Carlos López-Fanjul

Complutense University of Madrid, Madrid, Madrid, Spain

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Publications (45)140.05 Total impact

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    ABSTRACT: For a quantitative trait under stabilizing selection, the effect of epistasis on its genetic architecture and on the changes of genetic variance caused by bottlenecking were investigated using theory and simulation. Assuming empirical estimates of the rate and effects of mutations and the intensity of selection, we assessed the impact of two-locus epistasis (synergistic/antagonistic) among linked or unlinked loci on the distribution of effects and frequencies of segregating loci in populations at the mutation-selection-drift balance. Strong pervasive epistasis did not modify substantially the genetic properties of the trait and, therefore, the most likely explanation for the low amount of variation usually accounted by the loci detected in genome-wide association analyses is that many causal loci will pass undetected. We investigated the impact of epistasis on the changes in genetic variance components when large populations were subjected to successive bottlenecks of different sizes, considering the action of genetic drift, operating singly (D), or jointly with mutation (MD) and selection (MSD). An initial increase of the different components of the genetic variance, as well as a dramatic acceleration of the between-line divergence, were always associated with synergistic epistasis but were strongly constrained by selection. This article is protected by copyright. All rights reserved.
    Evolution 04/2014; DOI:10.1111/evo.12413 · 4.66 Impact Factor
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    C López-Fanjul, A García-Dorado
    Heredity 04/2011; 106(4):535-6. DOI:10.1038/hdy.2010.110 · 3.80 Impact Factor
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    R. G. Ruano, L. S. Silvela, C. Lopez-Fanjul, MA Toro
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    ABSTRACT: In Tribolium castaneum, a fitness-related trait was defined as the total weight loss (TWL) of a vial after 21 days, due to the activity of a single pair of parents. TWL showed considerable inbreeding depression and strong heterosis. A significant increase in the heritability of TWL was observed with inbreeding. The results are compatible with the genetic variation of TWL in the base population being generated by segregation of a number of recessive alleles at low frequencies.ResumenCambio de la varianza aditiva de un componente principal de la eficacia biológica en Tribolium castenum con consanguinidadElcarácter “merma” (TWL) se define en Tribolium castaneum como la pérdida de peso que experimenta un pocillo de cristal que contiene medio alimenticioyuna pareja de Padre, debida a la actividad de esa pareja durante 21 días. Es, por tanto, un importante componente de la eficacia biológica. El caracter mostró considerable depresión por consanguinidad, fuerte heterosis en cruzamientosy, además, un significativo incremento de su heredabilidad con el del coeficiente de consanguinidad. Estos resultados son compatibles con que la variación genética del carácter en la población base se debiera a la scgregación de alelos recesivos con bajas frecuencias iniciales.ZusammenfassungÄnderung der additiv-genetischen Varianz eines Fitness-Merkmals bei Inzucht von Tnboleum castaneumDas Fitness Merkmal wurde als Gewichtsverlust (TWL) eines Zuchtgefäßes mit Elternpaar und ihren Nachkommen in 21 Tagen definiert. TWL zeigte beträchtliche Inzuchtdepression und Heterosis. Durch Inzucht ergab sich eine signifikante Steigerung der additiv-genetischen Varianz. Die Ergebnisse machen die Annahme wahrscheinlich, daß in der Ausgangsgeneration die genetische Variabilität von TWL durch Spaltung rezessiver Allele geringer Häufigkeit verursacht wurde.
    Journal of Animal Breeding and Genetics 02/2011; 113(1‐6):93 - 97. DOI:10.1111/j.1439-0388.1996.tb00594.x · 2.06 Impact Factor
  • C. López-Fanjul
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    ABSTRACT: Fisher's theorem of natural selection implies that the population genetic variance of quasi-neutral traits should be mostly additive. In the case of fitness component traits, however, that variance would be characterised by a substantial contribution from non-additive loci. In parallel, Robertson's theorem states that selection will change the population mean of a trait proportionally to the magnitude of the genetic correlation between that trait and fitness, which should be weak for quasi-neutral traits or strong for the mean fitness components. Drosophila data from inbreeding and artificial selection experiments are discussed within that theoretical framework. In addition, the process of regeneration by mutation of the genetic variance of a quasi-neutral trait (abdominal bristle number) in a Drosophila population initially homozygous at all loci has been analysed. After 485 generations of mutation accumulation, the levels of additive variance found in this population closely approached those commonly observed in laboratory populations. Furthermore, these values, together with previously reported estimates for natural populations, could be jointly explained by a model assuming weak causal stabilising selection.
