OMIA 000483-9913 : Polled/Horns in Bos taurus

In other species: goat , sheep , domestic yak , bighorn sheep , water buffalo , kouprey

Possibly relevant human trait(s) and/or gene(s) (MIM number): 110100

Mendelian trait/disorder: yes

Mode of inheritance: Autosomal

Considered a defect: no

Key variant known: yes

Year key variant first reported: 2012

Cross-species summary: There is substantial variation in the extent of horn growth, making classification difficult. However, in general, the presence or absence of horns can be attributed to the action of two alleles at an autosomal locus. The type of gene action varies considerably between and among species.

Species-specific name: Haplotype BHP; Haplotype HHP; Haplotype JHP

Species-specific symbol: BHP; HHP; JHP

Species-specific description: The absence of horns (polledness) is of substantial benefit in cattle, from an economic and welfare point of view: bruising due to horns is eliminated, and the stress associated with de-horning is avoided. (Information complied by Ulrika Tjälldén and Vanja Kinch, Uppsala, March 1998).

In addition to naturally occurring variants for this trait, variants have been created artificially: Genetically-modifed organism; GMO.

History: In cattle, one of the first Mendelian traits to attract attention was the presence/absence of horns. Polled cattle had long been recognised, e.g. having been described by Herodotus (c450BC) and Tacitus (c90AD). The inherited nature of this trait was well recognised (but not understood) long before the rediscovery of Mendelism (see, e.g. Darwin 1859 [p. 14]; Darwin 1868 [vol ii, p. 316]). In 1902, polledness was one of the first six animal traits (and the first cattle trait) to be shown to have Mendelian inheritance (Bateson and Saunders, 1902). In 1906, the American agricultural polymath W.J. Spillman (who is not only regarded as a founding father of agricultural economics, but also independently rediscovered Mendelism while crossing strains of wheat!) published a paper in Science (Spillman 1906a) and another in the newly-founded Journal of Heredity (Spillman 1906b), providing convincing evidence that the presence/absence of horns is a Mendelian trait, with polled being dominant to horned. This trait soon became a classic Mendelian trait, cited in many textbooks. Indeed, as delightfully recorded by Crow (1992), this trait even attracted the attention of the Nobel-prize winning physicist Erwin Schrödinger, who wrote two letters to J.B.S Haldane in 1945, in relation to “the hornless cattle problem”. In these letters, Schrödinger derived an equation that predicts the frequency of horned offspring in a closed herd after any number of generations of complete selection against horned bulls, but with no selection on cows.

Nothing much was added to our knowledge of this trait until the first wave of genomics tools provided sufficient microsatellite markers to enable Georges et al. (1993) to map the presence/absence of horns to within a recombination fraction of 13% with two markers on chromosome BTA1 (see Mapping section). To present readers, such “loose” linkage might seem to be not worthy of much celebration. At the time, however, this result was sufficiently important and novel to warrant publication in Nature Genetics. Subsequent progress in mapping is summarised in the Mapping section. (Most of the wording under this heading is from Nicholas, F.W. (2012; Mendelian Inheritance in Cattle, chap 2 [pp. 11-19] in Bovine Genomics [ed. J. Womack], Wiley-Blackwell, Ames, Iowa.)

As noted by Allais-Bonnet et al. (2013), the "Celtic" mutation (See Molecular section below) must be very old: "fixation of the Celtic mutation in ancient Nordic and British breeds and segregation of this allele in Icelandic cattle, which evolved independently from other Nordic breeds since the time of the Vikings, makes it possible to date back this mutation to at least one millenary. Because of historical relationships between the Vikings and the ancient Germans and between the ancient Germans and the Scythians, this mutation could be even older and explain the polledness of the cattle raised by these people . . . , which Herodotus, in the fifth century BC, mistakenly attributed to the extreme cold of the modern day Ukrainian steppes."

In a landmark paper that presents the final results of a project initially reported by Tan et al. (2013), Carlson et al. (2016) described "the use of genome editing using transcription activator-like effector nucleases (TALENs) to introgress the putative PC POLLED allele [see Molecular section below] into the genome of bovine embryo fibroblasts to try and produce a genotype identical to what is achievable using natural mating, but without the attendant genetic drag and admixture". Importantly, they later commented that "To our knowledge this is the first empirical validation of a putative causative allele in livestock, and this report provides evidence that PC, a sequence variant duplication of unknown function in a genomic region with no known or predicted coding or noncoding genes, is causative for polled. Genetic improvement of livestock using TALENs, or other genome-editing methods, establishes an alternative to transgenic (genetic modification)-based methods for genetic improvement of livestock using variation already present in species without the admixture that can result from classic breeding methods. . . . This solution to the dehorning problem would be economically viable and could potentially improve the welfare of cattle in the United States."

