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A model is described for the biometrical analysis of interaction behavioural characteristics. The model is illustrated by analysis of the mating speed of Drosophila pseudoobscura derived from crosses of fast and of slow mating strains generated through artificial selection.
We measured the effect of larval density on thorax length, development time, sex ratio and a measure of total fitness, using strains of Drosophila melanogaster artificially selected for increased thorax length, control lines otherwise cultured in an identical way, and the base stock from which the lines had been derived. We used the addition experimental design (Mather & Caligari, 1981). No genotype-environment interaction was observed when comparing the reduction in thorax length of 'large' and 'control' lines with increasing larval density for any culture series, i.e. rank ordering of genotypes and additive genetic variances remained the same in all the environments tested. In contrast, the reduction in thorax length for the base stock as density increased was proportionally smaller than that of the 'large' and 'control' lines. Development time increased more rapidly with larval density in the 'large' lines than in the 'controls' or base stock. Sex ratio was unaffected by larval density but thorax length and the development time of females were more affected than those of males by increasing larval density. The estimate of total fitness showed clear evidence of gene-environment interaction for the effect of body size on fitness, with genetically large individuals at an increasing disadvantage with increasing larval density.
Although the genetic basis of multiple disease resistance (MDR) is poorly understood, it is of great value for understanding the evolution of disease resistance in natural plant populations and for increasing crop yields in agriculture. In Brassica rapa, we studied genetic correlations among levels of disease resistance to three fungal pathogens: Peronospora parasitica, Albugo candida and Leptosphaeria maculans. A large, replicated quantitative genetics experiment used artificial selection on resistance to individual pathogens, and examined correlated responses to selection for resistance to other, unselected pathogens. Data from 9518 plants, each measured simultaneously for resistance to three fungal pathogens, showed heritable genetic variation for resistance to each pathogen and a positive genetic correlation between resistance to P. parasitica and L. maculans. This indicates that some resistance genes provide defence against fundamental characteristics common to two taxonomic orders of fungal pathogens. Conceivably, such MDR could contribute to a durable defence that might not be easily circumvented by rapidly evolving fungal pathogens.
The fruitfly, Drosophila melanogaster, can be transfected with P-elements and induced to overexpress a transfected gene whose impact on lifespan can be measured. Here, it is reported that in previous experiments a transfected gene was not expressed. This suggested a new statistical analysis indicating that (1) the size of the insert, the position and the interactions of the insert with the genetic backgrounds into which the P-element are inserted have effects on lifespan similar to those attributed to overexpression; (2) these effects occur without expression of the transfected gene; and (3) effects of interactions with backgrounds and effects of positions are as large as responses to six generations of strong directional artificial selection. Reports of effects of overexpression of transfected genes on lifespan in Drosophila melanogaster may be experimental artefacts. Credible experiments on the phenotypic effects of transgenesis need proper controls for the effects of insert size and position and should estimate the magnitudes of interactions of treatment with genetic backgrounds.
Cryptic genetic variation is the dark matter of biology: it is variation that is not normally seen, but that might be an essential source of physiological and evolutionary potential. It is uncovered by environmental or genetic perturbations, and is thought to modify the penetrance of common diseases, the response of livestock and crops to artificial selection and the capacity of populations to respond to the emergence of a potentially advantageous macro-mutation. We argue in this review that cryptic genetic variation is pervasive but under-appreciated, we highlight recent progress in determining the nature and identity of genes that underlie cryptic genetic effects and we outline future research directions.
Substantial advances have been made in the genetic improvement of agriculturally important animal and plant populations through artificial selection on quantitative traits. Most of this selection has been on the basis of observable phenotype, without knowledge of the genetic architecture of the selected characteristics. However, continuing molecular genetic analysis of traits in animal and plant populations is leading to a better understanding of quantitative trait genetics. The genes and genetic markers that are being discovered can be used to enhance the genetic improvement of breeding stock through marker-assisted selection.
