Utilization of farm animal genetic resources in a changing agro-ecological environment in the Nordic countries

REVIEW ARTICLE published: 25 Feb ua y 2015 doi: 10.3389/ gene.2015.00052 U iliza ion o a m animal gene ic esou ces in a changing ag o-ecological en i onmen in he No dic coun ies Juha Kan anen1,2*, Pe e Lø endahl3,E ling S andbe g4,Emma Ey ho sdo i 5,Meng-Hua Li1,6, Anne Ke unen-P æbel7,Pee Be g7and Theo Meuwissen8 1G een Technology, Na u al Resou ces Ins i u e Finland, Jokioinen, Finland 2Depa men o Biology, Uni e si y o Eas e n Finland, Kuopio, Finland 3Cen e o Quan i a i e Gene ics and Genomics, Depa men o Molecula Biology and Gene ics, Aa hus Uni e si y, Tjele, Denma k 4Depa men o Animal B eeding and Gene ics, Swedish Uni e si y o Ag icul u al Sciences, Uppsala, Sweden 5Facul y o Land and Animal Resou ces, Ag icul u al Uni e si y o Iceland, Reykja ik, Iceland 6Key Labo a o y o Animal Ecology and Conse a ion Biology, Ins i u e o Zoology, Chinese Academy o Sciences, Beijing, China 7No dGen – No dic Gene ic Resou ce Cen e , Aas, No way 8Depa men o Animal and Aquacul u al Sciences, No wegian Uni e si y o Li e Sciences, Aas, No way Edi ed by: Ino Cu ik, Uni e si y o Zag eb, C oa ia Re iewed by: Shaolin Wang, Uni e si y o Vi ginia, USA Mahdi Saa chi, Iowa S a e Uni e si y, USA *Co espondence: Juha Kan anen, G een Technology, Na u al Resou ces Ins i u e Finland, Mylly ie 1, FI-31600 Jokioinen, Finland e-mail: [email p o ec ed] Li es ock p oduc ion is he mos impo an componen o no he n Eu opean ag icul u e and con ibu es o and will be a ec ed by clima e change. Ne e heless, he ole o a m animal gene ic esou ces in he adap a ion o new ag o-ecological condi ions and mi iga ion o animal p oduc ion’s e ec s on clima e change has been inadequa ely discussed despi e he e being se e al impo an associa ions be ween animal gene ic esou ces and clima e change issues. The sus ainabili y o animal p oduc ion sys ems and u u e ood secu i y equi e access o a wide di e si y o animal gene ic esou ces. The e a e se e al gene ic ques ions ha should be conside ed in s a egies p omo ing adap a ion o clima e change and mi iga ion o en i onmen al e ec s o li es ock p oduc ion. Fo example, i may become impo an o choose among b eeds and e en among a m animal species acco ding o hei sui abili y o a u u e wi h al e ed p oduc ion sys ems. Some animals wi h use ul pheno ypes and geno ypes may be mo e use ul han o he s in he changing en i onmen . Robus animal b eeds wi h he po en ial o adap o new ag o-ecological condi ions and ole a e new diseases will be needed. The key issue in mi iga ion o ha m ul g eenhouse gas e ec s induced by li es ock p oduc ion is he educ ion o me hane (CH4) emissions om uminan s. The e a e di e ences in CH4 emissions among b eeds and among indi idual animals wi hin b eeds ha sugges a po en ial o imp o emen in he ai h ough gene ic selec ion. Cha ac e iza ion o b eeds and indi iduals wi h mode n genomic ools should be applied o iden i y b eeds ha ha e gene ically adap ed o ma ginal condi ions and o ge c i ical in o ma ion o b eeding and conse a ion p og ams o a m animal gene ic esou ces. We conclude ha pheno yping and genomic echnologies and adop ion o new b eeding app oaches, such as genomic selec ion in og ession, will p omo e b eeding o use ul cha ac e s in li es ock species. Keywo ds: adap a ion, animal gene ic esou ces, clima e change, genomics, genomic selec ion, li es ock, me hane, mi iga ion INTRODUCTION S udies on impac s o clima e change on p ima y indus ies in he No dic coun ies (Denma k, Finland, Iceland, No way, and Swe- den) ha e ocused mainly on ag icul u al p oduc i i y, land use, poli ical issues, and wa e esou ce a ailabili y (e.g., Olesen and Bindi, 2002; Cisca e al., 2011; Hakala e al., 2011; Olesen e al., 2011; Höglind e al., 2013). Di e en clima e change scena ios and adap a ion s a egies o no he n Eu opean condi ions ha e also been discussed (e.g., Benes ad, 2005; Olesen e al., 2011). In s udies and epo s, clima e change and li es ock issues ha e been only modes ly conside ed e en hough li es ock p oduc ion is he mos impo an sec o in no he n Eu opean ag icul u e as mea- su ed by he o al alue o p oduc ion (e.g., Niemi and Ahls ed , 2014) and has e ec s on and is in luenced by clima e change. Domes ic animal gene ic esou ces o ood and ag icul u e in pa icula ha e no ye been adequa ely conside ed in s a egies o adap a ion o and mi iga ion o cu en global clima e changes (McMichael e al., 2007; Ho man, 2010) and issues on gene ic esou ces a e ypically ocussed on u u e plan b eeding scena ios (e.g., Cecca elli e al., 2010; Olesen e al., 2011). Howe e , The Global Plan o Ac ion o Animal Gene ic Resou ces (GPA), published by he Commission on Gene ic Resou ces o Food and Ag icul u e, lis s se e al associa ions be ween animal gene ic esou ces and clima e change (FAO, 2007). As poin ed ou by Ho man (2010) and Pilling and Ho man (2011), sus ainabili y and obus ness o animal p oduc ion sys ems and u u e ood www. on ie sin.o g Feb ua y 2015 | Volume 6 | A icle 52 |1 Kan anen e al. Clima e change and animal gene ic esou ces secu i y equi e accessibili y o a wide di e si y o animal gene ic esou ces. Animal gene ic esou ces a e de ined as gene ic di e - si y in domes ica ed animal species ha ing economic o o he socio-cul u al alues and ound among species, among animal