Inheritance: As described in the History section, this trait was the first cattle trait to be shown to be inherited in a Mendelian manner. This form of inheritance has been confirmed many times, e.g. Stookey and Goonewardene (1996). (Information complied by Ulrika Tjälldén and Vanja Kinch, Uppsala, March 1998); supplemented by FN 14th June 2013

Mapping: Georges et al. (1993) mapped this locus to the linkage group corresponding to chromosome BTA1, within a recombination fraction of 13% with two markers. Schmutz et al. (1995) provided the first estimate of physical location of the locus as being "within 1q12-14, very near the centromere of Chromosome (Chr) 1". By 2005, the region in which the locus is located was narrowed to a 1 Mb segment (Drögemüller et al., 2005). In early 2012, the region was narrowed further to 385 kb by Seichter et al. (2012). The BTA1 location of horns/polled in cattle bears no homology to the OAR10 location of horns/polled in sheep (OMIA 000483-9940), suggesting that the same trait in the two species is determined by different genes.

Mariasegaram et al. (2012) confirmed the location of this locus, and identified a 303bp allele of a new microsatellite (CSAFG29) that is stromngly associated with the polled trait in Australian Brahman cattle.

Using the Illumina bovine 777K SNP beadchip on 15 homozygous polled cattle from nine breeds, together with 36 half-sibs), Allais-Bonnet et al. (2013) independently confirmed the mapping of this locus to the centromeric end of chromosome BTA1, in the region 1,693,164 to 2,018,403 (UMD3.1 bovine genome assembly).

In an independent study, Wiedemar et al. (2014) homozygosity-mapped the polled locus in 1,019 polled cattle from 14 different breeds, each of which had been genotyped at 777,962 SNP loci with the Illumina BovineHD BeadChip. Their analysis enabled them to refine "the critical region of the Simmental polled mutation to 212 kb and identified an overlapping region of 932 kb containing the Holstein polled mutation".

Rothammer et al. (2014) narrowed the Holstein-Friesian candidate region to 53kb.

Stafuzza et al. (2018) mapped "polledness in Nelore (Bos taurus indicus)" to the "centromeric region of chromosome 1 with 3.11 Mb size (BTA1: 878,631-3,987,104 bp)", suggesting that "the postulated African horn locus is located very close to the Polled locus described for taurine breeds or it may be a mutation in the same region, since the literature that suggests the existence of an African horn locus is based on inconsistent inheritance patterns and dates from the last century."

Molecular basis: Finally, after many frustrating years, mutations completely associated with the last of Bateson and Saunder's (1902) six initial Mendelian traits have been determined. Resequencing (by Medugorac et al., 2012) of the region mapped by Seichter et al. (2012), and subsequent investigation of putative DNA sequence variants, revealed two alleles completely associated with polledness in cattle of European origin: a 202bp indel in breeds of “Celtic” origin (called P[sub]C or (P[sub]202ID)) and a 260kb haplotype block marked by two indels and three SNPs in cattle of Friesian origin (called P[sub]F). Heterozygosity or homozygosity for either of these alleles, or heterozygosity for both alleles, is completely associated with polledness. Despite this major discovery by Medugorac et al. (2012), and as if to reinforce the intransigence of this iconic Mendelian trait, no gene can yet be named, because none of these mutations disrupts "any known coding sequence or a splice site, or an intronic region, or any known regulatory regions". We are still a long way from having a complete understanding of this enigmatic locus. Until then, in OMIA the polled "gene" is called POLLED.

Independent fine-mapping, followed by extensive sequencing in the candidate region, enabled Allais-Bonnet et al. (2013) to confirm their previously-reported "Celtic" (P[sub]C) mutation, but to specify it slightly differently as "a duplication of 212 bp replacing a segment of 10 bp (g.1706051_1706060 delins 1705834_1706045 dup" (UMD3.1 bovine genome assembly) and to confirm that it occurs in a region with no known genes or expressed sequence tags.

