in the New Hampshire broiler strain to 42-49 ppm iron, and the laying process sharpened this tendency. A drastic reduction in laying, accompanied by a lower egg iron content (28.9 ppm iron), was observed. 6. Iron data obtained in the present paper reflect the specific genetic adaptation to egg production of the shaver strain.

Animal Husbandry↗

Impact of sedation method on the diagnosis of hip and elbow dysplasia in Swedish dogs.

Our objective was to investigate the effect of sedation method on the screening result for hip and elbow dysplasia. The study was based on a questionnaire survey of routines for hip and elbow screening at Swedish veterinary clinics and results of hip and elbow status, for eight breeds (Bernese Mountain Dog, Boxer, German Shepherd Dog, Golden Retriever, Labrador Retriever, Newfoundland, Rottweiler, and Saint Bernard) recorded by the Swedish Kennel Club. In total 5877 and 5406 dogs examined for hip and elbow dysplasia, respectively, from January 2002 through March 2003 were included. We used logistic regression to examine whether the type of chemical restraint used for sedation affected the screening result for hip and elbow dysplasia. In addition to sedation method, the effects of veterinary clinic, sex, breed, and age at screening were studied. The type of chemical restraint used for sedation affected the screening result for hip but not for elbow dysplasia. Acepromazine gave less than half the odds of hip dysplasia compared with medetomidine and butorphanol (the most common method), medetomidine alone or xylazine. Females had about 25% higher odds for developing hip dysplasia whereas males had almost 40% higher odds for developing elbow dysplasia. Saint Bernard, Newfoundland and German Shepherd Dog had the highest odds of developing hip dysplasia, whereas Rottweiler and Labrador Retriever had the lowest odds. Boxer had the lowest risk for elbow dysplasia, followed by Labrador Retriever. Saint Bernard and Rottweiler had the highest odds of elbow dysplasia. Increasing age increased the odds of both hip and elbow dysplasia, by about 2.5% per month. Following the results in this study, recording of the type of chemical restraint used for sedation during hip screening has now become mandatory in Sweden. This makes it possible to account for the effect of sedation method in a model for prediction of breeding values for hip dysplasia.

Acepromazine↗

Genetic variation in fertility of heat-stressed male mice.

The damaging effects of heat stress on male fertility are evident in developing spermatozoa expressed in ejaculates 18-28 days post-stress in mice. Our objectives were to: (1) assess genetic variation in fertility of heat-stressed male mice and (2) determine response to selection for fertility after heat stress in male mice. Mature male mice were exposed to heat stress (35+/-1 degrees C; n=50) or control (21+/-1 degrees C; n=10) conditions for 24h (day 0) and then hemicastrated for tissue collection. Two periods of mating tests followed, period 1 (from days 3 to 11) when no reductions in fertility were anticipated, and period 2 (days 18-26) when variation in fertility was expected. Period 2 pregnant females were sacrificed in late gestation. Males were indexed by multiplying overall mean ovulation rate by pre-implantation survival and number of pregnant period 2 mates. The five highest and five lowest ranking males were identified as heat stress resistant and susceptible, respectively. Resistant males were 61.2units superior in the index, 57.5% greater in pregnancy rate, and 57.6 total fetuses greater than susceptible males. Progeny of resistant sires were superior to progeny of susceptible sires in estimated breeding value by 4.5units for the index, 4.1% for pregnancy rate, and 5.2 fetuses (P<0.0001). Heritability estimates for the index, pregnancy rate, and number of fetuses ranged from 0.09 to 0.13, suggesting male fertility following heat stress is heritable and responds to selection.

Animals↗

Individual variation in plasma estradiol-17beta and androgen levels during egg formation in the European starling Sturnus vulgaris: implications for regulation of yolk steroids.

While it is clear that maternal transfer of steroids to egg yolk can have significant effects on offspring phenotype, an unresolved question is whether females can facultatively adjust yolk hormone levels independently of their own plasma levels or whether yolk steroid levels are simply a direct consequence of temporal variation in the female's hormonal status. In part, this is because we lack detailed information about the day-to-day pattern of changes in plasma hormone levels during the laying cycle for non-domesticated birds. Here, we describe changes in plasma estradiol-17beta (E2) and androgens, throughout laying in relation to specific stages of ovarian follicular development in the European starling (Sturnus vulgaris). Plasma E2 levels increased rapidly from the onset of rapid yolk development (RYD) to reach maximum levels in birds with a complete follicle hierarchy (> or = 4 yolky follicles). However, levels decreased linearly throughout the later stages of follicle development returning to pre-breeding values before the final yolky follicle was ovulated. In females with > or = 4 yolky follicles there was 10-fold variation in plasma E2 levels among individual females, but this was not related to plasma levels of the main yolk precursor vitellogenin or to the total mass of yolky follicles developing at the time of blood sampling. In contrast to E2, plasma androgen levels showed only a very gradual linear decline throughout the laying cycle from pre-RYD to clutch completion. Furthermore, androgen levels showed less individual variability: 4-fold variation among females with > or = 4 yolky follicles, although this was also independent of our measures of reproductive function. Data on inter- and intra-individual variation in female hormone levels are important to set-up a priori predictions for, and interpretation of, studies of yolk hormone levels.