b eeds wi hin he species and in c yoconse ed ma e ial (emb yos and semen). Gene ic di e si y e e s o di e ences in allele e- quencies and allele combina ions among b eeds o a m animal species and he spec um o gene ic a ia ion wi hin he b eeds. In GPA, clima e change was widely ecognized as a majo challenge o ag icul u e and ood secu i y (FAO, 2007). GPA is based on achie emen s and common ag eemen s eached a he In e na ional Technical Con e ence on Animal Gene ic Resou ces held in In e laken, Swi ze land in 2007. I includes ou p io i y a eas, p o iding sugges ions and guidelines o cha ac e iza ion, sus ainable use and conse a ion o animal gene ic esou ces and ins i u ional capaci y building ela ed o hese issues. Howden e al. (2007) sugges ed se e al p ac ical app oaches ha could ad ance he po en ial o li es ock p oduc ion sys ems o adap o clima e change and educe g eenhouse gas (GHG) emissions. These included al e ed o a ion o pas u e, modi ica- ions o imes o g azing and iming o ep oduc ion, al e a ion o o age c ops, adequa e wa e supplies, use o supplemen a y eeds and concen a es, and educed need o win e housing in cold clima es. Va ious poli ical op ions a e also a ailable o egula e li es ock p oduc ion and consump ion o p oduc s o animal o igin, which can lead o a educ ion in GHG emissions (Ge be e al., 2010; Ga ne , 2011). Poli ical egula ion can diminish GHG emissions h ough axa ion and subsidies and by p omo ing new ene gy-sa ing echnologies and use o cleane and enewable uels, by c ea ing a po olio o p oduc s o pa icula ma ke s, and by in luencing consume beha io . One use ul way o diminish GHG is o educe mea consump ion, pa icula ly in ich coun ies (Ga ne , 2011). Howe e , he e a e se e al gene ic and animal b eeding ques- ions ha should be conside ed in clima e change s a egies (Ho man, 2010; Wall e al., 2010; B uce, 2013), such as choos- ing among b eeds and e en among species sui ed o changing ci cums ances (Seo e al., 2010). The e may be an inc eased demand o obus animal b eeds wi h he po en ial o adap o changes in en i onmen al condi ions and ole a e new li es ock diseases (Ho man, 2010). Cha ac e iza ion o b eeds wi h mod- e n genomic ools can be applied o iden i y b eeds ha ha e gene ically adap ed o ma ginal ci cums ances. The genomic da a also p o ide c i ical in o ma ion o conse a ion p og ams o a m animal gene ic esou ces. All hese gene ic issues we e exam- ined and discussed in he No dic Resea ch Ne wo k on Animal Gene ic Resou ces in he Adap a ion o Clima e Change (AnGR- No dicNET1). AnGR-No dicNET’s aims we e o p o ide ma e ial, esul s and conclusions o a No dic s a egy o he conse a ion, u iliza ion and in es iga ion o animal gene ic esou ces wi hin adap a ion and mi iga ion issues. AnGR-No dicNET was pa o he p og am “Clima e Change Impac s, Adap a ion and Mi iga- ion in No dic P ima y Indus ies,” which is a hema ic esea ch ne wo k p og am de eloped by he No dic Council o Minis e s (Ba ua e al., 2014). In his pape , we e iew some conclusions o 1h ps://si es.google.com/a/no dgen.o g/ang /home AnGR-No dicNET and he cu en knowledge o clima e change e ec s on he No dic ag o-ecosys ems and li es ock p oduc ion and gi e ecommenda ions o animal b eeding ha conside adap a ion and mi iga ion issues. Mo eo e , we discuss he alues o animal gene ic esou ces o u u e b eeding wo k. CHANGES IN AGROCLIMATIC CONDITIONS IN NORTHERN EUROPE Cu en clima e change, de ec ed as al e a ions in a mosphe ic composi ion, is mainly caused by human ac i i ies, e.g., he bu ning o ossil uels, u baniza ion, shi s in land use, ag icul- u al p ac ices, and li es ock p oduc ion (Meehl e al., 2007). N- e ilize p oduc ion and applica ion, on- a m use o ossil uels, clea ing o es s and o he land o g ow eed o animals and g aze li es ock, manu e managemen , manu e emissions and p ocess- ing, and anspo ing he end p oduc s a e examples o ac i i ies in li es ock p oduc ion ha p oduce GHG (Gill e al., 2010). The changes in a mosphe ic composi ion a ise om an h opogenic emissions o , e.g., ca bon dioxide (CO2), me hane (CH4) and ni ous oxide (N2O) (Ka l and T enbe h, 2003). Acco ding o FAO’s epo (S ein eld e al., 2006), globally 18 pe cen o an h opogenic GHG emissions a e a ibu able o ca le, sheep, goa s and o he domes ic uminan species, camels, ho ses, pigs and poul y. Howe e , he p opo ion o GHG coming om li es ock p oduc ion can a y na ionally and e en egionally depending on he densi y o li es ock popula ions and se e i y o impac s o li es ock p oduc ion on he en i onmen (Mi loehne , 2010). A e iew on GHG emissions om li es ock p oduc ion in he No dic coun ies is gi en elsewhe e (Åby e al., 2014), show- ing na ional emissions om ag icul u e gene ally being lowe han he global a e age. Animal p oduc ion based on uminan s p oduces CH4and N2O in he main while ha o monogas ic species p oduces N2O (Wall e al., 2010). The e is a isk ha li es ock- ela ed GHG emissions will inc ease in he u u e: he human popula ion will con inue o g ow and demand o animal p oduc s will inc ease bo h globally and in he No dic coun ies (Delgado, 2003; Åby e al., 2014; Ge land e al., 2014). This will lead o an inc ease in domes ic animal popula ions and he mi iga ion and adap a ion issues ela ed o li es ock p oduc ion will become inc easingly impo an . The ise o a e age annual su ace empe a u e, a ia ion in p ecipi a ion