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Domestication and plasticity of brain organization in mallards (Anas platyrhynchos).

The sizes and histological differentiation of structures in the central nervous system of wild and domestic ducks were compared using allometric methods. Whole brain volume is 14.3% less in domestic ducks than in wild birds, and the size of certain brain structures is more variable in domestic ducks than in the wild birds. These findings are consistent with results of studies in other domestic birds and in domestic mammals. Although the brain subdivision volumes are not reduced to the same extent, the relative composition of the brain is maintained in domestic ducks. The overall reduction in rhombencephalic parts of the brain (16.3%) is greater than that in the prosencephalon (13.5%). Within sensory centers the nuclei of the trigeminal system undergo the most marked decrease (21.3%). Optic (13.3%) and limbic (11.1%) structures are less reduced, and the olfactory regions do not show any differences in volume compared to these regions in wild mallards. Reductions in the volumes of brain structures in domesticated animals have been thought to be processes of functional adaptation controlled by artificial selection. Our observations are discussed with regard to hypotheses about trends in domestication and evolutionary plasticity.

Animals↗

Tau differences between short-day responsive and short-day nonresponsive white-footed mice (Peromyscus leucopus) do not affect reproductive photoresponsiveness.

In laboratory-bred rodent populations, intraspecific variation in circadian system organization is a known cause of individual variation in reproductive photoresponsiveness. The authors sought to determine whether circadian system variation accounted for individual variation in reproductive photoresponsiveness in a single, highly genetically variable population of Peromyscus leucopus recently derived from the wild. Running-wheel activity patterns of male and female mice, aged 70 to 90 days, from artificially selected lines of reproductively photoresponsive (R) and nonresponsive (NR) lines were monitored under short-day photoperiod (8 h light, 16 h dark), long-day photoperiod (16 h light, 8 h dark), and constant darkness (DD). NR mice displayed a significantly longer mean free-running period (24.08 h) in DD compared with R mice (23.75 h), due in large part to a difference between NR and R females (24.25 h vs. 23.74 h, respectively). All other entrainment characteristics (alpha, phase angle of activity) under short days, long days, and DD were similar between R and NR mice. Variation in free-running period and entrainment characteristics has been shown to affect photoresponsiveness in other rodent species by altering the manner in which the circadian system interprets short days. To determine whether variation in photoresponsiveness in P. leucopus is due to differences in free-running period instead of variation downstream from the central circadian clock in the pathway controlling photoresponsiveness, the authors exposed young R and NR mice to DD and measured the effect on reproductive organ development. If variation in free-running period affected how the circadian system of mice interpreted short days, then both R and NR mice exposed to DD should have exhibited a delay in gonadal development. Only R mice exhibited pubertal delay in DD. NR mice exhibited large paired testes, paired seminal vesicles, paired ovaries, and uterine weight typical of mice nonresponsive to short days, whereas R mice exhibited reproductive organ weight typical of mice responsive to short days. These data suggest that despite significant differences in free-running period between R and NR mice, individual variation in photoresponsiveness is not due to differences in how the circadian systems of R and NR mice interpret the LD cycle.

Animals↗

Independence of genetic geographical variation between photoperiodic diapause, circadian eclosion rhythm, and Thr-Gly repeat region of the period gene in Drosophila littoralis.

Drosophila littoralis is a latitudinally widespread European species of the Drosophila virilis group. The species has ample genetic variation in photoperiodism (adult diapause) and circadian rhythmicity (pupal eclosion rhythm), with adaptive latitudinal clines in both of them. The possible common genetic basis between the variability of photoperiodism and circadian rhythms was studied by a long-term crossing experiment. A northern strain (65 degrees N) having long critical day length (CDL = 19.9 h) for diapause, early phase of the entrained rhythm in LD 3:21 (psi(LD3:21) = 12.3 h), and short period (tau= 18.8 h) of the free-running rhythm for the eclosion rhythm was crossed with a southern strain (42 degrees N) having short CDL (12.4 h), late eclosion phase (psi(LD3:21) = 20.2 h), and long period (tau= 22.8 h). After 54 generations, including free recombination, artificial selection, and genetic drift, a novel strain resulted, having even more "southern" diapause and more "northern" eclosion rhythm characteristics than found in any of the geographical strains. The observed complete separation of eclosion rhythm characteristics from photoperiodism is a new finding in D. littoralis; in earlier studies followed for 16 generations, the changes had been mostly parallel. Evidently, the genes controlling the variability of the eclosion rhythm and photoperiodism in D. littoralis are different but closely linked. To test for the possible gene loci underlying the observed geographical variability, the period gene was studied in 10 strains covering all the known clock variability in D. littoralis. The authors sequenced the most suspected Thr-Gly region, which is known to take part in the adaptive clock variability in Drosophila melanogaster. No coding differences were found in the strains, showing that this region is not included in the adaptive clock variability in D. littoralis.

