Genetics of growth reaction norms in farmed rainbow trout

RESEARCH ARTICLE Gene ics o G ow h Reac ion No ms in Fa med Rainbow T ou Panya Sae-Lim 1 *, Han Mulde 2 , Bja ne Gje de 1 , Heikki Koskinen 3 , Ma ie Lillehamme 1 , An i Kause 4 1Aquacul u e and Gene ics, No ima, Oslo eien 1, Ås, No way, 2Animal B eeding and Genomics Cen e, Wageningen Uni e si y, Wageningen, he Ne he lands, 3Aquacul u e Uni , Na u al Resou ces Ins i u e Finland, Te o, Finland, 4Biome ical Gene ics, Na u al Resou ces Ins i u e Finland, Jokioinen, Finland *panya.sae-lim@no ima.no Abs ac Rainbow ou is a med globally unde di e se uncon ollable en i onmen s. Fish wi h low mac oen i onmen al sensi i i y (ES) o g ow h is impo an o h i e and g ow unde hese uncon ollable en i onmen s. The ES may e ol e as a co ela ed esponse o selec ion o g ow h in one en i onmen when he gene ic co ela ion be ween ES and g ow h is nonze o. The aims o his s udy we e o quan i y addi i e gene ic a iance o ES o body weigh (BW), de ined as he slope o eac ion no m ac oss b eeding en i onmen (BE) and p oduc- ion en i onmen (PE), and o es ima e he gene ic co ela ion ( g(in , sl) ) be ween BW and ES. To es ima e he i able a iance o ES, he cohe i abili y o ES was de i ed using selec- ion index heo y. The BW eco ds om 43,040 ainbow ou pe o ming ei he in eshwa e o seawa e we e analysed using a eac ion no m model. High addi i e gene ic a iance o ES (9584) was obse ed, in e ing ha gene ic changes in ES can be expec ed. The cohe i abili y o ES was ei he -0.06 (in e cep a PE) o -0.08 (in e cep a BE), sugges ing ha BW obse a ion in ei he PE o BE esul s in low accu acy o selec ion o ES. Ye , he g(in , sl) was nega i e (-0.41 o -0.33) indica ing ha selec ion o BW in one en i onmen is expec ed o esul in mo e sensi i e ish. To a oid an inc ease o ES while selec ing o BW, i is possible o ha e equal gene ic gain in BW in bo h en i onmen s so ha ES is main ained s able. In oduc ion The pe o mance o o ganisms is in luenced by he su ounding en i onmen al condi ions, leading o pheno ypically plas ic esponses o en i onmen al changes. Such plas ic esponses ha e been obse ed, o example, as adap i e plas ici y in he neck ee h o Daphnia (wa e leas) which de elops as a p o ec i e esponse o he chemical cues o a p eda o y Chaobo us p esen in he wa e [1]. In ish species, pheno ypic plas ici y has been explo ed especially om ecological and e olu iona y poin s o iew. The e is e idence o gene ic basis o pheno ypic plas ici y, o example in salmonids [2], T inidadian guppies (Poecilia e icula a)[3,4] and pup ishes (Cyp inodon ne adensis)[5]. PLOS ONE | DOI:10.1371/jou nal.pone.0135133 Augus 12, 2015 1/17 OPEN ACCESS Ci a ion: Sae-Lim P, Mulde H, Gje de B, Koskinen H, Lillehamme M, Kause A (2015) Gene ics o G ow h Reac ion No ms in Fa med Rainbow T ou . PLoS ONE 10(8): e0135133. doi:10.1371/jou nal. pone.0135133 Edi o : Gen Hua Yue, Temasek Li e Sciences Labo a o y, SINGAPORE Recei ed: Feb ua y 6, 2015 Accep ed: July 18, 2015 Published: Augus 12, 2015 Copy igh : © 2015 Sae-Lim e al. 