FREE SHIPPING over $40
OVER 10000 FIVE-STAR REVIEWS
5% OFF ALL SUBSCRIPTIONS
100% HAPPINESS GUARANTEE
Analyze Diet
0
PloS one2013; 8(2); e56497; doi: 10.1371/journal.pone.0056497

Expression levels of LCORL are associated with body size in horses.

Abstract: Body size is an important characteristic for horses of various breeds and essential for the classification of ponies concerning the limit value of 148 cm (58.27 inches) height at the withers. Genome-wide association analyses revealed the highest associated quantitative trait locus for height at the withers on horse chromosome (ECA) 3 upstream of the candidate gene LCORL. Using 214 Hanoverian horses genotyped on the Illumina equine SNP50 BeadChip and 42 different horse breeds across all size ranges, we confirmed the highly associated single nucleotide polymorphism BIEC2-808543 (-log(10)P = 8.3) and the adjacent gene LCORL as the most promising candidate for body size. We investigated the relative expression levels of LCORL and its two neighbouring genes NCAPG and DCAF16 using quantitative real-time PCR (RT-qPCR). We could demonstrate a significant association of the relative LCORL expression levels with the size of the horses and the BIEC2-808543 genotypes within and across horse breeds. In heterozygous C/T-horses expression levels of LCORL were significantly decreased by 40% and in homozygous C/C-horses by 56% relative to the smaller T/T-horses. Bioinformatic analyses indicated that this SNP T>C mutation is disrupting a putative binding site of the transcription factor TFIID which is important for the transcription process of genes involved in skeletal bone development. Thus, our findings suggest that expression levels of LCORL play a key role for body size within and across horse breeds and regulation of the expression of LCORL is associated with genetic variants of BIEC2-808543. This is the first functional study for a body size regulating polymorphism in horses and a further step to unravel the mechanisms for understanding the genetic regulation of body size in horses.
Get Full Text
Publication Date: 2013-02-13 PubMed ID: 23418579PubMed Central: PMC3572084DOI: 10.1371/journal.pone.0056497Google Scholar: Lookup
The Equine Research Bank provides access to a large database of publicly available scientific literature. Inclusion in the Research Bank does not imply endorsement of study methods or findings by Mad Barn.
LinkedInCopy
  • Journal Article
  • Research Support
  • Non-U.S. Gov't

Summary

This research summary has been generated with artificial intelligence and may contain errors and omissions. Refer to the original study to confirm details provided. Submit correction.

The research investigated how the expression levels of the LCORL gene are related to the body size in horses. The dominant gene locus controlling the height in horses was found upstream in the LCORL gene. The study verifies this fact by examining different horse breeds and the relationship between LCORL expression and body size.

Association between genetic coding and body size

  • The study focused on the strong correlation between LCORL gene expression and the size of horses. This was analyzed in a diverse set of 42 horse breeds. The size range of the horses in the study was extensive, thereby supporting the study’s conclusion on the influence of the LCORL gene on body size.
  • Analyses on the genome-wide scale pointed out the quantitative trait locus (the part of the genome which is associated with a particular phenotypic trait) responsible for height at withers (the ridge between the shoulder blades of an animal) was located on horse chromosome 3 near the LCORL gene.

LCORL gene expression

  • The study measured and compared the expression levels of LCORL and two neighbouring genes (NCAPG and DCAF16) using RT-qPCR (Real-Time Quantitative Reverse Transcription PCR). This method is widely used for measuring gene expression levels.
  • The results showed significant association between the relative LCORL expression levels and the size of the horses. In particular, the study found that horses with different BIEC2-808543 genotypes (expressions of a particular polymorphism) had distinct LCORL expression levels. This further solidifies the role of the LCORL gene’s expression in determining a horse’s size.

Role of Specific Single Nucleotide Polymorphism

  • In the heterozygous (two different alleles of the same gene) C/T-horses, LCORL expression was found to be decreased by 40% and in homozygous (two identical alleles of the same gene) C/C-horses, it was decreased by 56% in comparison to smaller T/T-horses.
  • The research suggests that a SNP (Single Nucleotide Polymorphism – a variation at a single site in DNA) T>C mutation disrupts a binding site of the transcription factor TFIID that plays a role in skeletal bone development.

Hence, the study wraps up suggesting that the LCORL gene’s expression levels play a pivotal role in regulating body size within and across different horse breeds. This study gives notable insights into understanding the genetic regulation of body size in horses.

Cite This Article

APA
Metzger J, Schrimpf R, Philipp U, Distl O. (2013). Expression levels of LCORL are associated with body size in horses. PLoS One, 8(2), e56497. https://doi.org/10.1371/journal.pone.0056497

Publication

PloS one
ISSN: 1932-6203
NlmUniqueID: 101285081
Country: United States
Language: English
Volume: 8
Issue: 2
Pages: e56497
PII: e56497

Researcher Affiliations

Metzger, Julia
  • Institute for Animal Breeding and Genetics, University of Veterinary Medicine Hannover, Hannover, Germany.
Schrimpf, Rahel
    Philipp, Ute
      Distl, Ottmar

        MeSH Terms

        • Animals
        • Body Size / genetics
        • Breeding
        • Chromosome Mapping
        • Chromosomes, Mammalian / genetics
        • Female
        • Gene Expression Profiling / veterinary
        • Gene Frequency
        • Genotype
        • Horses / genetics
        • Male
        • Molecular Sequence Data
        • Polymorphism, Single Nucleotide
        • Protein Isoforms / genetics
        • Reverse Transcriptase Polymerase Chain Reaction
        • Sequence Analysis, DNA
        • Trans-Activators / genetics

        Conflict of Interest Statement

        The authors have declared that no competing interests exist.