    01/2011; 1(03). DOI:10.1017/S2040470010005418
  • A. Caballero, C. López-Fanjul
    Journal of Animal Breeding and Genetics 04/2010; 104(1‐5):175 - 179. DOI:10.1111/j.1439-0388.1987.tb00121.x · 2.06 Impact Factor
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    ABSTRACT: In the C1 population of Drosophila melanogaster of moderate effective size ( approximately 500), which was genetically invariant in its origin, we studied the regeneration by spontaneous mutation of the genetic variance for two metric traits [abdominal (AB) and sternopleural (ST) bristle number] and that of the concealed mutation load for viability, together with their temporal stability, using alternative selection models based on mutational parameters estimated in the C1 genetic background. During generations 381-485 of mutation accumulation (MA), the additive variances of AB and ST approached the levels observed in standing laboratory populations, fluctuating around their expected equilibrium values under neutrality or under relatively weak causal stabilizing selection. This type of selection was required to simultaneously account for the observed additive variance in our population and for those previously reported in natural and laboratory populations, indicating that most mutations affecting bristle traits would only be subjected to weak selective constraints. Although gene action for bristles was essentially additive, transient situations occurred where inbreeding resulted in a depression of the mean and an increase of the additive variance. This was ascribed to the occasional segregation of mutations of large recessive effects. On the other hand, the observed non-lethal inbreeding depression for viability must be explained by the segregation of alleles of considerable and largely recessive deleterious effects, and the corresponding load concealed in the heterozygous condition was found to be temporally stable, as expected from tighter constraints imposed by natural selection.
    Genetics Research 04/2010; 92(2):91-102. DOI:10.1017/S001667231000011X · 2.20 Impact Factor
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    ABSTRACT: For different fitness mutational models, with epistasis introduced, we simulated the consequences of drift (D scenario) or mutation, selection, and drift (MSD scenario) in populations at the MSD balance subsequently subjected to bottlenecks of size N = 2, 10, 50 during 100 generations. No "conversion" of nonadditive into additive variance was observed, all components of the fitness genetic variance initially increasing with the inbreeding coefficient F and subsequently decreasing to zero (D) or to an equilibrium value (MSD). In the D scenario, epistasis had no appreciable effect on inbreeding depression and that on the temporal change of variance components was relevant only for high rates of strong epistatic mutation. In parallel, between-line differentiation in mean fitness accelerated with F and that in additive variance reached a maximum at F approximately 0.6-0.7, both processes being intensified by strong epistasis. In the MSD scenario, however, the increase in additive variance was smaller, as it was used by selection to purge inbreeding depression (N > or = 10), and selection prevented between-line differentiation. Epistasis, either synergistic or antagonistic (this leading to multiple adaptive peaks), had no appreciable effect on MSD results nor, therefore, on the evolutionary rate of fitness change.
    Genetics 07/2009; 183(1):299-313. DOI:10.1534/genetics.109.104893 · 4.87 Impact Factor
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    Carlos López-Fanjul, Almudena Fernández, Miguel A Toro
    Genetics 06/2007; 176(1):725-7. DOI:10.1534/genetics.106.062901 · 4.87 Impact Factor
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    ABSTRACT: The build up of an equilibrium between mutation, selection, and drift in populations of moderate size is an important evolutionary issue, and can be critical in the conservation of endangered populations. We studied this process in two Drosophila melanogaster populations initially lacking genetic variability (C1 and C2) that were subsequently maintained during 431 or 165 generations with effective population size N(e) approximately 500 (estimated by lethal complementation analysis). Each population originated synchronously to a companion set of full-sib mutation accumulation (MA) lines, C1 and MA1 were derived from an isogenic origin and C2 and MA2 from a single MA1 line at generation 265. The results suggest that both C1 and C2 populations were close to the mutation-selection-drift balance for viability and bristle traits, and are consistent with a 2.5-fold increase of the mutation rate in C2 and MA2. Despite this increase, the average panmictic viability in C2 was only slightly below that of C1, indicating that the expressed loads due to segregating deleterious mutation were small, in agreement with the low deleterious mutation rate (0.015-0.045) previously reported for the MA1 lines. In C1, the nonlethal inbreeding depression rate for viability was 30% of that usually estimated in segregating populations. The genetic variance for bristles regenerated in C1 and C2 was moderately smaller than the average value reported for natural populations, implying that they have accumulated a substantial adaptive potential. In light of neutral and selective predictions, these results suggest that bristle additive variance was predominantly due to segregation of mutations with deleterious effects of the order of 10(-3), and is consistent with relatively weak causal stabilizing selection (V(s) approximately 30).
