Local search in the housefly Musca domestica after feeding on sucrose

Local search behaviour of the housefly, Musca domestica, released by ingestion of sucrose, was recorded on a bitpad digitizer. Local search is characterized by an initial increase in turning rate and a decrease in locomotory rate followed by an exponential return to control (unfed) levels for both functions. Greater variation in the slope of the exponential return to control levels was observed between runs of different flies than between runs of an individual fly, suggesting a possible internal basis for locomotory and turning functions. The local search pattern, based on decrease in relative turning rate generates a spiral configuration. A zig-zag component increases path width and increases the chances of a fly relocating the drop residue. Looping patterns correlate with successive recontacts with the drop residue and associated changes in turn direction and stopping. The results are discussed with respect to sources of internal and external orientation information controlling local search.     

QUANTITATIVE GENETICS OF SURVIVAL AND GROWTH IN IMPATIENS CAPENSIS

When variation in life-history characters is caused by many genes of small effect, then quantitative-genetic parameters may quantify constraints on rate and direction of microevolutionary change. I estimated heritabilities and genetic correlations for 16 life-history and morphological characters in two populations of Impatiens capensis, a partially self-pollinating herbaceous annual. The Madison population had little or no additive genetic variance for any of these characters, while the Milwaukee population had significant narrowsense heritabilities and genetic correlations for several traits, including adult size, which is highly correlated with fitness. All genetic correlations among fitness components were positive, hence there is no evidence for antagonistic pleiotropy among these traits. Dissimilarity of heritabilities in the two populations supports theoretical predictions that long-term changes in genetic variance-covariance patterns may occur when population sizes are small and selection is strong, as may occur in many plant species.     

Search orientation of Musca domestica in patches of sucrose drops

ABSTRACT. The searching tactics of the housefly, Musca dormestica L. (Diptera: Muscidae), have been delineated from digitized pathways of flies walking in patches of sucrose drops arranged in linear (ROW) and hexagonal (HEX) arrays. The areas covered by flies in ROW and HEX patches are distinctly different, but flies seem not to employ different tactics for the two types of resource arrays. The number of drops located, if at least one drop is found, does not differ between ROW and HEX. Most quantitative measures of local search remain constant after the first interdrop interval, although feeding time decreases as flies sample successive drops. Local search intensifies after each drop is ingested, with locomotory rate decreasing and turning rate increasing, followed by decay of both measures toward ranging levels. Searching can be characterized by two movement tendencies resulting from specific, definable, locomotory functions: a forward-moving tendency is expressed by the fly as it leaves a resource in approximately the same direction as it arrives; and local search is characterized by looping, rather than straight walking, with a variable turning rate that generates a rough ‘zigzag’ superimposed on looping. The two movement tendencies, combined, allow flies to locate resources in a linear arrangement, because of the forward-moving tendency, and to locate resources not arranged in a linear array because of the ‘noisy’ loop. M. domestica does not appear to retain and use information gained from one patch of drops in another, so the search tactic of the fly seems therefore to be a compromise between straight movement and circular movement that may be adaptive for an environment subject to frequent changes in the spatial distribution of resources. Giving-up-time, the period between ingesting the last drop and leaving the patch, is a function of the rate of change in the transition from local search to ranging, which is constant in our experiments. If a fly does not encounter another drop while ranging, during which it walks relatively straight, the fly moves out of and away from the patch.     

Realized Sampling Variances of Estimates of Genetic Parameters and the Difference between Genetic and Phenotypic Correlations

A data set of 1572 heritability estimates and 1015 pairs of genetic and phenotypic correlation estimates, constructed from a survey of published beef cattle genetic parameter estimates, provided a rare opportunity to study realized sampling variances of genetic parameter estimates. The distribution of both heritability estimates and genetic correlation estimates, when plotted against estimated accuracy, was consistent with random error variance being some three times the sampling variance predicted from standard formulae. This result was consistent with the observation that the variance of estimates of heritabilities and genetic correlations between populations were about four times the predicted sampling variance, suggesting few real differences in genetic parameters between populations. Except where there was a strong biological or statistical expectation of a difference, there was little evidence for differences between genetic and phenotypic correlations for most trait combinations or for differences in genetic correlations between populations. These results suggest that, even for controlled populations, estimating genetic parameters specific to a given population is less useful than commonly believed. A serendipitous discovery was that, in the standard formula for theoretical standard error of a genetic correlation estimate, the heritabilities refer to the estimated values and not, as seems generally assumed, the true population values.     

