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). E ec o inco po a ing
g eenhouse gas emission cos s in o economic alues o ai s o in ensi e and
ex ensi e bee ca le b eeds. Li es . Sci. 158, 1–11. doi: 10.1016/j.li sci.2013.
09.009
Åby, B. A., Kan anen, J., and Meuwissen, T. (2014). Fu u e ends in li es ock
p oduc ion in he No dic coun ies wi h a changing clima e and human popula-
ion g ow h. Ac a Ag ic. Scand. A Anim. 64, 73–79. doi: 10.1080/09064702.2014.
950321
Axelsson, E., Ra nakuma , A., A end , M.-L., Maqbool, K., Webs e , M. T., Pe loski,
M., e al. (2013). The genomic signa u e o dog domes ica ion e eals adap a ion
o a s a ch- ich die . Na u e 495, 360–365. doi: 10.1038/na u e11837
Ba el, D. P. (2004). Mic oRNAs: genomics, biogenesis, mechanism, and unc ion.
Cell 116, 281–297. doi: 10.1016/S0092-8674(04)00045-5
Ba ua, S. K., Be g, P., B u oll, A., Cede be g, C., D inkwa e , K. F., Eide, A.,
e al. (2014). Clima e Change and P ima y Indus ies. Impac s, Adap a ion and
Mi iga ion in he No dic Coun ies (TemaNo d 2014: 552). Copenhagen: No dic
Council o Minis e s. doi: 10.6027/TN2014-552
Beaumon , M. A., and Balding, D. J. (2004). Iden i ying adap i e gene ic di e gence
among popula ions om genome scans. Mol. Ecol. 13, 969–980. doi: 10.1111/
j.1365-294X.2004.02125.x
Benes ad, R. E. (2005). Clima e change scena ios o no he n Eu ope om mul i-
model IPCC AR4 clima e simula ions. Geophys. Res. Le . 32, L17704. doi:
10.1029/2005GL023401
Be ns ein, L., Bosch, P., Canziani, O., Chen, Z., Ch is , R., Da idson, O., e al.
(2007). IPCC, 2007: Clima e Change 2007: Syn hesis Repo . Con ibu ion o
Wo king G oups I, II, and III o he Fou h Assessmen Repo o he In e go e n-
men al Panel on Clima e Change. Gene a: In e go e nmen al Panel on Clima e
Change, 52 p. +Appendix.
Bläue , A., and Kan anen, J. (2013). T ansi ion om hun ing o animal husband y
in Sou he n, Wes e n and Eas e n Finland: new da ed os eological e idence.
J. A chaeol. Sci. 40, 1646–1666. doi: 10.1016/j.jas.2012.10.033
Boadi, D., Benchaa , C., Chique e, J., and Massé, D. (2004). Mi iga ion s a egies o
educe en e ic me hane emissions om dai y cows: upda e e iew. Can. J. Anim.
Sci. 84, 319–335. doi: 10.4141/A03-109
Bo h, C., A emye , A. V., Blaauw, B., Cowie, R., J., Dekhuijzen, A. J., Ee a, T., e al.
(2004). La ge-scale geog aphical a ia ion con i ms ha clima e change causes
bi ds o lay ea lie . P oc. R. Soc. Lond. B 271, 1657–1662. doi: 10.1098/ spb.2004.
2770
B uce, A. (2013). The lo e o low me hane li es ock: co-p oducing echnology
and animals o educed clima e change impac . Li e Sci. Soc. Policy 9, 10. doi:
10.1186/2195-7819-9-10
Calus, M., G oen, A. F., and De Jong, G. (2002). Geno ype ×en i onmen
in e ac ion o p o ein yield in Du ch dai y ca le as quan i ied by di e en
models. J. Dai y Sci. 85, 3115–3123. doi: 10.3168/jds.S0022-0302(02)74399-3
Cappe , J. L., Cady, R. A., and Bauman, D. E. (2009). The en i onmen al impac o
dai y p oduc ion: 1944 compa ed wi h 2007. J. Anim. Sci. 87, 2160–2167. doi:
10.2527/jas.2009-1781
Ca on, Y., Ma ens, K., Lempe eu , L., Saege man, C., and Losson, B. (2014).