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, which pe mi s un es ic ed use, dis ibu ion, and ep oduc ion in any medium, p o ided he o iginal au ho and sou ce a e c edi ed. Da a A ailabili y S a emen : Na u al Resou ces Ins i u e Finland (Luke) own he da a unde lying his pape . Please send eques s o he da a o pe i. heinimaa@luke. i. Funding: This s udy is unded by No wegian Resea ch Council (NRC: 234144/E40), h p://www. o sknings ade .no/en/Home_page/1177315753906. The unde s had no ole in s udy design, da a collec ion and analysis, decision o publish, o p epa a ion o he manusc ip . Compe ing In e es s: The au ho s ha e decla ed ha no compe ing in e es s exis . Among animal b eede s, pheno ypic plas ici y is e med mac oen i onmen al sensi i i y (ES) [6]. I has been o in e es o animal b eede s because o i s connec ion wi h animal’s pe - o mance ac oss en i onmen s [7,8] and o he obus ness and wel a e o animals [9]. Fo a geno ype, such as a clone, amily, popula ion, o a species, mac oen i onmen al sensi i i y can be de ined by i s slope o eac ion no m ac oss en i onmen s. Assuming a linea eac ion no m, he deg ee o mac oen i onmen al sensi i i y can be quan i ied by he eg ession slope o a geno ype's pe o mance, such as g ow h, agains an en i onmen al g adien [10–12]. Rainbow ou Onco hynchus mykiss (Walbaum 1792) is one o he main ish species a med unde di e se en i onmen al condi ions ac oss con inen s. Rapid g ow h is one o he mos impo an ai s o p o i able ou a ming. Howe e , ish may no be able o main ain high g ow h when ea ing condi ions a e subop imal. The e o e, a mo e obus ish wi h high s abil- i y o g ow h is impo an o h i e unde a iable en i onmen al condi ions. To quan i y he po en ial o changing mac oen i onmen al sensi i i y h ough selec ion, an es ima e o gene ic a iance in mac oen i onmen al sensi i i y is equi ed. The gene ic a ia ion in he mac oen i onmen al sensi i i y is known as non-pa allel eac ion no ms, causing geno ype- by-en i onmen in e ac ion (GxE) [13]. E idences o GxE in g ow h o ainbow ou ha e been epo ed [14–20]. Howe e , so a mos s udies use mul i- ai model in which GxE is quan i ied as he gene ic co ela ion be ween he eco ds o he same ai measu ed in di e en en i onmen s. Such gene ic co ela ion exp esses he magni ude o e- anking o amilies wi h espec o hei b eeding alue, bu i does no p o ide an explana ion on how mac oen i on- men al sensi i i y can e ol e ac oss en i onmen s. The concep o mac oen i onmen al sensi- i i y has ne e been applied o b eeding in aquacul u e be o e. In Finland, he na ional b eeding p og amme o ainbow ou b eeds especially o imp o ed g ow h pe o mance in comme cial p oduc ion en i onmen a he Bal ic Sea [21]. Howe e , he s ock is also ea ed in inland eshwa e p oduc ion en i onmen s, and expo ed o Russia and Asia whe e he p oduc ion en i onmen di e s om Finland subs an ially. Hence, he mac oen i onmen al sensi i i y is conside ed as an impo an ai . The aims o his s udy we e wo- old. Fi s ly, we quan i y he gene ic a iance o ES, de ined as he slope o eac ion no m ac oss seawa e and eshwa e p oduc ion en i onmen s in Finland, using a eac ion no m model. Secondly, o s udy whe he selec ion o as g ow h in one en i onmen will change ES, we es ima e he gene ic co ela ion be ween ES and body weigh in one en i onmen , de ined as he in e