e en s, and he inc eased occu ence o ex eme wea he e en s, such as wa m pe iods, hea wa es and hea y ain all, a e examples o clima e change (Be ns ein e al., 2007), all o which ha e impac s on ag icul u e and li es ock p oduc ion. The In e na ional Panel on Clima e Change (IPCC) p esen ed scena ios on ends in clima e a iables occu ing du ing he 21s Cen u y i no success ul ac ions a e aken o diminish GHG emissions. Di e en biogeog aphic zones, which a e sepa a ed acco ding o clima ological, biogeog aphic and geological ac o s, a e assumed o expe ience di e en clima ic changes (Benes ad, 2005; Be ns ein e al., 2007; Peel e al., 2007) and clima e is chang- ing in sligh ly di e en ways also ac oss he No dic coun ies. Fo example, in Denma k, which belongs mos ly o he No h A lan ic biogeog aphic zone, cha ac e ized by a mild and humid clima e, he annual mean empe a u e is es ima ed o inc ease by +2°C du ing he 21s Cen u y, leading o d ie and ho e summe s. F on ie s in Gene ics | Li es ock Genomics Feb ua y 2015 | Volume 6 | A icle 52 |2 Kan anen e al. Clima e change and animal gene ic esou ces Also p ecipi a ion in win e is expec ed o inc ease (es ima ion o 0.5 mm/mon h pe decade). In No way, Sweden and Finland, which mos ly belong o he Bo eal o No h Alpine biogeog aphic zones, he annual mean empe a u e is expec ed o inc ease by >3°C o e he cou se o he 21s Cen u y. These egions a e cha ac e ized by a ma ked inc ease in annual p ecipi a ion (close o +1 mm/mon h pe decade), we e win e s and isks o loods. The numbe o days wi h snow co e and/o os will be ewe in he u u e and he snow condi ions will no be as eliable as now (Jylhä e al., 2008). In Iceland, a coun y in he A c ic bio- geog aphic zone wi h low empe a u es, ex eme annual a ia ion in sunligh and sho in ensi e g owing seasons, clima e change is al eady documen ed as a ec ing he dis ibu ion o mois u e, esul ing in shi s in dis ibu ion o plan s and wildli e animals. The annual su ace empe a u e is expec ed o inc ease by 2–4°C, mainly in win e , and p ecipi a ion will be as much as 20% highe in many a eas (Be ns ein e al., 2007). These gene al ou comes o clima e change will a y e en wi hin he no he n Eu opean biogeog aphic egions as e ealed by so- e med downscaled egional clima e models wi h a spa- ial esolu ion o 50 km o less (Benes ad, 2005 and e e ences he ein; Be ns ein e al., 2007). In egions wi h complex landscape s uc u es, e.g., ypical in No way, a p onounced local pa e n in empe a u e and ain all can be de ec ed. In he No dic egion, he s onges wa ming is es ima ed o he high moun ains in sou he n No way, and he in e io egions o Finland, Sweden and No way, which all a e impo an dai y p oduc ion a eas. The s onges ends in p ecipi a ion a e assumed in he egions o No way ha a e cha ac e ized by abundan sloping geog aphy. EFFECTS OF CLIMATE CHANGE ON LIVESTOCK PRODUCTION In he long un, and cu en ly o a ying ex en s, he clima e changes desc ibed will ha e a ious di ec and indi ec e ec s on li es ock p oduc ion (Na done e al., 2010). Ai empe a u e, humidi y, ai mo emen , and p ecipi a ion a e en i onmen al ac- o s ha a ec daily wea he condi ions and di ec ly a ec animal wel a e wi h he po en ial o c ea e hea s ess (Robinson, 2001; Na done e al., 2006). In he No dic coun ies, he uminan a m animal species (mainly small uminan s) g aze om sp ing o la e au umn ( eindee emain ou side all he ime, as do honeybees) and a e mo e subjec o he di ec e ec s o clima e change han he monogas ic species, o which a ming is mo e indus ial- ized. Animals can su e om occasional hea s ess du ing he summe season e en in no he n Eu ope. Ra agnolo e al. (2000) es ima ed, o example, ha when he empe a u e is +25°C and ela i e humidi y 50%, lac a ing cows a e ou side o hei op imal ambien empe a u e zone. When ela i e humidi y inc eases, he h eshold empe a u e dec eases. Se e al ene gy- equi ing phys- iological and me abolic unc ions, such as inc eased espi a ion, inc eased wa e in ake and educed eed in ake, a e needed o main ain op imal body empe a u e. These adap a ions, howe e , lead o lowe p oduc i i y and e ili y (Ra agnolo e al., 2000; De Rensis and Sca amuzzi, 2003; Wes , 2003; Na done e al., 2006). The e a e di e ences among species, among b eeds wi hin species, and among indi iduals wi hin b eeds ega ding hea s ess ole ance. The uminan s’ abili y o he mo egula e is ypically be e han ha o he monogas ics (Na done e al., 2006). In addi ion, mode n highly p oduc i e a m animal b eeds, which ypically show inc eased me abolic hea p oduc ion may ole a e ex eme clima ic condi ions less well han mode a e and low- ou pu b eeds (Na done e al., 2006; Ho man, 2010 and e e - ences he ein). Ra agnolo and Misz al (2002) showed ha he e is gene ic a ia ion among indi idual (Hols ein) cows in hei hea s ess sensi i i y, bo h wi h espec o milk yield and e ili y, and ha high-yielding cows we e mo e p one o dec ease hei p oduc ion when hea s essed. The hea s ess sensi i i y o milk yield was, howe e , no gene ically co ela ed o ha o e ili y, and he au ho s hypo hesize ha di e en me abolic and physiological p ocesses a e esponsible o hea ole ance o hese wo ai s. An al eady exis ing p oblem associa ed wi h clima e change ha has inc easingly un a o able e ec s on animal wel a e and li es ock p oduc ion is he occu ence and equency o animal diseases (Gale e al., 2009). Fo