    Evolution 04/2007; 61(3):653-65. DOI:10.1111/j.1558-5646.2007.00052.x · 4.66 Impact Factor
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    ABSTRACT: In a previous experiment, the effect of 255 generations of mutation accumulation (MA) on the second chromosome viability of Drosophila melanogaster was studied using 200 full-sib MA1 lines and a large C1 control, both derived from a genetically homogeneous base population. At generation 265, one of those MA1 lines was expanded to start 150 new full-sib MA2 lines and a new C2 large control. After 46 generations, the rate of decline in mean viability in MA2 was approximately 2.5 times that estimated in MA1, while the average degree of dominance of mutations was small and nonsignificant by generation 40 and moderate by generation 80. In parallel, the inbreeding depression rate for viability and the amount of additive variance for two bristle traits in C2 were 2-3 times larger than those in C1. The results are consistent with a mutation rate in the line from which MA2 and C2 were derived about 2.5 times larger than that in MA1. The mean viability of C2 remained roughly similar to that of C1, but the rate of MA2 line extinction increased progressively, leading to mutational collapse, which can be ascribed to accelerated mutation and/or synergy after important deleterious accumulation.
    Genetics 06/2006; 173(1):267-77. DOI:10.1534/genetics.106.056200 · 4.87 Impact Factor
  • Carlos López-Fanjul, Almudena Fernández, Miguel A Toro
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    ABSTRACT: The effect of population bottlenecks on the components of the genetic variance/covariance generated by n neutral independent additive x additive loci has been studied theoretically. In its simplest version, this situation can be modelled by specifying the allele frequencies and homozygous effects at each locus, and an additional factor measuring the strength of the n-th order epistatic interaction. The variance/covariance components in an infinitely large panmictic population (ancestral components) were compared with their expected values at equilibrium over replicates randomly derived from the base population, after t bottlenecks of size N (derived components). Formulae were obtained giving the derived components (and the between-line variance) as functions of the ancestral ones (alternatively, in terms of allele frequencies and effects) and the corresponding inbreeding coefficient F(t). The n-th order derived component of the genetic variance/covariance is continuously eroded by inbreeding, but the remaining components may increase initially until a critical F(t) value is attained, which is inversely related to the order of the pertinent component, and subsequently decline to zero. These changes can be assigned to the between-line variances/covariances of gene substitution and epistatic effects induced by drift. Numerical examples indicate that: (1) the derived additive variance/covariance component will generally exceed its ancestral value unless epistasis is weak; (2) the derived epistatic variance/covariance components will generally exceed their ancestral values unless allele frequencies are extreme; (3) for systems showing equal ancestral additive and total non-additive variance/covariance components, those including a smaller number of epistatic loci may generate a larger excess in additive variance/covariance after bottlenecks than others involving a larger number of loci, provided that F(t) is low. Our results indicate that it is unlikely that the rate of evolution may be significantly accelerated after population bottlenecks, in spite of occasional increments of the derived additive variance over its ancestral value.
