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Developmental Constraint Through Negative Pleiotropy in the Zygomatic Arch. Christopher J Percival

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Developmental Constraint Through Negative Pleiotropy in the Zygomatic Arch. Christopher J Percival The Jackson Laboratory The Mouseion at the JAXlibrary Faculty Research 2018 Faculty Research 1-27-2018 Developmental constraint through negative pleiotropy in the zygomatic arch. Christopher J Percival Rebecca Green Charles C Roseman Daniel M. Gatti The Jackson Laboratory, [email protected] Judy Morgan The Jackson Laboratory, [email protected] See next page for additional authors Follow this and additional works at: https://mouseion.jax.org/stfb2018 Part of the Life Sciences Commons, and the Medicine and Health Sciences Commons Recommended Citation Percival, Christopher J; Green, Rebecca; Roseman, Charles C; Gatti, Daniel M.; Morgan, Judy; Murray, Stephen A.; Donahue, Leah Rae; Mayeux, Jessica M; Pollard, K Michael; Hua, Kunjie; Pomp, Daniel; Marcucio, Ralph; and Hallgrímsson, Benedikt, "Developmental constraint through negative pleiotropy in the zygomatic arch." (2018). Faculty Research 2018. 38. https://mouseion.jax.org/stfb2018/38 This Article is brought to you for free and open access by the Faculty Research at The ousM eion at the JAXlibrary. It has been accepted for inclusion in Faculty Research 2018 by an authorized administrator of The ousM eion at the JAXlibrary. For more information, please contact [email protected]. Authors Christopher J Percival, Rebecca Green, Charles C Roseman, Daniel M. Gatti, Judy Morgan, Stephen A. Murray, Leah Rae Donahue, Jessica M Mayeux, K Michael Pollard, Kunjie Hua, Daniel Pomp, Ralph Marcucio, and Benedikt Hallgrímsson This article is available at The ousM eion at the JAXlibrary: https://mouseion.jax.org/stfb2018/38 Percival et al. EvoDevo (2018) 9:3 https://doi.org/10.1186/s13227-018-0092-3 EvoDevo RESEARCH Open Access Developmental constraint through negative pleiotropy in the zygomatic arch Christopher J. Percival1* , Rebecca Green2,3,4, Charles C. Roseman5, Daniel M. Gatti6, Judith L. Morgan6, Stephen A. Murray6, Leah Rae Donahue6, Jessica M. Mayeux7, K. Michael Pollard7, Kunjie Hua8, Daniel Pomp8, Ralph Marcucio9 and Benedikt Hallgrímsson2,3,4* Abstract Background: Previous analysis suggested that the relative contribution of individual bones to regional skull lengths difer between inbred mouse strains. If the negative correlation of adjacent bone lengths is associated with genetic variation in a heterogeneous population, it would be an example of negative pleiotropy, which occurs when a genetic factor leads to opposite efects in two phenotypes. Confrming negative pleiotropy and determining its basis may reveal important information about the maintenance of overall skull integration and developmental constraint on skull morphology. Results: We identifed negative correlations between the lengths of the frontal and parietal bones in the midline cranial vault as well as the zygomatic bone and zygomatic process of the maxilla, which contribute to the zygomatic arch. Through gene association mapping of a large heterogeneous population of Diversity Outbred (DO) mice, we identifed a quantitative trait locus on chromosome 17 driving the antagonistic contribution of these two zygomatic arch bones to total zygomatic arch length. Candidate genes in this region were identifed and real-time PCR of the maxillary processes of DO founder strain embryos indicated diferences in the RNA expression levels for two of the candidate genes, Camkmt and Six2. Conclusions: A genomic region underlying negative pleiotropy of two zygomatic arch bones was identifed, which provides a mechanism for antagonism in component bone lengths while constraining overall zygomatic arch length. This type of mechanism may have led to variation in the contribution of individual bones to the zygomatic arch noted across mammals. Given that similar genetic and developmental mechanisms may underlie negative correlations in other parts of the skull, these results provide an important step toward understanding the developmental basis of evolutionary variation and constraint in skull morphology. Keywords: Craniofacial, Skull, Micro-computed tomography, Morphometrics, Integration, QTL analysis, Diversity Outbred, RT-PCR, Mus musculus *Correspondence: [email protected]; [email protected] 1 Department of Anthropology, Stony Brook University, Stony Brook, NY, USA 4 Department of Cell Biology and Anatomy, University of Calgary, Calgary, AB, Canada Full list of author information is available at the end of the article © The Author(s) 2018. This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver (http://creativecommons.org/ publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated. Percival et al. EvoDevo (2018) 9:3 Page 2 of 16 Background correlations in relative size [16]. Here, we explicitly tested Te skull is a complex structure that supports and pro- whether pairs of adjacent linear distances within cranial tects tissues critical for survival, including the brain, vault, zygomatic arch, and cranial base are negatively cor- sense organs, and masticatory apparatus. While a wide related. In cases where they were, we performed genome- range of skull morphology has evolved across mam- wide mapping in a large heterogeneous population of malian species, fundamental patterns of skull bone Diversity Outbred (DO) mice to identify genomic regions integration are generally conserved [1–4]. Tis shared driving negative pleiotropy. Te DO mice are derived morphological pattern refects conserved tissue origins from the CC founder strains. Each mouse carries a high [5], ossifcation patterns [6–8], and strong selective pres- degree of heterozygosity and a unique combination of sure for an adequately integrated and functional head alleles [17, 18], which is ideal for high-resolution genetic [2, 9]. Te same factors that reinforce the development mapping. Although genome-wide mapping has been per- of an integrated head can serve as developmental con- formed on DO mice for other phenotypes such as blood straints on the directions that evolution can take [10, 11]. measurements and body composition [17–19], this is the Understanding the genetic basis for this integration and frst study that addresses craniofacial morphology. developmental constraint is critical for illuminating how Previous quantitative trait locus (QTL) analyses of genes and development can infuence processes of evo- mouse craniofacial variation have illustrated the ubiquity lution. Here, we investigate the genetic basis for nega- of pleiotropy in the genetics of craniofacial form [20–22] tive correlations between adjacent skull bones in a large and identifed candidate genes associated with variation genetically heterogeneous population of mice. Tese in multivariate measures of craniofacial shape in a vari- negative correlations may serve as developmental con- ety of mouse populations [23–27], including another out- straints that support overall integration of the head while bred mouse sample [28]. Te DO mice may prove more also allowing for signifcant variation in the relative size valuable than previously analyzed mouse crosses or back- of individual bones within subregions. Identifying genetic crosses, because they represent a strongly genetically het- factors underlying this type of developmental constraint erogeneous population that has been outbred for more is important for understanding the basis of morphologi- than 10 generations, leading to relatively high genomic cal integration and will illuminate critical connections mapping resolution. In addition, haplotype variation at between developmental and evolutionary processes. a given SNP can be tied back to the complete genomic Random pairs of linear distances across the skull are sequence of eight diverse inbred founder strains. expected to display positive correlations driven by the Within our mapping results, we expect that a locus overall growth of the integrated head. Negative correla- driving negative pleiotropy will have a signifcant haplo- tions between raw linear distance measurements are rare type efect on the linear distances of each adjacent bone. and can be evidence of scale-independent negative plei- Furthermore, we expect that variation in founder strain otropy, which exists when genetic variation leads to an haplotypes at this locus will be associated with oppo- opposite phenotypic efect on two traits [10, 12]. Within site phenotypic efects for each bone (i.e., one bone gets the craniofacial skeleton, a pair of negatively correlated longer when the other gets shorter). Tird, if this locus and adjacent bone lengths provides a relatively simple represents a developmental constraint on the morphol- system within which to search for genes underlying nega- ogy of the combined length of both bones, we expect that tive pleiotropy. Although not as commonly discussed, local haplotype variation will have no efect on overall negative pleiotropy is fundamentally diferent than antag- regional length. Identifying a genomic region associated onistic pleiotropy, which occurs when a genetic factor with negative pleiotropy and candidate genes within it is contributes a positive ftness efect for at least one trait a critical frst step in understanding an important genetic and a
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