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The Role of Domestication and Breeding on Gluten Intolerance

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The Role of Domestication and Breeding on Gluten Intolerance C HAPTER 15 Cereals Taxonomy: The Role of Domestication and Breeding on Gl ten !ntolerance María J. Giménez1, Javier Gil-Humanes2, Juan B. Alvarez3, Francisco Barro1 1 Instituto e A!ricultura "osteni#le, $"I$, $%r o#a, "&ain. 2 'epartment o( Genetics, Cell Biolo!), an 'evelo&ment an Center (or Genome *n!ineerin!, +niversit) o( Minnesota, Minnea&olis, Minnesota, +nited "tates. 3 'epartamento e Genética, *scuela ,écnica "u&erior e In!eniería A!ron%mica ) e Montes, Cam&us e -a#anales, +niversi a e $%r o#a, $%r o#a, "&ain. m.!a/01ias.csic.es, !il22/101umn.edu, .#[email protected], (#arro1ias.csic.es 'oi3 4tt&355 x. oi.or!51/.36205oms.203 Ho" to cite this cha#ter Giménez MJ, Gil-Humanes J, Alvarez JB, Barro F. Cereals Taxonomy: The Role of Domestication and Breeding on Gluten Intolerance. In Arranz E, Fernández-Bañares F, Rosell CM, Rodri o !, "eña A#, edi$ors. Advances in the Understanding of Gluten Related athology and the !volution of Gluten"#ree #oods. Bar%elona, #&ain: (mnia#cien%e) *+,-. &. ./0--*1. 763 M.J. Giménez, J. Gil-Humanes, J.B. Alvarez, F. Barro A $ s t r a c t "tora!e &roteins o( 84eat, r)e, #arle) an , to a lesser e2tent, oats contain e&ito&es res&onsi#le (or tri!!erin! t4e celiac isease 9$':. In recent eca es an increase (re;uenc) o( $' 4as #een o#serve , an t4ou!4 t4e reasons (or t4is increase are unclear, mo ern &lant #ree in! 4as attracte criticism attri#utin! to t4e ne8 varieties a &art o( t4e res&onsi#ilit) in 8orsenin! t4e ata o( &revalence. <4eat is one o( t4e most im&ortant cro&s 8orl 8i e, &resentin! #ot4 4i!4 a a&ta#ilit) to i((erent environments an )iel s. ,4e omestication o( 84eat is t4e result o( a &revious natural inters&eci(ic 4)#ri ization (irst #et8een i&loi , an t4en #et8een i&loi an tetra&loi s&ecies t4at resulte in 4e2a&loi 84eat. ,4e ol (armers #e!an to select t4e traits t4at 8ere #etter a a&te to t4e use. In t4e 2/t4 centur) t4e 84eat #ree in! 4a its !reat a vance an mo ern varieties 8ere evelo&e . ,4e !lia in-relate !enes, res&onsi#le (or tri!!erin! $', 4ave no a a&tive value an t4ere(ore, i( t4e to2icit) o( 84eat 8as increase urin! t4e &rocess o( omestication an #ree in! t4is 8oul 4ave #een ma e unconsciousl). 'urin! t4e &rocess of natural 4)#ri ization t4e num#er o( !lia in-relate !enes 8as increase . Brea 84eat, r)e, an Ae. tauschii 4ave t4e 4i!4est num#er o( $' e&ito&es &er !ene, an it seems t4at in #rea 84eat, t4is 4i!4 num#er o( e&ito&es is e2&laine #) t4e ' !enome (rom Ae. tauschii. 'urin! t4e &rocess o( omestication an #ree in!, t4e num#er o( $' e&ito&es &er !ene i not increase an even ecrease in some cases. Key"ords $ereal omestication, 84eat #ree ing, &rolamins, gluten, immunotoxicit). 