        References

        This article includes 41 references
        1. Signer-Hasler H, Flury C, Haase B, Burger D, Simianer H, Leeb T, Rieder S. A genome-wide association study reveals loci influencing height and other conformation traits in horses.. PLoS One 2012;7(5):e37282.
          pmc: PMC3353922pubmed: 22615965doi: 10.1371/journal.pone.0037282google scholar: lookup
        2. Makvandi-Nejad S, Hoffman GE, Allen JJ, Chu E, Gu E, Chandler AM, Loredo AI, Bellone RR, Mezey JG, Brooks SA, Sutter NB. Four loci explain 83% of size variation in the horse.. PLoS One 2012;7(7):e39929.
          pmc: PMC3394777pubmed: 22808074doi: 10.1371/journal.pone.0039929google scholar: lookup
        3. Gudbjartsson DF, Walters GB, Thorleifsson G, Stefansson H, Halldorsson BV, Zusmanovich P, Sulem P, Thorlacius S, Gylfason A, Steinberg S, Helgadottir A, Ingason A, Steinthorsdottir V, Olafsdottir EJ, Olafsdottir GH, Jonsson T, Borch-Johnsen K, Hansen T, Andersen G, Jorgensen T, Pedersen O, Aben KK, Witjes JA, Swinkels DW, den Heijer M, Franke B, Verbeek AL, Becker DM, Yanek LR, Becker LC, Tryggvadottir L, Rafnar T, Gulcher J, Kiemeney LA, Kong A, Thorsteinsdottir U, Stefansson K. Many sequence variants affecting diversity of adult human height.. Nat Genet 2008 May;40(5):609-15.
          pubmed: 18391951doi: 10.1038/ng.122google scholar: lookup
        4. Lindholm-Perry AK, Sexten AK, Kuehn LA, Smith TP, King DA, Shackelford SD, Wheeler TL, Ferrell CL, Jenkins TG, Snelling WM, Freetly HC. Association, effects and validation of polymorphisms within the NCAPG - LCORL locus located on BTA6 with feed intake, gain, meat and carcass traits in beef cattle.. BMC Genet 2011 Dec 14;12:103.
          pmc: PMC3287254pubmed: 22168586doi: 10.1186/1471-2156-12-103google scholar: lookup
        5. Shriner D, Adeyemo A, Gerry NP, Herbert A, Chen G, Doumatey A, Huang H, Zhou J, Christman MF, Rotimi CN. Transferability and fine-mapping of genome-wide associated loci for adult height across human populations.. PLoS One 2009 Dec 22;4(12):e8398.
          pmc: PMC2792725pubmed: 20027299doi: 10.1371/journal.pone.0008398google scholar: lookup
        6. Soranzo N, Rivadeneira F, Chinappen-Horsley U, Malkina I, Richards JB, Hammond N, Stolk L, Nica A, Inouye M, Hofman A, Stephens J, Wheeler E, Arp P, Gwilliam R, Jhamai PM, Potter S, Chaney A, Ghori MJ, Ravindrarajah R, Ermakov S, Estrada K, Pols HA, Williams FM, McArdle WL, van Meurs JB, Loos RJ, Dermitzakis ET, Ahmadi KR, Hart DJ, Ouwehand WH, Wareham NJ, Barroso I, Sandhu MS, Strachan DP, Livshits G, Spector TD, Uitterlinden AG, Deloukas P. Meta-analysis of genome-wide scans for human adult stature identifies novel Loci and associations with measures of skeletal frame size.. PLoS Genet 2009 Apr;5(4):e1000445.
          pmc: PMC2661236pubmed: 19343178doi: 10.1371/journal.pgen.1000445google scholar: lookup
        7. van de Pol C, Sloet van Oldruitenborgh-Oosterbaan MM. Measuring the height of ponies at the withers: influence of time of day, water and feed withdrawal, weight-carrying, exercise and sedation.. Vet J 2007 Jul;174(1):69-76.
          pubmed: 17182264doi: 10.1016/j.tvjl.2006.10.023google scholar: lookup
        8. Ricard A. Heritability of jumping ability and height of pony breeds in France. Livest Prod Sci 89: 243–251.
        9. Van Bergen H, Van Arendonk J. Genetic parameters for linear type traits in Shetland Ponies. Livest Prod Sci 36: 273–284.
        10. Miglior F, Pagnacco G, Samore AB. A total merit index for the Italian Haflinger horse using breeding values predicted by a multi-trait animal model. Proc 6th WCGALP 24: 416–419.
        11. Stock KF, Distl O. Genetic correlations between conformation traits and radiographic findings in the limbs of German Warmblood riding horses.. Genet Sel Evol 2006 Nov-Dec;38(6):657-71.
          pmc: PMC2689269pubmed: 17129565doi: 10.1186/1297-9686-38-6-657google scholar: lookup
        12. Arnason T. Genetic studies on conformation and performance of Icelandic toelter horses.1. Estimation of non-genetic effects and genetic parameters. Acta Agriculturae Scandinavica 34: 409–427.
        13. Curtis GC, Grove-White D, Ellis RN, Argo CM. Height measurement in horses and ponies: optimising standard protocols.. Vet Rec 2010 Jul 24;167(4):127-33.
          pubmed: 20656991doi: 10.1136/vr.c3722google scholar: lookup