A similar approach enabled Allais-Bonne et al. (2013) to reduce the list of candidate mutations for the "Friesian" (P[sub]F) allele to "three SNP (g.1764239T>C, g.1768587C>A, g.1855898G>A) and a large duplication of 80,128 bp (g.1909352_1989480dup)" (UMD3.1 bovine genome assembly), again in a region with no known genes or expressed sequence tags.

Based on histological and gene-expression studies, Allais-Bonne et al. (2013) proposed that "the ectopic expression of a lincRNA in PC/p horn buds as a probable cause of horn bud agenesis" and "provide[d] evidence for an involvement of OLIG2, FOXL2 and RXFP2 in horn bud differentiation, and draw a first link between bovine, ovine and caprine Polled loci."

The Holstein situation was further investigated by Glatzer et al. (2013), who reported "Sequencing of the positional candidate genes within the 1 Mb polled region and whole genome sequencing of Holsteins revealed a single nucleotide polymorphism (SNP) AC000158: g.1390292G>A within intron 3 of the interferon gamma receptor 2 gene (IFNGR2) in perfect co-segregation with polledness in Holsteins. This complete association was validated in 443 animals of the same breed." Interestingly, as also reported by Glatzer et al. (2013), in a selective-sweep study by Stella et al. (2010) "The SNP featuring the largest difference in allelic frequency between polled and horned animals was located within the interferon gamma receptor 2 gene (IFNGR2)".

Sequencing and additional genotyping enabled Wiedemar et al. (2014) to confirm the 212bp indel "Celctic" mutation P[sub]C as causal for polled in Simmentals and other European beef breeds. They also "identified a total of 182 sequence variants as candidate mutations for polledness in Holstein cattle, including an 80 kb genomic duplication and three SNPs reported before [see above]." Also, the authors reported that "For the first time we showed that hornless cattle with scurs are obligate heterozygous for one of the polled mutations."

By testing almost 6000 cattle, Rothammer et al. (2014) confirmed the 80kb duplication mentioned above (now called P(sub)80kbID) to be the "single causal mutation" in Holstein-Friesian cattle", i.e. to be P[sub]F. In addition, their results did not support the SNP association reported by Glatzer et al. (2013).

By introgressing the P(sub)C "allele into dairy cattle by genome editing [via TALEN-stimulated homology-dependent repair to produce four cell lines either homozygous or heterozygous for the [P.sub.C] allele]", followed by reproductive cloning to produce five live calves (all lacking horn buds), two of which were polled at 10 months, Carlson et al. (2016) provided "the first empirical validation of a putative causative allele in livestock, and this report provides evidence that [P.sub.C], a sequence variant duplication of unknown function in a genomic region with no known or predicted coding or noncoding genes, is causative for polled."

A third polled allele, the Mongolian allele, was discovered by Medugorac et al. (2017) in Mongolian Turano cattle. It is "a complex 219-bp duplication–insertion (P219ID) beginning at 1,976,128 bp and a 7-bp deletion and 6-bp insertion (P1ID) located 621 bp upstream of this position . . . . This rearrangement results in duplication of an 11-bp motif (5′-AAAGAAGCAAA-3′) that is entirely conserved among Bovidae . . . and that is also duplicated in the 80-kb duplication responsible for Friesian polledness". Intriguingly, this allele has been introgressed into yaks with which Mongolian Turano cattle have been herded for more than 1,500 years, via backcrossing female yak-cattle hybrids to male yaks (Medugorac et al., 2017). The suggested abbreviation for the Mongolian allele is P[sub]M (FN, in consultation with A. Capitan and I. Medugorac) or P[sub]219ID (Chen et al., 2017).

Chen et al. (2017) reported that "all three candidate mutations [P[sub]202ID, P[sub]80kbID and P[sub]219ID] have been simultaneously introgressed into Shuxuan cattle . . . . Furthermore, one polled animal still remains to carry none of the three candidate mutations, which suggests that further mutation(s) would also exist."

Clinical features: Compared to horned cattlle, polled cattle have a narrower skull, and the frontal eminence is more pronounced (Brenneman et al., (1996). (Information complied by Ulrika Tjälldén and Vanja Kinch, Uppsala, March 1998). Also, "polled cattle have abnormal hair growth on the eyelashes and eyelids, and that some bulls carrying the Celtic mutation have pending prepuce and abnormal preputial withdrawal (Allais-Bonnet et al., 2013)" (A. Capitan, pers. comm. to FN, 2017)

Breed: Numerous breeds.

