Association of polymorphisms in CAPN and CAST genes with physicochemical properties of beef: a review
DOI:
https://doi.org/10.21615/cesmvz.16.1.1Keywords:
bovine, pH, proteolytic system, sensory quality, tenderness, WHC, posmortem managementAbstract
Physicochemical properties such as tenderness, color, juiciness, flavor, marbling, pH and water retention capacity, are considered the most important attributes in the meat quality. Posmortem management influences the functioning of the proteolytic system of the enzymes calpains and calpastatins, which are encoded by CAPN1 and CAST genes, respectively. This review displays the associations of CAPN1 and CAST polymorphisms in order to explain the aforementioned physicochemical properties, which determine the characteristics relate to the beef quality. There are CAPN1 and CAST polymorphisms associated with particular physicochemical properties of meat, but not all polymorphisms are associated with the mentioned properties. The activity of enzymes proteolytic system varies according to peri and posmortem management. The presence or absence of a polymorphism varies between races. Therefore, the associations of single nucleotide polymorphisms with the physicochemical properties of meat mentioned in this review could help to improve the quality indicators of beef.
Downloads
References
Abril M, Campo MM, Önenç A, Sanudo C, Albertí P, Negueruela AI. Beef colour evolution as a function of ultimate pH. Meat Sci 2001; 58(1): 69–78. https://doi.org/10.1016/S0309-1740(00)00133-9
Albrecht E, Teuscher F, Ender K, Wegner J. Growth-and breed-related changes of muscle bundle structure in cattle. J Anim Sci 2006: 84(11): 2959–2964. https://doi.org/10.2527/jas.2006-345
Ángel-Marín PA, Cardona-Cadavid H, Cerón-Muñoz MF. Genómica en la producción animal. Rev Colombiana Cienci Anim 2013; 5(2): 497–518.
Avilés C, Juárez M, Peña F, Domenech V, Clemente I, Molina A. Association of single nucleotide polymorphisms in CAPN1 and CAST genes with beef tenderness from Spanish commercial feedlots. Czech J Anim Sci 2013; 58(10): 479–487. https://doi.org/10.17221/6997-CJAS
Beak SH, Park SJ, Fassah DM, Kim HJ, Kim M, Jo C, et al. Relationships among carcass traits, auction price, and image analysis traits of marbling characteristics in Korean cattle beef. Meat Sci 2021; 171: 108268. https://doi.org/10.1016/j.meatsci.2020.108268
Braz CU, Taylor JF, Decker JE, Bresolin T, Espigolan R, Garcia DA, et al. Polymorphism analysis in genes associated with meat tenderness in Nelore cattle. Meta Gene 2018; 18: 73–78. https://doi.org/10.1016/j.mgene.2018.08.002
Calvo JH, Iguácel LP, Kirinus JK, Serrano M, Ripoll G, Casasús I, et al. A new single nucleotide polymorphism in the calpastatin (CAST) gene associated with beef tenderness. Meat Sci 2014; 96(2): 775–782. https://doi.org/10.1016/j.meatsci.2013.10.003
Cantet RJC, Gualdrón-Duarte JL, Munilla-Leguizamón S. Selección Genómica. Rev Arg Prod Anim 2008; 28(2): 133–136.
Carvalho ME, Eler JP, Bonin MN, Rezende FM, Biase FH, Meirelles FV, et al. Genotypic and allelic frequencies of gene polymorphisms associated with meat tenderness in Nellore beef cattle. Genet Mol Res 2017; 16(1): gmr16018957. https://doi.org/10.4238/gmr16018957
Casas E, White S N, Wheeler TL. Effects of calpastatin and μ-calpain markers in beef cattle on tenderness traits. J Anim Sci 2006; 84(3): 520–525. https://doi.org/10.2527/2006.843520x
Casas E, White SN, Riley DG, Smith TPL, Brenneman RA, Olson TA, et al. (2005). Assessment of single nucleotide polymorphisms in genes residing on chromosomes 14 and 29 for association with carcass composition traits in Bos indicus cattle. J Anim Sci 2005; 83(1): 13–19. https://doi.org/10.2527/2005.83113x
Cassar-Malek I, Picard B. Expression marker-based strategy to improve beef quality. Sci World J 2016; 2016: 1–11. https://doi.org/10.1155/2016/2185323
Cassar-Malek I, Picard B, Bernard C, Hocquette JF. Application of gene expression studies in livestock production systems: a European perspective. Aust J Exp Agric 2008; 48(7): 701–710. https://doi.org/10.1071/EA08018
Castro S, Ríos M, Ortiz Y, Manrique C, Jiménez A, Ariza F. Association of single nucleotide polymorphisms in CAPN1, CAST and MB genes with meat color of Brahman and crossbreed cattle. Meat Sci 2016; 117: 44–49. https://doi.org/10.1016/j.meatsci.2016.02.021
Chasco J, Lizaso G, Beriain MJ, Horcada A, Goirraiz C, Hernandez B, et al. Efecto de la Maduracion a Vacio en el Color de la Carne de Ternera de Raza Pirenaica. En: VI Jornadas Produccion Animal. Zaragoza: AIDA ITEA 1995; 16: 621–623.