cep o eac ion no m. In addi ion, we de i ed he cohe i abili y o ES. Al hough, he gene ic co a iance ma ix om eac ion no m and mul i- ai models is in e changeable [7,22–23], he eac ion no m model is chosen as he me hod in his s udy because i p o ides he gene ic pa ame e s o ES and body weigh di ec ly wi hou in e changing. To be able o compa e ou esul s wi h p e ious s udies, we exploi his in e changeable p ope y o calcula e gene ic a iance in ES and i s gene ic co ela ion wi h in e cep in aquacul u e GxE s udies ha all ha e used a mul i- ai model. Ma e ials and Me hods E hics S a emen All p ocedu es in ol ing animals we e app o ed by he animal ca e commi ee o he Na u al Resou ces Ins i u e Finland. To enhance animal wel a e and amelio a e su e ing du ing all ish handling, he ish we e always i s anaes he ized using MS-222. Da a sou ce All ish used in his s udy we e ob ained om he Finnish na ional b eeding p og amme. B eeding candida es a e held a he Te o ish a m in cen al Finland ( eshwa e nucleus Gene ics o Mac oen i onmen al Sensi i i y PLOS ONE | DOI:10.1371/jou nal.pone.0135133 Augus 12, 2015 2/17 s a ion) and he sibs o he b eeding candida es a e es ed a comme cial sea s a ions loca ed a he Bal ic Sea. The pheno ypic da a had 53,638 eco ds o body weigh a agging om ou yea classes and belonged o wo subpopula ions, one wi h yea classes o 1996 and 1999 and he o he wi h 1997 and 2000. Bo h o hese subpopula ions we e es ablished om he pa en s o yea class 1993. Si es we e ma ed o dams using ei he pa e nal nes ed ma ing o pa ial ac- o ial ma ing designs. Each yea class consis ed o 94 o 197 ull-sib amilies es ablished om he ma ing o 37 o 95 si es wi h 79 o 129 dams. A e ha ching, inge lings om he same ull-sib amily we e held in one o mo e amily anks un il he inge lings eached agging size (mean body weigh o app oxima ely 50 g). Du ing he agging, ull-sibs om each amily we e andomly sampled and di ided in o wo o h ee ba ches ha we e ea ed ei he a he eshwa e nucleus s a ion (de ined as “b eeding en i onmen ”o BE) o a one o wo seawa e s a ions (de ined as “p oduc ion en i onmen ” o PE) a he Bal ic Sea. When he ish we e 2-yea -old, hey we e indi idually weighed a BE ( ai : BW BE , in g) and PE ( ai : BW PE , in g) s a ions. The o al numbe o eco ds analysed was 22,175 indi iduals o BW BE and 20,865 indi iduals o BW PE (Table 1). The a e age BW BE (SD) and BW PE (SD) we e 1094 (363.9) g. and 1050.0 (334.5) g, espec i ely. The pedi- g ee was aced back o he pa en s ( he ounde s) o he 1990 yea class. The ances o s back o he ounde popula ion o he 1990 yea class we e included in he pedig ee. Gene ic Analysis Reac ion no m model. A eac ion no m model was used o es ima e gene ic (co) a iance o ES ( eg ession slope) o body weigh s eco ded on 2-yea -old ish. (Co) a iance compo- nen s o all analyses we e es ima ed using es ic ed maximum likelihood in ASReml e sion 3.0 [24]. App oxima e s anda d e o s we e calcula ed wi h ASReml ollowing Fishe e al. [25]. In addi ion o he analysis o obse ed body weigh s, he analysis was also pe o med wi h log- ans o med body weigh s. This was o es he hypo hesis ha gene ic a iance in ES may be in luenced by a scale e ec , ypically obse