Quantitative genetic analysis of brain size variation in sticklebacks: support for the mosaic model of brain evolution

The mosaic model of brain evolution postulates that different brain regions are relatively free to evolve independently from each other. Such independent evolution is possible only if genetic correlations among the different brain regions are less than unity. We estimated heritabilities, evolvabilities and genetic correlations of relative size of the brain, and its different regions in the three-spined stickleback (Gasterosteus aculeatus). We found that heritabilities were low (average h² = 0.24), suggesting a large plastic component to brain architecture. However, evolvabilities of different brain parts were moderate, suggesting the presence of additive genetic variance to sustain a response to selection in the long term. Genetic correlations among different brain regions were low (average r_G = 0.40) and significantly less than unity. These results, along with those from analyses of phenotypic and genetic integration, indicate a high degree of independence between different brain regions, suggesting that responses to selection are unlikely to be severely constrained by genetic and phenotypic correlations. Hence, the results give strong support for the mosaic model of brain evolution. However, the genetic correlation between brain and body size was high (r_G = 0.89), suggesting a constraint for independent evolution of brain and body size in sticklebacks.     

Age trends in genetic parameters for growth and wood density in <Emphasis Type="Italic">Eucalyptus globulus</Emphasis>

Genetic parameters for stem diameter and wood density were compared at selection (4–5 years) and harvest (16–17 years) age in an open-pollinated progeny trial of Eucalyptus globulus in Tasmania (Australia). The study examined 514 families collected from 17 subraces of E. globulus. Wood density was assessed on a subsample of trees indirectly using pilodyn penetration at both ages and directly by core basic density at harvest age. Significant additive genetic variance and narrow-sense heritabilities ( h_\textop² h_{\text{op}}^2 ) were detected for all traits. Univariate and multivariate estimates of heritabilities were similar for each trait except harvest-age diameter. Comparable univariate estimates of selection- and harvest-age heritabilities for diameter masked changes in genetic architecture that occurred with stand development, whereby the loss of additive genetic variance through size-dependent mortality was countered by the accentuation of additive genetic differences among survivors with age. Regardless, the additive genetic (r _a) and subrace (r _s) correlations across ages were generally high for diameter (0.95 and 0.61, respectively) and pilodyn penetration (0.77 and 0.96), as were the correlations of harvest-age core basic density with selection- and harvest-age pilodyn (r _a −0.83, −0.88; r _s −0.96, −0.83). While r _s between diameter and pilodyn were close to zero at both ages, there was a significant change in r _a from adverse at selection age (0.25) to close to zero (−0.07) at harvest age. We argue that this change in the genetic correlation reflects a decoupling of the genetic association of growth and wood density with age. This result highlights the need to validate the use of selection-age genetic parameters for predicting harvest-age breeding values.     

Linear and threshold analysis of direct and maternal genetic effects for secondary sex ratio in Iranian buffaloes

The objective of this study was to estimate variance components and genetic parameters for secondary sex ratio (SSR) in Iranian buffaloes. Calving records from April 1995 to June 2010 comprising 15,207 calving events from the first three lactations of 1066 buffalo herds of Iran were analyzed using linear and threshold animal models to estimate variance components, heritabilities and genetic correlations between direct and maternal genetic effects for SSR. Linear and threshold animal models included direct and maternal genetic effects with covariance between them and maternal permanent environmental effects were implemented by Gibbs sampling methodology. Posterior means of direct and maternal heritabilities and repeatability for SSR obtained from linear animal model were 0.15, 0.10, and 0.17, respectively. Threshold estimates of direct and maternal heritabilities and repeatability for SSR were 0.48, 0.27, and 0.52, respectively. The results showed that the correlations between direct and maternal genetic effects of SSR were negative and high in both models. In addition, the ratios of maternal permanent environmental variance were low. Exploitable genetic variation in SSR can take advantage of sexual dimorphism for economically important traits which may facilitate greater selection intensity and thus greater response to selection, as well as reducing the replacement costs. Threshold animal model may be applied in selection programs where animals are to be genetically ranked for female rate.     

