blup-analysis of genetic trends in the dohne merino …

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BLUP-ANALYSIS OF GENETIC TRENDS IN THE DOHNE MERINO H.C. de Klerk and H.J. Heydenrych* Dohne Agricultural Research Institute, Stutterheim and ‘University of Stellenbosch, Stellenbosch, Republic of South Africa SUMMARY The Dohne Merino is a South African bred mutton-woolled sheep specifically bred for the sour-grassveld areas. Although phenotypically well documented, little was known about its genetic development in economically important traits until the advent of BLUP- methodology. This paper describes genetic trends in greasy fleece mass and body mass traits over the whole development period and three distinct phases of development. Genetic progress was slow but obvious differences in the response gradients occurred in the three developmental phases. W ithin phases the different traits measured reacted similarly. INTRODUCTION The Dohne Merino is one of several South African sheep breeds (also the Afrino, Dormer, Dorper, Vandor and Van Rooy) which were specially developed for certain harsh agricultural environments, while at the same time satisfying specific market requirements. The aim in developing the Dohne Merino was to evolve a mutton-woolled sheep that was adapted to the climatic and veld conditions in the sour-grassveld areas in the Eastern parts of South Africa (Kotze, 1951). Further requirements were a high fertility, a high lamb growth rate, good quality wool without hair, coloured fibres or kemp, and the ability to lamb in autumn as well as in spring. Commencing in 1939 at the Dohne Research Station, German Merino (presently South African Mutton Merino) rams were mated to Merino ewes to produce half-bred lambs from which the Dohne Merino was developed. Although based on objective measurements and scientific methods, the development of the Dohne Merino was not executed as a selection experiment with a genetic control, but as a comprehensive breeding and selection process, while also considering the future acceptability of the breed to the farming industry. Therefore there was no scientific procedure available to measure genetic progress in the economic traits of the breed until BLUP-methodology became accessible for general use. The purpose of this paper is to discuss some genetic trends in the Dohne Merino as a typical example of the application of BLUP-methodology to study the genetic history of a new farm animal breed during its initial stages of development. 77

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B L U P -A N A L Y S IS O F G E N E T IC T R E N D S IN T H E D O H N E M E R IN O

H .C . d e K le rk an d H .J . H e y d e n ry c h *D o h n e A g r ic u ltu ra l R e s e a rc h In s titu te , S tu t te rh e im a n d ‘ U n iv e rs i ty o f S te lle n b o sc h ,

S te lle n b o sc h , R e p u b lic o f S o u th A fr ic a

S U M M A R Y

T h e D o h n e M e r in o is a S o u th A fr ic a n b re d m u tto n -w o o lle d s h e e p sp e c if ic a lly b r e d fo r th e so u r-g ra ssv e ld a re a s . A lth o u g h p h e n o ty p ic a lly w e ll d o c u m e n te d , l i t t le w a s k n o w n a b o u t its g e n e tic d e v e lo p m e n t in e co n o m ic a lly im p o r ta n t t r a i ts u n t il th e a d v e n t o f B L U P - m e th o d o lo g y . T h is p a p e r d e sc rib e s g e n e tic t r e n d s in g re asy f le e c e m a ss a n d b o d y m ass tra i ts o v e r th e w h o le d e v e lo p m e n t p e r io d a n d th re e d is tin c t p h a s e s o f d e v e lo p m e n t . G e n e t ic p ro g re s s w as s lo w b u t o b v io u s d if fe re n c e s in th e r e sp o n se g r a d ie n ts o c c u r re d in th e th re e d e v e lo p m e n ta l p h a se s . W ith in p h a se s th e d if fe re n t t r a i ts m e a s u re d r e a c te d s im ila rly .