New insigh in lymnaeid snails (Mollusca, Gas opoda) as in e media e hos s
o Fasciola hepa ica (T ema oda, Digenea) in Belgium and Luxembou g. Pa asi .
Vec o s 7:66. doi: 10.1186/1756-3305-7-66
Cecca elli, S., G ando, S., Maa ougui, M., Michael, M., Slash, M., Haghpa as , R.,
e al. (2010). Plan b eeding and clima e change. J. Ag ic. Sci. 148, 627–637. doi:
10.1017/S0021859610000651
Cisca , J.-C., Iglesias, A., Feyen, L., Szabó, L., Van Regemo e , D., Amelung B., e al.
(2011). Physical and economic consequences o clima e change in Eu ope. PNAS
108, 2678–2683. doi: 10.1073/pnas.1011612108
De Rensis, F., and Sca amuzzi, R. J. (2003). Hea s ess and seasonal e ec s on
ep oduc ion in he dai y cow—a e iew. The iogenology 60, 1139–1151. doi:
10.1016/S0093-691X(03)00126-2
Delgado, C. L. (2003). Rising consump ion o mea and milk in de eloping
coun ies has c ea ed a new ood e olu ion. J. Nu . 133, 3907S–3910S.
www. on ie sin.o g Feb ua y 2015 | Volume 6 | A icle 52 |7
Kan anen e al. Clima e change and animal gene ic esou ces
Du y, S., Shackel on, L. A., and Holmes, E. C. (2008). Ra es o e olu iona y
change in i uses: pa e ns and de e minan s. Na . Re . Gene . 9, 267–276. doi:
10.1038/n g2323
Falcone , D. S., and MacKay, T. F. C. (1996). In oduc ion o Quan i a i e Gene ics.
Ha low: Longman G oup L da.
FAO. (2007). Global Plan o Ac ion o Animal Gene ic Resou ces and In e laken
Decla a ion. Rome: Food and Ag icul u e O ganiza ion o he Uni ed Na ions.
Feil, R., and F aga, M. F. (2012). Epigene ics and he en i onmen : eme ging
pa e ns and implica ions. Na . Re . Gene . 13, 97–109. doi: 10.1038/n g3142
F icho , E., Scho ille, S. D., Boucha d, G., and F ancois, O. (2013). Tes ing o
associa ions be ween loci and en i onmen al g adien s using la en ac o mixed
models. Mol. Biol. E ol. 30, 1687–1699. doi: 10.1093/molbe /ms 063
Gale, P., D ew, T., Phipps L. P., Da id, G., and Woold idge, M. (2009). The e ec
o clima e change on he occu ence and p e alence o li es ock diseases in
G ea B i ain: a e iew. J. Appl. Mic obiol. 106, 1409–1423. doi: 10.1111/j.1365-
2672.2008.04036.x
Ga ne , T. (2011). Whe e a e he bes oppo uni ies o educing g eenhouse gas
emissions in he ood sys em (including he ood chain)? Food Policy 36, S23–
S32. doi: 10.1016/j. oodpol.2010.10.010
Ga nswo hy, P. C., C aigon, J., He nandez-Med ano, J. H., and Saunde s, N.
(2012). Va ia ion among indi idual dai y cows in me hane measu emen s made
on a m du ing milking. J. Dai y Sci. 95, 3181–3189. doi: 10.3168/jds.2011-4606
Ge be , P., Key, N., Po e , F., and S ein eld, H. (2010). Policy op ions in add essing
li es ock’s con ibu ion o clima e change. Animal 4, 393–406. doi: 10.1017/
S1751731110000133
Ge land, P., Ra e y, A. E., Še ˇ
cíko á, H., Li, N., Gu, D., and Spoo enbe g, T. (2014).
Wo ld popula ion s abiliza ion unlikely his cen u y. Science 346, 234–237. doi:
10.1126/science.1257469
Gill, M., Smi h, P., and Wilkinson, J. M. (2010). Mi iga ing clima e change: he ole
o domes ic li es ock. Animal 4, 323–333. doi: 10.1017/S1751731109004662
Gomes da Sil a, R. (1973). Imp o ing opical bee ca le by simul aneous selec ion
o weigh and hea ole ance. He i abili ies and co ela ions o he ai . J. Anim.