Amino Acid Sequence↗

Geographic distribution of haplotype diversity at the bovine casein locus.

The genetic diversity of the casein locus in cattle was studied on the basis of haplotype analysis. Consideration of recently described genetic variants of the casein genes which to date have not been the subject of diversity studies, allowed the identification of new haplotypes. Genotyping of 30 cattle breeds from four continents revealed a geographically associated distribution of haplotypes, mainly defined by frequencies of alleles at CSN1S1 and CSN3. The genetic diversity within taurine breeds in Europe was found to decrease significantly from the south to the north and from the east to the west. Such geographic patterns of cattle genetic variation at the casein locus may be a result of the domestication process of modern cattle as well as geographically differentiated natural or artificial selection. The comparison of African Bos taurus and Bos indicus breeds allowed the identification of several Bos indicus specific haplotypes (CSN1S1*C-CSN2*A2-CSN3*AI / CSN3*H) that are not found in pure taurine breeds. The occurrence of such haplotypes in southern European breeds also suggests that an introgression of indicine genes into taurine breeds could have contributed to the distribution of the genetic variation observed.

Animals↗

Reduction of inbreeding in commercial females by rotational mating with several sire lines.

A mating system to reduce the inbreeding of commercial females in the lower level was examined theoretically, assuming a hierarchical breed structure, in which favorable genes are accumulated in the upper level by artificial selection and the achieved genetic progress is transferred to the lower level through migration of males. The mating system examined was rotational mating with several closed sire lines in the upper level. Using the group coancestry theory, we derived recurrence equations for the inbreeding coefficient of the commercial females. The asymptotic inbreeding coefficient was also derived. Numerical computations showed that the critical factor for determining the inbreeding is the number of sire lines, and that the size of each sire line has a marginal effect. If four or five sire lines were available, rotational mating was found to be quite an effective system to reduce the short- and long-term inbreeding of the commercial females, irrespective of the effective size of each sire line. Oscillation of the inbreeding coefficient under rotational mating with initially related sire lines could be minimized by avoiding the consecutive use of highly related lines. Extensions and perspectives of the system are discussed in relation to practical application.

Animals↗

Automated measurement of Drosophila wings.

BACKGROUND: Many studies in evolutionary biology and genetics are limited by the rate at which phenotypic information can be acquired. The wings of Drosophila species are a favorable target for automated analysis because of the many interesting questions in evolution and development that can be addressed with them, and because of their simple structure. RESULTS: We have developed an automated image analysis system (WINGMACHINE) that measures the positions of all the veins and the edges of the wing blade of Drosophilid flies. A video image is obtained with the aid of a simple suction device that immobilizes the wing of a live fly. Low-level processing is used to find the major intersections of the veins. High-level processing then optimizes the fit of an a priori B-spline model of wing shape. WINGMACHINE allows the measurement of 1 wing per minute, including handling, imaging, analysis, and data editing. The repeatabilities of 12 vein intersections averaged 86% in a sample of flies of the same species and sex. Comparison of 2400 wings of 25 Drosophilid species shows that wing shape is quite conservative within the group, but that almost all taxa are diagnosably different from one another. Wing shape retains some phylogenetic structure, although some species have shapes very different from closely related species. The WINGMACHINE system facilitates artificial selection experiments on complex aspects of wing shape. We selected on an index which is a function of 14 separate measurements of each wing. After 14 generations, we achieved a 15 S.D. difference between up and down-selected treatments. CONCLUSION: WINGMACHINE enables rapid, highly repeatable measurements of wings in the family Drosophilidae. Our approach to image analysis may be applicable to a variety of biological objects that can be represented as a framework of connected lines.

