Houle publications.

Publications

Most Recent

Valvo, J. J., J. D. Aponte, M. J. Daniel, K. Dwinell, H. Rodd, D. Houle, and K. A. Hughes. 2021. Using Delaunay triangulation to sample whole-specimen color from digital images. Ecology and Evolution DOI: 10.1002/ece3.7992. We present a spatially explicit technique for evaluating the entire color spectra of 2-D specimens that treates points as homologous when they have a particular relationship to landmarks and semi-landmarks detectable on the specimen. We apply this to color variation in Trinidadian guppies, finding little evidence for previously supported patterns of color variation.
Sztepanacz, J. L., and D. Houle. 2021. Allometry constrains the evolution of sexual dimorphism in Drosophila across 33 million years of divergence. Evolution. https://doi.org/10.1111/evo.14200 Dimorphism in wing shape is remarkably similar across the genus Drosophila Differences in shape dimorphism are largely due to allometry of with wing size, although we also detected significant among-species variation in non-allometric dimorphism. This pattern is consistent with constraint due to the relative lack of genetic variation in shape dimoprhism found by Sztepanacz and Houle (2019)
Houle, D., G. H. Bolstad, and T. F. Hansen. 2020. Fly wing evolutionary rate is a near-isometric function of mutational variation. BioRxiv. https://www.biorxiv.org/content/10.1101/2020.08.27.268938v1 Using simulations, we confirm that Jiang and Zhang's (2020) finding that our method to determine the scaling relationship between mutational, standing genetic and among-species variation, used in Houle et al. (2017) is highly biased. Jiang and Zhang's proposed method is also biased. We propose and validate a new method where a third, statistically independent matrix is used as a standard that yields nearly unbiased estimates. Analyses using the Q methods confirm that the conclusions of Houle et al. (2017) were correct.
Houle, D., and C. Cheng. 2021. Predicting the evolution of sexual dimorphism in gene expression. Molecular Biology and Evolution. https://doi.org/10.1093/molbev/msaa329 We estimated the G matrix for gene expression in Drosophila melanogaster using existing data on gene expression in DGRP lines. Using the results of Cheng and Houle (2020) we show that the evolution of changes in dimorphism of gene experession in this sepcies is about equally likely to result from concordant selection as it is to result from antagonsitic selection.
Houle, D., and L. T. Jones. 2021. Micro-evo-devo, Pp. 1047-1060 in L. Nuño de la Rosa, and G. B. Müller, eds. Evolutionary Developmental Biology - A Reference Guide, Springer. https://doi.org/10.1007/978-3-319-32979-6_189 We review how microevolutionary stuides of the evolution of development can supplement the prevaling focus of evo-devo on macroevolutionary changes
Cheng, C., and D. Houle. 2020. Predicting multivariate responses to direct and indirect selection on sexual dimorphism. American Naturalist 196:391-405. DOI: 10.1086/710353. We studied the genetic variation of homologous traits in separate sexes by rotating the phenotype space to consist of the average trait value and the differences between sexes. We show that in the early stages of the evolution of sexual dimorphims, dimoprhism is much more likely to evolve as an indirect response of trait dimorphism to concordant selection on the two sexes than to direct responses to antagonstic selection. This may require reinterpretation of the adaptive significance of many sexual dimorphisms.
Sztepanacz, J. L., and D. Houle. 2019. Cross-sex genetic covariances limit the evolvability of wing-shape within and among species of Drosophila. Evolution 73:1617-1633. DOI: 10.1111/evo.13788. We have previously demonstrated that all twenty studied aspects of sex-averaged wing shape are genetically variable. Here we show that the sex-specific genetic variation is quite limited. Only six additional dimensions have statistically supported genetic variation when the phenotypes of each sex are considered together.
Houle, D., L. T. Jones, R. Fortune, and J. L. Sztepanacz. 2019. Why does allometry evolve so slowly? Integrative and Comparative Biology 59:1429-1440. DOI:10.1093/icb/icz099 We consider various hypotheses to explain the slow rate of evolution of allometric relationships. Selection experiments and direct measurements rule out the possiblity of constraints due to a lack of genetic variation. A plausible cause of constraint is that variants affecting allometry have widespread pleiotropic effects so that alterations in allometric relationships of one aspect of form have deleterious effects on many traits.
Pitchers, W., J. Nye, E. J. Márquez, A. Kowalski, I. Dworkin, and D. Houle. 2019. A Multivariate Genome-Wide Association Study of Wing Shape in Drosophila melanogaster. Genetics 211:1429-1447. DOI: 10.1534/genetics.118.301342. We report a fully multivariate genome-wide association study of wing shape. We find robust evidence that at least 500 SNPs affect wing shape in a population of DGRP genotypes. Some of these SNPs have effects that conform to the effects of RNAi knockdowns at loci near these SNPs. The implicated loci are concentrated in known developmental pahtways important for wing developments
Matamoro-Vidal, A., Y. Huang, I. Salazar-Ciudad, O. Shimmi and D. Houle. 2018. Quantitative morphological variation in the developing Drosophila wing. G3. 8:2399-2409. We extend the use of morphometrics of the wing to include the shape of wing poutch in third-instar larvae. This allows us to diagnose when genotypes affect wing shape.