Associated gene:

Symbol Description Species Chr Location OMIA gene details page Other Links
POLL polled Bos taurus - no genomic information (-..-) POLL Homologene, Ensembl, NCBI gene

Variants

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Breed(s) Variant Phenotype Gene Allele Type of Variant Reference Sequence Chr. g. or m. c. or n. p. Verbal Description EVA ID Year Published PubMed ID(s) Acknowledgements
Polled, Celtic allele POLL P[sub]C OR P[sub]202ID complex rearrangement UMD3.1 1 g.1706051_1706060 delins170583 "a duplication of 212 bp replacing a segment of 10 bp" in a region with no known genes or expressed sequence tags. Celtic = breeds of European origin, e.g. Charolais 2013 23717440
Friesian Polled, Friesian allele POLL P[sub]F OR P(sub)80kbID duplication UMD3.1 1 g.1909352_1989480dup 2013 23717440
Mongolian Turano Polled, Mongolian allele POLL P[sub]M OR P[sub]219ID complex rearrangement UMD3.1 1 g.1976128_?? "a complex 219-bp duplication-insertion (P219ID) beginning at 1,976,128 bp and a 7-bp deletion and 6-bp insertion (P1ID) located 621 bp upstream of this position . . . . This rearrangement results in duplication of an 11-bp motif (5'-AAAGAAGCAAA-3') that is entirely conserved among Bovidae . . . and that is also duplicated in the 80-kb duplication responsible for Friesian polledness" 2017 28135247