Sci. 37, 637–642.
Guis, H., Caminade, C., Cal e e, C., Mo se, A. P., T an, A., and Baylis, M. (2012).
Modelling he e ec s o pas and u u e clima e on he isk o blue ongue
eme gence in Eu ope. J. R. Soc. In e ace 9, 339–350. doi: 10.1098/ si .2011.0255
Guo, X., Wang, Z., Li, N., Qiu, F., Xu, Z., Yan, D., e al. (2014). Whole genome
sequencing o six dog b eeds om con inuous al i udes e eals adap a ion o
high-al i ude hypoxia. Gen. Res. 24, 1308–1315. doi: 10.1101/g .171876.113
Hakala, K., Hannukkala, A. O., Huusela-Veis ola, E., Jalli, M., and Pel onen-Sainio,
P. (2011). Pes s and diseases in a changing clima e: a majo challenge o Finnish
c op p oduc ion. Ag ic. Food Sci. 20, 3–14. doi: 10.2137/145960611795163042
Hansen Axelsson, H., Fikse, W. F., Ka go, M., Sø ensen, A. C., Johansson, K., and
Rydhme , L. (2013). Genomic selec ion using indica o ai s o educe he en i-
onmen al impac o milk p oduc ion. J. Dai y Sci. 96, 5306–5314. doi: 10.3168/
jds.2012-6041
Hansen Axelsson, H., Thomasen, J. R., Sø ensen, A. C., Rydhme , L., Ka go, M.,
Johansson, K., e al. (2015). B eake en p ices o eco ding o indica o ai s
o educe he en i onmen al impac o milk p oduc ion. J. Anim. B eed. Gene .
132, 30–41. doi: 10.1111/jbg.12106
Ha ison, P. W., W igh , A. E., and Mank, J. E. (2012). The e olu ion o gene
exp ession and he ansc ip ome-pheno ype ela ionships. Semin. Cell De .
Biol. 23, 222–229. doi: 10.1016/j.semcdb.2011.12.004
Hie ala, P., Wol o á, M., Wol , J., Kan anen, J., and Juga, J. (2014). Economic alues
o p oduc ion and unc ional ai s, including esidual eed in ake, in Finnish
milk p oduc ion. J. Dai y Sci. 97, 1092–1106. doi: 10.3168/jds.2013-7085
Hill, W. G. (2000). Main enance o quan i a i e gene ic a ia ion in animal b eed-
ing p og ammes. Li es . P od. Sci. 63, 99–109. doi: 10.1016/S0301-6226(99)
00115-3
Ho man, I. (2010). Clima e change and he cha ac e iza ion, b eeding and con-
se a ion o animal gene ic esou ces. Anim. Gene . 41(Suppl. 1), 32–46. doi:
10.1111/j.1365-2052.2010.02043.x
Höglind, M., Tho sen, S. M., and Semeno , M. A. (2013). Assessing unce ain ies
in impac o clima e change on c ass p oduc ion in No he n Eu ope using
ensembles o global clima e models. Ag ic. Fo es Me eo ol. 170, 103–113. doi:
10.1016/j.ag o me .2012.02.010
Hook, S. E., W igh , A. G., and McB ide, B. W. (2010). Me hanogens: me hane
p oduce s o he umen and mi iga ion s a egies. A chaea 2010, 945785. doi:
10.1155/2010/945785
Howden, S. M., Soussana, J.-F., Tubiello, F. N., Chhe i, N., Dunlop, M., and
Meinke, H. (2007). Adap ing ag icul u e o clima e change. PNAS 104, 19691–
19696. doi: 10.1073/pnas.0701890104
Jiang, Y., Xie, M., Chen, W., Talbo , R., Maddox, J., Fa au , T., e al. (2014). The
sheep genome illumina es biology o he umen and lipid me abolism. Science
344, 1168–1173. doi: 10.1126/science.1252806
Johnson, K. A., and Johnson, D. E. (1995). Me hane emissions om ca le. J. Anim.