    Journal of Theoretical Biology 04/2006; 239(2):161-71. DOI:10.1016/j.jtbi.2005.08.042 · 2.30 Impact Factor
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    ABSTRACT: The genetic variability for RAPDs band pattern was studied in a set of 157 mutation accumulation (MA) lines of Drosophila melanogaster. These MA lines were derived from the same isogenic base population and subsequently maintained by full-sib mating during 132 generations. The ancestral pattern of the original isogenic base can be unambiguously established as the consensus pattern of the MA lines and, because these lines are expected to be homozygous, dominance for band pattern is not a concern. Only repeatable changes in band pattern were considered. The number of ancestral bands detected implies that nine-nucleotide targets are enough for repeatable PCR amplification. Compared with the ancestral pattern, one MA line lost one band and two MA lines gained a new one. These results can be accounted for by the insertion of transposable elements occurring at a rate 0.07 < i < 0.21 per whole haploid genome and generation. This range is typical for Drosophila and consistent with the previously observed mobility for the roo family, supporting the generality of previous estimates of spontaneous mutation rates for morphological and fitness traits based on these MA lines. The sequence of one of the new bands suggests that the Idefix family is also active in the lines.
    Journal of Heredity 09/2005; 96(5):576-81. DOI:10.1093/jhered/esi076 · 1.97 Impact Factor
  • Carlos López-Fanjul, Almudena Fernández, Miguel A Toro
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    ABSTRACT: The effect of population bottlenecks on the components of the genetic covariance generated by two neutral independent epistatic loci has been studied theoretically (additive, covA; dominance, covD; additive-by-additive, covAA; additive-by-dominance, covAD; and dominance-by-dominance, covDD). The additive-by-additive model and a more general model covering all possible types of marginal gene action at the single-locus level (additive/dominance epistatic model) were considered. The covariance components in an infinitely large panmictic population (ancestral components) were compared with their expected values at equilibrium over replicates randomly derived from the base population, after t consecutive bottlenecks of equal size N (derived components). Formulae were obtained in terms of the allele frequencies and effects at each locus, the corresponding epistatic effects and the inbreeding coefficient Ft. These expressions show that the contribution of nonadditive loci to the derived additive covariance (covAt) does not linearly decrease with inbreeding, as in the pure additive case, and may initially increase or even change sign in specific situations. Numerical examples were also analyzed, restricted for simplicity to the case of all covariance components being positive. For additive-by-additive epistasis, the condition covAt > covA only holds for high frequencies of the allele decreasing the metric traits at each locus (negative allele) if epistasis is weak, or for intermediate allele frequencies if it is strong. For the additive/dominance epistatic model, however, covAt > covA applies for low frequencies of the negative alleles at one or both loci and mild epistasis, but this result can be progressively extended to intermediate frequencies as epistasis becomes stronger. Without epistasis the same qualitative results were found, indicating that marginal dominance induced by epistasis can be considered as the primary cause of an increase of the additive covariance after bottlenecks. For all models, the magnitude of the ratio covAt/covA was inversely related to N and t.
    Evolution 09/2004; 58(8):1655-63. DOI:10.1554/04-130 · 4.66 Impact Factor
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    Carlos López-Fanjul, Almudena Fernández, Miguel A Toro
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    ABSTRACT: For neutral additive genes, the quantitative index of population divergence (Q(ST)) is equivalent to Wright's fixation index (F(ST)). Thus, divergent or convergent selection is usually invoked, respectively, as a cause of the observed increase (Q(ST) > F(ST)) or decrease (Q(ST) < F(ST)) of Q(ST) from its neutral expectation (Q(ST) = F(ST)). However, neutral nonadditive gene action can mimic the additive expectations under selection. We have studied theoretically the effect of consecutive population bottlenecks on the difference F(ST) - Q(ST) for two neutral biallelic epistatic loci, covering all types of marginal gene action. With simple dominance, Q(ST) < F(ST) for only low to moderate frequencies of the recessive alleles; otherwise, Q(ST) > F(ST). Additional epistasis extends the condition Q(ST) < F(ST) to a broader range of frequencies. Irrespective of the type of nonadditive action, Q(ST) < F(ST) generally implies an increase of both the within-line additive variance after bottlenecks over its ancestral value (V(A)) and the between-line variance over its additive expectation (2F(ST)V(A)). Thus, both the redistribution of the genetic variance after bottlenecks and the F(ST) - Q(ST) value are governed largely by the marginal properties of single loci. The results indicate that the use of the F(ST) - Q(ST) criterion to investigate the relative importance of drift and selection in population differentiation should be restricted to pure additive traits.
