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Sensitivity of MST neurons to optic flow stimuli. II. Mechanisms of response selectivity revealed by small-field stimuli.

1. In these experiments we examined the receptive field mechanisms that support the optic flow field selective responses of neurons in the dorsomedial region of the medial superior temporal area (MSTd). Our experiments tested the predictions of two hypotheses of optic flow field selectivity. The direction mosaic hypothesis states that these receptive fields contain a set of planar direction-selective subfields that match the local directions of motion within optic flow fields. The vector field hypothesis states that these receptive fields are uniquely sensitive to distributed properties of planar, circular, or radial optic flow fields. 2. Experiments using large-field stimuli revealed that some neurons showed changes in optic flow field selectivity depending on the position of the stimulus in the receptive field; these are position-dependent responses. However, other neurons maintained the same optic flow field selectivities in spite of changes in stimulus position; these are position-invariant responses. We have used the position dependence or invariance of optic flow field selectivity as a way of testing the direction mosaic and vector field hypotheses. Position dependence is more consistent with the direction mosaic hypothesis, whereas position invariance is more consistent with the vector field hypothesis. 3. To test for position effects, we examined the optic flow field selectivity of small subfields within the large receptive fields of 160 MSTd neurons. First, we centered small-field optic flow stimuli of various sizes over the same position in the receptive field. Most MSTd neurons showed decreasing response amplitude with decreasing stimulus size but maintained optic flow field selectivity. 4. We then placed small-field stimuli at various positions within the large receptive field of these MSTd neurons. Position-invariant response selectivity was most prominent in single-component neurons, suggesting that they were more consistent with the vector field hypothesis. Position-dependent response selectivity was most prominent in triple-component neurons, suggesting that they were more consistent with the direction mosaic hypothesis. However, the variations in planar direction preference throughout the receptive field of these triple-component neurons were not consistent with a direction mosaic explanation of the large-field circular or radial selectivity observed. 5. Small-field position studies also demonstrated the existence of zones within the receptive field in which either direction-selective inhibitory or direction-selective excitatory responses predominated. The degree of overlap between these zones increased from nonselective to triple- to double- and finally to single-component neurons.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Effects of mass selection for increased weight at two ages on growth rate and carcass composition of Duroc-Landrace pigs.

Duroc boars from a line previously selected over five generations for 200-d weight and those from a randomly selected control line were mated to Landrace sows either from a line previously selected for increased 70-d weight or from a randomly selected pedigree control line. From these matings, 900 pigs were farrowed to examine the effects of crossing lines of pigs mass selected for weight at two ages on growth rate, survival, and carcass composition. A greater (P less than .01) percentage of pigs farrowed survived birth from control-line sows (.974) than from select-line sows (.914). Of those pigs born alive, a greater (P less than .05) percentage of pigs out of control-line sows survived to 21 d (.893) than out of select-line sows (.829). Pigs sired by select-line boars weighed 2.1 kg heavier (P less than .05) at 70 d than pigs sired by control-line boars. Pigs out of select-line sows weighed .11 kg less (P less than .10) at birth and .3 kg less (P less than .10) at 21 d of age but grew .026 kg/d faster (P less than .10) from 70 d to slaughter, weighed 3.9 kg more at 165 d of age (P less than .05), and reached 100 kg 7.0 d sooner (P less than .05) than pigs out of control-line sows. Carcasses from barrows sired by select-line boars had .29 cm more (P less than .10) fat at the 10th-rib than carcasses from barrows sired by control-line boars. Marbling scores were .31 unit greater (P less than .05) and muscle color scores were .25 unit greater (P less than .10) for carcasses from pigs out of select-line sows than for carcasses from pigs out of control-line sows. Selection for increased 70-d weight decreased age at 100 kg without increasing fat deposition. However, survival rates up to 100 kg were reduced. Mass selection for 200-d weight effectively increased 70-d weight, but fat thickness at 100 kg also increased.

Adipose Tissue↗

Response to 13 generations of selection for increased 8-week body weight in lines of mice carrying a sheep growth hormone-based transgene.