Search orientation in adultDrosophila melanogaster: Responses of rovers and sitters to resource dispersion in a food patch

Drosophila melanogasteradults were employed in single resource patches of varying density and size and in a multiple-patch array to determine the degree to which resource dispersion influences searching success. Individuals from rover and sitter selected lines, with extreme genotypes for local search duration, are not as successful as control-line (wild-type) flies in locating sucrose drops in single patches varying in size and density. The number of new drops located differed significantly between fly lines in all patch types, except in a high-density patch, and within each fly line over the different patch sizes and densities. The similarities in number of drops found by rovers and sitters in all patch types are not reflected in the time periods spent searching. In the multiple-patch array sitters never left the central patch, whereas most rovers and con-trol-line flies found additional patches. The proximate explanations for the success or failure of the three fly lines in different patch sizes and densities relate to the looping locomotor pattern characterizing local search in D. melanogaster.The reactivation of searching each time a drop is ingested or revisited keeps an individual in the immediate vicinity of the last encountered resource. Flies from the selected lines, each exhibiting extreme types of locomotor patterns, leave patches relatively unexploited because local search consists either of rapid, nearly linear movement away from a drop in rovers or of relatively long bouts of local search in sitters, which promotes revisiting rather than locating new drops. Control-line flies locate more drops than either rovers or sitters and in less time than sitters, suggesting that their intermediate phenotype for search behavior allows for more flexibility in searching in various patch sizes and resource densities. The results are discussed with reference to environmental and physiological factors that may modify searching behavior and, possibly, enhance the survival of individuals with extreme genotypes.     

On the sampling variance of intraclass correlations and genetic correlations.

Widely used standard expressions for the sampling variance of intraclass correlations and genetic correlation coefficients were reviewed for small and large sample sizes. For the sampling variance of the intraclass correlation, it was shown by simulation that the commonly used expression, derived using a first-order Taylor series performs better than alternative expressions found in the literature, when the between-sire degrees of freedom were small. The expressions for the sampling variance of the genetic correlation are significantly biased for small sample sizes, in particular when the population values, or their estimates, are close to zero. It was shown, both analytically and by simulation, that this is because the estimate of the sampling variance becomes very large in these cases due to very small values of the denominator of the expressions. It was concluded, therefore, that for small samples, estimates of the heritabilities and genetic correlations should not be used in the expressions for the sampling variance of the genetic correlation. It was shown analytically that in cases where the population values of the heritabilities are known, using the estimated heritabilities rather than their true values to estimate the genetic correlation results in a lower sampling variance for the genetic correlation. Therefore, for large samples, estimates of heritabilities, and not their true values, should be used.     

Genetic parameters for body weight,longissimus muscle depth and fat depth for Suffolk sheep in the Czech Republic

In this study, heritabilities and (co)variance components for body weight at 100 days (BW), muscle depth (MD) and fat depth (FD) were estimated for Suffolk, the most common sheep breed in the Czech Republic. Data from 1996 to 2004 were extracted from the sheep recording database of the Czech Sheep and Goat Breeding Association. Genetic parameters were estimated using multivariate animal models, including both direct and maternal genetic effects and permanent environmental effects. Average values for BW, MD and FD were 27.91 kg, 25.5 mm and 3.3 mm, respectively. Direct and maternal heritability for BW were 0.17 and 0.08, respectively, and direct heritabilities were 0.16 for MD and 0.08 for FD. Maternal heritability estimates for ultrasonic measurements were generally low. Direct genetic correlations between BW and MD and maternal genetic correlations between BW and MD were positive and favourable. Both direct genetic correlations between BW and FD and maternal genetic correlations between BW and FD were negative, but not significantly different from zero. The favourable genetic correlations between BW and MD make ultrasound measurements a valuable tool in breeding programs focusing on growth and carcass characteristics.     

Estimating heritabilities and genetic correlations: comparing the 'animal model' with parent-offspring regression using data from a natural population