Individuals of all vertebrate species differ consistently in their reactions to mildly stressful challenges. These typical reactions, described as personalities or coping strategies, have a clear genetic basis, but the structure of their inheritance in natural populations is almost unknown. We carried out a quantitative genetic analysis of two personality traits (exploration and boldness) and the combination of these two traits (early exploratory behaviour). This study was carried out on the lines resulting from a two-directional artificial selection experiment on early exploratory behaviour (EEB) of great tits (Parus major) originating from a wild population. In analyses using the original lines, reciprocal F(1) and reciprocal first backcross generations, additive, dominance, maternal effects ands sex-dependent expression of exploration, boldness and EEB were estimated. Both additive and dominant genetic effects were important determinants of phenotypic variation in exploratory behaviour and boldness. However, no sex-dependent expression was observed in either of these personality traits. These results are discussed with respect to the maintenance of genetic variation in personality traits, and the expected genetic structure of other behavioural and life history traits in general.
The amplification of genes encoding an insecticide-detoxifying esterase (E4) in the peach-potato aphid Myzus persicae is one of the few examples where this genetic phenomenon has been shown to be involved in the response of an intact higher organism to artificial selection. Here we report quantitative and qualitative studies of the repeat units (amplicons) containing the E4 genes in a highly resistant aphid clone. Initial studies to quantify esterase sequences showed a 5-11-fold increase in resistant aphids compared with susceptible aphids, suggesting the presence of 10-22 gene copies per diploid genome. A more incisive analysis by pulsed-field gel electrophoresis confirmed the presence of about 12 copies of the E4 gene and showed them to be on about 24 kb amplicons, arranged as a tandem array of direct repeats. This, together with previous results from crossing experiments and with recent in situ hybridization studies, confirms that the E4 gene amplification in this aphid clone is heterozygous at a single locus. However, these data show that the gene amplification alone cannot account for the approx. 60 times higher levels of E4 protein and its mRNA present in this aphid clone, and therefore resistance must involve changes in both esterase gene copy number and gene expression.
Crop plants were domesticated by prehistoric farmers through artificial selection to provide a means of feeding the human population. This article discusses the developmental genetics of crop domestication and improvement, including the historical framework and recent approaches in maize and other grasses. In many cases, selecting for a plant form that correlates with productivity involves controlling meristem activity. In the domestication of modern maize from its progenitor Zea mays ssp. parviglumis, QTL (quantitative trait loci) mapping, genetics and population genomics approaches have identified several genes that contain signatures of selection. Only a few genes involved in the derivation of the highly productive maize ear have been identified, including teosinte glume architecture1 and ramosa1. Future prospects hinge on forward and reverse genetics, as well as on other approaches from the developing discipline of evo-devo (evolutionary developmental biology).
Genetic variabilities within and between Finnish populations of Golden Retrievers, German Shepherds, Wirehaired Dachshunds, Pembroke Welsh Corgis and Bedlington Terriers were quantified with microsatellite allele numbers, observed heterozygosities, expected heterozygosities, FIS estimates, FST estimates and DS distances. In a sample of 50 individuals from each, breed and ten polymorphic loci, the highest genetic diversity was exhibited in the Wirehaired Dachshunds and lowest in the Bedlington Terriers. Although statistically significant deviations from the Hardy-Weinberg (H-W) equilibrium were observed, they occurred at an unexpectedly low frequency. Most strikingly, the extremely small Bedlington Terrier population displayed genotypes in H-W proportions in all investigated loci. The H-W deviations always occurred with positive FIS estimates, which, on average, were not larger than values reported for free-living canids. Genetic differentiation between the breeds was very large. As a comparison, present estimates were, on average, over two times higher than previously observed between breeds of sheep, and over two times higher than the highest estimates reported between human populations. Moreover, the highest DS distances were only slightly lower than the lowest values inferred between humans and chimpanzees. Severe bottlenecks in the recent past of the examined breeds were not statistically supported. The presented data imply genetic isolation and intense artificial selection in the history of these breeds of dogs.
Parasitoids are the most important natural enemies of many insect species. Larvae of many Drosophila species can defend themselves against attack by parasitoids through a cellular immune response called encapsulation. The paper reviews recent studies of the evolutionary biology and ecological genetics of resistance in Drosophila, concentrating on D. melanogaster. The physiological basis of encapsulation, and the genes known to interfere with resistance are briefly summarized. Evidence for within- and between-population genetic variation in resistance from isofemale line, artificial selection and classical genetic studies are reviewed. There is now firm evidence that resistance is costly to Drosophila, and the nature of this cost is discussed, and the possibility that it may involve a reduction in metabolic rate considered. Comparative data on encapsulation and metabolic rates across seven Drosophila species provides support for this hypothesis. Finally, the possible population and community ecological consequences of evolution in the levels of host resistance are examined.