Cheong HS, Yoon DH, Park BL, Kim LH, Bae JS, Namgoong S, et al. A single nucleotide polymorphism in CAPN1 associated with marbling score in Korean cattle. BMC genet 2008; 9(1): 33. https://doi.org/10.1186/1471-2156-9-33
Chiang WC, Chen YM, Lin SL, Wu KD, Tsai TJ. Bradykinin enhances reactive oxygen species generation, mitochondrial injury, and cell death induced by ATP depletion —A role of the phospholipase C-Ca2+ pathway. Free Radic Biol Med 2007; 43(5): 702–710. https://doi.org/10.1016/j.freeradbiomed.2007.04.032
Chriki S, Renand G, Picard B, Micol D, Journaux L, Hocquette JF. Meta-analysis of the relationships between beef tenderness and muscle characteristics. Livest Sci 2013; 155(2-3): 424–434. https://doi.org/10.1016/j.livsci.2013.04.009
Chung H, Shin S, Chung E. Effects of genetic variants for the bovine calpain gene on meat tenderness. Mol Biol Rep 2014; 41(5): 2963–2970. https://doi.org/10.1007/s11033-014-3152-3
Ciobanu DC, Bastiaansen JW, Lonergan SM, Thomsen H, Dekkers JC, Plastow GS, et al. New alleles in calpastatin gene are associated with meat quality traits in pigs. J Anim Sci 2004; 82(10): 2829–2839. https://doi.org/10.2527/2004.82102829x
Curi RA, Chardulo LAL, Mason MC, Arrigoni MDB, Silveira AC, De Oliveira HN. Effect of single nucleotide polymorphisms of CAPN1 and CAST genes on meat traits in Nellore beef cattle (Bos indicus) and in their crosses with Bos taurus. Anim Genet 2009; 40(4): 456–462. https://doi.org/10.1111/j.1365-2052.2009.01859.x
de Oliveira LG, Delgado EF, Steadham EM, Huff-Lonergan E, Lonergan SM. Association of calpain and calpastatin activity to postmortem myofibrillar protein degradation and sarcoplasmic proteome changes in bovine Longissiumus lumborum and Triceps brachii. Meat Sci 2019; 155: 50–60. https://doi.org/10.1016/j.meatsci.2019.04.015
Dear TN, Meier NT, Hunn M, Boehm T. Gene structure, chromosomal localization, and expression pattern of Capn12, a new member of the calpain large subunit gene family. Genomics 2000; 68(2): 152–160. https://doi.org/10.1006/geno.2000.6289
Ellies-Oury MP, Dumont R, Perrier G, Roux M, Micol D, Picard B. Effect of age and carcass weight on quality traits of m. rectus abdominis from Charolais heifers. Animal 2017; 11(4): 720–727. https://doi.org/10.1017/S1751731116001907
Elsik CG, Tellam RL, Worley KC. The genome sequence of taurine cattle: a window to ruminant biology and evolution. Science 2009; 324(5926): 522–528. https://doi.org/10.1126/science.1169588
Gagaoua M, Picard B, Soulat J, Monteils V. Clustering of sensory eating qualities of beef: Consistencies and differences within carcass, muscle, animal characteristics and rearing factors. Livest Sci 2018; 214: 245–258. https://doi.org/10.1016/j.livsci.2018.06.011
Gagaoua M, Terlouw EC, Picard B. The study of protein biomarkers to understand the biochemical processes underlying beef color development in young bulls. Meat Sci 2017; 134: 18–27. https://doi.org/10.1016/j.meatsci.2017.07.014
Gagaoua M, Terlouw EC, Micol D, Boudjellal A, Hocquette JF, Picard B. Understanding early post-mortem biochemical processes underlying meat color and pH decline in the Longissimus thoracis muscle of young Blond d’Aquitaine bulls using protein biomarkers. J Agri Food Chem 2015; 63(30): 6799–6809. https://doi.org/10.1021/acs.jafc.5b02615