Quantitative genetic parameters are nowadays more frequently estimated with restricted maximum likelihood using the 'animal model' than with traditional methods such as parent-offspring regressions. These methods have however rarely been evaluated using equivalent data sets. We compare heritabilities and genetic correlations from animal model and parent-offspring analyses, respectively, using data on eight morphological traits in the great reed warbler (Acrocephalus arundinaceus). Animal models were run using either mean trait values or individual repeated measurements to be able to separate between effects of including more extended pedigree information and effects of replicated sampling from the same individuals. We show that the inclusion of more pedigree information by the use of mean traits animal models had limited effect on the standard error and magnitude of heritabilities. In contrast, the use of repeated measures animal model generally had a positive effect on the sampling accuracy and resulted in lower heritabilities; the latter due to lower additive variance and higher phenotypic variance. For most trait combinations, both animal model methods gave genetic correlations that were lower than the parent-offspring estimates, whereas the standard errors were lower only for the mean traits animal model. We conclude that differences in heritabilities between the animal model and parent-offspring regressions were mostly due to the inclusion of individual replicates to the animal model rather than the inclusion of more extended pedigree information. Genetic correlations were, on the other hand, primarily affected by the inclusion of more pedigree information. This study is to our knowledge the most comprehensive empirical evaluation of the performance of the animal model in relation to parent-offspring regressions in a wild population. Our conclusions should be valuable for reconciliation of data obtained in earlier studies as well as for future meta-analyses utilizing estimates from both traditional methods and the animal model.     

Laboratory Estimates of Heritabilities and Genetic Correlations in Nature

A lower bound on heritability in a natural environment can be determined from the regression of offspring raised in the laboratory on parents raised in nature. An estimate of additive genetic variance in the laboratory is also required. The estimated lower bounds on heritabilities can sometimes be used to demonstrate a significant genetic correlation between two traits in nature, if their genetic and phenotypic correlations in nature have the same sign, and if sample sizes are large, and heritabilities and phenotypic and genetic correlations are high.     

Heritability of hemostasis phenotypes and their correlation with type 2 diabetes status in Mexican Americans

Hypercoagulation often occurs in type 2 diabetes, suggesting pleiotropy of the genes that influence disease liability and hemostasis-related phenotypes. To better understand the relationship between hemostasis and diabetes, we first used maximum-likelihood methods to estimate the relative contribution of additive genetic, measured environmental, and shared household effects to the normal variance of 16 hemostasis-related traits in 813 individuals participating in the San Antonio Family Heart Study. We estimated moderate to high heritabilities (0.20-0.60) for each phenotype. Von Willebrand factor (VWF), thrombin activatable fibrinolysis inhibitor, activated protein C (APC) ratio, factor V, and prothrombin time had heritabilities greater than 0.50. The correlation between type 2 diabetes status and the hemostasis-related traits was then partitioned into genetic and environmental components using bivariate variance-components methods. Significant (p < or = 0.05) positive genetic correlations (0.37-0.51) occurred with factors II and VIII, VWF, total protein S (tPS), and tissue factor pathway inhibitor. Significant negative genetic correlations were estimated for activated partial thromboplastin time (-0.49) and APC ratio (-0.38). By contrast, significant environmental correlations occurred only with factor II (-0.40) and tPS (-0.31). Our results suggest that genes are important contributors to the normal variation in hemostasis-related traits and that genes influencing hemostasis-related traits pleiotropically influence diabetes risk.     

Estimation of genetic parameters for test day records of dairy traits in the first three lactations

Application of test-day models for the genetic evaluation of dairy populations requires the solution of large mixed model equations. The size of the (co)variance matrices required with such models can be reduced through the use of its first eigenvectors. Here, the first two eigenvectors of (co)variance matrices estimated for dairy traits in first lactation were used as covariables to jointly estimate genetic parameters of the first three lactations. These eigenvectors appear to be similar across traits and have a biological interpretation, one being related to the level of production and the other to persistency. Furthermore, they explain more than 95% of the total genetic variation. Variances and heritabilities obtained with this model were consistent with previous studies. High correlations were found among production levels in different lactations. Persistency measures were less correlated. Genetic correlations between second and third lactations were close to one, indicating that these can be considered as the same trait. Genetic correlations within lactation were high except between extreme parts of the lactation. This study shows that the use of eigenvectors can reduce the rank of (co)variance matrices for the test-day model and can provide consistent genetic parameters.     