References


Note: the references are listed in reverse chronological order (from the most recent year to the earliest year), and alphabetically by first author within a year.
2018 Stafuzza, N.B., Silva, R.M.O., Peripolli, E., Bezerra, L.A.F., Lôbo, R.B., Magnabosco, C.U., Di Croce, F.A., Osterstock, J.B., Munari, D.P., Lourenco, D.A.L., Baldi, F. :
Genome-wide association study provides insights into genes related with horn development in Nelore beef cattle. PLoS One 13:e0202978, 2018. Pubmed reference: 30161212. DOI: 10.1371/journal.pone.0202978.
2017 Chen, S.Y., Liu, L., Fu, M., Zhang, G.W., Yi, J., Lai, S.J., Wang, W. :
Simultaneous introgression of three POLLED mutations into a synthetic breed of Chinese cattle. PLoS One 12:e0186862, 2017. Pubmed reference: 29053739. DOI: 10.1371/journal.pone.0186862.
Medugorac, I., Graf, A., Grohs, C., Rothammer, S., Zagdsuren, Y., Gladyr, E., Zinovieva, N., Barbieri, J., Seichter, D., Russ, I., Eggen, A., Hellenthal, G., Brem, G., Blum, H., Krebs, S., Capitan, A. :
Whole-genome analysis of introgressive hybridization and characterization of the bovine legacy of Mongolian yaks. Nat Genet 49:470-475, 2017. Pubmed reference: 28135247. DOI: 10.1038/ng.3775.
2016 Carlson, D.F., Lancto, C.A., Zang, B., Kim, E.S., Walton, M., Oldeschulte, D., Seabury, C., Sonstegard, T.S., Fahrenkrug, S.C. :
Production of hornless dairy cattle from genome-edited cell lines. Nat Biotechnol 34:479-81, 2016. Pubmed reference: 27153274. DOI: 10.1038/nbt.3560.
Randhawa, I.A., Khatkar, M.S., Thomson, P.C., Raadsma, H.W. :
A Meta-Assembly of Selection Signatures in Cattle. PLoS One 11:e0153013, 2016. Pubmed reference: 27045296. DOI: 10.1371/journal.pone.0153013.
Scheper, C., Wensch-Dorendorf, M., Yin, T., Dressel, H., Swalve, H., König, S. :
Evaluation of breeding strategies for polledness in dairy cattle using a newly developed simulation framework for quantitative and Mendelian traits. Genet Sel Evol 48:50, 2016. Pubmed reference: 27357942. DOI: 10.1186/s12711-016-0228-7.
Segelke, D., Täubert, H., Reinhardt, F., Thaller, G. :
Considering genetic characteristics in German Holstein breeding programs. J Dairy Sci 99:458-67, 2016. Pubmed reference: 26601581. DOI: 10.3168/jds.2015-9764.
2015 Tetens, J., Wiedemar, N., Menoud, A., Thaller, G., Drögemüller, C. :
Association mapping of the scurs locus in polled Simmental cattle--evidence for genetic heterogeneity. Anim Genet 46:224-5, 2015. Pubmed reference: 25645725. DOI: 10.1111/age.12237.
Wiener, D.J., Wiedemar, N., Welle, M.M., Drögemüller, C. :
Novel Features of the Prenatal Horn Bud Development in Cattle (Bos taurus). PLoS One 10:e0127691, 2015. Pubmed reference: 25993643. DOI: 10.1371/journal.pone.0127691.
2014 Rothammer, S., Capitan, A., Mullaart, E., Seichter, D., Russ, I., Medugorac, I. :
The 80-kb DNA duplication on BTA1 is the only remaining candidate mutation for the polled phenotype of Friesian origin. Genet Sel Evol 46:44, 2014. Pubmed reference: 24993890. DOI: 10.1186/1297-9686-46-44.
Spurlock, D.M., Stock, M.L., Coetzee, J.F. :
The impact of 3 strategies for incorporating polled genetics into a dairy cattle breeding program on the overall herd genetic merit. J Dairy Sci 97:5265-74, 2014. Pubmed reference: 24913645. DOI: 10.3168/jds.2013-7746.
Wiedemar, N., Tetens, J., Jagannathan, V., Menoud, A., Neuenschwander, S., Bruggmann, R., Thaller, G., Drögemüller, C. :
Independent polled mutations leading to complex gene expression differences in cattle. PLoS One 9:e93435, 2014. Pubmed reference: 24671182. DOI: 10.1371/journal.pone.0093435.
2013 Allais-Bonnet, A., Grohs, C., Medugorac, I., Krebs, S., Djari, A., Graf, A., Fritz, S., Seichter, D., Baur, A., Russ, I., Bouet, S., Rothammer, S., Wahlberg, P., Esquerré, D., Hoze, C., Boussaha, M., Weiss, B., Thépot, D., Fouilloux, M.N., Rossignol, M.N., van Marle-Köster, E., Hreiðarsdóttir, G.E., Barbey, S., Dozias, D., Cobo, E., Reversé, P., Catros, O., Marchand, J.L., Soulas, P., Roy, P., Marquant-Leguienne, B., Le Bourhis, D., Clément, L., Salas-Cortes, L., Venot, E., Pannetier, M., Phocas, F., Klopp, C., Rocha, D., Fouchet, M., Journaux, L., Bernard-Capel, C., Ponsart, C., Eggen, A., Blum, H., Gallard, Y., Boichard, D., Pailhoux, E., Capitan, A. :