        14. Sadek MH, Al-Aboud AZ, Ashmawy AA. Factor analysis of body measurements in Arabian horses.. J Anim Breed Genet 2006 Dec;123(6):369-77.
          pubmed: 17177691doi: 10.1111/j.1439-0388.2006.00618.xgoogle scholar: lookup
        15. Brooks SA, Makvandi-Nejad S, Chu E, Allen JJ, Streeter C, Gu E, McCleery B, Murphy BA, Bellone R, Sutter NB. Morphological variation in the horse: defining complex traits of body size and shape.. Anim Genet 2010 Dec;41 Suppl 2:159-65.
          pubmed: 21070291doi: 10.1111/j.1365-2052.2010.02127.xgoogle scholar: lookup
        16. Distl O, Schröder W, Dierks C, Klostermann A. Genome-wide association studies for performance and conformation traits in Hanoverian warmblood horses. 9th Dorothy Russel Havemeyer Foundation, International Equine Genome Mapping Workshop Oak Ridge Conference Center, Chaska, Minnesota: Distl, O. pp. 12.
        17. Schröder W. Athletic performance and conformation in Hanoverian warmblood horses - population genetic and genome-wide association analyses [cumulative thesis]. Hannover: University of Veterinary Medicine.
        18. Morsci NS, Schnabel RD, Taylor JF. Association analysis of adiponectin and somatostatin polymorphisms on BTA1 with growth and carcass traits in Angus cattle.. Anim Genet 2006 Dec;37(6):554-62.
          pubmed: 17121600doi: 10.1111/j.1365-2052.2006.01528.xgoogle scholar: lookup
        19. Tatarakis A, Margaritis T, Martinez-Jimenez CP, Kouskouti A, Mohan WS 2nd, Haroniti A, Kafetzopoulos D, Tora L, Talianidis I. Dominant and redundant functions of TFIID involved in the regulation of hepatic genes.. Mol Cell 2008 Aug 22;31(4):531-543.
          pubmed: 18722179doi: 10.1016/j.molcel.2008.07.013google scholar: lookup
        20. Saastamoinen A. Heritabilities for Body Size and Growth Rate and Phenotypic Correlations among Measurements in Young Horses. Acta Agricult Scand 40: 377–389.
        21. Hill EW, McGivney BA, Gu J, Whiston R, Machugh DE. A genome-wide SNP-association study confirms a sequence variant (g.66493737C>T) in the equine myostatin (MSTN) gene as the most powerful predictor of optimum racing distance for Thoroughbred racehorses.. BMC Genomics 2010 Oct 11;11:552.
          pmc: PMC3091701pubmed: 20932346doi: 10.1186/1471-2164-11-552google scholar: lookup
        22. Hill EW, Gu J, Eivers SS, Fonseca RG, McGivney BA, Govindarajan P, Orr N, Katz LM, MacHugh DE. A sequence polymorphism in MSTN predicts sprinting ability and racing stamina in thoroughbred horses.. PLoS One 2010 Jan 20;5(1):e8645.
          pmc: PMC2808334pubmed: 20098749doi: 10.1371/journal.pone.0008645google scholar: lookup
        23. Sawadogo M, Roeder RG. Interaction of a gene-specific transcription factor with the adenovirus major late promoter upstream of the TATA box region.. Cell 1985 Nov;43(1):165-75.
          pubmed: 4075392doi: 10.1016/0092-8674(85)90021-2google scholar: lookup
        24. Orphanides G, Lagrange T, Reinberg D. The general transcription factors of RNA polymerase II.. Genes Dev 1996 Nov 1;10(21):2657-83.
          pubmed: 8946909doi: 10.1101/gad.10.21.2657google scholar: lookup
        25. Yang MQ, Laflamme K, Gotea V, Joiner CH, Seidel NE, Wong C, Petrykowska HM, Lichtenberg J, Lee S, Welch L, Gallagher PG, Bodine DM, Elnitski L. Genome-wide detection of a TFIID localization element from an initial human disease mutation.. Nucleic Acids Res 2011 Mar;39(6):2175-87.
          pmc: PMC3064768pubmed: 21071415doi: 10.1093/nar/gkq1035google scholar: lookup
        26. Nakatani Y, Horikoshi M, Brenner M, Yamamoto T, Besnard F, Roeder RG, Freese E. A downstream initiation element required for efficient TATA box binding and in vitro function of TFIID.. Nature 1990 Nov 1;348(6296):86-8.
          pubmed: 2234067doi: 10.1038/348086a0google scholar: lookup
        27. Rodan GA, Harada S. The missing bone.. Cell 1997 May 30;89(5):677-80.
          pubmed: 9182754doi: 10.1016/s0092-8674(00)80249-4google scholar: lookup
        28. St-Arnaud R, Quelo I. Transcriptional coactivators potentiating AP-1 function in bone.. Front Biosci 1998 Aug 1;3:d838-48.
          pubmed: 9682038doi: 10.2741/a327google scholar: lookup