Animals↗

Ribosomal intergenic spacer (IGS) length variation across the Drosophilinae (Diptera: Drosophilidae).

BACKGROUND: The intergenic spacer of the ribosomal genes in eukaryotes (IGS) contains duplications of the core transcription promoter. The number of these duplicated promoters, as measured by the IGS length, appears to be correlated with growth rate and development time in several distantly related taxa. In the present study, we examined IGS length variation across a number of species of Drosophila to determine the amount of variation in this trait across different evolutionary time scales. Furthermore, we compared the usefulness of two methods commonly used to determine IGS length: Southern Blot Hybridization (SB) and Polymerase Chain Reaction (PCR). RESULTS: Our results show broad variation in IGS length across the genus Drosophila, but closely related species had similar IGS lengths. Our results also suggest that PCR tends to underestimate the true IGS size when the size is greater than 5 kb, and that this degree of underestimation is greater as the IGS size increases. CONCLUSION: Broad variation in IGS length occurs across large evolutionary divergences in the subfamily Drosophilinae. Although average IGS length has been shown to evolve rapidly under artificial selection, closely related taxa generally have similar average IGS lengths. Our comparison of methods suggests that without previous knowledge of the DNA sequence of the IGS and flanking regions, both methods be used to accurately measure IGS length.

Animals↗

Early rearing environment impacts cerebellar growth in juvenile salmon.

The size and structure of an animal's brain is typically assumed to result from either natural or artificial selection pressures over generations. However, because a fish's brain grows continuously throughout life, it may be particularly responsive to the environmental conditions the fish experiences during development. Salmon are an ideal model system for studying these effects because natural habitats differ significantly from the hatchery environments in which these fish are frequently reared. For example, in the wild, salmon alevins (i.e. yolk-sac fry) are buried in the gravel, while hatchery environments lack this structural component. We show that the simple manipulation of adding stones to a standard rearing tank can dramatically alter the growth of specific brain structures in steelhead salmon alevins (Oncorhynchus mykiss). We found that alevins reared with stones grew brains with significantly larger cerebella than genetically similar fish reared in conventional tanks. This shift to a larger cerebellar size was, in turn, accompanied by changes in locomotory behaviors--behaviors that correlate strongly to the function of this brain region. We next show that hatchery fish reared in a more naturalistic setting in the wild had significantly larger brains than their lab-reared counterparts. However, relative cerebellar volumes were similar between wild-reared alevins and those reared in the complex treatment in the laboratory. Together our results indicate that, within the first three weeks of life, variation in rearing environment can result in brain differences that are commonly attributed to generations of selection. These results highlight the need to consider enrichment strategies when designing captive rearing facilities for both conservation and laboratory use.

Animals↗

Phenotypic plasticity and experimental evolution.