IN T R O D U C T IO N

T h e D o h n e M e r in o is o n e o f sev e ra l S o u th A fr ic a n sh e e p b r e e d s (a ls o th e A fr in o , D o rm e r , D o rp e r , V a n d o r a n d V a n R o o y ) w h ich w e re sp e c ia lly d e v e lo p e d fo r c e r ta in h a rsh a g r ic u ltu ra l e n v iro n m e n ts , w h ile a t th e s a m e tim e sa tis fy in g sp e c if ic m a rk e t re q u ire m e n ts . T h e a im in d e v e lo p in g th e D o h n e M e rin o w as to ev o lv e a m u tto n -w o o lle d s h e e p th a t w as a d a p te d to th e c lim a tic a n d v e ld c o n d itio n s in th e so u r-g ra ssv e ld a r e a s in th e E a s te r n p a r ts o f S o u th A fr ic a (K o tz e , 1951). F u r th e r r e q u ire m e n ts w e re a h ig h fe rtil ity , a h ig h lam b g ro w th ra te , g o o d q u a li ty w o o l w ith o u t h a ir , c o lo u re d f ib re s o r k e m p , a n d th e a b ility to lam b in a u tu m n as w ell a s in sp rin g .

C o m m e n c in g in 1939 a t th e D o h n e R e se a rc h S ta tio n , G e r m a n M e r in o (p re s e n t ly S o u th A fr ic a n M u tto n M e r in o ) r a m s w e re m a te d to M e rin o e w es to p ro d u c e h a lf -b re d la m b s fro m w hich th e D o h n e M e r in o w a s d e v e lo p e d .

A lth o u g h b a se d o n o b je c tiv e m e a su re m e n ts a n d sc ien tif ic m e th o d s , th e d e v e lo p m e n t o f th e D o h n e M e r in o w a s n o t e x e c u te d as a s e le c tio n e x p e r im e n t w ith a g e n e t ic c o n tro l , b u t as a c o m p re h e n s iv e b re e d in g a n d s e le c tio n p ro c e ss , w h ile a lso c o n s id e r in g th e fu tu re a c c e p ta b ility o f th e b r e e d to th e fa rm in g in d u stry . T h e r e f o r e th e r e w a s n o sc ien tif ic p ro c e d u re a v a ila b le to m e a s u re g e n e tic p ro g re s s in th e e c o n o m ic t r a i t s o f th e b r e e d u n til B L U P -m e th o d o lo g y b e c a m e a c c ess ib le fo r g e n e ra l u se .

T h e p u rp o se o f th is p a p e r is to d iscuss so m e g e n e tic t r e n d s in th e D o h n e M e r in o a s a typ ica l e x a m p le o f th e a p p lic a t io n o f B L U P -m e th o d o lo g y to s tu d y th e g e n e t ic h is to ry o f a new fa rm a n im a l b r e e d d u r in g its in it ia l s ta g es o f d e v e lo p m e n t.

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M A T E R IA L A N D M E T H O D S

A ll D o h n e M e r in o sh e e p b o rn d u r in g th e d e v e lo p m e n t p e r io d 1948 to 1985 (a p p ro x im a te ly 10 000 in d iv id u a ls ) w e re in c lu d e d in th is s tu d y . D a ta in c lu d e d o b se rv a tio n s o n b o d y m ass a t b ir th , a n d a t 100 days, 180 d ay s a n d 18 m o n th s o f a g e , as w ell as g re asy f le e c e m ass a t th e l a t te r ag e .

T h e B L U P A n im a l M o d e l P ro g ra m o f th e S o u th A fr ic a n D e p a r tm e n t o f A g r ic u ltu re an d W a te r S u p p ly , as a d a p te d a n d f u r th e r d e v e lo p e d by D e lp o r t (1 9 8 9 ) w as u se d to a n a ly se th e d a ta .

A m ix e d m o d e l w a s u se d to d e sc r ib e th e o b se rv a tio n s . F o llo w in g B la ir (1 9 8 1 ) w h o w o rk e d w ith s im ila r sh e e p d a ta , a ll f irs t o rd e r in te ra c t io n s w e re e x c lu d ed fro m th e m o d e l w h ich is g iv en in th e fo llo w in g e q u a t io n .