Drosophila melanogaster is attacked by parasitoids that develop internally in the larva. They can defend themselves by a cellular immune response (host resistance), although this can be disabled by parasitoid countermeasures (parasitoid virulence). D. melanogaster and its parasitoids are an excellent system in which to study coevolution experimentally. We designed an experiment to compare changes in resistance and virulence in replicate populations of flies and parasitoids maintained together for approximately 10 fly (five parasitoid) generations. The experiment had three treatments each with three replicates: (A) no parasitoids (B) outbred parasitoids (C) partially inbred parasitoids. Host resistance increased in treatments B and C but there was no difference between these treatments. Parasitoid virulence appeared not to change during the experiment. Host larvae in treatments B and C fed at lower rates than those in A, evidence of a trade-off between resistance and larval competitive ability. We found no evidence for local adaptation, as hosts from the different replicates of treatment C performed no differently against parasitoids from the same and other replicates. Also, we found no evidence for the evolution of behavioural traits in the host that could lead to lower probabilities of being attacked. Comparing the evolution of host resistance in these seminatural settings with that in artificial selection experiments provides insight into how the conflicting selection pressures on host resistance interact.
The length of the female's primary sperm-storage organ, the seminal receptacle, has undergone rapid divergence within the Drosophila genus. Quantitative genetic analysis of seminal receptacle length was carried out on two laboratory strains of Drosophila melanogaster that had undergone artificial selection for both increased and decreased organ length. Realized heritabilities were 0.176 and 0.270 for the two experiments. Parental strains, F1, F1r (reciprocal), F2, backcross, and backcross reciprocal generations were used in a line-cross (generation means) analysis. This analysis revealed that additive, dominance, and additive-by-dominance epistasis contributed significantly to the means. No significant maternal effects were found. Variance analysis indicated that a completely additive model was adequate to explain the variances observed in these lines. Castle-Wright minimal estimates of 5.25 and 1.91, segregating loci responsible for mean differences, were found for the two respective experiments. There were significant positive correlations between additive effects of seminal receptacle length and thorax length in both experiments. The correlated evolution of sperm and seminal receptacle length is discussed.
In many organisms metamorphosis allows for an ecologically important habitat-shift from water to land. However, in some salamanders an adaptive life cycle mode has evolved that is characterized by metamorphic failure (paedomorphosis); these species remain in the aquatic habitat throughout the life cycle. Perhaps the most famous example of metamorphic failure is the Mexican axolotl (Ambystoma mexicanum), which has become a focal species for developmental biology since it was introduced into laboratory culture in the 1800s. Our previous genetic linkage mapping analysis, using an interspecific crossing design, demonstrated that a major gene effect underlies the expression of metamorphic failure in laboratory stocks of the Mexican axolotl. Here, we repeated this experiment using A. mexicanum that were sampled directly from their natural habitat at Lake Xochimilco, Mexico. We found no significant association between the major gene and metamorphic failure when wild-caught axolotls were used in the experimental design, although there is evidence of a smaller genetic effect. Thus, there appears to be genetic variation among Mexican axolotls (and possibly A. tigrinum tigrinum) at loci that contribute to metamorphic failure. This result suggests a role for more than one mutation and possibly artificial selection in the evolution of the major gene effect in the laboratory Mexican axolotl.
Ectoparasites are abundant in natural communities, can have pronounced deleterious fitness consequences to their host and are important vectors of transmissible parasitic disease. Yet very few studies have estimated the magnitude of heritable genetic variation underlying resistance against ectoparasitism, which significantly limits our ability to predict the evolution of this ecologically important character. The present paper reports results of artificial selection for increased resistance in Drosophila nigrospiracula against ectoparasitic, haematophagous mites, Macrocheles subbadius. In this system, which occurs naturally in the Sonoran Desert of North America, ectoparasitism significantly damages the expression of host fitness traits, including longevity, fecundity and male mating success. In the present study, resistance, which was modelled as a threshold trait, responded significantly to selection applied on either sex. Realized heritability, calculated as a mean across four replicates, was estimated to be 0.152 +/- 0.014 (SE). The heritability estimate from selection on males did not differ from that on females, but both estimates differed significantly from zero. This documented presence of additive genetic variation for resistance, coupled with knowledge of the fitness consequences of ectoparasitism, indicates that the host population possesses significant evolutionary potential. Selection was applied on the pre-attachment phase, thereby targeting behavioural forms of defence. This study therefore establishes parallels between insects and other animals in their ability to protect themselves and evolve behavioural defences against ectoparasites.