Natural or artificial selection that favors higher values of a particular trait within a given population should engender an evolutionary response that increases the mean value of the trait. For this prediction to hold, the phenotypic variance of the trait must be caused in part by additive effects of alleles segregating in the population, and also the trait must not be too strongly genetically correlated with other traits that are under selection. Another prediction, rarely discussed in the literature, is that directional selection should favor alleles that increase phenotypic plasticity in the direction of selection, where phenotypic plasticity is defined as the ability of one genotype to produce more than one phenotype when exposed to different environments. This prediction has received relatively little empirical attention. Nonetheless, many laboratory experiments impose selection regimes that could allow for the evolution of enhanced plasticity (e.g. desiccation trials with Drosophila that last for several hours or days). We review one example that involved culturing of Drosophila on lemon for multiple generations and then tested for enhanced plasticity of detoxifying enzymes. We also review an example with vertebrates that involves selective breeding for high voluntary activity levels in house mice, targeting wheel-running behavior on days 5+6 of a 6-day wheel exposure. This selection regime allows for the possibility of wheel running itself or subordinate traits that support such running to increase in plasticity over days 1-4 of wheel access. Indeed, some traits, such as the concentration of the glucose transporter GLUT4 in gastrocnemius muscle, do show enhanced plasticity in the selected lines over a 5-6 day period. In several experiments we have housed mice from both the Selected (S) and Control (C) lines with or without wheel access for several weeks to test for differences in plasticity (training effects). A variety of patterns were observed, including no training effects in either S or C mice, similar changes in both the S and C lines, greater changes in the S lines but in the same direction in the C lines, and even opposite directions of change in the S and C lines. For some of the traits that show a greater training effect in the S lines, but in the same direction as in C lines, the greater effect can be explained statistically by the greater wheel running exhibited by S lines ('more pain, more gain'). For others, however, the differences seem to reflect inherently greater plasticity in the S lines (i.e. for a given amount of stimulus, such as wheel running/day, individuals in the S lines show a greater response as compared with individuals in the C lines). We suggest that any selection experiment in which the selective event is more than instantaneous should explore whether plasticity in the appropriate (adaptive) direction has increased as a component of the response to selection.

Adaptation, Physiological↗

Phenotypic characterization of mice bred for high and low peak bone mass.

In humans, peak bone mineral density (BMD) is a highly heritable trait and a strong determinant of subsequent osteoporotic fracture risk. To identify the genetic factors responsible for variation in peak BMD, investigators have turned to animal models. In this study we examined the heritability of BMD acquisition and characterized differences in skeletal geometry, histomorphometry, and biomechanical competence between two lines of mice artificially selected for extremes of peak whole body BMD. F2 progeny from a cross between C57BL/6 and DBA/2 inbred strains was used as the foundation population to develop lines selected for either high or low BMD. Whole body BMD was measured by dual-energy X-ray absorptiometry (DXA). By the third generation of selection, highest-scoring BMD (HiBMD) mice exhibited 14% greater peak BMD than lowest-scoring BMD (LoBMD) mice. The mean realized heritability of peak BMD was 36%. Femoral shaft cortical area and thickness and vertebral cancellous bone volume (BV) were significantly greater (16-30%) in the HiBMD line compared with the LoBMD line. Mean cancellous bone formation rates (BFRs) were 35% lower in HiBMD mice compared with LoBMD mice. Failure load and stiffness in the femoral shaft, femoral neck, and L6 vertebrae were all substantially greater (by 25-190%) in HiBMD mice. Thus, these divergently selected murine lines serve to illustrate some of the means by which genetic mechanisms can affect skeletal structure and remodeling. Identification of the individual genes influencing peak BMD in this experimental system will likely reveal some of the genetic determinants of overall bone strength.

Analysis of Variance↗

Synthetic taxonomy of Rosa races using ACT-STATIS.

Fifteen Rosa cultivated races were described by means of phenotypic frequencies (11 tables). Two groups of correlated contingency tables were identified by ACT-STATIS (Analyse Conjointe de Tableaux-Structuration de Tableaux à Trois Indices de la Statistique) interstructure analysis. Three data sets appeared to be independent from the others. Typologies of races were obtained after ACT-STATIS compromise analyses for the two groups of correlated tables, and after Principal Component Analyses for the independent data sets. Each typology was original and variously influenced by genealogical structure, mutation or artificial selection pressures. A weighted synthesis was attempted in order to build a taxonomy of races taking into account these diversity factors. The good agreement between the resulting classification and the assumptions about the history of Rosa domestication advocated for a wider utilization of ACT-STATIS and RV coefficient when the relationships between individuals or populations have to be studied on the basis of their similarities.

Classification↗

An analysis of genetic diversity across the maize genome using microsatellites.