^ i jk lm f1 + a i + b j + c k + ^1 + ^m + -e ijk lm

w h e re Y jjy m is a n in d iv id u a l p h e n o ty p ic o b se rv a tio n ,

ju is th e p o p u la t io n m e a n

aj is th e sex e f fe c t (m a le o r f e m a le )

b j is th e fixed a g e o f d a m e ffe c t ( m a tu re o r im m a tu re )

cj. is th e fixed b i r th s ta te e ffe c t (s in g le o r m u ltip le )

d j is th e fixed a v e ra g e b ir th y e a r e ffe c t, f re e f ro m g e n e tic tre n d

f™ is th e r a n d o m e f fe c t o f th e m 1*1 a n im a l in th e i1*1 y e a r (p re d ic te d b r e e d in g v a lu e ) , an d

e ijk lm ^ le r a n d o m re s id u a l e f fe c t fo r e a c h o b se rv a tio n .

A s su ch , th e m o d e l r e p re s e n ts a s in g le t r a i t a n im a l m o d e l in w hich b r e e d in g v a lu e s a re p re d ic te d fo r a ll a n im a ls .

D u e to th e la rg e n u m b e r o f e q u a t io n s in v o lv ed , i te ra t iv e p ro c e d u re s w e re e m p lo y e d . B la ir an d P o lia k (1 9 8 4 ) c o n s id e re d so lu tio n s c o n v e rg e d w h e n th e c o n v e rg e n c e c r i te r io n w as less th a n 0,0001 kg (s h e e p d a ta ) fo r b o th fix ed a n d ra n d o m e ffec ts . In th is s tu d y i te r a t io n s w e re lim ite d to 3 0 0 ro u n d s a n d th e n c o n s id e re d a s co n v e rg e d .

G e n e t ic t r e n d s w e re o b ta in e d by a v e ra g in g th e b re e d in g v a lu e s o f a n im a ls b o rn w ith in sp ec ific y e a rs (e x c lu d in g b a s e p o p u la t io n a n im a ls ) . R e g re s s io n s w e re e m p lo y e d to d e te rm in e th e g e n e tic t r e n d o v e r th e e n t i r e p e r io d as w ell a s fo r th r e e d is t in c t p h a se s . T h e s e w e re th e in itia l p h a s e (1 948 - 1955) w h e n sm a ll n u m b e rs o f c ro s sb re d a n im a ls w e re a v a ila b le , th e e x p a n s io n p h a s e (1 9 5 6 - 1975), a n d th e p e r fo rm a n c e te s tin g p h a s e (1 9 7 6 -1985) d u r in g w h ich s e le c tio n w a s p r im a r ily b a s e d o n p e r fo rm a n c e in d ices .

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R E S U L T S

T h e in it ia l m e a n s a n d g e n e t ic t r e n d g ra d ie n ts o f five p ro d u c tio n t r a i ts fo r th e to ta l p e r io d o f s tu d y (1 948 to 1985) a n d fo u r t r a i ts fo r th e th re e d e v e lo p m e n ta l p h a s e s a r e sh o w n in T a b le

T a b le 1 M e a n in it ia l m ass a n d g e n e tic t r e n d g ra d ie n ts (kg)