The purpose of this study was to evaluate selection in lines of transgenic mice. Two replicates of lines that either carried or did not carry the sheep metallothionein-1a sheep growth hormone transgene (oMt1a-oGH) were established. The host lines had been previously selected for rapid growth or selected randomly. Within-litter selection for increased 8-wk body weight was carried out for 13 generations. The frequency of oMt1a-oGH was monitored in all generations in the transgenic lines, but no genotypic information regarding the transgene was used as an aid to selection. The oMt1a-oGH was activated from weaning, at 3 wk, until 8 wk of age by adding ZnSO4 to the drinking water. Zinc stimulation of the transgene was not done during mating, gestation, or lactation. Data on body weights and weight gains were analyzed with a conventional mixed model and with an animal model. Genetic progress was achieved in all lines subjected to directional selection. In the control background, response to selection for 8-wk body weight was larger in the nontransgenic lines than in the transgenic lines, whereas no difference was found in the selected background. The frequency of the transgene was increased from the initial .5 to .62 in the randomly selected background but decreased to .04 in lines from a selected background. The REML estimates of variance components and genetic gain estimates varied greatly between the two methods. In general, there was better agreement between the realized heritability estimates and the heritability estimates obtained from the conventional mixed model analysis than between realized heritability estimates and results obtained using the animal model. Favorable correlated responses were obtained for 3- and 6-wk body weights and on 3- to 6- and 6- to 8-wk weight gains. Correlated responses to selection were larger in the selected than in the nonselected background but were not affected by the presence of the transgene. Results suggest that constructs similar to the oMt1a-oGH, which allow tight regulation, may be successfully incorporated into commercial livestock and should have larger effects in populations that have not been subject to selection.

Animals↗

Genomic regions involved in response to grain yield selection at high and low nitrogen fertilization in maize.

In order to validate the role of genomic regions involved in nitrogen use efficiency and detected in a population of recombinant inbred lines (RIL), we have applied from the same population a recurrent selection for adaptation to low N-input (N0) and to high N-input (N1). Variation of allele frequency at neutral marker during the two cycles of recurrent selection may provide information about markers linked to QTLs. Significant temporal variation of allele frequency was investigated using the test of Waples, which tests the hypothesis of genetic drift versus selection. Most genomic regions (12/19) responding to selection were detected for selection at high N-input and only two were common to selection at high and low N-inputs. This was consistent with the greater grain yield response to selection observed for the population selected under high N-input compared with the population selected under low N-input, when they were evaluated at high N-fertilization. In contrast, when they were evaluated at low N-input both types of selection gave similar yield. As was expected, in the first cycle we observed selection of markers linked to grain yield QTLs. In the course of the second cycle three situations were observed: the confirmation of most regions already selected in C1 including all C1 regions overlapping with grain yield QTLs; the non-confirmation of some C1 regions (2/9); and the identification of new genomic zones (10/17). The detected marker-QTL associations revealed the consistency of the involvement of some traits, such as root architecture and glutamine synthetase activity, which would be of major importance for grain yield setting whatever the nitrogen fertilization.

Alleles↗

Radical amino acid change versus positive selection in the evolution of viral envelope proteins.

To detect positive selection in protein-coding sequence evolution, the ratio of the nonsynonymous to synonymous substitution rate (K(A)/K(S)) is commonly used. When this ratio is higher than 1, positive selection on nonsynonymous changes is considered to have occurred. However, the question of what kinds of amino acid change are likely to be involved in positive selection has not been well studied, though intuitively it seems that radical changes frequently occur in positively selected changes. To address this question, we examined chemically radical and conservative replacements in the evolution of hepatitis C virus (HCV) protein sequences. In the envelope region, 34 positively and 440 negatively selected sites were identified by the K(A)/K(S) ratio. Radical and conservative changes were compared between the two types of selected sites using two methods. First, the numbers of radical and conservative replacements were counted at the positively and negatively selected sites according to three kinds of chemical classifications. In all three classifications, the resulting ratios of the two numbers were not statistically different for the two types of selected sites (P>0.05). Second, the distribution of chemical changes was compared between the two types of selected sites using two kinds of chemical distances. The distributions of the two chemical distances were not statistically different for the two types of selected sites (P>0.05). These results indicate that the ratio of chemically radical and conservative changes is similar for positively and negatively selected sites in the envelope protein of HCV or, in other words, there is no correlation between radical change and positive selection in the evolution of this protein.

