Experimental Evolution of a Pheromone Signal

ABSTRACTSexual signals are important in speciation, but understanding their evolution is complex as these signals are often composed of multiple, genetically interdependent components. To understand how signals evolve, we thus need to consider selection responses in multiple components and account for the genetic correlations among components. One intriguing possibility is that selection changes genetic correlations between components under selection and those not under selection, reducing evolutionary constraints. However, this hypothesis remains largely untested empirically. In this study, we investigate the evolutionary response of the multi-component female sex pheromone blend of the moth Heliothis subflexa to 10 generations of artificial selection. We observed a selection response of about 3/4s of a phenotypic standard deviation in the components under selection. Interestingly, other pheromone components that are biochemically and genetically linked to the components under selection did not change. We found that this component-specific selection response was likely facilitated due to reduced genetic coupling between the component under selection and the components not under selection, resulting from changes in genetic covariances within the 10 generations of selection. Our findings provide rare empirical support for an intriguing mechanism by which a sexual signal can respond to selection without possible constraints from indirect selection responses.SUMMARY STATEMENTThis study reveals a mechanism that can facilitate evolution of sex pheromone components under selection independent of other components of the sex pheromone that are not under selection.

Detection of chromosomal segments underlying scrotal circumference in ram lambs and age at onset of puberty in ewe lambs

Scrotal circumference (SC) is highly correlated with total sperm production and quality of the produced semen. In ewe lambs, puberty is an important reproductive trait and extreme delay in reaching puberty will have a negative effect on breeding efficiency. To identify genomic regions (QTL) underlying variation in SC and age at onset of puberty in ewe lambs (AP), a whole genome scan was performed with 169 microsatellites covering the ovine autosomes. Progeny (360 animals) from six half-sib families in a population of Kermani sheep, an indigenous fat tailed sheep breed in south-east of Iran, were recorded for SC and AP. Linkage analysis using the interval mapping approach indicated a total of 21 QTL for the studied traits on nine chromosomes (OAR1, 2, 3, 5, 6, 9, 11, 18 and 24), including a 5% genome wide significant QTL for SC on OAR5. There was evidence for the overlap of QTL affecting SC and AP on OAR5 and OAR9. The size of the QTL effect ranged from 0.8 to 1.6 in units of the phenotypic standard deviation. Although QTL for testis size in ram lambs have been mapped, this is the first report of QTL for age at onset of puberty in ewe lambs. Pursuing the underlying causes of these QTL may shed light on the molecular aspects of male and female reproductive development in sheep.

Genetics of early conception and its relationship to growth traits in red deer (Cervus elaphus)

The genetics of early conception success and live-animal growth were studied in five herds of red deer in New Zealand. Conception date (CD) was used as the criterion of success in seasonally mated hinds, with 2493 mating records available. Liveweights analysed were weaning weight, yearling weight, 15-month weight and mature weight (hinds only). CD and liveweights were analysed using restricted maximum likelihood procedures with an animal model, including all available pedigree records. Under the management conditions applied, CD had a phenotypic standard deviation of 7.9 days, a repeatability across years of 0.29 ± 0.03 and a direct (univariate) heritability of 0.20 ± 0.06. Regression procedures using DNA markers to adjust the data for genetic differences resulting from an animal’s ancestral region of origin (mainly western vs eastern European) had little effect on the parameter estimates above. Direct heritability estimates for the four weight traits were 0.38 ± 0.03, 0.49 ± 0.02, 0.48 ± 0.04 and 0.46 ± 0.04, respectively, while the genetic correlations between CD and these traits (e.g. using 1763 paired records of CD with yearling weight) were –0.24 ± 0.11, –0.24 ± 0.09, –0.16 ± 0.10 and –0.04 ± 0.09, respectively. Selection for earlier CD would be successful and compatible with selection for higher juvenile weights.