Novel insights into the bovine polled phenotype and horn ontogenesis in Bovidae. PLoS One 8:e63512, 2013. Pubmed reference: 23717440. DOI: 10.1371/journal.pone.0063512.
Druet, T., Pérez-Pardal, L., Charlier, C., Gautier, M. :
Identification of large selective sweeps associated with major genes in cattle. Anim Genet 44:758-62, 2013. Pubmed reference: 23859468. DOI: 10.1111/age.12073.
Glatzer, S., Merten, N.J., Dierks, C., Wöhlke, A., Philipp, U., Distl, O. :
A Single Nucleotide Polymorphism within the Interferon Gamma Receptor 2 Gene Perfectly Coincides with Polledness in Holstein Cattle. PLoS One 8:e67992, 2013. Pubmed reference: 23805331. DOI: 10.1371/journal.pone.0067992.
Hötzel, M.J., Sneddon, J.N. :
The role of extensionists in Santa Catarina, Brazil, in the adoption and rejection of providing pain relief to calves for dehorning. J Dairy Sci 96:1535-48, 2013. Pubmed reference: 23332854. DOI: 10.3168/jds.2012-5780.
Ramey, H.R., Decker, J.E., McKay, S.D., Rolf, M.M., Schnabel, R.D., Taylor, J.F. :
Detection of selective sweeps in cattle using genome-wide SNP data. BMC Genomics 14:382, 2013. Pubmed reference: 23758707. DOI: 10.1186/1471-2164-14-382.
Stock, M.L., Baldridge, S.L., Griffin, D., Coetzee, J.F. :
Bovine dehorning: assessing pain and providing analgesic management. Vet Clin North Am Food Anim Pract 29:103-33, 2013. Pubmed reference: 23438402. DOI: 10.1016/j.cvfa.2012.11.001.
Tan, W., Carlson, D.F., Lancto, C.A., Garbe, J.R., Webster, D.A., Hackett, P.B., Fahrenkrug, S.C. :
Efficient nonmeiotic allele introgression in livestock using custom endonucleases. Proc Natl Acad Sci U S A 110:16526-31, 2013. Pubmed reference: 24014591. DOI: 10.1073/pnas.1310478110.
2012 Mariasegaram, M., Harrison, B.E., Bolton, J.A., Tier, B., Henshall, J.M., Barendse, W., Prayaga, K.C. :
Fine-mapping the POLL locus in Brahman cattle yields the diagnostic marker CSAFG29. Anim Genet 43:683-8, 2012. Pubmed reference: 22497221. DOI: 10.1111/j.1365-2052.2012.02336.x.
Medugorac, I., Seichter, D., Graf, A., Russ, I., Blum, H., Göpel, K.H., Rothammer, S., Förster, M., Krebs, S. :
Bovine polledness - an autosomal dominant trait with allelic heterogeneity. PLoS One 7:e39477, 2012. Pubmed reference: 22737241. DOI: 10.1371/journal.pone.0039477.
Seichter, D., Russ, I., Rothammer, S., Eder, J., Förster, M., Medugorac, I. :
SNP-based association mapping of the polled gene in divergent cattle breeds. Anim Genet 43:595-8, 2012. Pubmed reference: 22497248. DOI: 10.1111/j.1365-2052.2011.02302.x.
2011 Gottardo, F., Nalon, E., Contiero, B., Normando, S., Dalvit, P., Cozzi, G. :
The dehorning of dairy calves: practices and opinions of 639 farmers. J Dairy Sci 94:5724-34, 2011. Pubmed reference: 22032397. DOI: 10.3168/jds.2011-4443.
2010 Mariasegaram, M., Reverter, A., Barris, W., Lehnert, SA., Dalrymple, B., Prayaga, K. :
Transcription profiling provides insights into gene pathways involved in horn and scurs development in cattle. BMC Genomics 11:370, 2010. Pubmed reference: 20537189. DOI: 10.1186/1471-2164-11-370.
Stella, A., Ajmone-Marsan, P., Lazzari, B., Boettcher, P. :
Identification of selection signatures in cattle breeds selected for dairy production. Genetics 185:1451-61, 2010. Pubmed reference: 20479146. DOI: 10.1534/genetics.110.116111.
Wohlke, A., Pommerien, NJ., Distl, O. :
A synaptojanin 1 (SYNJ1) single nucleotide polymorphism not responsible for polledness in German Holstein, Limousin, Charolais and Pinzgauer cattle. Anim Genet 41:335-6, 2010. Pubmed reference: 19922587. DOI: 10.1111/j.1365-2052.2009.02009.x.
2008 Cargill, EJ., Nissing, NJ., Grosz, MD. :
Single nucleotide polymorphisms concordant with the horned/polled trait in Holsteins. BMC Res Notes 1:128, 2008. Pubmed reference: 19063733. DOI: 10.1186/1756-0500-1-128.