Amino Acid Substitution↗

Change in sexual size dimorphism as a correlated response to selection on fecundity.

Fecundity selection is often suggested as the main causal factor underlying the prevalence of female-biased sexual size dimorphism (SSD), but this assumption has not been empirically tested. We selected female Drosophila melanogaster for increased or decreased fecundity (eggs laid over a single 18-h period, between days 5 and 7 posteclosion) for 20 generations, to see what effect this would have on SSD in three morphological traits (thorax width, abdomen width and thorax length). A direct response to fecundity selection was found in the downward direction (16.6%), whereas the response to upward selection (5.7%) was not statistically significant. Significant sex by selection interaction terms in the ANOVAs for thorax width and abdomen width indicate that the two sexes responded differently. Females usually showed a greater correlated response than males. In lines selected for increased fecundity, the correlated response in females for thorax and abdomen width was greater than the direct response in standard deviation units. SSD generally increased with selection for increased fecundity, but showed no consistent trend with selection for decreased fecundity. These results support the general hypothesis that SSD can evolve rapidly in response to fecundity selection. Selection on fecundity also produced correlated responses in life history traits. Downward selection resulted in flies that had lower viability and longevity, and both directions of selection were associated with an increase in development time.

Animals↗

Inference from clines stabilized by frequency-dependent selection.

Frequency-dependent selection against rare forms can maintain clines. For weak selection, s, in simple linear models of frequency-dependence, single locus clines are stabilized with a maximum slope of between square root of s/square root of 8 sigma and square root of s/square root of 12 delta, where sigma is the dispersal distance. These clines are similar to those maintained by heterozygote disadvantage. Using computer simulations, the weak-selection analytical results are extended to higher selection pressures with up to three unlinked genes. Graphs are used to display the effect of selection, migration, dominance, and number of loci on cline widths, speeds of cline movements, two-way gametic correlations ("linkage disequilibria"), and heterozygote deficits. The effects of changing the order of reproduction, migration, and selection, are also briefly explored. Epistasis can also maintain tension zones. We show that epistatic selection is similar in its effects to frequency-dependent selection, except that the disequilibria produced in the zone will be higher for a given level of selection. If selection consists of a mixture of frequency-dependence and epistasis, as is likely in nature, the error made in estimating selection is usually less than twofold. From the graphs, selection and migration can be estimated using knowledge of the dominance and number of genes, of gene frequencies and of gametic correlations from a hybrid zone.

Computer Simulation↗

Genetic and statistical analyses of strong selection on polygenic traits: what, me normal?

We develop a general population genetic framework for analyzing selection on many loci, and apply it to strong truncation and disruptive selection on an additive polygenic trait. We first present statistical methods for analyzing the infinitesimal model, in which offspring breeding values are normally distributed around the mean of the parents, with fixed variance. These show that the usual assumption of a Gaussian distribution of breeding values in the population gives remarkably accurate predictions for the mean and the variance, even when disruptive selection generates substantial deviations from normality. We then set out a general genetic analysis of selection and recombination. The population is represented by multilocus cumulants describing the distribution of haploid genotypes, and selection is described by the relation between mean fitness and these cumulants. We provide exact recursions in terms of generating functions for the effects of selection on non-central moments. The effects of recombination are simply calculated as a weighted sum over all the permutations produced by meiosis. Finally, the new cumulants that describe the next generation are computed from the non-central moments. Although this scheme is applied here in detail only to selection on an additive trait, it is quite general. For arbitrary epistasis and linkage, we describe a consistent infinitesimal limit in which the short-term selection response is dominated by infinitesimal allele frequency changes and linkage disequilibria. Numerical multilocus results show that the standard Gaussian approximation gives accurate predictions for the dynamics of the mean and genetic variance in this limit. Even with intense truncation selection, linkage disequilibria of order three and higher never cause much deviation from normality. Thus, the empirical deviations frequently found between predicted and observed responses to artificial selection are not caused by linkage-disequilibrium-induced departures from normality. Disruptive selection can generate substantial four-way disequilibria, and hence kurtosis; but even then, the Gaussian assumption predicts the variance accurately. In contrast to the apparent simplicity of the infinitesimal limit, data suggest that changes in genetic variance after 10 or more generations of selection are likely to be dominated by allele frequency dynamics that depend on genetic details.