2015-2017

Houle, D., G. H. Bolstad, K. Van Der Linde, and T. F. Hansen. 2017. Mutation predicts 40 million years of fly wing evolution. Nature in press. DOI:10.1038/nature23473. We demonstrate that the rate of evolution of wing shape in the family Drosophilidae is highly correlated with both the mutational and standing genetic variation within one species. The rate of evolution is very low, calling into question whether this can be explained by the amount of mutational variation constraining the rate of adaptation. An alternative explanation is that mutation and standing variation are shaped by the same selective forces that determine the rate of among species evolution.
Hollis, B., D. Houle and T.J. Kawecki 2016. Evolution of postcopulatory molecular interactions in Drosophila populations lacking sperm competition. Journal of Evolutionary Biology. 29:77-85.
Bolstad, G. H., J. A. Cassara, E. Marquez, T. F. Hansen, K. Van Der Linde, D. Houle, and C. Pelabon. 2015. Proceedings of the National Academy of Sciences 112:13284-13289 doi: 10.1073/pnas.1505357112 Continuing our series of selection experiments on apparently constrained traits, we demonstrate both genetic variation and strong selection favoring allometry. Artificial selection drove the allometry of a wing vein beyond the range of that found in the genus Drosophila in one year, but strong countervailing natural selection rapidly returned the population to its original allometry.
Ray, R. P., A. Matamoro-Vidal, P. S. Ribiero, N. Tapon, D. Houle, I. Salazar-Ciudad, and B. J. Thompson. 2015. Patterned Anchorage to the Apical Extracellular Matrix Defines Tissue Shape in the Developing Appendages of Drosophila. Developmental Cell, 34:310-322. doi: 10.1016/j.devcel.2015.06.019. The Drosophila wing is stretched into its adult shape by contraction of the tissues connecting the wing to the body pulling against the attachment of the distal wing to the cuticle . Our co-authors have demonstrated genes that influence this process, and our cell-based model reproduces mutant effects on wing shape.
Houle, D., and K. Meyer. 2015. Estimating sampling error of evolutionary statistics based on genetic covariance matrices using maximum likelihood. Journal of Evolutionary Biology 28:1542-1549. doi: 10.1111/jeb.12674. Programs, data. We describe a rapid, maximum likelihood-based method for assessing the error in G matrices and statistics that depend on them. The method yields similar results to other more computationally intensive methods.
Houle, D., and E. Marquez. 2015. Linkage Disequilibrium and Inversion-Typing of the Drosophila melanogaster Genome Reference Panel. G3 doi:10.1534/g3.115.019554. We calculated linkage disequilbrium between all variable sites in this set of sequenced lines from a single population, and the list of correlated sites is available from the Supplementary Information. The pattern of LD also reveals the inversion genotypes of the lines, in some cases contradicting previous analyses.
Matamoro-Vidal, A., I. Salazar-Ciudad, and D. Houle. 2015. Making quantitative morphological variation from basic developmental processes: where are we? The case of the Drosophila wing. Developmental Dynamics DOI: 10.1002/DVDY.24255. We review the major developmental events that shape the Drosophila wing, and consider how these processes influence variation of the adult wing.