Gill JL, Bishop SC, McCorquodale C, Williams JL, Wiener P. Association of selected SNP with carcass and taste panel assessed meat quality traits in a commercial population of Aberdeen Angus-sired beef cattle. Genet Sel Evol 2009; 41(1): 36. https://doi.org/10.1186/1297-9686-41-36
Goll DE, Thompson VF, Li H, Wei WE I, Cong J. The calpain system. Physiol Rev 2003; 83: 731–801. https://doi.org/10.1152/physrev.00029.2002
Heaton MP, Harhay GP, Bennett GL, Stone RT, Grosse WM, Casas E, et al. Selection and use of SNP markers for animal identification and paternity analysis in US beef cattle. Mamm Genome 2002; 13(5): 272–281. https://doi.org/10.1007/s00335-001-2146-3
Hocquette JF, Gondret F, Baéza E, Médale F, Jurie C, Pethick DW. Intramuscular fat content in meat-producing animals: development, genetic and nutritional control, and identification of putative markers. Animal 2010; 4(02): 303–319. http://dx.doi.org/10.1017/S1751731109991091
Hou G, Huang M, Gao X, Li J, Gao H, Ren H, et al. Association of Calpain 1 (CAPN1) and HRSP12 allelic variants in beef cattle with carcass traits. Afr J Biotechnol 2011; 10(63): 13714–13718. https://doi.org/10.5897/AJB11.338
Huang Z, Hoffmann FW, Norton RL, Hashimoto AC, Hoffmann PR. Selenoprotein K is a novel target of m-calpain, and cleavage is regulated by Toll-like receptor-induced calpastatin in macrophages. J Biol Chem 2011; 286(40): 34830–34838. https://doi.org/10.1074/jbc.M111.265520
Hwang IH, Thompson JM. The interaction between pH and temperature decline early postmortem on the calpain system and objective tenderness in electrically stimulated beef longissimus dorsi muscle. Meat Sci 2001; 58(2): 167–174. https://doi.org/10.1016/S0309-1740(00)00147-9
Juszczuk-Kubiak E, Sakowski T, Flisikowski K, Wiciñska K, Oprzadek J, Rosochacki SJ. Bovine mu-calpain (CAPN1) gene: new SNP within intron 14. J Appl Genet 2004; 45(4): 457–460.
Kappes SM, Keele JW, Stone RT, McGraw RA, Sonstegard TS, Smith TP, et al. A second-generation linkage map of the bovine genome. Genome Res 1997; 7(3): 235–249. https://doi.org/10.1101/gr.7.3.235
KÖK S, Atalay S. The Use of various SNPs in CAST and CAPN1 genes to determine the meat tenderness in Turkish grey cattle. Kafkas Üniv Vet Fak Derg 2018; 24(1): 1–8. https://doi.org/10.9775/kvfd.2017.17617
Konarska M, Kuchida K, Tarr G, Polkinghorne RJ. Relationships between marbling measures across principal muscles. Meat Sci 2017; 123: 67–78. https://doi.org/10.1016/j.meatsci.2016.09.005
Kristensen L, Purslow PP. The effect of ageing on the water-holding capacity of pork: role of cytoskeletal proteins. Meat Sci 2001; 58(1): 17–23. https://doi.org/10.1016/S0309-1740(00)00125-X
Leal-Gutiérrez JD, Jiménez-Robayo LM. Análisis computacional del efecto de polimorfismos de genes del sistema μ-calpaína/calpastatina sobre la calidad de la carne bovina. Rev Med Vet Zoot 2015; 62(1): 50-66. http://dx.doi.org/10.15446/rfmvz.v62n1.49385
Leal-Gutiérrez JD, Jiménez-Robayo LM, Ariza M, Manrique C, López J, Martínez C, et al. Polimorfismos de los genes CAPN1, CAST, DES, PRKAG3 y RYR1 asociados a la capacidad de retención de agua en crudo y cocinado en carne de bovino en cruces Bos indicus y Bos taurus en Colombia. Arch Zootec 2015; 64(245): 29–35.