2007 Prayaga KC :
Genetic options to replace dehorning in beef cattle — a review Australian Journal of Agricultural Research 58:1-8, 2007.
2006 Wunderlich, KR., Abbey, CA., Clayton, DR., Song, Y., Schein, JE., Georges, M., Coppieters, W., Adelson, DL., Taylor, JF., Davis, SL., Gill, CA. :
A 2.5-Mb contig constructed from Angus, Longhorn and horned Hereford DNA spanning the polled interval on bovine chromosome 1. Anim Genet 37:592-4, 2006. Pubmed reference: 17121607. DOI: 10.1111/j.1365-2052.2006.01538.x.
2005 Drögemüller, C., Wöhlke, A., Leeb, T., Distl, O. :
A 4 Mb high resolution BAC contig on bovine chromosome 1q12 and comparative analysis with human chromosome 21q22. Comp Funct Genomics 6:194-203, 2005. Pubmed reference: 18629192. DOI: 10.1002/cfg.476.
Drögemüller, C., Wöhlke, A., Mömke, S., Distl, O. :
Fine mapping of the polled locus to a 1-Mb region on bovine chromosome 1q12. Mamm Genome 16:613-20, 2005. Pubmed reference: 16180143. DOI: 10.1007/s00335-005-0016-0.
2004 Roman, A. :
L’élevage bovin en Egypte antique. Bull Soc Fr Hist Méd Sci Vét 3:35-45, 2004.
2000 Brockmann, G.A., Martin, J., Teuscher, F., Schwerin, M. :
Marker controlled inheritance of the polled locus in Simmental cattle [German] Archiv fur Tierzucht-Archives of Animal Breeding 43:207-212, 2000.
Kantanen, J., Olsaker, I., Brusgaard, K., Eythorsdottir, E., Holm, LE., Lien, S., Danell, B., Adalsteinsson, S. :
Frequencies of genes for coat colour and horns in Nordic cattle breeds. Genet Sel Evol 32:561-76, 2000. Pubmed reference: 14736370. DOI: 10.1051/gse:2000137.
Lamminger, A., Hamann, H., Rohrmoser, G., Rosenberger, E., Krausslich, H., Distl, O. :
Relationship between polledness and traits used in the breeding objectives for German Fleckvieh [German] Zuchtungskunde 72:325-339, 2000.
1999 Goonewardene, L.A., Pang, H., Berg, R.T., Price, M.A. :
A comparison of reproductive and growth traits of horned and polled cattle in three synthetic beef lines Canadian Journal of Animal Science 79:123-127, 1999.
Goonewardene, L.A., Price, M.A., Liu, M.F., Berg, R.T., Erichsen, C.M. :
A study of growth and carcass traits in dehorned and polled composite bulls Canadian Journal of Animal Science 79:383-385, 1999.
1997 Harlizius, B., Tammen, I., Eichler, K., Eggen, A., Hetzel, D.J.S. :
New markers on bovine chromosome 1 are closely linked to the polled gene in Simmental and Pinzgauer cattle Mammalian Genome 8:255-257, 1997. Pubmed reference: 9096105.
Kommisrud, E., Steine, T. :
Semen collection, semen production and fertility rates in polled vs horned bulls in norway Reproduction in Domestic Animals 32:221-223, 1997.
1996 Brenneman, R.A., Davis, S.K., Sanders, J.O., Burns, B.M., Wheeler, T.C., Turner, J.W., Taylor, J.F. :
The polled locus maps to BTA1 in a Bos indicus x Bos taurus cross Journal of Heredity 87:156-161, 1996. Pubmed reference: 8830095.
Lundrigan, B. :
Morphology of horns and fighting behavior in the family bovidae Journal of Mammalogy 77:462-475, 1996.
Picard, K., Festabianchet, M., Thomas, D. :
The cost of horniness - heat loss may counter sexual selection for large horns in temperate bovids Ecoscience 3:280-284, 1996.
Roberts, S.C. :
The evolution of hornedness in female ruminants [Review] Behaviour 133:399-442, 1996.
Stookey, J.M., Goonewardene, L.A. :
A comparison of production traits and welfare implications between horned and polled beef bulls Canadian Journal of Animal Science 76:1-5, 1996.
1995 Coghlan, A. :
How now, hornless cow New Scientist 148:24, 1995.
Hoffsis, G. :
Surgical (cosmetic) dehorning in cattle Veterinary Clinics of North America - Food Animal Practice 11:159-169, 1995.
Krausslich, H., Rohrmoser, G. :
Breeding a line of polled German Simmentals Zuchtungskunde 67:15-24, 1995.
Schmutz, S.M., Marquess, F.L.S., Berryere, T.G., Moker, J.S. :
DNA marker-assisted selection of the polled condition in Charolais cattle Mammalian Genome 6:710-713, 1995. Pubmed reference: 8563169.