One of the most striking examples of plant hairs is the single-celled epidermal seed trichome of cultivated cotton. The developmental morphology of these commercial "fibers" has been well-characterized in Gossypium hirsutum, but little is known about the pattern and tempo of fiber development in wild Gossypium species, all of which have short, agronomically inferior fiber. To identify developmental differences that account for variation in fiber length, and to place these differences in a phylogenetic context, we conducted SEM studies of ovules at and near the time of flowering, and generated growth curves for cultivated and wild diploid and tetraploid species. Trichome initiation was found to be similar in all taxa, with few notable differences in trichome density or early growth. Developmental profiles of the fibers of most wild species are similar, with fiber elongation terminating at about two weeks post-anthesis. In contrast, growth is extended to three weeks in the A- and F-genome diploids. This prolonged elongation period is diagnosed as a key evolutionary event in the origin of long fiber. A second evolutionary innovation is that absolute growth rate is higher in species with long fibers. Domestication of species is associated with a further prolongation of elongation at both the diploid and allopolyploid levels, suggesting the effects of parallel artificial selection. Comparative analysis of fiber growth curves lends developmental support to previous quantitative genetic suggestions that genes for fiber "improvement" in tetraploid cotton were contributed by the agronomically inferior D-genome diploid parent.
STATEMENT OF PROBLEM: There is limited scientific information on the relationship between tooth shade and skin color. This lack of knowledge may impact the ability of the prosthodontist to select artificial teeth that complement the facial complexion of the patient. PURPOSE: This observational study explored the possibility of a relationship between tooth shade and skin color. MATERIALS AND METHODS: A total of 119 individuals aged 18 to 80 years participated in this observational study. Two investigators calibrated for examining tooth shade performed all examinations. A Vita-Lumin shade guide was used to examine either the maxillary right or left central incisor, one of which had to be restoration- and caries-free for study inclusion. Tooth shades were divided into 4 categories according to value, and skin tones were divided into 4 categories (fair, fair/medium, medium, and dark) with the use of L'Oreal True Illusion compact makeup shades as a guide. Categorical modeling with chi-square analysis and Fisher's exact tests was used to analyze the data (alpha=.05). RESULTS: No interaction among age, skin color, and tooth shade or gender, skin color, and tooth shade was found; however, age was associated with tooth shade (P<.001). Older persons were more likely to have teeth with lower values (darker). Among those aged 60 and older, 85% had teeth in the medium and low range values compared with 17% of those younger than 31. Significant tooth shade differences were discovered among patients with different skin colors (P<.05). Among those with low values (darker teeth), 50% were of fair complexion and 17% were of dark complexion (P<.001). CONCLUSION: Within the limitations of this study, tooth shade value and skin color were inversely related. Older adults were more likely to have darker teeth (lower value).
Wildlife management involves regulation of population numbers of wild vertebrate species. In some cases there are too many animals and in others there are too few. Genetic issues arise in both instances. The historical and genetic evidence for the number of mammals that were in the founder populations of successful colonizing species in Australia and New Zealand is reviewed here. Small numbers have often given rise to large populations, despite the concomitant loss of genetic variability. Restriction of the number of over-abundant and pest species by either physical or chemical methods frequently constitutes very strong artificial selection, which leads to rapid genetic change; an example of major importance in the two countries is sodium monofluoroacetate (compound 1080). Pathogenic agents, surgical sterilization, hormonal contraceptives and translocation have all been used with varying degrees of success. The strengths and weaknesses of these techniques are assessed. A method that has received much attention is immunocontraception. We argue that this attempt to use the animals' own immune system to modulate reproduction is incompatible with the basic biological function of protection against infectious disease. Immune function genes are highly variable in vertebrates, and so often genetic change in the population subjected to immunocontraception is likely to be even more rapid than is the case with lethal agents. Selection for failure to respond to the immunocontraceptive will occur, and will change immune function in general. Poor scientific description of ecosystem complexity makes it difficult to predict the consequences of immunocontraception on wildlife populations.