767 Cereals Taxonomy: The Role o Domes"i#a"ion and Bree$in% on Glu"en In"olerance 1& !ntrod ction ,4e c4an!e in t4e 4uman iet urin! t4e =eolit4ic Revolution 4as #een associate 8it4 a general ecline in 4ealt4 in some areas1. $eliac isease 9$': is one of iseases t4at emerge in t4at &erio 2,3 #ut es&ite of #ein! kno8n since ancient times, its 4istor) is relativel) recent. ,4e (irst re(erences concernin! t4e intake of certain (oo s ma) #e 4armful o not ap&ear until t4e late nineteent4 centur), an t4e (irst #reakt4rou!4 came after <orl <ar II, 8it4 t4e emonstration of t4e role of gluten as t4e agent res&onsi#le (or triggerin! t4e isease7. In t4e &ast 0/ )ears, t4e kno8le ge about $' 4as improve signi(icantl), resultin! in a #etter un erstan in! of t4e isease &at4ogenesis, iagnosis, an t4erap)?. @rogress in t4e un erstan in! of t4e isease inclu es t4e recognition of its autoimmune nature, its genetic #asis, an t4e i enti(ication of immunogenic gluten (ragments causin! $' in man) &atients0. ,4e e(inition of t4e isease an t4e iagnostic criteria, 4ave un ergone c4an!es as all questions concernin! t4e $' 4ave #ecome clearer. ,4us, t4e gui elines (or t4e iagnosis of $' recommen e #) t4e *uropean "ociet) (or @ae iatric Gastroenterology He&atology an =utrition 9*"@GHA=:, (irst &u#lis4e in 1970, 4ave #een revise t8ice. As e(ine in t4e last gui elines of 2012A, B$' is an immune-me iate s)stemic isor er elicite #) gluten an relate &rolamins in geneticall) susce&ti#le in ivi uals an c4aracterize #) t4e &resence of a variable com#ination of gluten- e&en ent clinical mani(estations, $D-s&eci(ic anti#o ies, HCA-'D2 or HCA-'DE 4aplot)&es, an enteropat4). $'-s&eci(ic anti#o ies comprise autoanti#o ies against ,G2, inclu in! en omysial anti#o ies 9*MA:, an anti#o ies against eami ate (orms of gliadin &e&ti es 9DG@:F. ,4e &revalence of $' is 1% of t4e general &opulation in 8estern countries #ut varies (rom countr) to countr). Ho8ever, an increase (requenc) of $' in recent ecades 4as #een observe , 84ic4 can #e &artl) attri#ute to t4e advent of serological testin! an increase &u#lic a8areness in some countriesE. ,4e reasons (or t4is increase are unclear, #ut several 4)&ot4eses as 76? M.J. Giménez, J. Gil-Humanes, J.B. Alvarez, F. Barro 4ygiene6 an t4e risin! consumption of cereals, es&eciall) 84eat 9or its erivatives:10, 4ave #een &ropose amon! ot4ers. ,4e &ossi#le roles attri#ute to mo ern &lant #ree in! in 8orsenin! (igures of &revalence of $' are iscusse in t4is c4apter. '& Taxonomy and Domestication of Cereals ,4e transition &rocess (rom 4unter-gat4erin! to a se entar), agriculture-#ase 4uman societ) starte aroun 12,/// )ears ago11. $ereals cultivation an t4eir elaborate &ro ucts 4ave &la)e an essential role in t4e evelopment of 4uman societies, an no8adays re&resent an important element in most of t4e i((erent cultures. Arc4aeological evi ences s4o8 t4at 4umans 4arveste t4e 8il (orms of cereals (rom natural stan s #e(ore t4e) starte to eli#erate, cultivate an omesticate cereals, 84ic4 involve t4e selection an mo i(ication of important traits suc4 as see size an rac4is sti((ness in t4e (irst cultivate (iel s. ,4e eas) carr) an storage, toget4er 8it4 t4e 4i!4 content in car#oh) rates an &roteins, are some of t4e c4aracteristics associate to t4e (irst &lants t4at 8ere cultivate an omesticate . $ereals are t4e omestic variants of t4e s&ecies (rom t4e Gramineae (amil) 9Poaceae Barn4art:. ,4e &aleobotanical recor s suggest t4at t4is (amil) 8as generate about ?