2013-2014

Pelabon,C., C. Firmat, G. H. Bolstad, K. L. Voje, D. Houle, J. Cassara,A. Le Rouzic and T. F. Hansen 2014. Evolution of morphological allometry. Annals of the New York Academy of Sciences. 1320:58-75. doi: 10.1111/nyas.12470. Suprisingly little is know about the evolution of allometry. Many experiments have addressed the evolution of shape, rather than allometry.
Hollis, B., Houle, D., Yan, Z., Kawecki, T. J. and Keller, L. 2014. Evolution under monogamy feminizes gene expression in Drosophila melanogaster. Nature Communications 5:3482. When flies mate in single pairs, with no opportunity for competition or choice, gene expression in males becomes more like that of females.
Meyer, K., and D. Houle , 2013. Sampling based approximation of confidence intervals for functions of genetic covariance matrices. Proceedings of the Association for Advances in Animal Breeding, 20: 523-527. Conference in Napier, New Zealand. We demonstrate a simple, fast method for generating confidence intervals for stastics based on estimates of genetic variance-covariance matrices. It performs very well in simulations. See Houle and Meyer 2015 for follow-up.
Schrider, D., D.Houle, M. Lynch and M. W. Hahn. 2013. Rates and genomic consequences of spontaneous mutational events in Drosophila melanogaster. Genetics. 194:937-954. We document the mutation rates to a wide variety of changes, including larger deletions and insertions that have been neglected in earlier analyses. We show the existence of multiple mutation events, where one event leads to changes at multiple base pairs that are not contiguous in the DNA sequence.
Houle, D. and J. Fierst. 2013. Properties of spontaneous mutational variance and covariance for wing size and shape in Drosophila melanogaster. Evolution 67:1116-1130. Supplementary Information We studied spontaneous mutational variance and covariance for wing size and shape, finding that mutation affects many independent aspects of shape. We propose new exploratory comparisons of the differences and similarities between covariance matrices.