Detecting quantitative trait loci affecting beef tenderness on bovine chromosome 7 near calpastatin and lysyl oxidase

From a study of 3 large half-sib families of cattle, we describe linkage between DNA polymorphisms on bovine chromosome 7 and meat tenderness. Quantitative trait loci (QTL) for Longissimus lumborum peak force (LLPF) and Semitendonosis adhesion (STADH) were located to this map of DNA markers, which includes the calpastatin (CAST) and lysyl oxidase (LOX) genes. The LLPF QTL has a maximum lodscore of 4.9 and allele substitution of approximately 0.80 of a phenotypic standard deviation, and the peak is located over the CAST gene. The STADH QTL has a maximum lodscore of 3.5 and an allele substitution of approximately 0.37 of a phenotypic standard deviation, and the peak is located over the LOX gene. This suggests 2 separate likelihood peaks on the chromosome. Further analyses of meat tenderness measures in the Longissimus lumborum, LLPF and LL compression (LLC), in which outlier individuals or kill groups are removed, demonstrate large shifts in the location of LLPF QTL, as well as confirming that there are indeed 2 QTL on bovine chromosome 7. We found that both QTL are reflected in both LLPF and LLC measurements, suggesting that both these components of tenderness, myofibrillar and connective tissue, are detected by both measurements in this muscle.

Genetic response to selection on reindeer calf weights

<p>Selection response was investigated after 11 years of selection in the herding district of Ruvhten Sijte in Sweden (63&deg;N,12&deg;E) by comparing the weights of calves owned by herders applying selection with the weights of calves owned by herders not applying selection. Three owner groups in Ruvhten Sijte have selected calves at the autumn slaughter every year since 1986 by retaining heavy calves. They own approximately 45% of the reindeer in Ruvhten Sijte. The other owners do not select on recorded weights, and all animals mix during the mating season. The recorded data on calf weights (1986 to 1997) consisted of 12 484 observations, with mean selection differentials of 2.9 kg and 4.6 kg for female and male calves, respectively (phenotypic standard deviation equal to 4.4 kg). The female generation interval was 6.4 years. In 1996 and 1997, autumn live weights of calves owned by herders applying selection (n = 971) were compared with the weights of calves owned by herders not applying selection (n = 1125). The observed difference was 0.67 kg (total mean = 42.6 kg), of which 0.32 kg was due to differences in dam age structure between the two types of herds, therefore the estimated difference in selection response was 0.35 kg. The realized heritability was 0.2. The potential response that would have been realized if the selected part of the herd had been isolated from gene flow was predicted to be 2.0 kg. In conclusion, the introduction of a selection programme applied by all owners in a herding district would result in an increase in calf weights of approximately 2 kg after two generations of selection.</p><p>&nbsp;</p>

A terminal-sire index for selecting rams

A terminal-sire index for selecting rams was developed. It combines genetic evaluations for growth traits and carcass characteristics measured ultrasonically on live animals into a single criterion. Weightings for component traits are averages from the indexes of four slightly different breeding goals, determined using economic values and parameters from the literature. The weightings for breeding values of component traits are -1.45 for birth weight, +1.86 for weight at 50 d of age, +2.27 for gain from 50 to 100 d, -0.51 for ultrasonic fat depth, and +1.36 for ultrasonic loin muscle depth, in phenotypic standard deviation units. Selection that is based on the index will increase growth while simultaneously decreasing fat and increasing muscle. Key words: Breeding strategies, carcass characteristics, growth, selection, sheep

Pleiotropic Model of Maintenance of Quantitative Genetic Variation at Mutation-Selection Balance

A pleiotropic model of maintenance of quantitative genetic variation at mutation-selection balance is investigated. Mutations have effects on a metric trait and deleterious effects on fitness, for which a bivariate gamma distribution is assumed. Equations for calculating the strength of apparent stabilizing selection (Vs) and the genetic variance maintained in segregating populations (VG) were derived. A large population can hold a high genetic variance but the apparent stabilizing selection may or may not be relatively strong, depending on other properties such as the distribution of mutation effects. If the distribution of mutation effects on fitness is continuous such that there are few nearly neutral mutants, or a minimum fitness effect is assumed if most mutations are nearly neutral, VG increases to an asymptote as the population size increases. Both VG and Vs are strongly affected by the shape of the distribution of mutation effects. Compared with mutants of equal effect, allowing their effects on fitness to vary across loci can produce a much higher VG but also a high Vs (Vs in phenotypic standard deviation units, which is always larger than the ratio VP/Vm), implying weak apparent stabilizing selection. If the mutational variance Vm is ∼10-3 Ve (Ve, environmental variance), the model can explain typical values of heritability and also apparent stabilizing selection, provided the latter is quite weak as suggested by a recent review.