        29. Weedon MN, Lango H, Lindgren CM, Wallace C, Evans DM, Mangino M, Freathy RM, Perry JR, Stevens S, Hall AS, Samani NJ, Shields B, Prokopenko I, Farrall M, Dominiczak A, Johnson T, Bergmann S, Beckmann JS, Vollenweider P, Waterworth DM, Mooser V, Palmer CN, Morris AD, Ouwehand WH, Zhao JH, Li S, Loos RJ, Barroso I, Deloukas P, Sandhu MS, Wheeler E, Soranzo N, Inouye M, Wareham NJ, Caulfield M, Munroe PB, Hattersley AT, McCarthy MI, Frayling TM. Genome-wide association analysis identifies 20 loci that influence adult height.. Nat Genet 2008 May;40(5):575-83.
          pmc: PMC2681221pubmed: 18391952doi: 10.1038/ng.121google scholar: lookup
        30. Lettre G, Jackson AU, Gieger C, Schumacher FR, Berndt SI, Sanna S, Eyheramendy S, Voight BF, Butler JL, Guiducci C, Illig T, Hackett R, Heid IM, Jacobs KB, Lyssenko V, Uda M, Boehnke M, Chanock SJ, Groop LC, Hu FB, Isomaa B, Kraft P, Peltonen L, Salomaa V, Schlessinger D, Hunter DJ, Hayes RB, Abecasis GR, Wichmann HE, Mohlke KL, Hirschhorn JN. Identification of ten loci associated with height highlights new biological pathways in human growth.. Nat Genet 2008 May;40(5):584-91.
          pmc: PMC2687076pubmed: 18391950doi: 10.1038/ng.125google scholar: lookup
        31. Kunieda T, Park JM, Takeuchi H, Kubo T. Identification and characterization of Mlr1,2: two mouse homologues of Mblk-1, a transcription factor from the honeybee brain(1).. FEBS Lett 2003 Jan 30;535(1-3):61-5.
          pubmed: 12560079doi: 10.1016/s0014-5793(02)03858-9google scholar: lookup
        32. Wade CM, Giulotto E, Sigurdsson S, Zoli M, Gnerre S, Imsland F, Lear TL, Adelson DL, Bailey E, Bellone RR, Blöcker H, Distl O, Edgar RC, Garber M, Leeb T, Mauceli E, MacLeod JN, Penedo MC, Raison JM, Sharpe T, Vogel J, Andersson L, Antczak DF, Biagi T, Binns MM, Chowdhary BP, Coleman SJ, Della Valle G, Fryc S, Guérin G, Hasegawa T, Hill EW, Jurka J, Kiialainen A, Lindgren G, Liu J, Magnani E, Mickelson JR, Murray J, Nergadze SG, Onofrio R, Pedroni S, Piras MF, Raudsepp T, Rocchi M, Røed KH, Ryder OA, Searle S, Skow L, Swinburne JE, Syvänen AC, Tozaki T, Valberg SJ, Vaudin M, White JR, Zody MC, Lander ES, Lindblad-Toh K. Genome sequence, comparative analysis, and population genetics of the domestic horse.. Science 2009 Nov 6;326(5954):865-7.
          pmc: PMC3785132pubmed: 19892987doi: 10.1126/science.1178158google scholar: lookup
        33. Hall JG, Pagon RA, Wilson KM. Rothmund-Thomson syndrome with severe dwarfism.. Am J Dis Child 1980 Feb;134(2):165-9.
          pubmed: 7352442doi: 10.1001/archpedi.1980.02130140039013google scholar: lookup
        34. Moore GP, Panaretto BA, Robertson D. Effects of epidermal growth factor on hair growth in the mouse.. J Endocrinol 1981 Feb;88(2):293-9.
          pubmed: 6970792doi: 10.1677/joe.0.0880293google scholar: lookup
        35. Christmann L. Zuchtwertschätzung für Merkmale der Stutbuchaufnahme und der Stutenleistungsprüfung im Zuchtgebiet Hannover. Dissertation, Georg-August Universität Göttingen.
        36. Hamann H, Distl O. Genetic variability in Hanoverian warmblood horses using pedigree analysis.. J Anim Sci 2008 Jul;86(7):1503-13.
          pubmed: 18310493doi: 10.2527/jas.2007-0382google scholar: lookup
        37. Wheelan SJ, Church DM, Ostell JM. Spidey: a tool for mRNA-to-genomic alignments.. Genome Res 2001 Nov;11(11):1952-7.
          pmc: PMC311166pubmed: 11691860doi: 10.1101/gr.195301google scholar: lookup
        38. Johnson M, Zaretskaya I, Raytselis Y, Merezhuk Y, McGinnis S, Madden TL. NCBI BLAST: a better web interface.. Nucleic Acids Res 2008 Jul 1;36(Web Server issue):W5-9.
          pmc: PMC2447716pubmed: 18440982doi: 10.1093/nar/gkn201google scholar: lookup
        39. Livak KJ, Schmittgen TD. Analysis of relative gene expression data using real-time quantitative PCR and the 2(-Delta Delta C(T)) Method.. Methods 2001 Dec;25(4):402-8.
          pubmed: 11846609doi: 10.1006/meth.2001.1262google scholar: lookup
        40. Purcell S, Neale B, Todd-Brown K, Thomas L, Ferreira MA, Bender D, Maller J, Sklar P, de Bakker PI, Daly MJ, Sham PC. PLINK: a tool set for whole-genome association and population-based linkage analyses.. Am J Hum Genet 2007 Sep;81(3):559-75.
          pmc: PMC1950838pubmed: 17701901doi: 10.1086/519795google scholar: lookup
        41. Bradbury PJ, Zhang Z, Kroon DE, Casstevens TM, Ramdoss Y, Buckler ES. TASSEL: software for association mapping of complex traits in diverse samples.. Bioinformatics 2007 Oct 1;23(19):2633-5.
          pubmed: 17586829doi: 10.1093/bioinformatics/btm308google scholar: lookup