example, he sp ead o Blue ongue disease i us and Schmallenbe g i us is e iden ly associa ed wi h clima e change (Guis e al., 2012). Blue ongue disease, which is a i al disease in uminan s ansmi ed by bloodsucking midges (Culicoides spp.), has been ound in Denma k, No way and Swe- den, bu no cases o da e ha e been de ec ed in Finland and Iceland. Schmallenbe g i us has sp ead in all No dic coun ies excep Iceland2. The end is ha u u e ag oen i onmen s will be mo e a o able o se e al diseases han he p esen -day en i- onmen s. Global wa ming and incidence o ex eme me eo olog- ical e en s (d ough s and inc eased ain all) will c ea e, o may al eady ha e c ea ed, a o able mic oen i onmen s o a ious i uses, hei ec o species and ungal and bac e ial pa hogens. The densi y o insec ec o s may also inc ease as a esul o changes in annual beha io al cycles o mig a o y bi ds (Gale e al., 2009 and e e ences he ein). Global wa ming may shi iming when insec i o ous bi ds mig a e and nes , leading o he loss o synch ony be ween nes ing and peak ood abundance o mig a o y bi ds (Bo h e al., 2004). One addi ional challenge is ha pa hogens ypically ha e spe- ci ic cha ac e s ha allow apid sp ead. Fo example, RNA- i uses ha e a high mu a ion a e and can adap o new ci cums ances quickly (Du y e al., 2008; Gale e al., 2009 and e e ences he ein). New i al ec o -bo ne diseases may no necessa ily o igina e om close geog aphic egions bu may come om dis an egions, as he his o y o Blue ongue disease demons a es (Guis e al., 2012). In gene al, Culicoides spp. a e a majo h ea o animal wel a e by sp eading i uses ha cause se ious diseases (Gale e al., 2009). Midges can ansmi pa hogens and diseases o li es ock species om wild species, as exempli ied by Epizoo ic Hemo hagic Disease ha has sp ead om wild dee o li es ock (Sa ini e al., 2011). Midges a e no he only in e eb a es ha ec o many li es ock diseases. Ticks, mosqui oes and lymnaeid snails also ansmi ex emely ha m ul diseases o li es ock (Sco and Smi h, 1994; Randolph, 2009; Ca on e al., 2014). Mo eo e , i is expec ed ha he inc eased annual empe a u e, milde win e s and highe ain all will imp o e de elopmen al success o helmin h pa asi es, such as gas oin es inal nema odes and lukes, which will ha e mo e p onounced nega i e e ec s on he wel a e 2h p://www.no d isk.dk/ www. on ie sin.o g Feb ua y 2015 | Volume 6 | A icle 52 |3 Kan anen e al. Clima e change and animal gene ic esou ces o g azing ca le and sheep, and li es ock p oduc ion in gene al ( an Dijk e al., 2010). As a esul o clima e change, animal eeding s a egies in he No dic coun ies may need modi ying. Clima e change will ha e posi i e impac s on he “domes ic” p oduc ion o odde plan s in he No dic coun ies. Plan g ow h, yield and he p oduc- ion o c op and pas u e species will bene i om inc eases in a mosphe ic CO2concen a ion, a wa me clima e and a longe g owing season. Howe e , hese ag oclima ic changes will also b ing new challenges wi h he expansion o new weeds, insec pes s and plan diseases o he no he n Eu opean egions and p oblems wi h o e win e ing o pe ennial odde plan s (Tubiello e al., 2007; Hakala e al., 2011; Olesen e al., 2011; Höglind e al., 2013). Mo e chemical con ol in plan p o ec ion, esis an cul i a s and plan o a ions will be needed o o e come he nega i e e ec s o clima e change in plan p oduc ion (Cecca elli e al., 2010; Hakala e al., 2011). New a ie ies ha ole a e inc eased p ecipi a ion and annual odde plan s, such as maize (Zea mays L.), migh be commonly cul i a ed by he end o 21s Cen u y also in Scandina ia and Finland (Olesen e al., 2011). Howe e , he No dic cul i a ion adi ions o pe ennial o age g asses a e likely o con inue (a leas in he no he nmos and eas e n egions), such as pe ennial yeg ass (Lolium pe enne L.) and imo hy (Phleum p a ense L.) because pe ennial plan s ha a e ela i ely ole an o less op imal o e win e ing condi ions will be a o ed e en unde he changing ag oclima ic condi ions (Höglind e al., 2013). In he No dic coun ies, impo ed odde , mainly p o eins and ce eal concen a es, is impo an , pa icula ly in p oduc ion based on monogas ic species (Åby e al., 2014). Feed ading exis s bo h among EU-coun ies and non-Eu opean coun ies, e.g., B azil. I is sugges ed ha global wa ming and ex eme me eo ological e en s will dec ease c op yields and ag icul u al p oduc i i y in he sou he n coun ies, leading o educed a ail- abili y and inc eased p ices o g ains o animal eeds in he u u e (Wheele and Reynolds, 2013). This calls o imp o emen o sel -su iciency in odde p oduc ion in he No dic coun ies o u u e ood-secu i y. Such sel -su iciency can be imp o ed by using mo e e ilize s, pes con ol chemicals and o he inpu s in odde p oduc ion, and h ough plan b eeding and changes in land-use. Howe e , de o es a ion o new land o cul i a ion o odde plan s may ace a ious es ic ions owing o in e na ional poli ical ag eemen s and o en i onmen al easons. This may lead o he u iliza ion o less p oduc i e ma ginal land o odde p oduc ion and pas u es and could p o ide possibili ies o u ilize low-inpu b eeds and suppo hei conse a ion (Sæ he e al., 2006). In ecen yea s he end has been in he opposi e di ec ion and ewe pas u es han p e iously ha e been used o eed ca le (Åby e al., 2014). The socio-economic app oaches and subsidy policies should be de eloped in o de o make he use o low-inpu b