2010-2012

Function-Valued Traits Working Group (Stinchcombe et al.). 2012. Genetics and evolution of function-valued traits: understanding environmentally responsive phenotypes. Trends in Ecology and Evolution 27:637-647. pdf This working group review details why treating traits as functions, rather than sets of multivariate data, can aid analysis and interpretation.
Marquez, E. J., R. Cabeen, R. P. Woods, D. Houle. 2012. The measurement of local variation in shape. Evolutionary Biology 39:419-439. pdf Understanding changes in organismal shape requires examining local expansions and contractions of one form relative to another. We investigate the use of alternative interpolation functions to infer such changes. Lory is our software package for visualizing such changes.
Houle, D. C. Pelabon, G. P. Wagner, T. F. Hansen. 2011. Measurement and meaning in biology. Quarterly Review of Biology 86:1-32. pdf Measurement theory examines the relationship between aspects of the natural world and the numbers we use to represent them. It helps us recognize what statements about our measurements can be meaningful. Every scientist should be familiar with measurement theory, but our review of evolutionary publications reveals many errors that are violations of measurement principles.
Carter, A. J. R., and D. Houle 2011. Artificial selection reveals heritable variation for developmental instability. Evolution 65:3558-3564. pdf A long tern-selection experiment for wing asymmetry in Drosophila melanogaster shows responses for both increased and (in one replicate) decreased symmetry. This is the first robust evidence for genetic variance in asymmetry segregating in natural populations.
Hansen, T. F., C. Pelabon, and D. Houle. 2011. Heritability is not evolvability. Evolutionary Biology 38:258-277. pdf We delve more deeply into the reasons that heritability is unrelated to the ability to respond to selection. Our review of estimates published since Houle 1992 reveals the same lack of correlation between mean and variance-standardized measures of genetic variance.
Hollis, B., and D. Houle. 2011.Populations with elevated mutation load do not benefit from the operation of sexual selection. Journal of Evolutionary Biology 24:1918-1926. pdf We found that sexual selection did not increase the fitness of long-term selection lines whose mutation load was enhanced by mutagenesis. This argues that sexual conflict is important.
Le Rouzic, A., D. Houle, and T. F. Hansen. 2011. A modelling framework for the analysis of artificial-selection time series. Genetics Research 93:155-173. pdf Proposes informative analyses of artificial selection experiments. Some of these simply describe patterns in the responses, such as the rate of decrease in response over time, while others fit biological models including processes such as drift, mutation, and epistasis. Introduces a software package in R to perform these analyses.
Houle, D., D. R. Govindaraju and S. W. Omholt , 2010 Phenomics: the next challenge. Nature Reviews Genetics 11: 855-866. pdf We review the advantages of doing phenomics, high-throughput phenotyping, and the progress towards making phenomics practical.
Van der Linde, K., D. Houle, G. Spicer, and S. J. Steppan. 2010. A supermatrix analysis of the family Drosophilidae. Genetics Research 92:25-38. pdf We analyse a supermatrix covering a large number of species, including many closely related genera to Drosophila. We confirm the polyphyletic nature of the genus, and provide improved resolution in some parts of the phylogeny.
Fan, Y., D. Houle, W. Mio. 2010 Learning Metrics for Shape Classification and Discrimination. 20th International Conference on Pattern Recognition ICPR 2010: 2652-2655. pdf We propose a family of shape metrics that generalize the classical Procrustes distance by attributing weights to general linear combinations of landmarks. We develop an algorithm to learn a metric that is optimally suited to a given shape classification problem.
Pelabon, C., T. F. Hansen, A. J. R. Carter, and D. Houle. 2010. Evolution of variation and variability under fluctuating, stabilizing, and disruptive selection.Evolution 64:1912-1925. pdf Fluctuating and stabilizing selection decreased phenotypic variance and disruptive selection increased it, but all forms of selection increased fluctuating asymmetry. This suggests that canalization and developmental stability have different genetic bases.
Houle, D. 2010. Numbering the hairs on our heads: The shared challenge and promise of phenomics. Proceedings of the National Academy of Sciences USA. 107:1793-1799. doi:10.1073/pnas.0906195106. PMID: 19858477 pdf Phenomics, the rapid characterization of phenotypes, is a natural and inevitable complement to genomics.