Sci. 73, 2483–2492.
Joos , S., Bonin, M. W., B u o d, M., Desp és, L., Cono d, C., E ha d , G., e al.
(2007). A spa ial analysis me hod (SAM) o de ec candida e loci o selec ion:
owa ds a landscape genomics app oach o adap a ion. Mol. Ecol. 16, 3955–3969.
doi: 10.1111/j.1365-294X.2007.03442.x
Jylhä, K., F onzek, S., Tuomen i a, H., Ca e , T. R., and Ruos eenoja, K. (2008).
Changes in os , snow and Bal ic Sea ice by he end o he wen y- i s cen u y
based on clima e model p ojec ions o Eu ope. Clim. Change 86, 441–462. doi:
10.1007/s10584-007-9310-z
Kan anen, J., Edwa ds, C. J., B adley, D. G., Viinalass, H., Thessle , S., I ano a, Z.,
e al. (2009). Ma e nal and pa e nal genealogy o Eu asian au ine ca le (Bos
au us). He edi y 103, 404–415. doi: 10.1038/hdy.2009.68
Kan anen, J., Olsake , I., Adals einsson, S., Sandbe g, K., Ey ho sdo i , E.,
Pi honen, K., e al. (1999). Tempo al changes in gene ic a ia ion o No h
Eu opean ca le b eeds. Anim. Gene . 30, 16–27. doi: 10.1046/j.1365-2052.1999.
00379.x
Kan anen, J., Olsake , I., Holm, L.-E., Lien, S., Vilkki, J., B usgaa d, K., e al. (2000).
Gene ic di e si y and popula ion s uc u e o 20 No h Eu opean ca le b eeds.
J. He ed. 91, 446–457. doi: 10.1093/jhe ed/91.6.446
Ka l, T. R., and T enbe h, K. E. (2003). Mode n global clima e change. Science 302,
1719–1723. doi: 10.1126/science.1090228
Kolmodin, R., S andbe g, E., Madsen, P., Jensen, J., and Jo jani, H. (2002). Geno-
ype by en i onmen in e ac ion in No dic dai y ca le s udied by use o eac ion
no ms. Ac a Ag ic. Scand. A Anim. 52, 11–24. doi: 10.1080/09064700252806380
Kos ia, S. (2000). Genomic E olu ion and Di e si y in A iodac yla. Ph.D. he-
sis, Depa men o Biosciences, Di ision o Gene ics, Uni e si y o Helsinki,
Helsinki. 41.
Lassen, J., Ga nswo hy, P. C., Chagunda, M. G. G., Negussie, E., Lø endahl, P.,
and de Haas, Y. (2014). “P og ess wi h gene ic selec ion o low me hane ai s
in dai y cows,” in P oceedings, 10 h Wo ld Cong ess o Gene ics Applied o Li e-
s ock P oduc ion, Vancou e . A ailable a : h ps://asas.o g/docs/de aul -sou ce/
wcgalp-p oceedings-o al/036_pape _9384_manusc ip _600_0.pd ?s sn=2
Lassen, J., Lø endahl, P., and Madsen, J. (2012). Accu acy o nonin asi e b ea h
me hane measu emen s using Fou ie ans o m in a ed me hods on indi id-
ual cows. J. Dai y Sci. 95, 890–898. doi: 10.3168/jds.2011-4544
Lee, J.-R., Hong, C. P., Moon, J.-W., Jung, Y.-D., Kim, D.-S., Kim, T.-H., e al.
(2014). Genome-wide analysis o DNA me hyla ion pa e ns in ho se. BMC
Genom. 15:598. doi: 10.1186/1471-2164-15-598
Li, M.-H., Tapio, I., Vilkki, J., I ano a, Z., Kiselyo a, T., Ma zano , N., e al. (2007).
Gene ic s uc u e o ca le popula ions (Bos au us) in no he n Eu asia and
he neighbo ing Nea Eas e n egions: implica ions o b eeding s a egies and
conse a ion. Mol. Ecol. 16, 3839–3853. doi: 10.1111/j.1365-294X.2007.03437.x
Li, M., Tian, S., Jin, L., Zhou, G., Li, Y., Zhang, Y., e al. (2013). Genomic analyses
iden i y dis inc pa e ns o selec ion in domes ica ed pigs and Tibe an wild
boa s. Science 45, 1431–1439. doi: 10.1038/ng.2811
Lien, S., Kan anen, J., Olsake , I., Holm, L.-E., Ey ho sdo i , E., Sandbe g, K., e al.