Leal-Gutiérrez JD, Jiménez-Robayo LM, Ariza M, Manrique C, López J, Martínez C, et al. Efecto del tipo genético y la maduración sobre la retención de agua en carne de toros castrados. Arch Zootec 2014; 63(243): 409–418.
Lee B, Yoon S, Lee Y, Oh E, Yun YK, Do Kim B, et al. Comparison of marbling fleck characteristics and objective tenderness parameters with different marbling coarseness within longissimus thoracis muscle of high-marbled Hanwoo steer. Korean J Food Sci An 2018; 38(3): 606–614. https://doi.org/10.5851/kosfa.2018.38.3.606
Lee SH, Kim SC, Chai HH, Cho SH, Kim HC, Lim D, et al. Mutations in calpastatin and μ-calpain are associated with meat tenderness, flavor and juiciness in Hanwoo (Korean cattle): Molecular modeling of the effects of substitutions in the calpastatin/μ-calpain complex. Meat Sci 2014; 96(4): 1501–1508. https://doi.org/10.1016/j.meatsci.2013.11.026
Li J, Zhang LP, Gan QF, Li JY, Gao HJ, Yuan ZR, et al. Association of CAST gene polymorphisms with carcass and meat quality traits in Chinese commercial cattle herds. Asian Austral J Anim 2010; 23(11): 1405-1411. https://doi.org/10.5713/ajas.2010.90602
Li X, Ekerljung M, Lundström K, Lundén A. Association of polymorphisms at DGAT1, leptin, SCD1, CAPN1 and CAST genes with color, marbling and water holding capacity in meat from beef cattle populations in Sweden. Meat Sci 2013; 94(2): 153–158. https://doi.org/10.1016/j.meatsci.2013.01.010
Liu X, Usman T, Wang Y, Wang Z, Xu X, Wu M, et al. Polymorphisms in epigenetic and meat quality related genes in fourteen cattle breeds and association with beef quality and carcass traits. Asian Austral J Anim 2015; 28(4): 467–475. https://doi.org/10.5713/ajas.13.0837
Mancini RA, Hunt M. Current research in meat color. Meat Sci 2005; 71(1): 100–121. https://doi.org/10.1016/j.meatsci.2005.03.003
Marsh BB, Ringkob TP, Russell RL, Swartz DR, Pagel LA. Effects of early-postmortem glycolytic rate on beef tenderness. Meat Sci 1987; 21(4): 241–248. https://doi.org/10.1016/0309-1740(87)90061-1
Martínez CA, Manrique C, A Elzo M. Cattle genetic evaluation: a historical perception. Rev Colom Cienc Pecua 2012; 25(2): 293–311.
Melody JL, Lonergan SM, Rowe LJ, Huiatt TW, Mayes MS, Huff-Lonergan E. Early postmortem biochemical factors influence tenderness and water-holding capacity of three porcine muscles. J Anim Sci 2004; 82(4): 1195–1205. https://doi.org/10.2527/2004.8241195x
Morón-Fuenmayor O, Araujo-Febres O, Pietrosemoli S, Gallardo N, Sulbarán B, Peña J. Efecto de la castración sobre la composición físico-química y características sensoriales en carne de bovinos mestizos comerciales. Rev Fac Agron (LUZ) 2010; 27: 594-606.
Motter MM, Corva PM, Marrube G, Miquel MC, Papaleo Mazzuco J, Villarreal EL, et al. Asociación de dos marcadores del gen de la calpastatina con variables productivas de novillos Brangus engordados en pasturas. Revista Argentina de Producción Animal 2013; 33(1): 21–29.
O’keeffe M, Hood DE. Biochemical factors influencing metmyoglobin formation on beef from muscles of differing colour stability. Meat Sci 1982; 7(3): 209–228. https://doi.org/10.1016/0309-1740(82)90087-0
Oliete B, Carballo JA, Monserrat L, Varela A, Moreno T, Sánchez L. Variación del color de la carne en la raza Rubia Gallega, Holstein Frisian y su cruce. En: Efecto del tiempo de conservación. XII Congreso Zootecnia 2002; pp. 609–612.