How domestication bottlenecks and artificial selection shaped the amount and distribution of genetic variation in the genomes of modern crops is poorly understood. We analyzed diversity at 462 simple sequence repeats (SSRs) or microsatellites spread throughout the maize genome and compared the diversity observed at these SSRs in maize to that observed in its wild progenitor, teosinte. The results reveal a modest genome-wide deficit of diversity in maize relative to teosinte. The relative deficit of diversity is less for SSRs with dinucleotide repeat motifs than for SSRs with repeat motifs of more than two nucleotides, suggesting that the former with their higher mutation rate have partially recovered from the domestication bottleneck. We analyzed the relationship between SSR diversity and proximity to QTL for domestication traits and observed no relationship between these factors. However, we did observe a weak, although significant, spatial correlation for diversity statistics among SSRs within 2 cM of one another, suggesting that SSR diversity is weakly patterned across the genome. Twenty-four of 462 SSRs (5%) show some evidence of positive selection in maize under multiple tests. Overall, the pattern of genetic diversity at maize SSRs can be explained largely by a bottleneck effect with a smaller effect from selection.

DNA, Plant↗

Physiological bases of genetic differences in cannibalism behavior of the confused flour beetle Tribolium confusum.

Physiological causes of genetic differences in cannibalism were examined to gain a better understanding of constraints on behavior evolution. Cannibalism has complex population level consequences in Tribolium confusum, including dramatic effects on population size. Laboratory strains with low and high cannibalism rates, obtained through inbreeding, have maintained distinct levels of cannibalism for over two decades even in the absence of artificial selection to maintain the differences. Why strains differ in their cannibalism rates was examined by measuring: (1) the nutritional benefit from cannibalism in both nutritionally good and poor environments, and (2) the possibility that eggs are an important source of water. How strains achieve differences in cannibalism was examined by testing for differences between strains in their ability to find eggs and in their tendency to eat eggs. Beetles from both strains survive equally well in a nutritionally good environment, but they accomplish this in different ways. The low cannibalism strain has high survivorship with and without cannibalism. The high cannibalism strain has low survivorship when not fed eggs and survivorship equivalent to the low cannibalism strain when fed eggs, suggesting it compensates for poor nutritional adaptation by eating eggs. The strains also differ in feeding behavior; beetles from the high cannibalism strain have a higher appetite for eggs. Beetles from the two strains did not differ in locomotor activity, search efficiency, or need for water. The observed behavioral and nutritional differences may contribute to the maintenance of different levels of cannibalism.

Animals↗

Evolution of cadmium resistance in Daphnia magna.

A diverse laboratory population of Daphnia magna Straus was established by combining individuals from eight sources. Artificial selection for increased resistance to the acute toxic effects of cadmium was performed, and after eight generations, the average median effective concentration increased from 61 to 180 microg/L. No differences in life span, offspring production, time to first brood, number of offspring in the first brood, or intrinsic rate of population increase (r) were observed between the cadmium-adapted population and the controls under ideal conditions or under conditions of temperature or feeding ration stress, but cadmium-adapted daphnids were smaller. Control and cadmium-adapted populations were equally sensitive to copper and malathion, but the cadmium-adapted population was less sensitive to lead and more sensitive to phenol. Analysis of amplified fragment-length polymorphisms indicated a significant decrease in genetic diversity in the cadmium-adapted population. Although the evolved cadmium resistance would allow adapted populations to exist in areas where cadmium concentrations would be toxic to unadapted populations, the decreased genetic diversity, smaller size, and increased sensitivity to at least one other toxicant could reduce the probability of long-term survival even in the absence of future cadmium exposure.

Adaptation, Physiological↗

Quantitative- and molecular-genetic effects on animal well-being: adaptive mechanisms.