eeds in animal p oduc ion a ealis ic op ion o a me s. I appea s ha highe yields o odde and pas u e plan s will lead o inc eased p o i abili y o animal p oduc ion in he No dic coun ies (Cisca e al., 2011). The No dic li es ock p oduc ion sys ems, howe e , ha e o cope wi h a ious challenging ci cum- s ances in he u u e, e.g., o imp o e sel -su iciency in odde p oduc ion, as well as o mi iga e ha m ul en i onmen al e ec s caused by hei p oduc ion. Li es ock p oduc ion is a subs an ial sou ce o GHG and he e is an u gen need o modi y he p o- duc ion sys ems. Di e si y in p oduc ion sys ems may inc ease, which calls o ma ching he geno ypes o each sys em. The use o a m animal gene ic esou ces and animal b eeding play a ole in his con ex in inding solu ions o new challenges and making li es ock p oduc ion mo e en i onmen ally iendly. CHARACTERIZATION OF ANIMALS’ CH4PRODUCTION The key issue associa ed wi h nega i e en i onmen al impac s induced by li es ock p oduc ion and mi iga ion o GHG e ec s is he educ ion o CH4emissions om uminan s, especially om bee and dai y ca le (Ma in e al., 2010; Wall e al., 2010). The e ha e been se e al me hods used o measu e CH4concen a ions, such as gas ch oma og aphy, mass spec oscopy, and a unable lase diode echnique (Johnson and Johnson, 1995). Cu en ly i is common o use au oma ic ad anced echnology based on in a ed de ec o s, ei he in espi a ion chambe s o wi h mo e ecen ly de eloped me hods in eeding s a ions wi h au oma ic milking obo s (e.g., Ga nswo hy e al., 2012; Lassen e al., 2012, 2014). The use o espi a ion chambe s gi es highly accu a e measu emen s, bu he capaci y is limi ed o a ew animals pe week. The eeding s a ion me hods a e less accu a e bu ha e highe capaci y, up o 60 cows pe week pe uni , making hem sui able o gene ic s udies a a pilo scale (Lassen e al., 2014). Mic obial e men a ion o eed in he umen p oduces sho - chain a y acids, such as ace a e, p opiona e and bu y a e, which a e used as he animal’s ene gy sou ce. This e men a ion esul s in high le els o en e ic CH4(Ma in e al., 2010). I should be poin ed ou ha manipula ion o eeding a ec ing umen mic obial popula ions is one o he main app oaches o dec easing he le els o CH4emissions (Boadi e al., 2004; Hook e al., 2010; Ma in e al., 2010). Fo example, inc easing he ene gy densi y o he die dec eases CH4p oduc ion pe uni o diges ible ene gy consumed (Ya es e al., 2000). Howe e , his would mean an inc ease in ce eals and o he high ene gy componen s in ca le eed a ions. This can be conside ed as unwan ed in e ms o esou ce u iliza ion in ood p oduc ion o a g owing human popula ion and would also mean ha he No dic coun ies become mo e dependen on impo ed eeds u . In addi ion o he manipula ion o he animals’ die s, selec i e b eeding wo k is he o he p incipal means used o mi iga e GHG emissions (Wall e al., 2010; B uce, 2013). Feeding expe imen s in ca le and sheep indica ed ha he e a e a ia ions among indi idual animals in he p oduc ion o CH4when hey a e ed he same die s. In addi ion, as e iewed by Wall e al. (2010), he e exis s a ia ion in CH4emissions among indi idual ca le and among b eeds, sugges ing po en ial o imp o emen o he ai h ough gene ic selec ion. Howe e , Ma in e al. (2010) we e less op imis ic; hey concluded ha epea abili y o he successi e measu emen s has been low in expe imen s and is hea ily dependen on die and physiological s age o he animals. Cha ac e iza ion o indi idual animal CH4emissions o gene ic selec ion is an u gen ma e . The COST-ac ion p ojec METHAGENE ocuses on he ha moniza ion o CH4measu e- men echniques and de elops app oaches o inco po a ing CH4 F on ie s in Gene ics | Li es ock Genomics Feb ua y 2015 | Volume 6 | A icle 52 |4 Kan anen e al. Clima e change and animal gene ic esou ces emissions in o na ional b eeding s a egies. Taking in o accoun ha me hane is a p oduc o umen mic obial e men a ion p ocesses ha a e di ec ly a ec ed by die , be e unde s anding o animal genome in e ac ion wi h own umen mic obiome unde a ious eeding condi ions needs o be aken in o accoun in d awing mi iga ion s a egies. These subjec s a e add essed in a numbe o na ional and in e na ional esea ch p ojec s (e.g., EU-FP7-p ojec RUMINOMICS; REMRUM in Denma k; Rumen Mic obial Genomics Ne wo k). Resul s a e expec ed om hese p ojec s in he nea u u e. CHARACTERIZATION OF ANIMALS’ ENVIRONMENTAL ADAPTATION I an animal popula ion su i es, is p oduc i e and ep oduces in a gi en en i onmen , we can say ha his popula ion com- p ises sui able, adap ed pheno ypes o ha en i onmen . The adap a ions, such as disease and hea esis ance, wa e sca ci y ole ance and abili y o cope wi h poo quali y eed, a e aluable cha ac e is ics o a b eed and ha e impo ance when mi iga ing and adap ing o en i onmen al changes (Ho man, 2010; Mi kena e al., 2010). B eeds can become adap ed o speci ic en i onmen s h ough na u al and a i icial selec ion. “Adap a ion ai s” a e complex and o en polygenically con olled (P i cha d e al., 2011). The in e ac ions be ween geno ypes and en i onmen s a e ypically examined in li es ock species using quan i a i e gene ics app oaches (Falcone and MacKay, 1996). Dense SNP-ma ke s and nex -gene a ion-sequencing (NGS) echnology can also be used o sea ch o adap a ion pa e ns and selec ion oo p in s in animal genomes ha esul om long- e m na u al and a i icial selec ion (Ha ison e