2009-2007

Gomulkiewicz, R., and D. Houle. 2009. Demographic and genetic constraints on evolution. American Naturalist 174:E218-E229. PDF. The necessity to avoid extinction when subject to directional selection imposes a constraint on evolution, even when there is genetic variance allowing response to selection.
Van der Linde, K. and D. Houle. 2009. Inferring the nature of allometry from geometric data. Evolutionary Biology 36:311-322.PDF. We discuss problems with the generalized Procrustes approach registration to of morphological data when biological hypotheses need to be tested. We suggest two modifications to the registration procedure using subsets of the data that can improve the fit to some biological hypotheses.
Carter, A. J. R., T. M. Weier, D. Houle. 2009. Effect of inbreeding on fluctuating asymmetry of wing veins in two laboratory strains of Drosophila melanogaster. Heredity 102: 177-190.PDF.
Hollis, B., J. Fierst, and D. Houle. 2009. Sexual selection accelerates the elimination of a deleterious mutant in Drosophila melanogaster. Evolution 63:324-333.PDF. A null allele at alcohol dehydrogenase is a bad gene, suggesting that many other deleterious alleles could render males less attractive or less competitive for mates.
Tomaiuolo, M., R. Bertram and D. Houle. 2008.Enzyme isoforms may increase phenotypic robustness. Evolution 62:2868-2883. PDF. We showed that having two forms of an enzyme can increase the range of environments over which an individual functions properly. This provides a novel hypothesis for the divergence of gene function in evolution.
Hansen, T. F., and D. Houle. 2008.Measuring and comparing evolvability and constraint in multivariate characters. Journal of Evolutionary Biology21:1201-1219. PDF. We derive multivariate measures of the ability to evolve, the degree of conflict between the evolvability of character combinations, and measures of differences among populations based on these measures. This copy includes corrections to published version. The published Appendices contains several errors, and these in turn affect the approximations for mean conditional evolvability, (Eq. 5), mean respondability, (p. 1206), mean autonomy, (Eq. 6), and mean response difference, (Eq. 8). Thanks to John Stinchcombe for pointing out the problem.
Haag-Liautard, C., N. Coffey, D. Houle, M. Lynch, B. Charlesworth and P. D. Keightley. 2008. Direct estimation of the mitochondrial DNA mutation rate in D. melanogaster. PLOS Biology 6:1706-1714. PDF. We showed that the spontaneous mutation rate of mitochodria is higher than in the nuclear genome, and that there are peculiar biases in the transition probabilities when a mutation occurs.
Van der Linde, K. and D. Houle. 2008b. Response to OÕGrady et al.: The potential and peril of the supertree approach. Insect Systematics and Evolution 39:281-286. PDF. OÕGrady et al. criticized van der Linde and Houle 2008a for various reasons. We more fully explain the methods used, and show that their criticisms are largely misunderstandings of what we did, or are misconceptions about the nature of super-tree analyses.
Van der Linde, K. and D. Houle. 2008a. A supertree analysis and literature review of the genus Drosophila and closely related genera (Diptera, Drosophilidae). Insect Systematics and Evolution39:241-268. PDF. We use a supertree analysis to estimate the phylogeny of the subfamily Drosophilinae, which includes the Drosophila melanogaster. Our results suggest that the genus Drosophila is paraphyletic.
Houle, D. 2007. A dispatch from the multivariate frontier. Journal of Evolutionary Biology. 20:22-23.
Haag-Liautard, C., Dorris, X. Maside, S. Macaskill, L. Halligan, D. Houle, D. Charlesworth, P. D. Keightley. 2007. Direct estimation of per nucleotide and genomic deleterious mutation rates in Drosophila. Nature 445: 82-85. We estimated the spontaneous mutation rate of the nuclear genome using mutation accumulation lines from my lab.