Fourier Analysis↗

Population, evolutionary and genomic consequences of interference selection.

Weakly selected mutations are most likely to be physically clustered across genomes and, when sufficiently linked, they alter each others' fixation probability, a process we call interference selection (IS). Here we study population genetics and evolutionary consequences of IS on the selected mutations themselves and on adjacent selectively neutral variation. We show that IS reduces levels of polymorphism and increases low-frequency variants and linkage disequilibrium, in both selected and adjacent neutral mutations. IS can account for several well-documented patterns of variation and composition in genomic regions with low rates of crossing over in Drosophila. IS cannot be described simply as a reduction in the efficacy of selection and effective population size in standard models of selection and drift. Rather, IS can be better understood with models that incorporate a constant "traffic" of competing alleles. Our simulations also allow us to make genome-wide predictions that are specific to IS. We show that IS will be more severe at sites in the center of a region containing weakly selected mutations than at sites located close to the edge of the region. Drosophila melanogaster genomic data strongly support this prediction, with genes without introns showing significantly reduced codon bias in the center of coding regions. As expected, if introns relieve IS, genes with centrally located introns do not show reduced codon bias in the center of the coding region. We also show that reasonably small differences in the length of intermediate "neutral" sequences embedded in a region under selection increase the effectiveness of selection on the adjacent selected sequences. Hence, the presence and length of sequences such as introns or intergenic regions can be a trait subject to selection in recombining genomes. In support of this prediction, intron presence is positively correlated with a gene's codon bias in D. melanogaster. Finally, the study of temporal dynamics of IS after a change of recombination rate shows that nonequilibrium codon usage may be the norm rather than the exception.

Animals↗

Effect of long-term divergent selection on growth characteristics in Japanese quail.

The current study was undertaken to examine the effect of long-term selection for 4-wk BW on growth characteristics in divergent lines of Japanese quail and their control. Growth rate was significantly higher in males than females in all the lines. There was a significant increase in growth rate of the females selected for increased 4-wk BW over the control females, as well as a significant decline in growth rate of males selected for decreased 4-wk BW compared to their control counterparts. It appeared that selection for increased 4-wk BW did not alter the rate of growth in the males compared to the control males; however, in the females, selection for increased 4-wk BW resulted in an increase in growth rate. On the other hand, selection for decreased 4-wk BW resulted in a decline in growth rate in males but not in the females. The dynamics of the growth curve parameters indicate that selection for decreased 4-wk BW shifted the growth curve for females as well as altering the trajectories of growth in both sexes. However, selection for increased 4-wk BW only resulted in altering the trajectory of growth in the males. Long-term selection resulted in asymmetry of response in the low and high lines. In addition, different genes may respond differently to the same selection pressure in opposite directions. The use of the 4-parameter Richards model to analyze growth data from such an experiment provided a better understanding of how selection can alter the rate and trajectory of growth to affect the genetically determined growth potential of quail. Consequently selection for increased or decreased 4-wk BW affected the sexes differently.

Animals↗

Effect of forward and reverse selection for ethanol-induced locomotor response on other measures of ethanol sensitivity.

BACKGROUND: Ethanol sensitivity may be a predictor of genetic predisposition to ethanol abuse. To examine ethanol sensitivity in rodents, two lines of mice were bred in replicate for high (FAST-1 and -2) and low (SLOW-1 and -2) locomotor stimulant responses to ethanol. After large differences between the lines developed and further response to selection seemed to have arrested, reverse selection was initiated by breeding the slowest FAST mice with each other and the fastest SLOW mice with each other. The reverse-selected lines, named r-FAST and r-SLOW, were virtually equally sensitive to the stimulant effects of the selection dose of ethanol after 16 generations of reverse selection. METHODS: These experiments used this unique genetic model to examine two responses, putatively genetically correlated with sensitivity to ethanol stimulation: handling-induced convulsions during chronic ethanol withdrawal, and ethanol-induced hypothermia. Ethanol clearance, for which a small difference was previously found between the FAST and SLOW lines, was also examined. RESULTS: Handling-induced convulsions during chronic ethanol withdrawal were significantly greater in both FAST lines compared with both SLOW lines. Reverse selection eliminated the difference between the replicate 1 lines but did not alter the difference between the replicate 2 lines. Ethanol-induced hypothermia was greater in both SLOW lines compared with the FAST lines. This difference was significantly reduced by reverse selection in r-SLOW mice only. Ethanol clearance rates were similar among all lines and replicates. CONCLUSIONS: These data demonstrate the usefulness of reverse-selected lines for examining putatively genetically correlated traits. Changes in the correlated traits demonstrated the existence of persistent trait-relevant genetic heterogeneity and some genetic overlap between the correlated traits and the selection trait. Absence of a change after reverse selection suggests that trait-relevant genetic heterogeneity was eliminated by forward selection or, alternatively, that the trait was not influenced by genes associated with successful reverse selection.

