Manipulating genetic architecture to reveal fitness relationships

This is a sel -a chi ed e sion o an o iginal a icle. This e sion may di e om he o iginal in pagina ion and ypog aphic de ails. Au ho (s): Ti le: Yea : Ve sion: Copy igh : Righ s: Righ s u l: Please ci e he o iginal e sion: CC BY 4.0 h ps://c ea i ecommons.o g/licenses/by/4.0/ Manipula ing gene ic a chi ec u e o e eal i ness ela ionships © The Au ho s, 2014. Published e sion Ke ola, Ta mo; Bo a ynski, Zbigniew; Ko iaho, Janne Saka i Ke ola, T., Bo a ynski, Z., & Ko iaho, J. S. (2014). Manipula ing gene ic a chi ec u e o e eal i ness ela ionships. P oceedings o Pee age o Science, 2014(Janua y), A icle e1. h ps://doi.o g/10.14726/p ocpos.2014.e1 2014 Manipula ing gene ic a chi ec u e o e eal i ness ela ionships Ta mo Ke olaTa mo Ke ola11 ,, Zbyszek Bo a yńskiZbyszek Bo a yński11,2,2 && Janne S. Ko iahoJanne S. Ko iaho11,3,3 1Depa men o Biological and En i onmen al Science, P.O. Box 35, 40014, Uni e si y o Jy äskylä, Finland. [email p o ec ed] 2CIBIO, Cen o de In es igação em Biodi e sidade e Recu sos Gené icos, Uni e sidade do Po o, Campus Ag á io de Vai ão, 4485-661, Vai ão, Po ugal. 3Na u al His o y Museum, P.O. Box 35, FI-40014 Uni e si y o Jy äskylä, Finland. Ci a ion: Ke ola T e al. (2014) Manipula ing gene ic a chi ec u e o e eal i ness ela ionships. P ocPoS 1:e1 DOI: 10.14726/p ocpos.2014.e1 Public Pa on #1: T ai s a e o en claimed ( a he han shown) o be closely ela ed o i ness. Some imes his leads o con as ing p edic ions o he di ec ion o he associa ion be ween he ai alue and i ness, a p ime example o which is he deba e o e how (Da winian) i ness links wi h he a e o ene gy u no e a es (see: Bu on e al. 2011, o de ini ions). To esol e his deba e Bu on e al. (2011) sugges ed manipula ion o ai s and measu ing changes in i ness. In his pe spec i e, we p opose ha we should do he opposi e – manipula e gene ic a chi ec u e unde lying i ness and measu e he ai s esponse. E olu iona y heo ies equen ly ocus on he ole o esou ce alloca ion and esou ce ade-o s in de e mining i ness ai s (Nilsson 2002;Tomkins e al. 2004;Bo a yński and Ko eja 2010;Ke ola & Ko iaho 2009). One common empi ical measu e o esou ce use is es ing me abolic a e, and he e a e wo a enues o hinking in how es ing me abolic a e is ela ed o i ness. High es ing me abolic a e is ei he assumed o allow highe a es o ene gy u no e when needed, o high es ing me abolic a e is expec ed o be nega i ely ela ed o i ness as high me abolic a e pe se can es ic he use o limi ed ene gy on i ness ai s (Nilsson 2002;Bo a yński and Ko eja 2010;Ke ola & Ko iaho 2009). In e es ingly, he e a e only a hand ul o s udies ha ha e ho oughly add essed he alidi y o hese hypo heses and mos o he e idence is co ela i e (Bu on e al. 2011 see also: Whi e & Kea ney 2012). To o e come he limi a ion o co ela i e s udies Bu on e al. (2011) igh ly ad oca ed a manipula i e app oach. Howe e , hey sugges ed using a i icial selec ion expe imen s o ho monal manipula ions o modi y es ing me abolic a e and hen measu ing changes in i ness o de e mine he ou come o hese manipula ions. Due o di icul ies in measu ing i ness, labo ious p ocedu es in ol ed in p oduc ion o selec ion lines and po en ial con ounding ac o s included in ho monal manipula ions, hese me hods may be less han op imal. We ully ag ee ha manipula ions a e called o , bu p opose manipula ing gene ic a chi ec u e unde lying i ness and measu ing he ai s alues, a he han manipula ing es ing me abolic a e and hen measu ing i ness. FREE TO PUBLISH - FREE TO ACCESS - PUBLIC PATRONAGE P oceedings o Pee age o Science | Janua y 2014 | e1 ©Au ho , All Righ s Rese ed. Pu chase a Public Pa on license: h ps://www.pee ageo science.o g/ p oceedings/ge license.php?ms=14 1 o 3 One e y po en , bu unde used, i ness manipula ion is based on di ec ional dominance and e ealing hidden ha m ul ecessi e alleles by inb eeding. This me hod elies on a ew well es ablished p inciples (Falcone & Mackay 1996;Lynch & Walsh 1998). By de ini ion, i ness is unde di ec ional selec ion, which is impo an as condi ions o he build-up o di ec ional dominance would o he wise no exis . Wi h a neu al ai , he