al., 2012; Guo e al., 2014; L e al., 2014). Geno ype by en i onmen in e ac ion (G ×E) means ha geno ypes eac di e en ly o en i onmen al changes (Falcone and MacKay, 1996). Fo example, geno ype A can pe o m be e and display supe io i ness in high al i ude egions han geno ype B, while a sea le el geno ype B is he supe io pheno ype. O geno ype A pe o ms be e in bo h en i onmen s bu he di - e ence be ween he wo geno ypes is la ge in one en i onmen han in ano he . G ×E has been an ac i e esea ch ield in animal b eeding and quan i a i e gene ics. I he e is in o ma- ion a ailable on pe o mance o animals o e a wide ange o en i onmen s, i is possible o use a eac ion no m app oach o es ima ing b eeding alues. The eac ion no m can p edic he pe o mance o an indi idual in an en i onmen he animal has no been in (Calus e al., 2002; Kolmodin e al., 2002). Reac ion no ms ha e up ill now mainly been es ima ed using adi ional quan i a i e gene ics, bu he e is no heo e ical eason why hey could no also be es ima ed using molecula gene ic in o ma ion, which would p obably lead o g ea e accu acy in es ima ing b eeding alues o young animals (Sil a e al., 2014). F om a genomics poin o iew, adap a ions o animal b eeds o en i onmen s o die s a e ypically associa ed wi h s uc u al and unc ional genomic a ia ions (Axelsson e al., 2013; Li e al., 2013; Guo e al., 2014; L e al., 2014). Dense whole-genome SNP-chips and NGS applica ions, such as whole genome and mRNA sequencing, analysis o egula o y (miRNAs) elemen s, and DNA me hyla ion p o iles o epigene ic analysis, can be used o in es iga e gene ic backg ound o adap a ions in li es ock b eeds and species (Ba el, 2004; P i cha d e al., 2011; Feil and F aga, 2012; Ha ison e al., 2012; Jiang e al., 2014; Lee e al., 2014). Pai wise compa isons be ween closely ela ed axa ( o example b eeds o igina ing om di e en en i onmen s) p o ide a powe ul app oach o iden i ying loci ha show di e gence be ween popula ions and which may ha e been unde posi i e selec ion (Ha ison e al., 2012; Li e al., 2013; Jiang e al., 2014; L e al., 2014). The e is a body o di e en obus s a is ical and bioin o ma ics me hods o de ec ing selec ion signa u es (e.g., Beaumon and Balding, 2004; Joos e al., 2007; F icho e al., 2013; Wol , 2013 and many o he s) ha ha e been success ully used in genome-wide SNP and genomic sequence s udies (e.g., Guo e al., 2014; L e al., 2014). Measu es o CH4concen a ions om uminan s and he cha ac e iza ion o indi iduals and b eeds using mode n genomic, biome ical and bioin o ma ic ools play a pi o al ole in he implemen a ion o he s a egic p io i y a eas o GPA (FAO, 2007), hough he e is s ill a need o documen he ma ginal e ec o including CH4emissions in b eeding schemes selec ing o e iciency and p oduc i i y. Wi h his new in o ma ion we will unde s and be e cha ac e is ics o a m animal gene ic esou ces and can de elop animal b eeding and sus ainable u iliza ion o gene ic esou ces ha will make li es ock p oduc ion mo e en i- onmen al iendly. BREEDING GOALS CONSIDERING CLIMATE CHANGE Mi iga ion h ough selec ion e e s o b eeding animals ha ha e high p oduc i i y and e iciency, e ili y, good heal h, obus ness and ha p oduce less GHG (Boadi e al., 2004; Wall e al., 2010; B uce, 2013; Hie ala e al., 2014). The b eeding goals o adap- a ion a e e y simila o hose o mi iga ion: in adap a ion o new en i onmen al ci cums ances and p oduc ion en i onmen s, we conside ha e ili y, eed con e sa ion a e and pa icula ly heal h ai s, a e e y impo an . As poin ed ou in se e al p e- ious pape s, he imp o emen in p oduc i i y (highe a e age milk and mea yields e c.) means ewe emissions pe p oduc . In addi ion, ewe animals a e needed o mee he demand o animal p oduc s (e.g., Boadi e al., 2004; Wall e al., 2010; B uce, 2013). Imp o ing e ili y, on he o he hand, means sho e unp oduc i e pe iods, and imp o ing cal ing and ma e nal ai s, diminishing emissions by imp o ing su i al o o sp ing. Majo p oduc ion ai s such as eed con e sion a e, e ili y, heal h and o he i ness ai s ha e been shown o ha e a gene ic componen , demons a ing ha he e a e possibili ies o imp o e hem ia selec ion. The No dic b eeding p og ams ha e ypically b oad b eeding goals and bo h p oduc ion and heal h cha ac e s a e conside ed (e.g., Miglio e al., 2005; Åby e al., 2013; Hie ala e al., 2014). T ai s impo an o mi iga ion and adap a ion a e ypically ei he di ec ly o indi ec ly conside ed in he No dic mul i ai b eeding schemes, which makes i easie o b eed animals ha a e needed o u u e li es ock p oduc ion (Åby e al., 2013). Cu en ly, e - ili y, heal h and o he i ness and unc ional ai s ha e ecei ed mo e a en ion in b eeding goals han p e iously (e.g., Hie ala e al., 2014). This end can be conside ed highly ecommendable because, o example, bo h e ili y and heal h ai s o dai y ca le www. on ie sin.o g Feb ua y 2015 | Volume 6 | A icle 52 |5 Kan anen e al. Clima e change and animal gene ic esou ces (and se e al o he a m animal species) ha e de e io a ed, espe- cially in popula ions whe e he ai s ha e no been conside ed in he o al b eeding alues (Lucy, 2001; Miglio e al., 2005). Good e ili y, e.g., in dai y cows, is known o co ela e nega i ely wi h gene ic me i o milk p oduc ion (Rauw e al., 1998; an de Waaij, 2004). One op ion o including CH4p oduc ion in a u u e b eeding p og am is o ca y ou di ec selec ion on