T r a i t I n i t ia lm a s s

T o ta lp e r io d

1948 - 55 1956 - 75 1976 - 85

B irth m ass 100 d ay

3 .57 0.005 - - -

m ass 180 day

22 .79 0.045 -0.053 0 .068 0 .0 3 2

m ass 18 m o n th

29.23 0 .059 -0 .052N o t

0 .098 0 .0 4 2

m assG re a s y

44 .30 0 .059 m e a su re d 0 .017 0 .129

fle e c em ass

3 .99 0 .004 -0.006 0.015 0.001

D IS C U S S IO N

A ll fo u r tra i ts s tu d ie d , sh o w slow b u t p o sitiv e re sp o n se t r e n d s o v e r th e to ta l p e r io d o f 37 y e a rs ( a b o u t 10 g e n e ra tio n s ) . I f th e in itia l m asse s a re c o n s id e re d , th e v a lu e fo r 18 m o n th m ass a p p e a rs re la tiv e ly sm a ll c o m p a re d to th e re g re ss io n s fo r b o d y m a ss a t 100 a n d 180 days o f ag e . T h is re s u l t is h o w e v e r in a g re e m e n t w ith th e h e r i ta b i l i t ie s o f 0 .3 9 a n d 0.53 fo r 100 a n d 180 d ay m ass , c o m p a re d to 0.31 fo r 18 m o n th b o d y m ass a s fo u n d by F o u r ie (1 981) fo r th e s a m e flock . T h e t r e n d g ra d ie n ts fo r b ir th m ass a n d g re a sy f le e c e m a ss a r e m u c h sm a lle r a n d o f th e sa m e o rd e r , a s a r e a lso th e ir in it ia l v a lu e s .

G e n e t ic im p ro v e m e n t to o k p la c e a t r a te s v a ry in g fro m 0 .1 % fo r g re a sy f le e c e m a s s to 0 .2 7 % fo r 18 m o n th b o d y m ass w h ich is a b o u t o n e te n th o f th e r a te p o s s ib le in a w e ll p la n n e d s in g le t r a i t s e le c tio n sy s tem .

Such s lo w p ro g re s s in in d iv id u a l c h a ra c te ris tic s is h o w e v e r a c c e p ta b le in th e p re s e n t s i tu a t io n w h e re th e d e v e lo p m e n t o f a n e w b re e d to o k p la c e a n d w h e re th e to ta l sh e e p w as c o n s id e re d , in c lu d in g m a n y e c o n o m ica lly im p o r ta n t t r a i ts a s w e ll a s b r e e d c h a ra c te r is tic s to b e ta k e n u p in th e s ta n d a rd s o f e x ec e lle n c e o f th e n e w b re e d .

C o m p a r is o n o f th e t r e n d s o v e r th e th r e e d if fe re n t d e v e lo p m e n ta l p h a s e s a r e m o re in te re s t in g . T w o b o d y m ass tra i ts (1 0 0 a n d 180 day m a ss) a s w e ll a s g re a sy f le e c e m ass sh o w e d a n e g a tiv e t r e n d d u r in g th e in itia l p h a se (1948 - 1955) p o ss ib ly b e c a u s e o f sm a ll n u m b e rs in v o lv e d a n d th e c o n se q u e n t in b re e d in g . B o d y m a ss a t 18 m o n th s o f a g e w as n o t m e a s u re d d u r in g th is p h a se .

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D u r in g th e e x p a n s io n p h a s e (1 9 5 6 - 1975) s e le c tio n w a s p r im a rily su b je c tiv e , w ith p e r fo rm a n c e d a ta p lay in g a m in o r ro le . H o w e v e r b o th th e 100 d ay a n d 180 d a y m a ss tra its sh o w e d a g r e a te r r e s p o n s e in th e se c o n d , th a n in th e th ird (p e r fo rm a n c e te s tin g ) p h a se (1 9 7 6 - 1985). T h is r e su lt c a n p o ss ib ly b e a sc rib e d to th e fa c t th a t a l th o u g h o n a su b je c tiv e b a s is , th e s e t r a i ts w e re s e le c te d fo r d ire c tly d u r in g th e se c o n d p h a s e , w h ile th e y w e re s e le c te d fo r in d ire c tly in th e th ird p h a s e w h e n p e r fo rm a n c e d a ta an d in d ic e s w e re a v a ila b le fo r 18 m o n th m ass a n d g re a sy f le e c e m ass , w h ich th e n b e c a m e th e d ire c tly s e le c te d tra its .