1994 Picard, K., Thomas, D.W., Festabianchet, M., Lanthier, C. :
Bovid Horns - An Important Site for Heat Loss During Winter Journal of Mammalogy 75:710-713, 1994.
1993 Georges, M., Drinkwater, R., King, T., Mishra, A., Moore, S.S., Nielsen, D., Sargeant, L.S., Sorensen, A., Steele, M.R., Zhao, X.Y., Womack, J.E., Hetzel, J. :
Microsatellite Mapping of a Gene Affecting Horn Development in Bos-Taurus Nature Genetics 4:206-210, 1993. Pubmed reference: 8348158. DOI: 10.1038/ng0693-206.
1992 Crow, J.F. :
Schrodinger, Erwin and the Hornless Cattle Problem Genetics 130:237-239, 1992. Pubmed reference: 1541388.
Steel, J. :
Identifying Horny Cattle Search 23:248, 1992.
1991 Estes, R.D. :
The Significance of Horns and Other Male Secondary Sexual Characters in Female Bovids Applied Animal Behaviour Science 29:403-451, 1991.
1990 Lange, H., Brem, G., Utz, J., Gottschalk, A., Karnbaum, B., Krausslich, H. :
Investigations on the polled condition in German Simmental cattle Bayer. Landwirtsch. Jahrb. 67:15-68, 1990.
1989 Krecek, J. :
Present Trends in Horned Cattle Production Biologizace a Chemizace Zivocisne Vyroby-Veterinaria 25:506-509, 1989.
Utz, J., Lange, H. :
[No more horns in the future? Breeding trial on polledness in Simmental cattle] Tierzuchter 41:104-105, 1989.
1982 Brem, G., Karnbaum, B., Rosenberg, E. :
Zur Vererbung der Hornlosigkeit beim Fleckvieh Bayer. Landwirtsch. Jahrb. 59:688-695, 1982.
1980 Frisch, J.E., Nishimura, H., Cousins, K.J., Turner, H.G. :
The inheritance and effect on production of polledness in four crossbred lines of beef cattle Animal Production 31:119-126, 1980.
1978 Long, C.R., Gregory, K.E. :
Inheritance of the horned, scurred and polled condition in cattle Journal of Heredity 69:395-400, 1978.
1958 Berge, S. :
[Varying dominance of hornlessness in cattle] Nordisk Jordbrugsforskning 40:105-110, 1958.
1955 Siegert, M. :
The horns of cattle and their origin with personal investigations on the structure and inheritance of scurs Veterinary Medicine Dissertation, Freie University, Berlin :31, 1955.
1952 Williams, H.D., Williams, T. :
The inheritance of horns and their modifications in polled Hereford cattle Journal of Heredity 43:267-272, 1952.
1936 White, W.T., Ibsen, H.L. :
Horn inheritance in Galloway-Holstein cattle crosses Journal of Genetics 32:33-49, 1936.
1935 Dove, W.F. :
The physiology of horn growth: A study of the morphogenesis, the interaction of tissues, and the evolutionary processes of a mendelian recessive character by means of transplantation of tissues Journal of Experimental Zoology 69:347-405, 1935.
1927 Auld, R.C.M. :
Polled and horned cattle Journal of Heredity 18:309-321, 1927.
1924 Cole, L.J. :
The Wisconsin experiment in cross-breeding cattle Proc World's Dairy Congress 2:1383-1388, 1924.
1916 Lloyd-Jones, O., Evvard, J,M. :
Inheritance of color and horns in blue-gray cattle. Iowa Agric Exp Sta, Ames 30:67-106a, 1916.
1906 Spillman, W.J. :
Mendel’s law in relation to animal breeding. Proceedings of the American Breeders’ Association 1:171-177, 1906.
Spillman, W.J. :
Application of Mendel's law to a practical problem in breeding cattle Journal of Heredity 2:173-177, 1906.
Spillman, WJ. :
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1902 Bateson, W., Saunders, E.R. :
The facts of heredity in the light of Mendel's discovery Reports to the Evolution Committee of the Royal Society 1:125-160, 1902.
1868 Darwin, C.R. :
The Variation of Animals and Plants under Domestication John Murray, London : Vol. II, p. 316, 1868.
1859 Darwin, C.R. :
On the Origin of Species by Means of Natural Selection John Murray, London :, 1859.
90 Tacitus :
The Germany. The Oxford translation revised with notes, section 5. http://ancienthistory.about.com/od/europe/l/bl_text_Tacitus_Germania.htm :, 90.
-450 Herodotus :
The History, Book 4. Translated by G. C. Macaulay (1890), section 29. http://www.sacred-texts.com/cla/hh/hh4020.htm :, -450.

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