Animals↗

Sexual selection and the risk of extinction in birds.

The relationship between sexual selection and extinction risk has rarely been investigated. This is unfortunate because extinction plays a key role in determining the patterns of species richness seen in extant clades, which form the basis of comparative studies into the role that sexual selection may play in promoting speciation. We investigate the extent to which the perceived risk of extinction relates to four different estimates of sexual selection in 1030 species of birds. We find no evidence that the number of threatened species is distributed unevenly according to a social mating system, and neither of our two measures of pre-mating sexual selection (sexual dimorphism and dichromatism) was related to extinction risk, after controlling for phylogenetic inertia. However, threatened species apparently experience more intense post-mating sexual selection, measured as testis size, than non-threatened species. These results persisted after including body size as a covariate in the analysis, and became even stronger after controlling for clutch size (two known correlates of extinction risk). Sexual selection may therefore be a double-edged process-promoting speciation on one hand but promoting extinction on the other. Furthermore, we suggest that it is post-mating sexual selection, in particular, that is responsible for the negative effect of sexual selection on clade size. Why this might be is unclear, but the mean population fitness of species with high intensities of post-mating sexual selection may be especially low if costs associated with multiple mating are high or if the selection load imposed by post-mating selection is higher relative to that of pre-mating sexual selection.

Animals↗

Correlated responses to selection for faster development and early reproduction in Drosophila: the evolution of larval traits.

Studies on selection for faster development in Drosophila have typically focused on the trade-offs among development time, adult weight, and adult life span. Relatively less attention has been paid to the evolution of preadult life stages and behaviors in response to such selection. We have earlier reported that four laboratory populations of D. melanogaster selected for faster development and early reproduction, relative to control populations, showed considerably reduced preadult development time and survivorship, dry weight at eclosion, and larval growth rates. Here we study the larval phase of these populations in greater detail. We show here that the reduction in development time after about 50 generations of selection is due to reduced duration of the first and third larval instars and the pupal stage, whereas the duration of the second larval instar has not changed. About 90% of the preadult mortality in the selected populations is due to larval mortality. The third instar larvae, pupae, and freshly eclosed adults of the selected populations weigh significantly less than controls, and this difference appears during the third larval instar. Thereafter, percentage weight loss during the pupal stage does not differ between selected and control populations. The minimum amount of time a larva must feed to subsequently complete development is lower in the selected populations, which also exhibit a syndrome of reduced energy expenditure through reduction in larval feeding rate, larval digging and foraging activity, and pupation height. Comparison of these results with those observed earlier in populations selected for adaptation to larval crowding and faster development under a different protocol from ours reveal differences in the evolved traits that suggest that the responses to selection for faster development are greatly affected by the larval density at which selection acts and on details of the selection pressures acting on the timing of reproduction.

Adaptation, Physiological↗

The impact of missing and erroneous genotypes on tagging SNP selection and power of subsequent association tests.