        Citations

        This article has been cited 66 times.
        1. Batcher K, Varney S, Raudsepp T, Jevit M, Dickinson P, Jagannathan V, Leeb T, Bannasch D. Ancient segmentally duplicated LCORL retrocopies in equids. PLoS One 2023;18(6):e0286861.
          doi: 10.1371/journal.pone.0286861pubmed: 37289743google scholar: lookup
        2. Park S. Height-Related Polygenic Variants Are Associated with Metabolic Syndrome Risk and Interact with Energy Intake and a Rice-Main Diet to Influence Height in KoGES. Nutrients 2023 Apr 4;15(7).
          doi: 10.3390/nᔇ1764pubmed: 37049604google scholar: lookup
        3. Zhi Y, Wang D, Zhang K, Wang Y, Geng W, Chen B, Li H, Li Z, Tian Y, Kang X, Liu X. Genome-Wide Genetic Structure of Henan Indigenous Chicken Breeds. Animals (Basel) 2023 Feb 19;13(4).
          doi: 10.3390/ani13040753pubmed: 36830540google scholar: lookup
        4. Kim YM, Ha SJ, Seong HS, Choi JY, Baek HJ, Yang BC, Choi JW, Kim NY. Identification of Copy Number Variations in Four Horse Breed Populations in South Korea. Animals (Basel) 2022 Dec 12;12(24).
          doi: 10.3390/ani12243501pubmed: 36552421google scholar: lookup
        5. Szmatoła T, Gurgul A, Jasielczuk I, Oclon E, Ropka-Molik K, Stefaniuk-Szmukier M, Polak G, Tomczyk-Wrona I, Bugno-Poniewierska M. Assessment and Distribution of Runs of Homozygosity in Horse Breeds Representing Different Utility Types. Animals (Basel) 2022 Nov 25;12(23).
          doi: 10.3390/ani12233293pubmed: 36496815google scholar: lookup
        6. Zhao XW, Wu J, Kishino H, Chen L. Massive Loss of Transcription Factors Promotes the Initial Diversification of Placental Mammals. Int J Mol Sci 2022 Aug 26;23(17).
          doi: 10.3390/ijms23179720pubmed: 36077118google scholar: lookup
        7. Pan B, Long H, Yuan Y, Zhang H, Peng Y, Zhou D, Liu C, Xiang B, Huang Y, Zhao Y, Zhao Z, E G. Identification of Body Size Determination Related Candidate Genes in Domestic Pig Using Genome-Wide Selection Signal Analysis. Animals (Basel) 2022 Jul 19;12(14).
          doi: 10.3390/ani12141839pubmed: 35883386google scholar: lookup
        8. Puttabyatappa M, Saadat N, Elangovan VR, Dou J, Bakulski K, Padmanabhan V. Developmental programming: Impact of prenatal bisphenol-A exposure on liver and muscle transcriptome of female sheep. Toxicol Appl Pharmacol 2022 Sep 15;451:116161.
          doi: 10.1016/j.taap.2022.116161pubmed: 35817127google scholar: lookup
        9. Finno CJ. Science-in-brief: Genomic and transcriptomic approaches to the investigation of equine diseases. Equine Vet J 2022 Mar;54(2):444-448.
          doi: 10.1111/evj.13549pubmed: 35133024google scholar: lookup
        10. Vosgerau S, Krattenmacher N, Falker-Gieske C, Seidel A, Tetens J, Stock KF, Nolte W, Wobbe M, Blaj I, Reents R, Kühn C, von Depka Prondzinski M, Kalm E, Thaller G. Genetic and genomic characterization followed by single-step genomic evaluation of withers height in German Warmblood horses. J Appl Genet 2022 May;63(2):369-378.
          doi: 10.1007/s13353-021-00681-wpubmed: 35028913google scholar: lookup
        11. Cao X, Cheng J, Huang Y, Lan X, Lei C, Chen H. Comparative Enhancer Map of Cattle Muscle Genome Annotated by ATAC-Seq. Front Vet Sci 2021;8:782409.
          doi: 10.3389/fvets.2021.782409pubmed: 34977215google scholar: lookup
        12. Liu X, Zhang Y, Liu W, Li Y, Pan J, Pu Y, Han J, Orlando L, Ma Y, Jiang L. A single-nucleotide mutation within the TBX3 enhancer increased body size in Chinese horses. Curr Biol 2022 Jan 24;32(2):480-487.e6.
          doi: 10.1016/j.cub.2021.11.052pubmed: 34906355google scholar: lookup
        13. Dadousis C, Somavilla A, Ilska JJ, Johnsson M, Batista L, Mellanby RJ, Headon D, Gottardo P, Whalen A, Wilson D, Dunn IC, Gorjanc G, Kranis A, Hickey JM. A genome-wide association analysis for body weight at 35 days measured on 137,343 broiler chickens. Genet Sel Evol 2021 Sep 8;53(1):70.
          doi: 10.1186/s12711-021-00663-wpubmed: 34496773google scholar: lookup
        14. Samaha G, Wade CM, Mazrier H, Grueber CE, Haase B. Exploiting genomic synteny in Felidae: cross-species genome alignments and SNV discovery can aid conservation management. BMC Genomics 2021 Aug 6;22(1):601.
          doi: 10.1186/s12864-021-07899-2pubmed: 34362297google scholar: lookup
        15. Rosengren MK, Sigurðardóttir H, Eriksson S, Naboulsi R, Jouni A, Novoa-Bravo M, Albertsdóttir E, Kristjánsson Þ, Rhodin M, Viklund Å, Velie BD, Negro JJ, Solé M, Lindgren G. A QTL for conformation of back and croup influences lateral gait quality in Icelandic horses. BMC Genomics 2021 Apr 14;22(1):267.
          doi: 10.1186/s12864-021-07454-zpubmed: 33853519google scholar: lookup
        16. Wang YM, Khederzadeh S, Li SR, Otecko NO, Irwin DM, Thakur M, Ren XD, Wang MS, Wu DD, Zhang YP. Integrating Genomic and Transcriptomic Data to Reveal Genetic Mechanisms Underlying Piao Chicken Rumpless Trait. Genomics Proteomics Bioinformatics 2021 Oct;19(5):787-799.
          doi: 10.1016/j.gpb.2020.06.019pubmed: 33631431google scholar: lookup
        17. Dzomba EF, Chimonyo M, Pierneef R, Muchadeyi FC. Runs of homozygosity analysis of South African sheep breeds from various production systems investigated using OvineSNP50k data. BMC Genomics 2021 Jan 6;22(1):7.