Androgens↗

Divergent selection on feather pecking behaviour in laying hens (Gallus gallus domesticus).

A selection experiment was initiated in 1996 in which selection for (HP line) and against (LP line) feather pecking was performed. The foundation stock was a White Leghorn layer strain established in 1970 and maintained since then as a random bred control line at the Institute. Six hatches were produced over three generations. At the age of 68 weeks (generation 0, 1996), 35 weeks (generation 1, 1997), 30 weeks (generation 2, 1998), and 27 weeks (generation 3, 1999) female birds were transferred to observation pens and their feather pecking behaviour was recorded. In each generation, 30 females and 8 males were selected from approximately 200 females and 60 males. The selection criterion was breeding value estimated by animal model on the trait 'number of bouts of feather pecking per bird per hour'.Feather pecking behaviour in adult hens was significantly higher in HP than in LP. In generation 2 the following was recorded: 3.10 versus 1.37 bouts per bird per hour (P<0.01), 7.04 versus 3.58 pecks per bird per hour (P<0.05) and the proportion of hens recorded feather pecking in the 180min observation period was 67 versus 56% (P<0.05). In generation 3 the following was recorded: 4.56 versus 0.63 bouts per bird per hour (P<0.001), 13.9 versus 2.51 pecks per bird per hour (P<0.001) and the proportion of hens recorded feather pecking in the 180min observation period was 75 versus 49% (P<0.001).In generation 3, plumage condition was better in LP on neck, breast, back, wings and tail, as well as overall (P<0.001). Body weight did not differ between lines in generation 2, but in generation 3, HP hens were on average heavier than LP hens at the age of 27 weeks (1435g versus 1371g, P<0.001).

Journal Article↗

Clines in polygenic traits.

This article outlines theoretical models of clines in additive polygenic traits, which are maintained by stabilizing selection towards a spatially varying optimum. Clines in the trait mean can be accurately predicted, given knowledge of the genetic variance. However, predicting the variance is difficult, because it depends on genetic details. Changes in genetic variance arise from changes in allele frequency, and in linkage disequilibria. Allele frequency changes dominate when selection is weak relative to recombination, and when there are a moderate number of loci. With a continuum of alleles, gene flow inflates the genetic variance in the same way as a source of mutations of small effect. The variance can be approximated by assuming a Gaussian distribution of allelic effects; with a sufficiently steep cline, this is accurate even when mutation and selection alone are better described by the 'House of Cards' approximation. With just two alleles at each locus, the phenotype changes in a similar way: the mean remains close to the optimum, while the variance changes more slowly, and over a wider region. However, there may be substantial cryptic divergence at the underlying loci. With strong selection and many loci, linkage disequilibria are the main cause of changes in genetic variance. Even for strong selection, the infinitesimal model can be closely approximated by assuming a Gaussian distribution of breeding values. Linkage disequilibria can generate a substantial increase in genetic variance, which is concentrated at sharp gradients in trait means.

Alleles↗

How is female mate choice affected by male competition?

The plethora of studies devoted to the topics of male competition and female mate choice belie the fact that their interaction remains poorly understood. Indeed, on the question of whether competition should help or hinder the choice process, opinions scattered throughout the sexual selection literature seem unnecessarily polarised. We argue, in the light of recent theoretical and empirical advances, that the effect of competition on mate choice depends on whether it results in the choosy sex attaining high breeding value for total fitness, considering both direct and indirect fitness benefits. Specifically, trade-offs may occur between different fitness benefits if some are correlated with male competitive ability whilst others are not. Moreover, the costs and benefits of mating with competitive males may vary in time and/or space. These considerations highlight the importance of injecting a life-history perspective into sexual selection studies. Within this context, we turn to the sexual selection literature to try to offer insights into the circumstances when competition might be expected to have positive or negative implications for pre-copulatory female choice. In this regard, we elaborate on three stages where competition might impact upon the choice process: (i) during mate detection, (ii) mate evaluation, and (iii) in dictating actual mating outcomes. We conclude by offering researchers several potentially rewarding avenues for future research.

Animals↗