/-A/ million )ears ago 9MHA:. <it4in t4is (amil), t4e main s&ecies of agronomical interest are (oun in t4e t4ree su#(amilies3 Ehrhartoideae Cin> 9rice:, Panicoideae Cin> 9maize, sugar cane, an sor!4um: an Pooideae Bent4am 9oat, 84eat, r)e, an #arle):, t4is last su#(amil) is (orme #) 15 tri#es, #ein! t4e tri#es Avenae 9oat: an Triticeae 9#arle), r)e, an 84eat: t4e most important 9Figure 1). Alt4ou!4 no8, s&ecies of #ot4 tri#es are name as cereals, &4ylogenetic relations4i&s suggest t4at t4e se&aration #et8een #ot4 grou&s #egan about 2/ MHA 12. ,4e se&aration 8it4in t4e Triticeae tri#e is more recent an 4as #een establis4e aroun t8o MHAI alt4ou!4 a recent stu ) suggests t4at t4ese s&eciation events mi!4t 4ave occurre along to t4e last ten million )ears13. 760 Cereals Taxonomy: The Role o Domes"i#a"ion and Bree$in% on Glu"en In"olerance Fi%ure '. (ri%in and rela"ionshi) o ma*or #ereals. Su, amily Pooideae Bentham -oa", .hea", rye, and ,arley/, #om)rise$ '0 "ri,es, ,eing "he "ri,es Avenae -oa"/ and Triticeae -,arley, rye, and .hea"/ "he mos" im)ortant. '&1& (at ,4e genus Avena inclu es cultivate s&ecies 8it4 i((erent &loi ) level. ,4ree cytogeneticall) in e&en ent stocks of Avena C. occurre urin! omestication 9A. sativa, A. strigosa, an A. abyssinica:, alt4ou!4 onl) A. sativa #ecame a &rinci&al cereal. Jats 8ere &robabl) evolve (rom 8ee ) t)&es t4at in(ecte 84eat an #arle) (iel s, an not un er t4e omestication as a crop14. ,4e 4exaploi A. sativa, common oat 92n K 0L K 72), is 8i el) cultivate , an its main uses are (or 4uman consumption 9oat meals an rolle oats: an livestoc> (ee . ,4e 8il relative of A. sativa is t4e 4exaploi 8il oat A. sterilis, 84ose (irst evi ences of cultivation ate M11,7// )ears #e(ore &resent 9BP:, in 84ic4 is t4ou!4t to #e a &re omestication cultivation &ractice15. Besi es, t4e (irst evi ences of omesticate A. sativa cultivation 4ave #een (oun in "acarovca, Mol avia 9M7,0//-7,400 BP:. 76A M.J. Giménez, J. Gil-Humanes, J.B. Alvarez, F. Barro '&'& Barley $ultivate #arle) 9Hordeum vulgare: 92n K 2L K 17I HH: constitutes one o( t4e (irst omesticate cro&s, an one o( t4e most im&ortant 8orl cro&s. It evolve (rom its 8il &ro!enitor H. spontaneum, 84ic4 8as ori!inate ?.? MHA. <il #arle) starte to #e 4arveste #) 4umans as earl) as ?/,/// )ears a!o10, an t4e main c4aracters associate 8it4 its omestication 8ere3 non-#rittle rac4is, increase see 8ei!4t, si2-ro8e ears an na>e see s. ,4e omestication o( 8il #arle) 4as #een escri#e in t4e literature to occur in several !eo!ra&4ical locations, 8it4 at least t4ree main centers o( omestication3 Fertile $rescent, $entral Asia, an ,i#et 9revie8e in Greco et al1A:.
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