          doi: 10.1186/s12864-020-07314-2pubmed: 33407115google scholar: lookup
        18. Xu J, Fu Y, Hu Y, Yin L, Tang Z, Yin D, Zhu M, Yu M, Li X, Zhou Y, Zhao S, Liu X. Whole genome variants across 57 pig breeds enable comprehensive identification of genetic signatures that underlie breed features. J Anim Sci Biotechnol 2020 Dec 3;11(1):115.
          doi: 10.1186/s40104-020-00520-8pubmed: 33292532google scholar: lookup
        19. Kvist L, Honka J, Niskanen M, Liedes O, Aspi J. Selection in the Finnhorse, a native all-around horse breed. J Anim Breed Genet 2021 Mar;138(2):188-203.
          doi: 10.1111/jbg.12524pubmed: 33226152google scholar: lookup
        20. Asadollahpour Nanaei H, Esmailizadeh A, Ayatollahi Mehrgardi A, Han J, Wu DD, Li Y, Zhang YP. Comparative population genomic analysis uncovers novel genomic footprints and genes associated with small body size in Chinese pony. BMC Genomics 2020 Jul 20;21(1):496.
          doi: 10.1186/s12864-020-06887-2pubmed: 32689947google scholar: lookup
        21. Al Abri MA, Holl HM, Kalla SE, Sutter NB, Brooks SA. Whole genome detection of sequence and structural polymorphism in six diverse horses. PLoS One 2020;15(4):e0230899.
          doi: 10.1371/journal.pone.0230899pubmed: 32271776google scholar: lookup
        22. Pu Y, Zhang Y, Zhang T, Han J, Ma Y, Liu X. Identification of Novel lncRNAs Differentially Expressed in Placentas of Chinese Ningqiang Pony and Yili Horse Breeds. Animals (Basel) 2020 Jan 11;10(1).
          doi: 10.3390/ani10010119pubmed: 31940795google scholar: lookup
        23. Srikanth K, Kim NY, Park W, Kim JM, Kim KD, Lee KT, Son JH, Chai HH, Choi JW, Jang GW, Kim H, Ryu YC, Nam JW, Park JE, Kim JM, Lim D. Comprehensive genome and transcriptome analyses reveal genetic relationship, selection signature, and transcriptome landscape of small-sized Korean native Jeju horse. Sci Rep 2019 Nov 13;9(1):16672.
          doi: 10.1038/s41598-019-53102-8pubmed: 31723199google scholar: lookup
        24. Moreira GCM, Poleti MD, Pértille F, Boschiero C, Cesar ASM, Godoy TF, Ledur MC, Reecy JM, Garrick DJ, Coutinho LL. Unraveling genomic associations with feed efficiency and body weight traits in chickens through an integrative approach. BMC Genet 2019 Nov 6;20(1):83.
          doi: 10.1186/s12863-019-0783-3pubmed: 31694549google scholar: lookup
        25. Raudsepp T, Finno CJ, Bellone RR, Petersen JL. Ten years of the horse reference genome: insights into equine biology, domestication and population dynamics in the post-genome era. Anim Genet 2019 Dec;50(6):569-597.
          doi: 10.1111/age.12857pubmed: 31568563google scholar: lookup
        26. Moreira GCM, Salvian M, Boschiero C, Cesar ASM, Reecy JM, Godoy TF, Ledur MC, Garrick D, Mourão GB, Coutinho LL. Genome-wide association scan for QTL and their positional candidate genes associated with internal organ traits in chickens. BMC Genomics 2019 Aug 22;20(1):669.
          doi: 10.1186/s12864-019-6040-3pubmed: 31438838google scholar: lookup
        27. Naqvi S, Godfrey AK, Hughes JF, Goodheart ML, Mitchell RN, Page DC. Conservation, acquisition, and functional impact of sex-biased gene expression in mammals. Science 2019 Jul 19;365(6450).
          doi: 10.1126/science.aaw7317pubmed: 31320509google scholar: lookup
        28. Velie BD, Lillie M, Fegraeus KJ, Rosengren MK, Solé M, Wiklund M, Ihler CF, Strand E, Lindgren G. Exploring the genetics of trotting racing ability in horses using a unique Nordic horse model. BMC Genomics 2019 Feb 4;20(1):104.
          doi: 10.1186/s12864-019-5484-9pubmed: 30717660google scholar: lookup
        29. Gurgul A, Jasielczuk I, Semik-Gurgul E, Pawlina-Tyszko K, Stefaniuk-Szmukier M, Szmatoła T, Polak G, Tomczyk-Wrona I, Bugno-Poniewierska M. A genome-wide scan for diversifying selection signatures in selected horse breeds. PLoS One 2019;14(1):e0210751.
          doi: 10.1371/journal.pone.0210751pubmed: 30699152google scholar: lookup
        30. Kim NY, Seong HS, Kim DC, Park NG, Yang BC, Son JK, Shin SM, Woo JH, Shin MC, Yoo JH, Choi JW. Genome-wide analyses of the Jeju, Thoroughbred, and Jeju crossbred horse populations using the high density SNP array. Genes Genomics 2018 Nov;40(11):1249-1258.
          doi: 10.1007/s13258-018-0722-0pubmed: 30099720google scholar: lookup
        31. Metzger J, Rau J, Naccache F, Bas Conn L, Lindgren G, Distl O. Genome data uncover four synergistic key regulators for extremely small body size in horses. BMC Genomics 2018 Jun 25;19(1):492.
          doi: 10.1186/s12864-018-4877-5pubmed: 29940849google scholar: lookup
        32. Tozaki T, Kikuchi M, Kakoi H, Hirota KI, Nagata SI. A genome-wide association study for body weight in Japanese Thoroughbred racehorses clarifies candidate regions on chromosomes 3, 9, 15, and 18. J Equine Sci 2017;28(4):127-134.
          doi: 10.1294/jes.28.127pubmed: 29270069google scholar: lookup
        33. Taye M, Lee W, Jeon S, Yoon J, Dessie T, Hanotte O, Mwai OA, Kemp S, Cho S, Oh SJ, Lee HK, Kim H. Exploring evidence of positive selection signatures in cattle breeds selected for different traits. Mamm Genome 2017 Dec;28(11-12):528-541.