B o d y m a ss a t 18 m o n th s w a s th e on ly t r a i t m e a su re d , w h ich sh o w e d a h ig h e r r e sp o n se d u r in g th e p e r fo rm a n c e te s tin g p h a s e th a n d u r in g th e e x p a n s io n p h a se . S tu d ie d o n its ow n, th is r e s u l t is q u ite p r e d ic ta b le b e c a u s e o f th e o b jec tiv e , p e r fo rm a n c e b a s e d s e le c tio n in th e th ird p h a se . O n th e o th e r h a n d g re a sy f le e c e m ass, th e o th e r t r a i t s u b je c te d to d ire c t o b je c tiv e s e le c tio n d u r in g th e th ir d p h a s e , r e a c te d v ery m u c h lik e 100 a n d 180 d a y b o d y m a ss in th a t i t sh o w e d a m u c h lo w e r r e sp o n se d u r in g th e th ird th a n in th e s e c o n d p h a se . T h is w as p o ss ib ly d u e to th e m o re d isc r im in a tin g se le c tio n fo r w o o l q u a lity w h ich a c c o m p a n ie d s e le c tio n fo r g re a sy f le e c e m ass d u r in g th e p e r fo rm a n c e te s tin g p h a s e . F o r in s ta n c e , m u c h g r e a te r e m p h a s is w a s p la c e d o n w o o l f in e n e ss a n d 23 m ic ro n s w as th e m a x im u m c u t o f f lim it d u r in g th is p h a se .

A n o th e r a sp e c t to b e c o n s id e re d w ith re g a rd to th e d is sa p o in tin g re s p o n s e d u r in g th e p e r fo rm a n c e te s tin g p h a s e is t h a t s o m e "u n re la te d " o u ts id e ra m s f ro m c o -o p e ra tin g flocks w e re u se d in th e s tu d d u r in g 1978, 1980, 1981 a n d 1982. G ra p h ic r e p re s e n ta t io n o f th e d a ta sh o w e d c o n s id e ra b ly h ig h e r v a r ia b il i ty in b re e d in g v a lu e s d u r in g th e th ird p h a s e , w h ich in d ic a te d th a t w h ile so m e o u ts id e ra m s w e re o f a h ig h e r g e n e tic m e r i t th a n th e h o m e b re d ra m s , o th e r s w e re o f a lo w e r s ta n d a rd . A ll in d ic a tio n s a re th a t th e b r e e d h a s n o t y e t re a c h e d a g e n e tic p la te a u a n d a B L U P -a n a ly s is a t a la te r s ta g e m ig h t v e ry w e ll in d ic a te a fu th e r su rg e o f g e n e tic r e sp o n se . I f f u r th e r n e w b re e d s w e re to b e d e v e lo p e d a t th is p o in t in tim e , o n e w o u ld h a v e b e e n in a m u c h b e t t e r p o s it io n to m a k e g o o d u s e o f p e r fo rm a n c e te s tin g a n d to m o n ito r its d e v e lo p m e n t by th e u se o f B L U P -te c h n o lo g y .

R E F E R E N C E S

B L A IR , H .T ., 1981. P h . D . D is s e r ta t io n . M assey U n iv ., N e w Z e a la n d

B L A IR , H .T . an d P O L L A K , E .J ., 1984. J . A n im . Sci. 58: 1090 - 1096

D E L P O R T , G .J ., 1989. P h . D . D is s e r ta tio n , U n iv . o f th e O ra n g e F re e S ta te , B lo e m fo n te in , R e p u b lic o f S o u th A frica .

F O U R IE , A .J ., 1981. M .Sc. A g ric . T h e s is . U n iv . o f S te lle n b o sc h , R e p u b lic o f S o u th A frica .

K O T Z E , J .J .J ., 1951. F a rm in g in S o u th A fric a , A p ril, 1951.

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