Genetic and phenotypic correlations in a natural population of song sparrows

We estimated heritabilities, and genetic and phenotypic correlations between beak and body traits in the song sparrow ( Melospiza melodia ). We compared these estimates to values for the same traits in the Galápagos finches, Geospiza (Boag, 1983; Grant, 1983). Morphological variance is low in the song sparrow, and our results show that genetic and phenotypic correlations are considerably lower than correlations in the morphologically more variable Geospiza. Comparison using a larger sample of Galapagos populations confirms the existence of an association between variance and correlation for phenotypic values. We suggest two possible explanations for this association. First, most traits studied are functionally related, and the joint evolution of variance and correlation may have resulted from stabilizing selection about a line of optimal allometry between traits. Alternatively, introgression between populations and species could have caused correlation and variance to evolve jointly. Both selection and introgression were probably influential in producing the observed pattern, but it is not possible to estimate their relative importance with current data. Genetic and phenotypic correlations were correlated in the song sparrow, but heritabilities of traits varied greatly. As a result, the genetic variance-covariance matrix for traits is not simply a constant multiple of the phenotypic matrix. Evolutionary response to natural selection cannot, therefore, be predicted from the measurement of phenotypic characteristics alone.     

Multivariate selection under adverse genetic correlations: impacts of population sizes and selection strategies on gains and coancestry in forest tree breeding

We examined gene models for two traits with and without antagonistic pleiotropy using a locus-based simulation model to investigate the effects of different population sizes, heritabilities and economic weights, using index selection, and index selection with optimum selection (OS), over 10 generations. Gene models included additive and dominance gene action, with equal and varying initial allele frequencies with and without pleiotropy for a fixed level of resources (i.e. founder sizes each generation of 40, 80 and 160 with progeny arrays that totaled 800 per generation). Pleiotropy (with an initial r _g of −0.5), reduced gain by ~8–10% when heritabilities for both traits were the same (0.2), relative to non-pleiotropic cases. When traits had different heritabilities (i.e. 0.2 and 0.4), gains in the lower heritability trait were substantially lower, especially with pleiotropy present. In general, OS with slightly larger population sizes could offset losses in gain, but rarely overrode the large effects of different heritabilities or economic weights. Pleiotropy increased response variance among traits, which was magnified when heritabilities were different. Identifying an appropriate weight on relatedness in the OS process is important to manage coancestry expectations around the loss of alleles (or fixation of recessive alleles) and to minimise response variance. The dynamics of selection intensity, drift, rate of coancestry build-up, response variance, etc. are complex for multi-trait selection; however, a few economically viable strategies could reduce the adverse effects of selecting against genetic correlations without drastically impairing gain.     

Estimating variance components and heritabilities in the wild: a case study using the ‘animal model’ approach

Using a genealogy containing over 1800 dams and nearly 400 sires (estimated by genetic paternity techniques), combined with maximum likelihood procedures and an ‘animal model’, we have estimated the heritabilities, genetic correlations and variance components of three morphometric traits in the Soay sheep (Ovis aries) on St Kilda, Scotland. This approach allows heritabilities to be estimated in natural populations that violate the assumptions of offspring–parent regression methods. Maternal (or paternal) effects can also be estimated under natural conditions. We demonstrate that all the traits, body weight, hind leg length and incisor arcade breadth, have low but significant heritabilities. Body weight, the trait that experiences the strongest selection, had the lowest heritability but the highest additive genetic coefficient of variation. An evolutionary response to selection is predicted. When maternal effects were not taken into consideration heritabilities were over‐estimated, although this effect was only significant in female offspring.     

Quantitative genetic parameters of agronomic and quality traits in a global germplasm collection reveal excellent breeding perspectives for Jatropha curcas L.

Jatropha curcas L. (jatropha) is an undomesticated plant, which has received great attention in recent years for its potential in biofuel production and in greening and rehabilitation of wastelands. Yet the absence of improved cultivars and the lack of agronomic knowledge are limiting factors for successful jatropha cultivation. The objectives of the present study were to investigate the perspectives of a worldwide jatropha breeding program and specifically to (i) estimate variance components and heritabilities for agronomic and quality traits in the early phase of cultivation; (ii) assess phenotypic and genetic correlations among those traits; and (iii) discuss strategies for breeding high yielding jatropha cultivars. Data on various traits was collected from 375 jatropha genotypes, which were tested at seven locations during the first 3 years of growth. The accumulated seed yields and the seed yields per harvest year differed significantly among the testing locations. The estimates of genetic and genotype‐by‐environment interaction variances were significant and estimates of heritabilities were high for all yield parameters. The estimates of genetic correlations indicated a strong association among yield parameters. Oil yield was strongly correlated with seed yield and only weakly with oil content in seeds. The perspectives of a jatropha breeding program are excellent. Improved cultivars, definition of favorable environmental factors and refinement of agronomic management practices are needed to secure sustainable jatropha cultivation.