ed o body weigh in ish species, i.e., inc easing a iance o BW wi h inc easing mean o BW. Fo ins ance, pa allel eac ion no ms o geno- ypes (no gene ic a iance o slopes) wi h di e en in e cep s a e in ac ansla ed in o di e - en magni udes o sensi i i y i change in body weigh is calcula ed as a pe cen age change in he ai mean. The log- ans o ma ion educes such scale e ec [26]. Table 1. Popula ion s uc u e. Subpopula ion I Subpopula ion II 1996 1999 1997 2000 Popula ion s uc u e Si es, dams 57, 129 37, 94 65, 79 95, 121 Si es pe dam, mean ( ange) 1.00 (1–1) 1.00 (1–1) 2.41 (1–3) 1.63 (1–3) Dams pe si e, mean ( ange) 2.26 (1–4) 2.54 (1–4) 2.93 (1–5) 2.06 (1–5) Full-sib amilies, amily anks 129, 129 94, 135 191, 259 197, 197 Numbe o fish wi h eco ds F eshwa e nucleus s a ion 4994 3084 8099 5998 Fish pe ull-sib amily 38.7 32.8 42.4 30.4 Seawa e s a ion 2573 2442 8351 7499 Fish pe ull-sib amily 19.9 26.0 43.7 38.1 doi:10.1371/jou nal.pone.0135133. 001 Gene ics o Mac oen i onmen al Sensi i i y PLOS ONE | DOI:10.1371/jou nal.pone.0135133 Augus 12, 2015 3/17 The eac ion no m model was: yhijklmn ¼bin þbslXhþYC SITE SEX MATijklþ am;in þam;slXhþcn;in þcn;slXhþehijklmn;ð1Þ whe e yis he obse a ion (body weigh o log body weigh ) o he m h indi idual. The β in and β sl a e he fixed eg ession coe ficien s o he popula ion in e cep (in ) and slope (sl), espec- i ely. The X h is he eg esso o he en i onmen s (X h = 0 and 1) in which he in e cep was placed a X h = 0. The fixed e ec YC×SITE×SEX×MAT was included in he model o co ec o he in e ac ion o he i h yea class (YC,i= 1996, 1997, 1999, 2000), he j h es s a ion (SITE,j= 1: BE, 2 o 4: sea- es s a ions), he k h sex (SEX,k= 1: male, 2: emale, o 9: unknown), and he l h ma u i y (MAT,l= 2: ma u e a 2-yea -old, 3: ma u e a 3-yea -old, 9: unknown). The ais he andom addi i e gene ic e ec o in e cep (in ) and slope (sl) o eac- ion no m, ain asl "# ~ MVN[0,AG RN ], whe e Ais he addi i e gene ic ela ionship ma ix, G RN is gene ic co a iance ma ix om he eac ion no m model, and MVN is mul i a ia e no - mal dis ibu ion. The c n is he andom ull-sib ank e ec (unique numbe s in di e en yea classes), explaining an e ec common o ull-sibs o he han addi i e gene ics ( ank e ec due o he sepa a e ea ing o he amilies p io o agging and non-addi i e gene ic e ec ), cin csl "# ~ MVN[0,IC RN ], whe e C RN is common en i onmen al co a iance ma ix and Iis he iden- i y ma ix. The e~N(0, Is2 e1 0 0Is2 e2 "# ) is he andom esidual e ec o an animal min en i- onmen hwi h o each en i onmen a di e en esidual a iance. The si e’s and o sp ing’s es ima ed b eeding alues (EBVs) o slope ob ained om model (1) we e used o illus a e he ange o addi i e gene ic alues o slope a ailable o selec ion. The magni ude and he sign o a gene ic co ela ion be ween he slope and in e cep , and gene ic a iance o he in e cep , can change depending on which en i onmen he in e cep is de ined. Hence, he model was un wice, ei he wi h PE (Xh1=0)o BE(Xh2= 0) as he in e cep en i onmen . To illus a e he co a iance be ween EBVs o slope and in e cep , si e’s EBVs o he slope when he in e cep we e placed a PE we e anked and a o al o fi een si es wi h he highes , close o ze o, and he