2006-2004

PŽlabon, C., T. F. Hansen, A. J. R. Carter and D. Houle. 2006. Response of fluctuating and directional asymmetry to selection on wing shape in Drosophila melanogaster. Journal of Evolutionary Biology. 19:764-776.PDF.
van der Linde, K., J. S. Birdsley, L. M. Alonso and D. Houle. 2006. First records of Zaprionus indianus (Diptera, Drosophilidae), a potential pest species, from Panama and the United States of America. Florida Entomologist 89:402-404. PDF. We report the first records for this species in North America. Z. indianus first appeared in south Florida in July, and in Tallahassee, Florida between July and September 2005.
Houle, D. and Kondrashov, A.S. 2006. Mutation. Pp. 32-48 in Evolutionary Genetics: Concepts and Case Studies, edited C.W. Fox and J. B. Wolf. Oxford University Press. PDF.
Rifkin, S. A., D. Houle, J. Kim, and K. P. White. 2005. A mutation accumulation assay reveals a broad capacity for rapid evolution of gene expression. Nature 438:220-223. This study of mutation-accumulation lines revealed that expression of 40% of the nearly 12,000 genes studied had been affected by mutation. The rate of evolution between species is much lower than the neutral expectation based on this variation, implicating stabilizing selection on expression.
Fuller, R. C., D. Houle, and J. Travis. 2005. Sensory bias as an explanation for the evolution of mate preferences. American Naturalist 166:437-446. PDF. We place the five major models for mate preference in the same quantitative genetic context, revealing that much of the discussion of these models is misguided. In particular, most workers do not understand the sensory bias model, and it has therefore never been tested.
Mezey, J. G., D. Houle, and S. V. Nuzhdin. 2005. Naturally segregating quantitative trait loci affecting wing shape of Drosophila melanogaster.Genetics 169:2101-2113. PDF. We confirmed that many genes have variation affecting wing shape within our study population. Furthermore some of these show genetic interactions with the hedgehog and decapentaplegic pathways.
Mezey, J., and D. Houle. 2005. The dimensionality of genetic variation for wing shape in Drosophila melanogaster. Evolution 59:1027-1038. PDF. We studied the inheritance of 20 aspects of wing shape within a single population, and were able to demonstrate genetic variation in at least 17 of these 20. This suggests that the evolution of wing shape is not constrained by a lack of genetic variation.
Hereford, J., T. F. Hansen and D. Houle. 2004. Comparing strengths of directional selection: how strong is strong? Evolution 58:2133-2143. PDF. Appendix giving references to reviewed papers has been dropped from the Evolution web site. The strength of selection standardized by the mean of the trait is directly comparable to the strength of selection on fitness. Review of published estimates suggests that selection is generally overestimated, but it is not possible to choose among the many possible explanations for this.
McCune, A.R., D. Houle, K. McMillan, R. Annable and A. S. Kondrashov. 2004. Two classes of deleterious recessive alleles in a natural population of zebrafish, Danio rerio. Proc. Roy. Soc. London B. 271:2025-2033. PDF. Inbreeding depression in viability is due in part to recessive lethals that act early in development, but also to later-acting smaller effect mutations.
Houle, D. and S.V. Nuzhdin. 2004. Mutation accumulation and the effect ofcopia insertions in Drosophila melanogaster. Genetical Research. 83:7-18. PDF.
Hansen, T. F., and D. Houle. 2004. Evolvability, stabilizing selection, and the problem of stasis. Pp. 130-150 in The Evolutionary Biology of Complex Phenotypes. M. Pigliucci and K. Preston, Eds. Oxford University Press. PDF. We discuss possible explanations for the frequently observed lack of evolutionary change over long periods of time. The widely accepted idea that stabilizing selection is responsible seems implausible, and is not supported by evidence. The alternative that there are constraints on variation should be taken more seriously.