          doi: 10.1007/s00335-017-9715-6pubmed: 28905131google scholar: lookup
        34. Metzger J, Nolte A, Uhde AK, Hewicker-Trautwein M, Distl O. Whole genome sequencing identifies missense mutation in MTBP in Shar-Pei affected with Autoinflammatory Disease (SPAID). BMC Genomics 2017 May 4;18(1):348.
          doi: 10.1186/s12864-017-3737-zpubmed: 28472921google scholar: lookup
        35. Xia J, Fan H, Chang T, Xu L, Zhang W, Song Y, Zhu B, Zhang L, Gao X, Chen Y, Li J, Gao H. Searching for new loci and candidate genes for economically important traits through gene-based association analysis of Simmental cattle. Sci Rep 2017 Feb 7;7:42048.
          doi: 10.1038/srep42048pubmed: 28169328google scholar: lookup
        36. Metzger J, Gast AC, Schrimpf R, Rau J, Eikelberg D, Beineke A, Hellige M, Distl O. Whole-genome sequencing reveals a potential causal mutation for dwarfism in the Miniature Shetland pony. Mamm Genome 2017 Apr;28(3-4):143-151.
          doi: 10.1007/s00335-016-9673-4pubmed: 27942904google scholar: lookup
        37. Tozaki T, Sato F, Ishimaru M, Kikuchi M, Kakoi H, Hirota KI, Nagata SI. Sequence variants of BIEC2-808543 near LCORL are associated with body composition in Thoroughbreds under training. J Equine Sci 2016;27(3):107-114.
          doi: 10.1294/jes.27.107pubmed: 27703405google scholar: lookup
        38. Kwak IY, Pan W. Gene- and pathway-based association tests for multiple traits with GWAS summary statistics. Bioinformatics 2017 Jan 1;33(1):64-71.
          doi: 10.1093/bioinformatics/btw577pubmed: 27592708google scholar: lookup
        39. Metzger J, Pfahler S, Distl O. Variant detection and runs of homozygosity in next generation sequencing data elucidate the genetic background of Lundehund syndrome. BMC Genomics 2016 Aug 2;17:535.
          doi: 10.1186/s12864-016-2844-6pubmed: 27485430google scholar: lookup
        40. Boitard S, Boussaha M, Capitan A, Rocha D, Servin B. Uncovering Adaptation from Sequence Data: Lessons from Genome Resequencing of Four Cattle Breeds. Genetics 2016 May;203(1):433-50.
          doi: 10.1534/genetics.115.181594pubmed: 27017625google scholar: lookup
        41. Bolormaa S, Hayes BJ, van der Werf JH, Pethick D, Goddard ME, Daetwyler HD. Detailed phenotyping identifies genes with pleiotropic effects on body composition. BMC Genomics 2016 Mar 12;17:224.
          doi: 10.1186/s12864-016-2538-0pubmed: 26968377google scholar: lookup
        42. Matika O, Riggio V, Anselme-Moizan M, Law AS, Pong-Wong R, Archibald AL, Bishop SC. Genome-wide association reveals QTL for growth, bone and in vivo carcass traits as assessed by computed tomography in Scottish Blackface lambs. Genet Sel Evol 2016 Feb 8;48:11.
          doi: 10.1186/s12711-016-0191-3pubmed: 26856324google scholar: lookup
        43. Liu Y, Duan X, Chen S, He H, Liu X. NCAPG is differentially expressed during longissimus muscle development and is associated with growth traits in Chinese Qinchuan beef cattle. Genet Mol Biol 2015 Dec;38(4):450-6.
          doi: 10.1590/S1415-475738420140287pubmed: 26692155google scholar: lookup
        44. Metzger J, Wöhlke A, Mischke R, Hoffmann A, Hewicker-Trautwein M, Küch EM, Naim HY, Distl O. A Novel SLC27A4 Splice Acceptor Site Mutation in Great Danes with Ichthyosis. PLoS One 2015;10(10):e0141514.
          doi: 10.1371/journal.pone.0141514pubmed: 26506231google scholar: lookup
        45. Takasuga A. PLAG1 and NCAPG-LCORL in livestock. Anim Sci J 2016 Feb;87(2):159-67.
          doi: 10.1111/asj.12417pubmed: 26260584google scholar: lookup
        46. Sahana G, Höglund JK, Guldbrandtsen B, Lund MS. Loci associated with adult stature also affect calf birth survival in cattle. BMC Genet 2015 May 3;16:47.
          doi: 10.1186/s12863-015-0202-3pubmed: 25935543google scholar: lookup
        47. Metzger J, Tonda R, Beltran S, Agueda L, Gut M, Distl O. Next generation sequencing gives an insight into the characteristics of highly selected breeds versus non-breed horses in the course of domestication. BMC Genomics 2014 Jul 4;15(1):562.
          doi: 10.1186/1471-2164-15-562pubmed: 24996778google scholar: lookup
        48. Lindholm-Perry AK, Kuehn LA, Oliver WT, Sexten AK, Miles JR, Rempel LA, Cushman RA, Freetly HC. Adipose and muscle tissue gene expression of two genes (NCAPG and LCORL) located in a chromosomal region associated with cattle feed intake and gain. PLoS One 2013;8(11):e80882.
          doi: 10.1371/journal.pone.0080882pubmed: 24278337google scholar: lookup
        49. Metzger J, Philipp U, Lopes MS, da Camara Machado A, Felicetti M, Silvestrelli M, Distl O. Analysis of copy number variants by three detection algorithms and their association with body size in horses. BMC Genomics 2013 Jul 18;14:487.
          doi: 10.1186/1471-2164-14-487pubmed: 23865711google scholar: lookup
        50. Jafari H, Abebe BK, Cong L, Ahmed Z, Zhaofei W, Sun M, Muhatai G, Chuzhao L, Dang R. Review: Genomic insights into the adaptive traits and stress resistance in modern horses. Stress Biol 2026 Jan 12;6(1):5.
          doi: 10.1007/s44154-025-00274-1pubmed: 41521281google scholar: lookup