Page BT, Casas E, Heaton MP, Cullen NG, Hyndman DL, Morris CA, et al. Evaluation of single-nucleotide polymorphisms in CAPN1 for association with meat tenderness in cattle. J Anim Sci 2002; 80(12): 3077–3085. https://doi.org/10.2527/2002.80123077x
Page BT, Casas E, Quaas RL, Thallman RM, Wheeler TL, Shackelford SD, et al. Association of markers in the bovine CAPN1 gene with meat tenderness in large crossbred populations that sample influential industry sires. J Anim Sci 2004; 82(12): 3474–3481. https://doi.org/10.2527/2004.82123474x
Park SJ, Beak SH, Da Jin Sol Jung SY, Kim IHJ, Piao MY, Kang HJ, et al. Genetic, management, and nutritional factors affecting intramuscular fat deposition in beef cattle—a review. Asian Austral J Anim 2018; 31(7): 1043–1061. https://doi.org/1043.10.5713/ajas.18.0310
Pearson AM, Wenham LM, Carse WA, McLeod K, Davey CL, Kirton AH. Observations on the contribution of fat and lean to the aroma of cooked beef and lamb. J Anim Sci 1973; 36(3): 511–515. https://doi.org/10.2527/jas1973.363511x
Pinto LFB, Ferraz JBS, Meirelles FV, Eler JP, Rezende FM, Carvalho ME, et al. Association of SNPs on CAPN 1 and CAST genes with tenderness in Nellore cattle. Genet Mol Res 2010; 9(3): 1431–1442. https://doi.org/10.4238/vol9-3gmr881
Pinto LF, Ferraz JB, Pedrosa VB, Eler JP, Meirelles FV, Bonin MN, et al. Single nucleotide polymorphisms in CAPN and leptin genes associated with meat color and tenderness in Nellore cattle. Gen Mol Res 2011; 10(3): 2057–2064. http://dx.doi.org/10.4238/vol10-3gmr1263
Pratiwi N, Maskur M, Priyanto R, Jakaria J. Novel SNP of calpain-1 (CAPN1) gene and its association with carcass and meat characteristics traits in Bali cattle. J Indones Trop Anim Agric 2016; 41(3): 109–116. https://doi.org/10.14710/jitaa.41.3.109-116
Priolo A, Micol D, Agabriel J. Effects of grass feeding systems on ruminant meat colour and flavour. A review. Anim Res 2001; 50(3): 185-200. https://doi.org/10.1051/animres:2001125
Raynaud P, Gillard M, Parr T, Bardsley R, Amarger V, Levéziel H. Correlation between bovine calpastatin mRNA transcripts and protein isoforms. Arch Biochem Biophys 2005; 440(1): 46–53. https://doi.org/10.1016/j.abb.2005.05.028
Reardon W, Mullen AM, Sweeney T, Hamill RM. Association of polymorphisms in candidate genes with colour, water-holding capacity, and composition traits in bovine M. longissimus and M. semimembranosus. Meat Sci 2010; 86(2): 270–275. https://doi.org/10.1016/j.meatsci.2010.04.013
Renand G, Picard B, Touraille C, Berge P, Lepetit J. Relationships between muscle characteristics and meat quality traits of young Charolais bulls. Meat Sci 2001; 59(1): 49–60. https://doi.org/10.1016/S0309-1740(01)00051-1
Renerre M, Anton M, Gatellier P. Autoxidation of purified myoglobin from two bovine muscles. Meat Sci 1992; 32(3): 331–342. https://doi.org/10.1016/0309-1740(92)90096-M
Rodriguez-Zas SL, Southey BR, Heyen DW, Lewin HA. Interval and composite interval mapping of somatic cell score, yield, and components of milk in dairy cattle. J Dairy Sci 2002; 85(11): 3081–3091. https://doi.org/10.3168/jds.S0022-0302(02)74395-6
Schenkel FS, Miller SP, Jiang Z, Mandell IB, Ye X, Li H, et al. Association of a single nucleotide polymorphism in the calpastatin gene with carcass and meat quality traits of beef cattle. J Anim Sci 2006; 84(2): 291–299. https://doi.org/10.2527/2006.842291x
Shin SC, Chung ER. Association of SNP marker in the thyroglobulin gene with carcass and meat quality traits in Korean cattle. Asian Aaustral J Anim 2006: 20(2): 172–177. https://doi.org/10.5713/ajas.2007.172