2003-2000

Houle, D., J.G. Mezey, P. Galpern and A. Carter. 2003. Automated measurement of Drosophila wings. BMC Evolutionary Biology. 3:25. ONLINE. This paper introduces our mostly automated system for measuring Drosophila wings.
Mezey, J. and D. Houle. 2003. Comparing G matrices: Are common principal components informative? Genetics 165:411-425. PDF. Proportional (or more similar) G matrices imply a modular organisation of phenotypes, where each module affects a phenotypic subspace orthogonal to other such modules. Powerful CPC analyses therefore can tell us something fundamental about genotype-phenotype maps.
Worley, A. C., D. Houle, and S. C. H. Barrett. 2003. PDF. Consequences of hierarchical allocation for the evolution of life-history traits. American Naturalist 161:153-167. We use numerical calculations to investigate when trade-offs can be revealed by selection in a multi-trade-off hierarchy.
Houle, D., and L. Rowe. 2003. Natural selection in a bottle. American Naturalist. 161:50-67. PDF. We suggest that the study of natural selection in laboratory situations can provide an essential test of our ability to understand and predict adaptation. We present data on the fitness landscape for age at maturity of my laboratory fly population.
Fuller, R. C. and D. Houle. 2003. Inheritance of developmental instability. Pp. 157-184 in Developmental Instability: Causes and Consequences, edited by M. Polak. Oxford University Press. PDF. Traditional designs for estimating genetic variation in fluctuating asymmetry have no power. Therefore, the general lack of studies showing significant heritability of FA is not informative.
Fuller, R. C., and D. Houle. 2002. Detecting variation in developmental instability by artificial selection on fluctuating asymmetry. Journal of Evolutionary Biology. 15:954-960. PDF. We investigate the power of selection experiments to detect genetic variation in the ability to produce a symmetrical body. Selection for decreased symmetry, which is rarely performed, is much more powerful than selection for increased symmetry.
Steppan, S. J., P. C. Phillips, and D. Houle. 2002. Comparative quantitative genetics: evolution of the G matrix. Trends in Ecology and Evolution17:320-327. PDF. We review approaches to the study of the additive genetic variance-covariance matrix, concluding that proper analytical and statistical tools are just now becoming available to test interesting hypotheses.
McCune, A. R, R. C. Fuller, A. A. Aquilina, R. M. Dawley, J. M. Fadool, D. Houle, J. Travis, and A. S. Kondrashov. 2002. Low number of genomic recessive lethals in two fish species. Science 296:2398-2401. PDF. We show that the average number of lethals carried in two outbred fish populations is similar to the number carried by Drosophila, which has a much smaller genome.
Houle, D, J. Mezey. and Galpern, P. 2002. Interpretation of the results of partial principal components analysis. Evolution 56:433-440. PDF. We show that a popular new technique for comparing variance-covariance matrices gives results that do not reflect biological intuition, and we urge caution in its use.
Houle, D. and A.S. Kondrashov. 2002. Coevolution of costly mate choice and condition-dependent display of good genes. Proceedings of the Royal Society of London, Biological Sciences, 269:97-104. PDF. We show that a relatively simple good-genes model in which female choice is relative can explain even extreme exaggeration of male traits by sexual selection.
Houle, D. 2001. The character problem in life history evolution. Pp. 109-140 in, The Character Concept in Evolutionary Biology. G. P. Wagner, ed. Academic Press. PDF. I argue that the most appropriate evolutionary definition of a character is a biological process that integrates the effects of many genetic loci. I argue that such processes are likely to exist and to form an appropriate target for analysis intermediate to black box quantitative genetics and fully detailed functional genomics.
Houle, D. 2000. A simple model of the relationship between asymmetry and developmental stability. Journal of Evolutionary Biology 13:720-730. PDF. I showed that the standard model for the development of asymmetry of morphological traits cannot readily explain the strong relationships between asymmetry and fitness that some claim to have observed.
Houle, D. 2000. Is there a g factor for fitness? Pp. 149-159 in The Nature of Intelligence, Novartis Foundation Symposium 233. John Wiley, London. PDF. I investigated the idea that variation in fitness components covary positively, as do the components of test-taking ability in humans. I conclude that the idea is plausible but cannot be confirmed because suitable data are lacking.
Leung, B., M. R. Forbes, and D. Houle. 2000. Fluctuating asymmetry as a bioindicator of stress: comparing efficacy of analyses involving multiple traits. Am. Nat. 155:101-115. PDF.