OBJECTIVE: Single nucleotide polymorphisms (SNPs) serve as effective markers for localizing disease susceptibility genes, but current genotyping technologies are inadequate for genotyping all available SNP markers in a typical linkage/association study. Much attention has recently been paid to methods for selecting the minimal informative subset of SNPs in identifying haplotypes, but there has been little investigation of the effect of missing or erroneous genotypes on the performance of these SNP selection algorithms and subsequent association tests using the selected tagging SNPs. The purpose of this study is to explore the effect of missing genotype or genotyping error on tagging SNP selection and subsequent single marker and haplotype association tests using the selected tagging SNPs. METHODS: Through two sets of simulations, we evaluated the performance of three tagging SNP selection programs in the presence of missing or erroneous genotypes: Clayton's diversity based program htstep, Carlson's linkage disequilibrium (LD) based program ldSelect, and Stram's coefficient of determination based program tagsnp.exe. RESULTS: When randomly selected known loci were relabeled as 'missing', we found that the average number of tagging SNPs selected by all three algorithms changed very little and the power of subsequent single marker and haplotype association tests using the selected tagging SNPs remained close to the power of these tests in the absence of missing genotype. When random genotyping errors were introduced, we found that the average number of tagging SNPs selected by all three algorithms increased. In data sets simulated according to the haplotype frequecies in the CYP19 region, Stram's program had larger increase than Carlson's and Clayton's programs. In data sets simulated under the coalescent model, Carlson's program had the largest increase and Clayton's program had the smallest increase. In both sets of simulations, with the presence of genotyping errors, the power of the haplotype tests from all three programs decreased quickly, but there was not much reduction in power of the single marker tests. CONCLUSIONS: Missing genotypes do not seem to have much impact on tagging SNP selection and subsequent single marker and haplotype association tests. In contrast, genotyping errors could have severe impact on tagging SNP selection and haplotype tests, but not on single marker tests.

Algorithms↗

Positive selection, relaxation, and acceleration in the evolution of the human and chimp genome.

For years evolutionary biologists have been interested in searching for the genetic bases underlying humanness. Recent efforts at a large or a complete genomic scale have been conducted to search for positively selected genes in human and in chimp. However, recently developed methods allowing for a more sensitive and controlled approach in the detection of positive selection can be employed. Here, using 13,198 genes, we have deduced the sets of genes involved in rate acceleration, positive selection, and relaxation of selective constraints in human, in chimp, and in their ancestral lineage since the divergence from murids. Significant deviations from the strict molecular clock were observed in 469 human and in 651 chimp genes. The more stringent branch-site test of positive selection detected 108 human and 577 chimp positively selected genes. An important proportion of the positively selected genes did not show a significant acceleration in rates, and similarly, many of the accelerated genes did not show significant signals of positive selection. Functional differentiation of genes under rate acceleration, positive selection, and relaxation was not statistically significant between human and chimp with the exception of terms related to G-protein coupled receptors and sensory perception. Both of these were over-represented under relaxation in human in relation to chimp. Comparing differences between derived and ancestral lineages, a more conspicuous change in trends seems to have favored positive selection in the human lineage. Since most of the positively selected genes are different under the same functional categories between these species, we suggest that the individual roles of the alternative positively selected genes may be an important factor underlying biological differences between these species.

Adaptation, Biological↗

Evaluation of expected response to selection for orthopedic health and performance traits in Hanoverian Warmblood horses.

OBJECTIVE: To determine whether selection schemes accounting for orthopedic health traits were compatible with breeding progress in performance parameters in Hanoverian Warmblood horses. ANIMALS: 5,928 horses. PROCEDURE: Relative breeding values (RBVs) were predicted for osseous fragments in fetlock (metacarpo- and metatarsophalangeal) and tarsal joints, deforming arthropathy in tarsal joints, and pathologic changes in distal sesamoid bones. Selection schemes were developed on the basis of total indices for radiographic findings (TIR), dressage (TID), and jumping (TIJ). Response to selection was traced over 2 generations of horses for dressage and jumping ability and all-purpose breeding. Development of mean RBVs and mean total indices in sires and prevalences of orthopedic health traits in their offspring were used to assess response to selection. RESULTS: Giving equal weight toTIR andTID, TIJ, or a combined index of 60% TID and 40% TIJ, 43% to 53% of paternal grandsires and 70% to 82% of descending sires passed selection. In each case, RBVs and total indices increased by as much as 9% in selected sires, when compared with all sires, and prevalences of orthopedic health traits in offspring of selected sires decreased relatively by as much as 16%. When selection was exclusively based on TID, TIJ, or TID and TIJ, percentages of selected sires were 44% to 66% in the first and 73% to 84% in the second generation and TID and TIJ increased by 9% to 10% and 19% to 23%, respectively. CONCLUSIONS AND CLINICAL RELEVANCE: Compared with exclusively performance-based selection, percentages of selected sires changed slightly and breeding progress in TID, TIJ, or TID and TIJ was only slightly decreased; however, prevalences of orthopedic health traits decreased in offspring of TIR-selected sires.