Smith TPL, Casas E, Rexroad Iii CE, Kappes SM, Keele JW. Bovine CAPN1 maps to a region of BTA29 containing a quantitative trait locus for meat tenderness. J Anim Sci 2000; 78(10): 2589–2594. https://doi.org/10.2527/2000.78102589x
Smith T, Thomas MG, Bidner TD, Paschal JC, Franke DE. Single nucleotide polymorphisms in Brahman steers and their association with carcass and tenderness traits. Genet Mol Res 2009; 8(1): 39–46. https://doi.org/10.4238/vol8-1gmr537
Song S, Zhang X, Hayat K, Liu P, Jia C, Xia S, et al. Formation of the beef flavour precursors and their correlation with chemical parameters during the controlled thermal oxidation of tallow. Food Chem 2011; 124(1): 203–209. https://doi.org/10.1016/j.foodchem.2010.06.010
Straadt IK, Rasmussen M, Andersen HJ, Bertram HC. Aging-induced changes in microstructure and water distribution in fresh and cooked pork in relation to water-holding capacity and cooking loss–A combined confocal laser scanning microscopy (CLSM) and low-field nuclear magnetic resonance relaxation study. Meat Sci 2007; 75(4): 687–695. https://doi.org/10.1016/j.meatsci.2006.09.019
Sun X, Wu X, Fan Y, Mao Y, Ji D, Huang B, et al. Effects of polymorphisms in CAPN1 and CAST genes on meat tenderness of Chinese Simmental cattle. Arch Anim Breed 2018: 61(4): 433–439. https://doi.org/10.5194/aab-61-433-2018
Van Ba H, Reddy BV, Hwang I. Role of calpastatin in the regulation of mRNA expression of calpain, caspase, and heat shock protein systems in bovine muscle satellite cells. In Vitro Cell Dev Biol Animal 2015; 51(5): 447–454. https://doi.org/10.1007/s11626-014-9849-8
Viitala SM, Schulman NF, de Koning DJ, Elo K, Kinos R, Virta A, et al. Quantitative trait loci affecting milk production traits in Finnish Ayrshire dairy cattle. J Dairy Sci 2003; 86(5): 1828–1836. https://doi.org/10.3168/jds.S0022-0302(03)73769-2
Wheeler TL, Cundiff LV, Koch RM. Effect of marbling degree on beef palatability in Bos taurus and Bos indicus cattle. J Anim Sci 1994; 72(12): 3145–3151. https://doi.org/10.2527/1994.72123145x
White SN, Casas E, Wheeler TL, Shackelford SD, Koohmaraie M, Riley DG, et al. A new single nucleotide polymorphism in CAPN1 extends the current tenderness marker test to include cattle of Bos indicus, Bos taurus, and crossbred descent. J Anim Sci 2005; 83(9): 2001–2008. https://doi.org/10.2527/2005.8392001x
Wulf DM, Emnett RS, Leheska JM, Moeller SJ. Relationships among glycolytic potential, dark cutting (dark, firm, and dry) beef, and cooked beef palatability. J Anim Sci 2002; 80(7): 1895–1903. https://doi.org/10.2527/2002.8071895x
Xin J, Zhang LC, Li ZZ, Liu XH, Jin HG, Yan CG. Association of polymorphisms in the calpain I gene with meat quality traits in Yanbian yellow cattle of China. Asian Austral J Anim 2011; 24(1): 9–16. https://doi.org/10.5713/ajas.2011.90407
Yamada T, Sasaki S, Sukegawa S, Yoshioka S, Takahagi Y, Morita M, et al. Association of a single nucleotide polymorphism in titin gene with marbling in Japanese Black beef cattle. BMC Res Notes 2009; 2(1): 78. https://doi.org/10.1186/1756-0500-2-78
Downloads
Published
How to Cite
Issue
Section
License
Copyright (c) 2021 CES Medicina Veterinaria y Zootecnia
This work is licensed under a Creative Commons Attribution-ShareAlike 4.0 International License.
Article metrics | |
---|---|
Abstract views | |
Galley vies | |
PDF Views | |
HTML views | |
Other views |