1999 and Earlier

Pletcher, S.D., D. Houle, and J.W. Curtsinger. 1999. The evolution of age-specific mortality rates in Drosophila melanogaster: divergence among unselected lines. Genetics 153:813-823. PDF.
Lynch, M., J. Blanchard, D. Houle, T. Kibota, S. Schultz, L. Vassilieva, and J. Willis. 1999. Perspective: Spontaneous deleterious mutation. Evolution53: 645-663.
Houle, D. 1998. High enthusiasm and low R-squared. Asymmetry, developmental stability, and evolution, by A.P. M¿ller and J. P. Swaddle.Evolution 52:1872-1876. PDF.
Houle, D. 1998. How should we explain variation in the genetic variance of traits? Genetica, 102/103:241-253. PDF. I tested a series of hypotheses about the differences in mean-standardized variation in traits. More genetically complex traits had larger mutational inputs and more standing genetic variation. I really like this paper.
Pletcher, S.D., D. Houle, and J.W. Curtsinger. 1998. Age-specific properties of spontaneous mutations affecting mortality in Drosophila melanogaster. Genetics, 148:287-303. PDF.
Houle, D., A. S. Kondrashov, L. Yu. Yampolsky, S. Caldwell, and P. L. Steponkus. 1997. The effect of cryopreservation on the lethal mutation rate in Drosophila melanogaster. Genetical Research, 69:209-213. PDF.
Houle, D., K. A. Hughes, S. Assimacopolous, and B. Charlesworth. 1997.The effects of spontaneous mutation on quantitative traits. II. Dominance of mutations with effects on life-history traits. Genetical Research, 70:27-34. PDF.
Houle, D. 1997. Comment on "A meta-analysis of the heritability of developmental stability" by Moller and Thornhill. J. Evol. Biol. 10:17-20. ONLINE.
Houle, D., B. Morikawa, and M. Lynch. 1996. Comparing mutational variabilities. Genetics, 143:1467-1483. PDF.
Rowe, L. and D. Houle. 1996. The lek paradox, condition dependence and genetic variance in sexually selected traits. Proc. Roy. Soc. London B, 263:1415-1421. PDF.
Houle, D., K. A. Hughes, D. K. Hoffmaster, J. Ihara, S. Assimacopolous, D. Canada, and B. Charlesworth. 1994. The effects of spontaneous mutation on quantitative traits. I. Variance and covariance of life history traits. Genetics,138:773-785. PDF.
Kondrashov, A. S. and D. Houle. 1994. Genotype-environment interactions and the estimation of the genomic mutation rate in Drosophila melanogaster. Proc. Roy. Soc. London B, 258:221-227. PDF.
Houle, D. 1994. Adaptive distance and the genetic basis of heterosis.Evolution 48:1410-1417. PDF.
Houle, D. 1992. Comparing evolvability and variability of quantitative traits.Genetics 130:195-204. PDF.Data set and references available here.
Houle, D., D. Hoffmaster, S. Assimacopolous, and B. Charlesworth. 1992.The genomic mutation rate for fitness in Drosophila. Nature 359:58-60. However, see Correction, Nature, 371:358.
Houle, D. 1991. Genetic covariance of life history traits: what genetic correlations are made of and why it matters. Evolution 45:630-648. PDF.
Zeng, Z.-B., D. Houle, and C. C. Cockerham. 1990. How informative is Wright's estimator of the number of genes affecting a quantitative character? Genetics 126:235-247. PDF.
Houle, D. 1989. Allozyme-associated heterosis in Drosophila melanogaster.Genetics 123:789-801. PDF.
Houle, D. 1989. The maintenance of polygenic variation in finite populations.Evolution 43:1767-1780. PDF.
Eanes, W. F., C. Wesley, J. Hey, D. Houle, and J. W. Ajioka. 1988. The fitness consequences of P-element insertion in Drosophila melanogaster. Genetical Research 52:17-26
Hey, J. and D. Houle. 1987. Habitat choice in the Drosophila affinis subgroup. Heredity 58:463-471.
Eanes, W. F., J. Hey, and D. Houle. 1985. Homozygous and hemizygous viability variation on the X chromosome of Drosophila melanogaster. Genetics 111:831-844.
Eanes, W. F., B. Bingham, J. Hey, and D. Houle. 1985. Targeted selection experiments and enzyme polymorphism: negative evidence for octanoate selection at the G6PD locus in Drosophila melanogaster. Genetics 109:379-391.

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