lowes EBVs o he slope we e chosen o plo ing he eac ion no m. Gene ic cha ac e is ics o mac oen i onmen al sensi i i y. The s ic sense o he i abil- i y o ES is he a io be ween addi i e gene ic a iance o a slope o pheno ypic a iance o he slope. Due o he lack o pheno ypic a iance o he slope, i is no possible o calcula e he he i- abili y o ES. Th ee al e na i e pa ame e s we e used he e o desc ibe gene ic cha ac e is ics o ES. Following Scheine [27], he i abili y o ES (h2 ES) was calcula ed as: h2 ES ¼s2 GxE s2 P;ANOVA ;ð2Þ whe e s2 GxE is geno ype by en i onmen in e ac ion a iance. The s2 GxE is equal o he s anda dized addi i e gene ic a iance o he slope (^ s2 a;sl ^ s2 X) ha is independen om di e en scales o an en i onmen al a iable (X). The ^ s2 Xis he a iance o X[28], i.e., ^ s2 Xis 0.5 in his s udy, as he pos- sible alues o Xin his s udy a e 0 and 1. The s2 P;ANOVA is o al pheno ypic a iance ac oss en i on- men s, in Scheine 's app oach calcula ed om an analysis o a iance (ANOVA) [27]. Because s2 P;ANOVA may no be a ailable om he eac ion no m model, we adop ed Scheine ’s he i abili y by Gene ics o Mac oen i onmen al Sensi i i y PLOS ONE | DOI:10.1371/jou nal.pone.0135133 Augus 12, 2015 4/17 eplacing he s2 P;ANOVA by ^ s2 P;To al ¼ðnBE 1Þ^ s2 PBW;BE þðnPE 1Þ^ s2 PBW;PE þnBE nPEðWBE WPE Þ2=ðnBE þnPE Þ ðnBE þnPE 1Þ  ,whe enis he numbe o animals wi h a eco d o an animal ai , ^ s2 PBW is pheno ypic a iance o he ai and Wis he mean. No e ha nei he s2 P;ANOVA no ^ s2 P;To al is he pheno ypic a iance o he en i- onmen al sensi i i y. Thus, h2 ES is mo e desc ip i e a he han a p edic i e pa ame e [28]. Fu - he mo e, he defini ion o he i abili y in Eq (2) does no coincide wi h he he i abili y being he eg ession o b eeding alue on pheno ype. Because he exp ession in Eq (1) is no p edic i e o esponse o selec ion, we de ined a sec- ond measu e called cohe i abili y ollowing selec ion index p inciples. The pheno ype (P)o an indi idual ha includes eac ion no m pa ame e s can be de ined as (1). We assume no co a i- ances among a,c, and e, because he e is no ela ionship among a,c, and e. The pheno ypic a - iance (s2 P) o a ai is: s2 P¼s2 a;in þ2Xsa;in ;sl þX2s2 a;sl þs2 c;in þ2Xsc;in ;sl þX2s2 c;sl þs2 eð3Þ Es ima ed addi i e gene ic e ec o slope (^ asl) is equal o he eg ession on Pde ia ed om he popula ion mean, o ^ asl ¼bðPmÞ. The eg ession coe ficien (b) o he b eeding alue o slope on pheno ype is: b¼co ðasl;PÞ s2 P¼sa;in ;sl þXs2 a;sl s2 Pð4Þ The bin Eq (4)is“cohe i abili y” o ES. The e m cohe i abili y is used ins ead o he i abil- i y because cohe i abili y de ines he inhe i ance o associa ion be ween ES and BW in one en i onmen . The addi i e gene ic co a iance be ween in e cep and slope changes along he le els o he en i onmen al ac o (X). Hence, he magni ude and sign o cohe i abili y is dependen on he alue o X. A nega i e cohe i abili y is possible i he absolu e o −σ a,in , sl is g ea e han Xs2 a;sl and/o Xis nega i e and absolu e Xs2 a;sl is g ea e han σ a,in , sl . The sign o he cohe i abili y explains he change in co ela ed esponse o ES when mass selec ion o highe pheno ypic alues is pe o med. When in e cep is placed o he en i onmen , in which selec