(1999). Compa ison o milk p o ein allele equencies in No dic ca le b eeds.
Anim. Gene . 30, 85–91. doi: 10.1046/j.1365-2052.1999.00434.x
Lucy, M. C. (2001). Rep oduc i e loss in high-p oducing dai y ca le: whe e will i
end? J. Dai y Sci. 84, 1277–1293. doi: 10.3168/jds.S0022-0302(01)70158-0
L , F.-H., Agha, S., Kan anen, J., Colli, L., S ucki, S., Kijas, J. W., e al. (2014).
Adap a ions o clima e-media ed selec i e p essu es in sheep. Mol. Biol. E ol.
31, 3324–3343. doi: 10.1093/molbe /msu264
Ma in, C., Mo ga i, D. P., and Do eau, M. (2010). Me hane mi iga ion in
uminan s: om mic obe o he a m scale. Animal 4, 351–365. doi: 10.1017/
S1751731109990620
McMichael, A. J., Powles, J. W., Bu e , C. D., and Uauy, R. (2007). Food, li es ock
p oduc ion, ene gy, clima e change, and heal h. Lance 370, 1253–1263. doi:
10.1016/S0140-6736(07)61256-2
Meehl, G. A., S ocke , T. F., Collins, W. D., F iedlings ein, P., Gaye, A. T., G ego y,
J. M., e al. (2007). “Global clima e p ojec ions,” in Clima e Change 2007: The
Physical Science Basis. Con ibu ion o Wo king G oup I o he Fou h Assessmen
F on ie s in Gene ics | Li es ock Genomics Feb ua y 2015 | Volume 6 | A icle 52 |8
Kan anen e al. Clima e change and animal gene ic esou ces
Repo o he In e go e nmen al Panel on Clima e Change, eds S. Solomon,
D. Qin, M. Manning, Z. Chen, M. Ma quis, K. B. A e y , e al. (Camb idge:
Camb idge Uni e si y P ess), 747–845.
Meuwissen, T. (1997). Maximizing he esponse o selec ion wi h a p ede ined a e
o inb eeding. J. Anim. Sci. 75, 934–940.
Miglio , F., Mui , B. L., and Van Doo maal, B. J. (2005). Selec ion indices in hols ein
ca le o a ious coun ies. J. Dai y Sci. 88, 1255–1263. doi: 10.3168/jds.S0022-
0302(05)72792-2
Mi kena, T., Duguma, G., Haile, A., Tibbo, M., Okeyo, A. M., Wu zinge , M., e al.
(2010). Gene ics o adap a ion in domes ic a m animals: a e iew. Li es . Sci.
132, 1–12. doi: 10.1016/j.li sci.2010.05.003
Mi loehne , F. M. (2010). Is he ising demand o animal p o ein uelling clima e
change? J. Anim. B eed. Gene . 127, 421–422. doi: 10.1111/j.1439-0388.2010.
00909.x
Na done, A., Ronchi, B., Lace e a, N., and Be nabucci, U. (2006). Clima ic e ec s
on p oduc i e ai s in li es ock. Ve . Res. Commun. 30(Suppl. 1), 75–81. doi:
10.1007/s11259-006-0016-x
Na done, A., Ronchi, B., Lace e a, N., Ranie i, M. S., and Be nabucci, U. (2010).
E ec s o clima e changes on animal p oduc ion and sus ainabili y o li es ock
sys ems. Li es . Sci. 130, 57–69. doi: 10.1016/j.li sci.2010.02.011
Niemi, J., and Ahls ed , J. (eds). (2014). Finnish Ag icul u e and Ru al Indus ies
2014. Jokioinen: MTT Economic Resea ch 115a.
Niemi, M., Bläue , A., Iso-Tou u, T., Nys öm, V., Ha jula, J., Taa i sainen, J.-P.,
e al. (2013). Mi ochond ial DNA and Y-ch omosomal di e si y in ancien
popula ions o domes ic sheep (O is a ies) in Finland: compa ison wi h
con empo a y sheep b eeds. Gene . Sel. E ol. 45, 2. doi: 10.1186/1297-9686
-45-2
Ødegå d, J., Sonesson, A. K., Yazdi, M. H., and Meuwissen, T. H. E. (2009).