Animals↗

Results from nine generations of selection for increased litter size in swine.

Direct selection for increased litter size was done for nine generations. The select line consisted of approximately 15 sires and 60 dams per generation, and selection was based on estimated breeding values for number of live pigs. A control line of approximately 10 sires and 30 dams was maintained with stabilizing selection. Heritabilities estimated in the select line using restricted maximal likelihood procedures, daughter-dam regression within sires, and half-sib analysis were 0.01, 0.04, and 0.00 for number of pigs born alive (NBA) and 0.02, 0.16, and 0.00 for total born per litter (TB). Corresponding estimates for the control line were 0.01, 0.06, and 0.23 and 0.02, 0.07, and 0.09 for NBA and TB, respectively. Realized heritabilities for NBA from multiple regression were 0.09 +/- 0.08 in the select line and 0.11 +/- 0.166 in the control line. Heritability estimated from regression of differences in response between lines on differences in cumulative selection differentials was 0.13 +/- 0.07. At Generation 9, litter sizes, estimated breeding values, and cumulative selection differentials were 0.86 (P < 0.05), 0.63 (P < 0.01), and 9.05 (P < 0.01) pigs larger for the select line than for the control line. Phenotypic differences between lines for TB, adjusted backfat (BF), and days to 104 kg (DAYS) were not significant. Genetic trends in the select line were 0.053 +/- 0.002 pigs/yr for NBA, 0.054 +/- 0.013 mm/yr for BF, and 0.398 +/- 0.110 d/yr for DAYS. Corresponding phenotypic trends were 0.145 +/- 0.051 pigs/yr, -0.012 +/- 0.089 mm per yr, and 0.307 +/- 0.278 d/yr, respectively. Genetic trends in the control line were -0.026 +/- 0.004 pigs/yr for NBA, 0.026 +/- 0.022 mm/yr for BF, and -0.532 +/- 0.182 d/yr for DAYS. Corresponding phenotypic trends were 0.001 +/- 0.085 pigs/yr, -0.043 +/- 0.147 mm/yr, and -0.519 +/- 0.462 d/yr, respectively. Litter size can be increased by direct selection using breeding values estimated from an animal model, in conjunction with rearing selected gilts in litters of 10 pigs or less.

Adipose Tissue↗

Selection on mothers and offspring: whose phenotype is it and does it matter?

Reproductive and early life-history traits can be considered aspects of either offspring or maternal phenotype, and their evolution will therefore depend on selection operating through offspring and maternal components of fitness. Furthermore, selection at these levels may be antagonistic, with optimal offspring and maternal fitness occurring at different phenotypic values. We examined selection regimes on the correlated traits of birth weight, birth date, and litter size in Soay sheep (Ovis aries) using data from a long-term study of a free-living population on the archipelago of St. Kilda, Scotland. We tested the hypothesis that selective constraints on the evolution of the multivariate phenotype arise through antagonistic selection, either acting at offspring and maternal levels, or on correlated aspects of phenotype. All three traits were found to be under selection through variance in short-term and lifetime measures of fitness. Analysis of lifetime fitness revealed strong positive directional selection on birth weight and weaker selection for increased birth date at both levels. However, there was also evidence for stabilizing selection on these traits at the maternal level, with reduced fitness at high phenotypic values indicating lower phenotypic optima for mothers than for offspring. Additionally, antagonistic selection was found on litter size. From the offspring's point of view it is better to be born a singleton, whereas maternal fitness increases with average litter size. The decreased fitness of twins is caused by their reduced birth weight; therefore, this antagonistic selection likely results from trade-offs between litter size and birth weight that have different optimal resolutions with respect to offspring and maternal fitness. Our results highlight how selection regimes may vary depending on the assignment of reproductive and early life-history traits to either offspring or maternal phenotype.

Animals↗