        51. Sharma M, Singh A, Kumar V, Olla N, Arora R, Sharma R, Mohan NH, Ahlawat S. Advances in Equine Genomics: Decoding the Genetic Architecture of Morphology, Performance, Behavior, and Adaptation. Mol Biotechnol 2025 Dec 19;.
          doi: 10.1007/s12033-025-01544-zpubmed: 41417456google scholar: lookup
        52. Steensma MJ, Doekes HP, Derks MFL, Ducro BJ. Genome-wide association study reveals candidate loci on ECA1 and ECA9 for withers height in Friesian horses. Anim Genet 2025 Oct;56(5):e70049.
          doi: 10.1111/age.70049pubmed: 41090464google scholar: lookup
        53. Arshad F, Jayaraman S, Talenti A, Owen R, Mohsin M, Mansoor S, Asif M, Prendergast J. A comprehensive water buffalo pangenome reveals extensive structural variation linked to population-specific signatures of selection. Gigascience 2025 Jan 6;14.
          doi: 10.1093/gigascience/giaf099pubmed: 40884803google scholar: lookup
        54. Asti V, Summer A, Ablondi M, Sartori C, Giontella A, Pilastro V, Mecocci S, Cappelli K, Mancin E, Oian A, Mantovani R, Capomaccio S, Sabbioni A. Selection signatures and inbreeding: exploring genetic diversity in five native horse breeds. BMC Vet Res 2025 May 16;21(1):346.
          doi: 10.1186/s12917-025-04794-wpubmed: 40380299google scholar: lookup
        55. Rodrigues FM, Majeres LE, Dilger AC, McCann JC, Cassady CJ, Shike DW, Beever JE. Characterizing differences in the muscle transcriptome between cattle with alternative LCORL-NCAPG haplotypes. BMC Genomics 2025 May 14;26(1):479.
          doi: 10.1186/s12864-025-11665-zpubmed: 40369436google scholar: lookup
        56. Buckley RM, Bilgen N, Harris AC, Savolainen P, Tepeli C, Erdoğan M, Serres Armero A, Dreger DL, van Steenbeek FG, Hytönen MK, Parker HG, Hale J, Lohi H, Çınar Kul B, Boyko AR, Ostrander EA. Analysis of canine gene constraint identifies new variants for orofacial clefts and stature. Genome Res 2025 May 2;35(5):1080-1093.
          doi: 10.1101/gr.280092.124pubmed: 40127928google scholar: lookup
        57. McGivney CL, McGivney BA, Farries G, Gough KF, Han H, Holtby AR, MacHugh DE, Katz LM, Hill EW. A genome-wide association study for recurrent laryngeal neuropathy in the Thoroughbred horse identifies a candidate gene that regulates myelin structure. Equine Vet J 2025 Jul;57(4):943-952.
          doi: 10.1111/evj.14461pubmed: 39791379google scholar: lookup
        58. Nazari-Ghadikolaei A, Fikse WF, Viklund ÅG, Mikko S, Eriksson S. Single-Step Genome-Wide Association Study of Factors for Evaluated and Linearly Scored Traits in Swedish Warmblood Horses. J Anim Breed Genet 2025 Sep;142(5):499-512.
          doi: 10.1111/jbg.12923pubmed: 39754479google scholar: lookup
        59. Wyler SC, Gahlot S, Bideyan L, Yip C, Dushime J, Chen B, Lee JJ, Tinajero A, Limboy C, Bordash S, Heaselgrave SR, Nguyen TN, Lee S, Bookout A, Lantier L, Fowlkes JL, You YJ, Fujikawa T, Elmquist JK. LCoRL Regulates Growth and Metabolism. Endocrinology 2024 Oct 30;165(12).
          doi: 10.1210/endocr/bqae146pubmed: 39467326google scholar: lookup
        60. Šimon M, Kaić A, Potočnik K. Unveiling Genetic Potential for Equine Meat Production: A Bioinformatics Approach. Animals (Basel) 2024 Aug 22;14(16).
          doi: 10.3390/ani14162441pubmed: 39199974google scholar: lookup
        61. Yang T, Wang M, Liu Y, Li Y, Feng M, Zhao C. A mutation in POLR2A gene associated with body size traits in Dezhou donkeys revealed with GWAS. J Anim Sci 2024 Jan 3;102.
          doi: 10.1093/jas/skae217pubmed: 39079013google scholar: lookup
        62. Yang Y, Wang C, Chen S, Liu Y, Jia H, Wang H, He D. Identifying candidate genes and biological pathways in muscle development through multi-tissue transcriptome comparisons between male and female geese. Sci Rep 2024 Jul 16;14(1):16474.
          doi: 10.1038/s41598-024-67560-2pubmed: 39014195google scholar: lookup
        63. Reich P, Möller S, Stock KF, Nolte W, von Depka Prondzinski M, Reents R, Kalm E, Kühn C, Thaller G, Falker-Gieske C, Tetens J. Genomic analyses of withers height and linear conformation traits in German Warmblood horses using imputed sequence-level genotypes. Genet Sel Evol 2024 Jun 13;56(1):45.
          doi: 10.1186/s12711-024-00914-6pubmed: 38872118google scholar: lookup
        64. Saito I, Nakamura K, Tozaki T, Hano K, Takasu M. Genetic characterization of Japanese native horse breeds by genotyping variants that are associated with phenotypic traits. J Equine Sci 2023 Dec;34(4):115-120.
          doi: 10.1294/jes.34.115pubmed: 38274555google scholar: lookup
        65. Choudhury MP, Wang Z, Zhu M, Teng S, Yan J, Cao S, Yi G, Liu Y, Liao Y, Tang Z. Genome-Wide Detection of Copy Number Variations Associated with Miniature Features in Horses. Genes (Basel) 2023 Oct 13;14(10).
          doi: 10.3390/genes14101934pubmed: 37895283google scholar: lookup
        66. Ricard A, Crevier-Denoix N, Pourcelot P, Crichan H, Sabbagh M, Dumont-Saint-Priest B, Danvy S. Genetic analysis of geometric morphometric 3D visuals of French jumping horses. Genet Sel Evol 2023 Sep 18;55(1):63.
          doi: 10.1186/s12711-023-00837-8pubmed: 37723416google scholar: lookup
        Subscribe to our newsletter
        Join 99,658 horse owners like you who receive our email updates on horse health and nutrition.