Domestic farm animals play an important role in meeting some basic needs of humankind, especially food and clothing. The aspects of genetic improvement programs in livestock production pertinent to animal welfare and animal well-being are reviewed. A link is made between the evolutionary processes of adaptation and domestication and animal well-being. Animal behavior is a component of all these. Thus, the genetics of behavior may provide clues to the well-being of farm animal populations, and it will also be of relevance to public opinion issues of animal welfare. Many expressions of behavior by domestic livestock may be influenced by those processes that change gene as well as genotypic frequencies such as inbreeding, drift, and artificial selection. The environment in which the individual lives will also play a role, along with the interaction between genotype and environment. Selection for or against such behaviors as aggressiveness, docility, response to stress, and certain sexual behaviors in some livestock species has often been successful. This points to the existence of additive genetic variation for behavior, and scope for the inclusion of behavioral traits into selection programs, if these measures are shown to be related to welfare. Negative relationships between behaviors associated with well-being and traits of economic importance have been reported in most livestock species. However, estimates of genetic parameters, especially genetic correlations between objective measures of well-being and production traits, are scarce. There have been no comprehensive studies of the welfare of transgenic animals reported in the scientific literature. Increased use of biotechnology in animal agriculture, coupled with greater public scrutiny of livestock industries, may precipitate decisions concerning the interface of behavior and genetics that need to be addressed before scientists can conduct appropriate experimental evaluations.

Adaptation, Physiological↗

Identifying single genes of large effect in quantitative traits using best linear unbiased prediction.

The recent discovery of a major gene for rapid postweaning growth has reinforced the hypothesis that other quantitative continuous traits may be influenced by single genes of large effect. However, most methods for the detection of such genes rely on the discovery of multimodality in the population frequency distribution. The complicating effects of environment and artificial selection make the identification of such genes with field-collected data a formidable problem. An index is proposed that may serve as an indicator that a major gene is segregating within a population. The index is based on the assumption that under polygenic inheritance, an offspring's deviation from the midparent average is smaller than the deviation from either parent. Whereas, for the Mendelian segregation of a major gene, the opposite would be expected. A proposed class of indices is then based on the ratio [O - .5(S + D)]k/([O - S]k/2 [O - D]k/2) where O, S and D are the additive genetic values of an offspring and its sire and dam estimated via best linear unbiased prediction. Values of the index greater than 1. would be indicative of major gene inheritance. Simulation of small populations indicates that the index is quite sensitive to the existence of segregating major genes even in the absence of multimodality of the phenotypic distribution. However, the index remains dependent on the accuracy of genetic value estimation.

Animals↗

Oncogenes and tumor suppressor genes.

The artificial selection of the directly acting or acute RNA tumor viruses for high transforming ability has led to the isolation of defective retroviral genomes that have picked up, by accidental recombination, some of the important genes that influence, trigger or regulate cell division. These genes belong to at least four functionally different groups. Each of them can contribute to tumor development and/or progression after activation by structural or regulatory changes. Growth factor genes may act as oncogenes following constitutive activation in a cell that normally responds to, but does not produce, the corresponding growth factor (the autocrine model, exemplified by sis). Growth factor receptors may be fixed in a state of continuous, faulty signalling by the truncation of their external, ligand binding portion (examples: erb-B, fms). Genes coding for proteins involved in signal transduction may be activated by point mutations in certain, important domains (example: the ras-family). DNA binding proteins, presumably involved in DNA replication may drive cell division after constitutive activation by retroviral insertion, chromosomal translocation or gene amplification (example: the myc-family).

Gene Amplification↗

Lactation: historical patterns and potential for manipulation.

The advent of biotechnology has made data on undomesticated mammals relevant to dairy science. Such data indicate the potential of lactation for modification, reveal genetic material available for use through bioengineering, help distinguish adaptive features from historical artifacts, and clarify limits on lactational diversity that date from early evolution. Evolutionary analysis indicates that a complex degree of lactation preceded divergence of the extant mammalian lineages during the Mesozoic Era. Although aspects of monotreme lactation appear to be ancestral for extant mammals, the marsupials and eutherians exhibit divergent specializations. Evidence is consistent with the idea that protolacteal glands evolved by combining features of skin gland populations into a new functional complex. Secretions of these ancestral glands may have had antimicrobial properties that protected the eggs or hatchlings and organic components that supplemented offspring nutrition. Following development of highly nutritious milks, evolution produced diversity in milk composition and function, milk output, length of lactation, mammary gland anatomy, and contributions of lactation to offspring nutrition. Certain marsupials are specialized in terms of functional independence and temporal plasticity of mammary tissues. Mammalian diversity indicates that artificial selection and physiological manipulation of domestic artiodactyls has only modestly exploited the potential of mammary glands as a nutritional source for humans.

Animals↗