Multiple Quantitative Trait Locus Analysis of Bovine Chromosome 6 in the Israeli Holstein Population by a Daughter Design

Nine Israeli Holstein sire families with 2978 daughters were analyzed for quantitative trait loci effects on chromosome 6 for five milk production traits by a daughter design. All animals were genotyped for 2 markers. The three families with significant effects were genotyped for up to 10 additional markers spanning positions 0–122 cM of BTA6. Two sires were segregating for a locus affecting protein and fat percentage near position 55 cM with an estimated substitution effect of 0.18% protein, which is equivalent to one phenotypic standard deviation. This locus was localized to a confidence interval of 4 cM. One of these sires was also heterozygous for a locus affecting milk, fat, and protein production near the centromere. The hypothesis of two segregating loci was verified by multiple regression analysis. A third sire was heterozygous for a locus affecting milk and protein percentage near the telomeric end of the chromosome. Possible candidates for the major quantitative gene near position 55 cM were determined by comparative mapping. IBSP and SSP1 were used as anchors for the orthologous region on human chromosome 4. Twelve genes were detected within a 2-Mbp sequence. None of these genes have been previously associated with lactogenesis.

Response to recurrent selection under small effective population size

A formula was derived for the prediction of the response to recurrent selection when the effective population size (Ne) is small. Usually, responses to selection have been estimated by Rs = icsigma²A/sigmaPh, where i, c, sigma²A, and sigmaPh stand for standardized selection differential, parental control, additive variance, and phenotypic standard deviation, respectively. This expression, however, was derived under the assumption of infinite population size. By introducing the effects of finite population size, the expression derived was Rs = [ic(sigma²A + deltaFD1)/sigmaPh] - DFID, where deltaF, ID and D1 are the changes in the inbreeding coefficient, the inbreeding depression, and the covariance of additive and homozygous dominance effects, respectively. Thus, the predicted responses to selection based on these expressions will be smaller than those based on the standard procedures for traits with a high level of dominance such as yield. Responses to five cycles of half-sib selection were predicted for maize by both expressions, considering that 100 progenies were evaluated and 10 S1 progenies were recombined, which corresponds to Ne = 10 for each cycle. The accumulated response to selection estimated with the new expression was about 47 and 28% smaller than that based on the standard expression for yield and plant height, respectively. Thus, the expression usually used overestimates the responses to selection, which is in agreement with reported results, because it does not take into account the effective population size that is generally small in recurrent selection programs

Continued selection of Romney sheep for resistance or susceptibility to nematode infection: estimates of direct and correlated responses

Divergent breeding lines of Romney sheep, selected as lambs for consistently high or low faecal worm egg count (FEC) following natural multi-species challenge by nematode parasites, were established in New Zealand at Wallaceville Animal Research Centre in 1979 and at Rotomahana Station in 1985. In 1988 the Rotomahana lines, including an unselected control line maintained under the same management conditions, were transferred to Tokanui Station where they remained for 4 years. In 1993 elite high and low FEC animals from Tokanui, along with the controls, were transferred to Wallaceville, where merged lines have since been managed together. Selection responses from the lines at Rotomahana and Tokanui, and from a further 5 years of divergent selection in the merged lines, are reported here. For the two most recent lamb crops (1996 and 1997 birth years), log-transformed FECs of the high and low lines were 1·27 and -1·46 phenotypic standard deviation units from the control. After back-transformation to the original scale, where the FEC for control line lambs averaged 1255 eggs per g, the means for the high and low lines were 3Ό5 and 0·27 times the control mean. Animal-model restricted maximum likelihood estimates of her it ability and repeatability for single-record FEC (following separate infections) were 0·28 (s.e. 0·02) and 0·42 (s.e. 0Ό1), respectively. Correlated responses in production traits include significantly decreased post-weaning weight gain and increased dags (breech soiling) in lambs, and decreased fleece weight in yearlings and ewes in the low FEC line, compared with those in the high line. However the low FEC line had proportionally 0·11 more lambs weaned per ewe mated than the high FEC line (F < 0·01). It is concluded firstly that selection for high or low FEC in Romney s has achieved an 11-fold difference between the divergent lines. Secondly, it will generally be necessary in a commercial environment to apply index selection for a combination of increased productivity, decreased FEC and possibly decreased dags, when potential candidates are recorded under conditions of nematode challenge.