ion is p ac ised on P,Xbecomes ze o, leading o: b¼sa;in ;sl s2 a;in þs2 c;in þs2 e;in ¼sa;in ;sl s2 Ph ;ð5Þ whe e s2 Phis he pheno ypic a iance o BW in he selec ion en i onmen h. This alue is equal o s2 P;in desc ibed abo e. Finally, o unde s and he po en ial gene ic esponse in ES, he accu acy ( IH ) o p edic ing b eeding alue o ES when a selec ion c i e ion is BW in one o he en i onmen s is equal o: IH ¼ffiffiffiffiffiffiffi bg s2 a;sl s¼ffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi ðsa;in ;sl þXs2 a;slÞ2 s2 Ps2 a;sl s¼sa;in ;sl þXs2 a;sl sPsa;sl ;ð6Þ whe e g is sa;in ;sl þXs2 a;sl. The Eq (6) is equi alen o he equa ion de i ed by Kolmodin and Bijma [29]. Cohe i abili y o ES changes depending on a deg ee and o ms o GxE. To demons a e he ela ionship be ween cohe i abili y and GxE in bo h o ms, i.e. geno ype e- anking and Gene ics o Mac oen i onmen al Sensi i i y PLOS ONE | DOI:10.1371/jou nal.pone.0135133 Augus 12, 2015 5/17 he e ogenei y o a iances, Eq (5) is ea anged as (see S1 Appendix): b¼ sa;ðE1;E2Þs2 a;in s2 P;in ¼ gsa;E1sa;E2s2 a;in s2 P;in ;ð7Þ whe e g is he gene ic co ela ion be ween ai s measu ed in wo di e en en i onmen s (E 1 and E 2 ). The g di e en om uni y indica es a p esen o geno ype e- anking. The g¼sa;ðE1;E2Þ sa;E1sa;E2 , whe e sa;ðE1;E2Þ,sa;E1and sa;E2a e addi i e gene ic co a iance and addi i e gene ic s anda d de ia ion in E 1 and E 2 , espec i ely. Assume ha he e is no he e ogenei y o addi i e gene ic a iances (s2 a;E1=s2 a;E2=s2 a;in ) and s2 P;E1=s2 P;E2=s2 P;in ,Eq(7) is equal o: b¼h2 gh2¼h2ð g1Þð8Þ Eq (8) shows eg ession o cohe i abili y on geno ype e- anking, whe e he slope and in e - cep is equal o he h 2 o a ai . I he h 2 = 0.3 and g a ies om -1 o 1, he magni ude o cohe i abili y, ega dless o he sign inc eases when he gene ic co ela ion di e s om he uni y and he cohe i abili y is a maximum when he gene ic co ela ion equals -1. Placing he in e cep (X= 0) in ei he E 1 o E 2 does no in luence he magni ude o he cohe i abili y (Fig 1). Assume he e is he e ogenei y o addi i e gene ic a iances (s2 a;E16¼ s2 a;E2), Eq (7) is equal o: b¼ðhE1hE2Þ gh2 in ð9Þ In con as o Eq (8), Eq (9) shows ha he e ogenei y o addi i e gene ic a iances esul s in di e en alues o cohe i abili y because h 2in changes, depending on he in e cep (X=0) which is placed in ei he E 1 o E 2 as shown in Fig 2. Calcula ion o eac ion no m pa ame e s Fo he in e cep o eac ion no ms (body weigh a he in e cep en i onmen ), he i abili y (h2 in ) and common en i onmen al e ec (c2 in ) we e calcula ed as: h2 in ¼^ s2 a;in =^ s2 P;in , c2 in ¼^ s2 c;in =^ s2 P;in ,whe e^ s2 P;in is equal o ^ s2 a;in þ^ s2 c;in þ^ s2 e;in , which is he pheno ypic a iance o BW in he in e cep en i onmen when X= 0. Fo he slope o eac ion no ms, he h2 ES was calcula ed using Eq (1) while he cohe i abili y was calcula ed using Eq (5), assuming ha in e cep is placed in he selec ion en i onmen (X=0)asi eflec s ac ual si ua ion o selec- i e b eeding in aquacul u e. The cohe i abili y was calcula ed wice, ei he ha ing BE o PE as he selec ion en i onmen . The gene ic co ela ion be ween in e cep and slope ( g(in , sl) ) was calcula ed as: gðin ;slÞ¼^ sain ;sl ffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi ^ s2 a;in ^ s2 a;sl p. Compa ison o p e ious s udies. The e a e no p e ious s udies using eac ion no m model o s udy en i onmen al sensi i i y in aquacul u e. Hence, o compa e he eac ion no m pa ame e s o he p esen s udy o he p e ious GxE s udies, (co) a iance componen s o he p e ious s udies calcula ed using mul i- ai model we e used o calcula e he (co) a iance componen s o eac ion no m pa ame e s (see S1 and S2 Appendixes). The h2 ES, cohe i abili y and g,(in ,sl) we e calcula ed. The choice o GxE pape s in aquacul u e species was based on he ollowing: g ow h ai s as he s udied ai , a leas 30 ull-sib amilies, and p o iding all he pa ame e s needed o he calcula ions. In o al, 17 s udies we e ound, he species co e ing A c ic cha (Sal elinus alpinus)[30], A lan ic cod (Gadus mo hua)[31], Common ca p Gene ics o Mac oen i onmen al Sensi i i y PLOS ONE | DOI:10.1371/jou nal.pone.0135133 Augus 12, 2015 6/17 (Cyp inus ca pio)[32], Eu opean whi efish (Co egonus la a e us)[33], Eu opean sea bass (Dicen a chus lab ax)[34], Nile ilapia (O eoch omis nilo icus)[35,36], Pacific whi e sh imp (Li openaeus annamei)[37], Rainbow ou [14,16–21,38], Shi anus ilapia (O eoch omis shi - anus)[39]. Resul s In he Finnish da a, GxE o BW exis ed in bo h o ms; e- anking as indica ing by g o BW be ween BE and PE was 0.73, and he e ogenei y o gene ic a iances (Table 2). Bo h phenom- ena induce gene ic a ia ion o ES. Gene ic a iance o mac oen i onmen al sensi i i y The addi i e gene ic a iance o slope o BW (9584) was conside able and he h2 in was mode a e in bo h en i onmen s (0.23 o PE and 0.25 o BE), he h2 ES was low (0.07) implying he addi- i e gene ic a iance o ES explains only a small p opo ion ela i e o o al pheno ypic a i- ance o BW ac oss en i onmen s. Simila ly, he cohe i abili y o ES was low and nega i e, i.e., -0.06 o PE and -0.08 o BE. Thus he accu acy o selec ion o ES o BW is e y low when applying indi idual selec ion o BW in one o he en i onmen s. Fig 1. Rela ionship be ween cohe i abili y and he gene ic co ela ion be ween en i onmen s. The inpu pa ame e s a e a ai wi h pheno ypic a iance o 1 and he i abili y o 0.3 which a e he same ac oss wo en i onmen s. The gene ic co ela ion anges om -1 o 1. doi:10.1371/jou nal.pone.0135133.g001 Gene ics o Mac oen i onmen al Sensi i i y PLOS ONE | DOI:10.1371/jou nal.pone.0135133 Augus 12, 2015 7/17 The magni ude o he i abili y o ES o log- ans o med BW was simila o he he i abili y o ES o obse ed BW. The addi i e gene ic a iance o slope o log- ans o med BW was 69% in PE and 65% in BE o he addi i e gene ic a iance o in e cep , ela i ely sligh ly highe han on he obse ed scale (57% in PE and 53% in BE). This indica es ha simple scale e ec s did no gene a e gene ic a ia ion o mac oen i onmen al sensi i i y. When PE was he in e cep , he slope EBVs o si es (-234.5 o 228.8) and animals (-210.6 o 199.8) anged om s ongly nega i e o posi i e (Fig 3). A posi i e slope implies ha EBVs o BW a e ele a ed in BE as compa ed o EBVs o BW in he in e cep en i onmen PE. I BE was he in e cep en i onmen , he EBVs o slope would change sign. Gene ic co ela ion be ween in e cep and slope The signi ican g(in , sl) (SE) be ween BW in a gi en en i onmen and ES anged om -0.33 (0.10) o -0.41 (0.10), depending on he en i onmen used as he in e cep en i onmen (Table 2). The si es wi h s eep slope EBVs had high in e cep (a PE o BE) (Fig 4). The si es wi h la slope EBV had low in e cep EBV. The nega i e co ela ions om log- ans o med Fig 2. Rela ionship be ween cohe i abili y, he e ogenei y o addi i e gene ic a iances and he gene ic co ela ion be ween en i onmen s. The inpu pa ame e s a e a ai ha has di e en magni udes o he i abili y; 0.1 (line wi h ci cles) and 0.5 (line wi h squa es) in wo di e en en i onmen s (E 1 o E 2 ) and pheno ypic a iances a e equal o 1. The gene ic co ela ion anged om -1 o 1. Line g aphs show ha he e ogenei y o addi i e gene ic a iances esul s in di e en alues o cohe i abili y because h 2in (0.1 o 0.5) changes, depending on he in e cep which is placed in ei he E 1 o E 2 . doi:10.1371/jou nal.pone.0135133.g002 Gene ics o Mac oen i onmen al Sensi i i y PLOS ONE | DOI:10.1371/jou nal.pone.0135133 Augus 12, 2015 8/17 da a (-0.40 o -0.42) emained simila o un ans o med da a. This shows ha apid g ow h in one en i onmen is gene ically ela ed o ele a ed sensi i i y ac oss en i onmen s. Gene ic pa ame e s calcula ed om he p e ious GxE s udies In he p e ious aquacul u e s udies on GxE in g ow h, he h2 ES anged om 0.010 o 0.207 (median = 0.110) while he cohe i abili y anged om -0.600 o 0.500 (median = -0.011; in e - cep a E 1 and = -0.078; in e cep a E 2 )(Fig 5). The g(in , sl) be ween g ow h ai s and ES a - ied om -1.00 o 0.94 (median = -0.386) as shown in Fig 6. Discussion Gene ic a ia ion o mac oen i onmen al sensi i i y Subs an ial addi i e gene ic a iance o mac oen i onmen al sensi i i y (ES) o bo h obse ed and log- ans o med body weigh (BW) indica es po en ial o gene ic esponse o selec ion on ES. A e he log- ans o ma ion o BW, he a iance componen s o ES we e educed bu h2 ES emained simila o he one es ima ed om he un ans o med da a. This indica es ha scale e ec s (high a iance depending on high mean) do no explain he gene ic e ec s o ES. Table 2. Va iance componen s and gene ic co ela ions be ween in e cep and slope om he eac- ion no m (RN) models. Pa ame e In e cep P oduc ion B eeding Body weigh s ^ a;in 216754.9 18040.0 s ^ a;sl 29584.3 9584.3 s ^ c;in 23041.1 3227.3 s ^ c;sl 23822.7 3822.7 s ^ e;in 253197.8 51092.8 s ^ P;To al 273168.5 73168.5 h2 in 0.23 (0.03) 0.25 (0.03) h2 ES 0.07 (0.03) 0.07 (0.03) c2 in 0.04 (0.01) 0.04 (0.01) Cohe i abili y -0.06 (0.02) -0.08 (0.02) g(in , sl) -0.33 (0.10) -0.41 (0.10) Log(body weigh ) s ^ a;in 20.016 0.017 s ^ a;sl 20.011 0.011 s ^ c;in 20.003 0.003 s ^ c;sl 20.004 0.004 s ^ e;in 20.092 0.071 s ^ P;To al 20.102 0.102 h2 in 0.15 (0.02) 0.18 (0.03) h2 ES 0.06 (0.02) 0.06 (0.02) c2 in 0.03 (0.01) 0.04 (0.01) g(in , sl) -0.40 (0.10) -0.42 (0.10) doi:10.1371/jou nal.pone.0135133. 002 Gene ics o Mac oen i onmen al Sensi i i y PLOS ONE | DOI:10.1371/jou nal.pone.0135133 Augus 12, 2015 9/17 9. Ellen E, S a L, Ui dehaag K, B om F, KlopčičM, Reen s R, e al. (2009) Robus ness as a b eeding goal and i s ela ion wi h heal h, wel a e and in eg i y. 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