he ai . Howe e , o do his, he e is a need o pheno ypic eco ding o di ec mea- su emen s o he ai in many uminan s in se e al he ds in o de o c ea e a e e ence popula ion o es ima e genomic b eeding alues (Hansen Axelsson e al., 2013, 2015). Fo his o happen he e is a need o de elop be e and cheape measu emen echniques (Hansen Axelsson e al., 2013). Mo e esea ch is in p og ess in his ield and some o i is suppo ed by he EU-COST p ojec METHAGENE. In he mean ime, we can imp o e he ai indi ec ly h ough selec ion o p oxy ai s ha a e co ela ed wi h CH4emissions pe uni o p oduc (e.g., milk yield, e ili y, eed e iciency, and longe i y o he animals; Cappe e al., 2009; B uce, 2013; Hie ala e al., 2014). Mo eo e , wi h he ad en o ela i ely cheap SNP-chips wi h ens o hund eds o housands o ma ke s, i is also possible o es ima e genomic b eeding alues o animals ha ha e no hemsel es, no hei close ela i es, li ed and p oduced in he en i onmen whe e hey o hei o sp ing a e expec ed o li e. S a ed di e en ly, i would be possible o ind ma ke s ha a e associa ed wi h pe o mance in condi ions ha we belie e we will ha e in he No dic coun ies in he coming decades i we can gene ically e alua e animals ha cu en ly li e unde such condi ions elsewhe e. AVAILABLE ANIMAL GENETIC RESOURCES Cu en ly he e a e h ee ypes o li es ock b eed a ailable in he No dic coun ies o u u e selec ion p og ams: (1) he majo comme cial b eeds, (2) he mino b eeds, which a e ypically na i e b eeds and ha a e also used in comme cial he ds bu mo e ypically in special p oduc ion si ua ions, and (3) endange ed b eeds, which a e also na i e b eeds and kep o ec ea ional pu poses and a ely o p oduc ion pu poses. The majo b eeds domina e p oduc ion sys ems and hey may possess impo an wi hin-b eed gene ic a ia ion o selec o adap a ion o new ag o-ecological condi ions and mi iga ion o ha m ul e ec s o animal p oduc ion on clima e change (e.g., Gomes da Sil a, 1973). I is e y impo an ha hese b eeds do no un in o inb eeding p oblems, o he wise in e io al e na i e b eeds, i hey s ill exis a ha ime, will ha e o be in oduced in o he p oduc ion sys em. Inb eeding p oblems ha e o be a oided also o he mino and endange ed b eeds in o de o main ain iabili y o e many u u e gene a ions. Long- e m selec ion expe imen s ha e shown ha managed popula ions can be sus ained wi hou signi ican loss o gene ic a ia ion o mo e han 100 gene a ions when he e ec i e popula ion size is main ained a 100 o mo e (Hill, 2000). Howe e , he e ec i e popula ion size o majo comme cial b eeds is ypically much less han 100 (Kan anen e al., 1999; Tabe le e al., 2008). Op imal con ibu ion heo y p o ides a amewo k o maximiz- ing esponse o selec ion while con olling he e ec i e popula- ion size (Meuwissen, 1997). So wa e o op imum con ibu ion selec ion exis s, bu imp o emen s a e needed in o de o add ess he di e en si ua ions ha occu in p ac ical b eeding schemes. The na i e b eeds ha e he longes adap a ion his o y o No dic en i onmen al and p oduc ion condi ions. These b eeds a e based on ancien animal popula ions ha sp ead o no h- e n Eu ope housands o yea s ago when he ansi ion om hun ing- ishing-ga he ing li elihoods o animal a ming and cul- i a ion began (Kan anen e al., 2000; Bläue and Kan anen, 2013; Niemi e al., 2013). The e o e, we a gue ha he No dic na i e b eeds, which ypically a e mino and endange ed b eeds, may possess s uc u al and unc ional genomic a ia ions o speci ic ai s, such as disease esis ances. Fo example, he na i e Finnca le display a high le el o polymo phism in he Majo -His ocompa ibili y-Complex sys em (mo e speci ically a he BoLA-DRB3 locus) ha con ols a majo pa o he immune sys em (Kos ia, 2000). The No dic na i e ca le b eeds exhibi allelic combina ions in he casein loci ha ha e a posi i e impac on p ocessing p ope ies o milk (Lien e al., 1999). In addi- ion, he e a e se e al anecdo es abou adap i e cha ac e s o na i e b eeds ha should be scien i ically s udied and c i ically e alua ed. Due o clima ic changes, he comme cial and widesp ead b eeds may show sho comings in some ai s, such as insu icien esis ance o a new disease o ole ance o o he en i onmen al s ess (Na done e al., 2006; Ho man, 2010). The mino and endange ed b eeds may possess genes ha code o speci ic ai s, such as disease esis ances, which may become desi ed by he majo b eed owne s, bu o which he majo b eed does no possess he necessa y gene ic a ia ion. Howe e , he majo b eeds can be selec ed o any desi ed ai , jus as he mino o endan- ge ed b eeds we e once selec ed o his ai , bu i may ake many gene a ions o es ablish he desi ed ai in he majo b eed. The majo b eed could bene i om alleles a ailable in mino and endange ed b eeds using c ossing and genomic in og ession, and genomic ma ke in o ma ion o in og ess a o able alleles, while keeping a o able alleles o p oduc ion ai s in he majo b eed (Ødegå d e al., 2009). I he ai is due o a single o a ew genes, such genes can be mapped and be in og essed in o he comme cial b eed (Ødegå d e al., 2009). Al hough o en success ul in plan b eeding, his app oach is o en no easible o li es ock because mos li es ock ai s a e complex, i.e., highly polygenic, and in og ession akes ∼5 gene a ions, which in li e- s ock migh easily be 10 yea s o mo e. C ossb eeding sys ems can be de ised ha a leas pa ly con ey he desi ed ai om he a e in o he comme cial b