    Genetics 08/2003; 164(4):1627-33. · 4.87 Impact Factor
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    ABSTRACT: Fecundity is usually considered as a trait closely connected to fitness and is expected to exhibit substantial nonadditive genetic variation and inbreeding depression. However, two independent experiments, using populations of different geographical origin, indicate that early fecundity in Drosophila melanogaster behaves as a typical additive trait of low heritability. The first experiment involved artificial selection in inbred and non-inbred lines, all of them started from a common base population previously maintained in the laboratory for about 35 generations. The realized heritability estimate was 0.151 +/- 0.075 and the inbreeding depression was very small and nonsignificant (0.09 +/- 0.09% of the non-inbred mean per 1% increase in inbreeding coefficient). With inbreeding, the observed decrease in the within-line additive genetic variance and the corresponding increase of the between-line variance were very close to their expected values for pure additive gene action. This result is at odds with previous studies showing inbreeding depression and, therefore, directional dominance for the same trait and species. All experiments, however, used laboratory populations, and it is possible that the original genetic architecture of the trait in nature was subsequently altered by the joint action of random drift and adaptation to captivity. Thus, we carried out a second experiment, involving inbreeding without artificial selection in a population recently collected from the wild. In this case we obtained, again, a maximum-likelihood heritability estimate of 0.210 +/- 0.027 and very little nonsignificant inbreeding depression (0.06 +/- 0.12%). The results suggest that, for fitness-component traits, low levels of additive genetic variance are not necessarily associated with large inbreeding depression or high levels of nonadditive genetic variance.
    Evolution 04/2003; 57(3):558-65. DOI:10.1111/j.0014-3820.2003.tb01547.x · 4.66 Impact Factor
  • Carlos López-Fanjul, Almudena Fernández, Miguel A Toro
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    ABSTRACT: The effect of population bottlenecks on the components of the genetic variance generated by two neutral independent epistatic loci has been studied theoretically (VA, additive; VD, dominant; VAA, additive x additive; VAD, additive x dominant; VDD; dominant x dominant components of variance). Nonoverdominance and overdominance models were considered, covering all possible types of marginal gene action at the single locus level. The variance components in an infinitely large panmictic population (ancestral components) were compared with their expected values at equilibrium, after t consecutive bottlenecks of equal size N (derived components). Formulae were obtained in terms of allele frequencies and effects at each locus and the corresponding epistatic value. An excess of VA after bottlenecks can be assigned to two sources: (1) the spatiotemporal changes in the marginal average effects of gene substitution alpha(i), which are equal to zero only for additive gene action within and between loci; and (2) the covariance between alpha2(i) and the heterozygosity at the loci involved, which is generated by dominance, with or without epistasis. Numerical examples were analyzed, indicating that an increase in VA after bottlenecks will only occur if its ancestral value is minimal or very small. For the nonoverdominance model with weak reinforcing epistasis, that increase has been detected only for extreme frequencies of the negative allele at one or both loci. With strong epistasis, however, this result can be extended to a broad range of intermediate frequencies. With no epistasis, the same qualitative results were found, indicating that dominance can be considered as the primary cause of an increase in VA following bottlenecks. In parallel, the derived total nonadditive variance exceeded its ancestral value (V(NA) = V(D) + V(AA) + V(AD) + V(DD)) for a range of combinations of allele frequencies covering those for an excess of VA and for very large frequencies of the negative allele at both loci. For the overdominance model, an increase in V(A) and V(NA) was respectively observed for equilibrium (intermediate) frequencies at one or both loci or for extreme frequencies at both loci. For all models, the magnitude of the change of V(A) and V(NA) was inversely related to N and t. At low levels of inbreeding, the between-line variance was not affected by the type of gene action. For the models considered, the results indicate that it is unlikely that the rate of evolution may be accelerated after population bottlenecks, in spite of occasional increments of the derived V(A) over its ancestral value.