In og ession o a majo QTL om an in e io in o a supe io popula ion using
genomic selec ion. Gene . Sel. E ol. 41, 38. doi: 10.1186/1297-9686-41-38
Olesen, J. E., and Bindi, M. (2002). Consequences o clima e change o Eu opean
ag icul u al p oduc i i y, land use and policy. Eu . J. Ag on. 16, 239–262. doi:
10.1016/S1161-0301(02)00004-7
Olesen, J. E., T nka, M., Ke sebaum, K. C., Skjel åg, A. O., Sequin, B., Pel onen-
Sainio, P., e al. (2011). Impac s and adap a ion o Eu opean c op p oduc ion
sys ems o clima e change. Eu . J. Ag on. 34, 96–112. doi: 10.1016/j.eja.2010.
11.003
Peel, M. C., Finlayson, B. L., and McMahon, T. A. (2007). Upda ed wo ld map o
he Köppen-Geige clima e classi ica ion. Hyd ol. Ea h Sys . Sci. 11, 1633–1644.
doi: 10.5194/hess-11-1633-2007
Pilling, D., and Ho man, I. (2011). Clima e Change and Animal Gene ic Resou ces
o Food and Ag icul u e: S a e o Knowledge, Risks and Oppo uni ies. Back-
g ound S udy Pape No. 53. Rome: Commission on Gene ic Resou ces o Food
and Ag icul u e.
P i cha d, J. K., Pick ell, J. K., and Coop, G. (2011). The gene ics o human
adap a ion: ha d sweeps, so sweeps, and polygenic adap a ion. Cu . Biol. 20,
R208–R215. doi: 10.1016/j.cub.2009.11.055
Randolph, S. E. (2009). Tick-bo ne disease sys ems eme ge om he shadows: he
beau y lies in molecula de ail, he message in epidemiology. Pa asi ology 136,
1403–1413. doi: 10.1017/S0031182009005782
Rauw, W. M., Kanis, E., Noo dhuizen-S assen, E. N., and G omme s, F. J. (1998).
Undesi able side e ec s o selec ion o high p oduc ion e iciency in a m
animals: a e iew. Li es . P od. Sci. 56, 15–33. doi: 10.1016/S0301-6226(98)
00147-X
Ra agnolo, O., and Misz al, I. (2002). E ec o hea s ess on non e u n a e in Hol-
s ein cows: gene ic analyses. J. Dai y Sci. 85, 3092–3100. doi: 10.3168/jds.S0022-
0302(02)74396-8
Ra agnolo, O., Misz al, I., and Hoogenboom, G. (2000). Gene ic componen o
hea s ess in dai y ca le, de elopmen o hea index unc ion. J. Dai y Sci. 83,
2120–2125. doi: 10.3168/jds.S0022-0302(00)75094-6
Robinson, P. J. (2001). On he De ini ion o a Hea Wa e. J. Appl. Me eo ol. Clim.
40, 762–775. doi: 10.1175/1520-0450(2001)040<0762:OTDOAH>2.0.CO;2
Sa ini, G., A onso, A., Mello , P., A adaib, I., Yadin, H., and Sanaa, M. (2011).
Epizoo ic hemo hagic disease. Res. Ve . Sci. 91, 1–17. doi: 10.1016/j. sc.2011.
05.004
Seo, S. N., McCa l, B. A., and Mendelsohn, R. (2010). F om bee o sheep
unde global wa ming? An analysis o adap a ion by li es ock species choice
in Sou h Ame ica. Ecol. Econ. 69, 2486–2494. doi: 10.1016/j.ecolecon.2010.07.
025
Sco , M. E., and Smi h, G. (1994). Pa asi ic and In ec ious Diseases: Epidemiology
and Ecology. San Diego: Academic P ess.