expec a ion is ha oughly equal numbe o alleles has posi i e and nega i e e ec s (Lynch & Walsh 1998, p.257, 270). When a ai wi h such gene ic backg ound is subjec ed o di ec ional selec ion o inc eased ai alue, ecessi e alleles inc easing he ai alue (due o di ec ion o selec ion: bene icial alleles) will be ixed apidly. Recessi e alleles ha dec ease ai alue (ha m ul ecessi es) will be much ha de o emo e because hey a e e ealed o selec ion only in a small subse o o sp ing in he popula ion. The e o e, he likelihood o ha m ul ecessi e alleles emains high in ai s unde di ec ional selec ion (Falcone & Mackay 1996;Lynch & Walsh 1998). When he e is di ec ional selec ion, di ec ional dominance de elops, and, when e ealed, ecessi e alleles will always change ai alues in he di ec ion opposing he long e m pas selec ion, i.e. away om i ness (Falcone & Mackay 1996;Lynch & Walsh 1998). The e o e, expe imen ally manipula ed inb eeding can be used o in e whe he he ocal ai is ela ed o i ness and i smalle o la ge ai alues ha e inc eased he i ness. Al hough, we ad oca e he use o inb eeding me hod o e ealing i ness associa ions, one needs o be cau ious when applying i . Fo example he ecessi e alleles in e y impo an i ness ai s may ha e been pu ged by s ong di ec ional selec ion, which can subs an ially lowe inb eeding dep ession and i is clea ha he me hod should no be applied on al eady inb ed popula ions, whe e inb eeding manipula ion does no cause inc eased inb eeding dep ession. Mo eo e , while inb eeding can be use ul in e ealing he cou se o pas di ec ional selec ion, i is also clea ha i ness may be a esul o op imiza ion o ai exp ession in se e al ai s (Falcone & Mackay 1996). Explo ing inb eeding on ai o ai in e ac ions can be used o de e mine whe he inb eeding is associa ed wi h concu en changes in o he ai s, as was o example done in Malle & Chippindale (2011) and in Ke ola & Ko iaho (2012), es ing also exis ence o non-linea selec ion behind he i ness. In he simples case, calcula ing an in e ac ion be ween wo (mean s anda dized) ai s p o ides a new index ha can be es ed o a linea associa ion wi h i ness. Fo example, i selec ion has been agains high me abolic a e in la ge indi iduals, and agains low me abolic a es in small indi iduals he magni ude o his index is inc eased when indi iduals a e inb ed. Thus, he exis ence o non-linea selec ion is no p e en ing he use o his me hod. On he con a y, i p o ides ex a in o ma ion on how ai s in e ac i ely a ec i ness, ex ending applicabili y o he me hod o mul iple ai s ha wo k in conce o be e i ness (Ke ola & Ko iaho 2012). The inb eeding me hod has been used ew imes be o e in he con ex o iden i ying sexually selec ed ai s (see: Co on e al. 2004), in compa ing he s eng h o selec ion on male and emale ai s (Malle & Chippindale 2011), and in de e mining whe he high main enance me abolism inc eases o dec eases i ness (Ke ola & Ko iaho 2009, 2012; Bo a yński e al. 2012). Cu en , albei limi ed, e idence sugges s ha inb eeding inc eases he main enance me abolism and g ea ly ele a es exp ession o main enance genes (Ke ola & Ko iaho 2009, 2012;Bo a yński e al. 2012;Ma ila e al. 2012;K is ensen e al. 2005). Thus, i seems ha high le els o main enance me abolism may be connec ed o dec eased i ness. Al hough i ness is he mos impo an concep in e olu iona y biology, i is a bugge o a concep no easily amenable o empi ical measu emen . To o e come he p oblem o measu emen , he inb eeding me hod desc ibed abo e is use ul and will p o ide impo an insigh s in o he na u e o he i ness ela ed ai s. Despi e he a e usage o inb eeding o esol ing he ai ’s ela ionship wi h i ness, he e is a la ge body o li e a u e sugges ing ha li e-his o y ai s ( ha a e closely ela ed o i ness) a e indeed a ec ed mo e by inb eeding han o example mo phological ai s ( e iewed in: DeRose & Ro 1999) showing ha inb eeding manipula ion is a po en , bu clea ly unde used, ool in e olu iona y biology. ACKNOWLEDGEMENTSACKNOWLEDGEMENTS We hank Academy o Finland o unding and R. Hegna o e ising he language. P oceedings o Pee age o Science | Janua y 2014 | e1 ©Au ho , All Righ s Rese ed. 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