eed. Al e na i ely, a Genomic Selec ion In og ession app oach can be employed (Ødegå d e al., 2009), whe e genomic selec ion is applied o a apid in oduc ion o a new ai in he comme cial b eed. In his p ocess, mino b eeds ha possess he ai ep esen a much mo e use ul esou ce han he endange ed b eeds because when c ossed wi h he majo b eed, hei o sp ing combine he desi ed ai wi h comme cial iabili y (since bo h pa en al b eeds a e comme cially iable). Examples o his si ua ion a e he use o No dic ed bulls on US-Hols ein cows o imp o e hei e ili y (conside ing No dic Reds as a mino b eed a he global ca le b eeding scale), he use o Chinese Meishan pigs o imp o e F on ie s in Gene ics | Li es ock Genomics Feb ua y 2015 | Volume 6 | A icle 52 |6 Kan anen e al. Clima e change and animal gene ic esou ces e ili y in some Eu opean pig b eeding p og ams and c ossing less e ile sheep b eeds wi h he highly p oli ic Finnsheep. In gene al, mino and endange ed b eeds ep esen a aluable esou ce o comme cial b eeding schemes o inc ease he a e a which desi able ai s can be es ablished in majo b eeds. Thus, o add ess u u e un o eseen p oduc ion challenges such as clima ic changes, which equi e new, desi ed ai s in majo b eeds, i is impo an o main ain a la ge numbe o mino b eeds ha a e imp o ed o specialized p oduc ion en i onmen s, and, o a lesse ex en , in endange ed b eeds. In all No dic coun ies he e a e na ional s a egies o conse e bo h in i o and in i o na i e b eeds and hei gene ic esou ces. Howe e , hese s a egies should be e ised, e.g., by conside ing he geog aphic dis ibu ion o a e b eeds wi hin he coun ies and s eng hening c yop ese a ion o gene ic ma e ials. Se e al na i e b eeds exis in ela i ely es ic ed local a eas and in he ou b eaks o se ious animal diseases he whole b eed o mos o i can be los . No dic b eeds ha e been p e iously analyzed o neu al gene ic ma ke s (e.g., Tapio e al., 2006, 2010; Li e al., 2007; Kan anen e al., 2009), bu mo e cha ac e iza ion o conse a ion alues and adap a ions is needed in o de o p omo e e icien use o gene ic esou ces in he u u e. In he AnG -No dicNET p ojec , a new measu e o aluing b eeds o conse a ion, e med “adap i i y co e age” has been de eloped (Wellman e al., 2014). This quan i ies how well a se o b eeds could be adap ed o wide ange o en i on- men s wi hin a limi ed imespan. In his quan i ica ion, adap- i i y co e age conside s bo h neu al and non-neu al gene ic a ia ion. CONCLUSION The No dic mul i ai b eeding p og ams o se e al animal b eeds conside di ec ly o indi ec ly ai s ha a e impo an o mi iga ion o en i onmen al e ec s o li es ock p oduc ion o ad ance animals’ adap a ion o new ag oecological condi ions. The impo an ai s in his con ex a e, o example, p oduc i i y in gene al, e ili y, eed con e sa ion a e and heal h. How- e e , e ili y, heal h and o he i ness ai s should ecei e mo e weigh and alue in animal b eeding o s eng hen adap a ion po en ial. Mo eo e , he b eeding p og ams should main ain high e ec i e popula ion sizes in o de o keep high gene ic a ia- ion in majo and mino b eeds. Including CH4p oduc ion o uminan s as a ai in b eeding p og ams needs mo e esea ch and he de elopmen o be e and cheape CH4measu emen echniques. In he u u e genomic selec ion and genomic selec ion in og ession app oaches may play pi o al oles, pa icula ly in “adap a ion b eeding.” Valuable alleles in e ms o adap a ion o clima e change can be in oduced in o majo b eeds om conse ed na i e b eeds h ough genomic selec ion in og ession b eeding. The e o e, in i o and in i o conse a ion o mino and endange ed b eeds, which a e ypically na i e b eeds, should be s eng hened and hei adap a ion ai s in es iga ed using mode n genomic and bioin o ma ics ools. AUTHOR CONTRIBUTIONS All au ho s ha e designed he e iew pape , d a ed he manusc ip and e ised i c i ically. All au ho s ha e app o ed he e sion o be published. ACKNOWLEDGMENTS The unding gi en by No dic Council o Minis e s, No dFo sk and No dGen—No dic Gene ic Resou ce Cen e is g ea ly acknowl- edged. REFERENCES Åby, B. A., Aass, L., Sehes ed, E., and Vangen, O. (2013). 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Con lic o In e es S a emen : The au ho s decla e ha he esea ch was con- duc ed in he absence o any comme cial o inancial ela ionships ha could be cons ued as a po en ial con lic o in e es . Recei ed: 31 Oc obe 2014; accep ed: 05 Feb ua y 2015; published online: 25 Feb ua y 2015. Ci a ion: Kan anen J, Lø endahl P, S andbe g E, Ey ho sdo i E, Li M-H, Ke unen- P æbel A, Be g P and Meuwissen T (2015) U iliza ion o a m animal gene ic esou ces in a changing ag o-ecological en i onmen in he No dic coun ies. F on . Gene . 6:52. doi: 10.3389/ gene.2015.00052 This a icle was submi ed o Li es ock Genomics, a sec ion o he jou nal F on ie s in Gene ics. Copy igh ©2015 Kan anen, Lø endahl, S andbe g, Ey ho sdo i , Li, Ke unen- P æbel, Be g and Meuwissen. This is an open-access a icle dis ibu ed unde he e ms o he C ea i e Commons A ibu ion License (CC BY). The use, dis ibu ion o ep oduc ion in o he o ums is pe mi ed, p o ided he o iginal au ho (s) o licenso a e c edi ed and ha he o iginal publica ion in his jou nal is ci ed, in acco dance wi h accep ed academic p ac ice. No use, dis ibu ion o ep oduc ion is pe mi ed which does no comply wi h hese e ms. www. on ie sin.o g Feb ua y 2015 | Volume 6 | A icle 52 |9