Sil a, F. F., Mulde , H. A., Knol, E. F., Lopes, M. S., Guima ães, S. E. F., Lopes, P.
S., e al. (2014). Si e e alua ion o o al numbe bo n in pigs using a genomic
eac ion no ms app oach. J. Anim. Sci. 92, 3825–3834. doi: 10.2527/jas.2013-
6486
S ein eld, H., Ge be , P., Wassenaa , T., Cas el, V., Rosales, M., and de Haan, C.
(2006). Li es ock’s Long Shadow. Rome: The Food and Ag icul u al O ganiza ion
o he Uni ed Na ions (FAO).
Sæ he , N. H., Sickel, H., No de haug, A., Sickel, M., and Vangen, O. (2006). Plan
and ege a ion p e e ences o a high and a mode a e yielding No wegian dai y
ca le b eed g azing semi-na u al moun ain pas u es. Anim. Res. 55, 363–387.
doi: 10.1051/anim es:2006033
Tabe le , P., Valen ini, A., Rezaei, H. R., Nade i, S., Pompanon, S., Neg ini, R., e al.
(2008). A e ca le, sheep, and goa s endange ed species? Mol. Ecol. 17, 275–284.
doi: 10.1111/j.1365-294X.2007.03475.x
Tapio, I., Vä , S., Bennewi z, J., Male iciu e, J., Fimland, E., G islis, Z., e al. (2006).
P io i iza ion o conse a ion o no he n Eu opean ca le b eeds based on
analysis o mic osa elli e da a. Conse . Biol. 20, 1768–1779. doi: 10.1111/j.1523-
1739.2006.00488.x
Tapio, M., Oze o , M., Tapio, I., To o, M. A., Ma zano , N., ´
Cinkulo , M.,
e al. (2010). Mic osa elli e-based gene ic di e si y and popula ion s uc u e
o domes ic sheep in no he n Eu asia. BMC Gene . 11:76. doi: 10.1186/1471-
2156-11-76
Tubiello, F. N., Soussana, J.-F., and Howden, S. M. (2007). C op and pas-
u e esponse o clima e change. PNAS 104, 19686–19690. doi: 10.1073/pnas.
0701728104
an de Waaij, E. H. (2004). A esou ce alloca ion model desc ibing consequences
o a i icial selec ion unde me abolic s ess. J. Anim. Sci. 82, 973–981.
an Dijk, J., Sa gison, N. D., Kenyon, F., and Skuce, P. J. (2010). Clima e change
and in ec ious disease helmin hological challenges o a med uminan s in
empe a e egions. Animal 4, 377–392. doi: 10.1017/S1751731109990991
Wall, E., Simm, G., and Mo an, D. (2010). De eloping b eeding schemes o assis
mi iga ion o g eenhouse gas emissions. Animal 4, 366–376. doi: 10.1017/
S175173110999070X
Wellman, R., Bennewi z, J., and Meuwissen, T. (2014). A uni ied app oach o
cha ac e ize and conse e adap i e and neu al gene ic di e si y in subdi ided
popula ions. Gene . Res. 96:e16. doi: 10.1017/S0016672314000196
Wes , J. W. (2003). E ec s o hea -s ess on p oduc ion in dai y ca le. J. Dai y Sci.
86, 2131–2144. doi: 10.3168/jds.S0022-0302(03)73803-X
Wheele , T., and Reynolds, C. (2013). P edic ing he isks om clima e change o
o age and c op p oduc ion o animal eed. Anim. F on . 3:36–41. doi: 10.2527/
a .2013-0006
Wol , J. B. (2013). P inciples o ansc ip ome analysis and gene exp ession quan-
i ica ion: an RNA-seq u o ial. Mol. Ecol. Res. 13, 559–572. doi: 10.1111/1755-
0998.12109
Ya es, C. M., Cammell, S. B., F ance, J., and Bee e , D. E. (2000). P edic ions o
me hane emissions om dai y cows using mul iple eg ession analysis. P oc. B .
Soc. Anim. Sci. 2000:94.
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.
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P æbel, Be g and Meuwissen. This is an open-access a icle dis ibu ed unde he
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