    Evolution 06/2002; 56(5):865-76. DOI:10.1554/0014-3820(2002)056[0865:TEOEOT]2.0.CO;2 · 4.66 Impact Factor
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    D Chavarrías, C López-Fanjul, A García-Dorado
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    ABSTRACT: The effect of 250 generations of mutation accumulation (MA) on the second chromosome competitive viability of Drosophila melanogaster was analyzed both in homozygous and heterozygous conditions. We used full-sib MA lines, where selection hampers the accumulation of severely deleterious mutations but is ineffective against mildly deleterious ones. A large control population was simultaneously evaluated. Competitive viability scores, unaffected by the expression of mutations in heterozygosis, were obtained relative to a Cy/L(2) genotype. The rate of decline in mean DeltaM approximately 0.1% was small. However, that of increase in variance DeltaV approximately 0.08 x 10(-3) was similar to the values obtained in previous experiments when severely deleterious mutations were excluded. The corresponding estimates of the mutation rate lambda > or = 0.01 and the average effect of mutations E(s) < or = 0.08 are in good agreement with Bateman-Mukai and minimum distance estimates for noncompetitive viability obtained from the same MA lines after 105 generations. Thus, competitive and noncompetitive viability show similar mutational properties. The regression estimate of the degree of dominance for mild-to-moderate deleterious mutations was approximately 0.3, suggesting that the pertinent value for new unselected mutations should be somewhat smaller.
    Genetics 06/2001; 158(2):681-93. · 4.87 Impact Factor
  • Carlos López-Fanjul, Almudena Fernández, Miguel A. Toro
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    ABSTRACT: The effect of population bottlenecks on the mean and the additive variance generated by two neutral independent epistatic loci has been studied theoretically. Six epistatic models, used in the analysis of binary disease traits, were considered. Ancestral values in an infinitely large panmictic population were compared with their expectations at equilibrium, after t consecutive bottlenecks of equal size N (derived values). An increase in the additive variance after bottlenecks (inversely related to N and t) will occur only if the frequencies of the negative allele at each locus are: (1) low, invariably associated to strong inbreeding depression; (2) high, always accompanied by an enhancement of the mean with inbreeding. The latter is an undesirable property, making the pertinent models unsuitable for the genetic analysis of disease. For the epistatic models considered, it is unlikely that the rate of evolution may be accelerated after population bottlenecks, in spite of occasional increments of the derived additive variance over its ancestral value.
    Theoretical Population Biology 09/2000; 58(1):49-59. DOI:10.1006/tpbi.2000.1470 · 1.53 Impact Factor
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    ABSTRACT: In Tribolium castaneum (CS) and T. confusum (CF), intra- and interspecific rates of homosexual mounting have been measured. The intraspecific results are compatible with the hypothesis of both species being sexually indiscriminate. However, the CF intraspecific rates were very high (35%-53% of mountings were homosexual), suggesting a lower sexual attractiveness, or a stronger rejection to being mounted, of CF females relative to conspecific males. CS males discriminate between species but, in interspecific contacts, preferentially mounted CF males rather than CF females. CF males do not discriminate between species, but the loss of sexual attractiveness of CF females, or their rejection to being mounted, may act as a precopulatory isolation mechanism.
    Heredity 09/2000; DOI:10.1046/j.1365-2540.2000.00741.x · 3.80 Impact Factor
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    Aurora García-Dorado, Jesus Fernández, Carlos López-Fanjul
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    ABSTRACT: Spontaneous mutations were allowed to accumulate over 209 generations in more than 100 lines, all of them independently derived from a completely homozygous population of Drosophila melanogaster and subsequently maintained under strong inbreeding (equivalent to full-sib mating). Traits scored were: abdominal (AB) and sternopleural (ST) bristle number, wing length (WL) and egg-to-adult viability (V). On two occasions--early (generations 93-122) and late (generations 169-209)--ANOVA estimates of the mutational variance and the mutational line x generation interaction variance were obtained. Mutational heritabilities of morphological traits ranged from 2 x 10(-4) to 2 x 10(-3) and the mutational coefficient of variation of viability was 0.01. For AB, WL and V, temporal uniformity of the mutational variance was observed. However, a fluctuation of the mutational heritability of ST was detected and could be ascribed to random genotype x environment interaction.
    Genetics Research 03/2000; 75(1):47-51. DOI:10.1017/S0016672399004267 · 2.20 Impact Factor

Publication Stats

826 Citations
140.05 Total Impact Points

Institutions

  • 1979–2014
    • Complutense University of Madrid
      • • Department of Genetics
      • • Department of Animal Production
      • • Facultad de Ciencias Biológicas
      Madrid, Madrid, Spain
  • 1992
    • University of Oviedo
      • Department of Functional Biology
      Oviedo, Asturias, Spain