Comparison of Finnish Landrace crossbred ewes with Columbia, Rambouillet, and Targhee on western range by Robert David Lewis A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science in Animal Science Montana State University © Copyright by Robert David Lewis (1984) Abstract: Effects of crossing 1/2 Finnish Landrace and 1/2 Rambouillet (FxR) rams with Columbia (C), Rambouillet (R), and Targhee (T) ewes on productivity traits under western range spring lambing management were evaluated. From 1975 to 1978, 563 whiteface (WF) ewes composed of C, R, and T, bred to either FxR or WF (of the same breed) rams produced 821 lambs. Ewes bred to FxR had similar levels of prolificacy and individual lamb weights at birth, 60 d, and 120 d (weaning) as those bred to WF rams. Lambs sired by FxR had 7.3% higher survivability to 60 d than those sired by WF (P<.01), resulting in 4.1 kg more lamb weaned per ewe lambing for ewes bred to FxR rams (P<.01). From 1976 to 1982, 146 FxR-sired ewes (1/4Fx) and 143 WF-sired ewes were exposed to Suffolk rams to lamb beginning at T yr of age through to 6 yr of age, for a total of 1264 lambing opportunities. At 1 yr of age 1/4Fx ewes had higher fertility (P<.01) than WF ewes (37.7$ vs 1.3$). Prolificacy (lambs per ewe lambing) for 1/4Fx ewes was 1.10 lambs, and total kg of lamb weaned per 1/4Fx ewe exposed was 14.0 kg. No differences in fertility were found between mature (ages 2 to 6 yr) 1/4Fx and WF ewes. Prolificacy for mature 1/4Fx was .36 lambs higher than that for WF ewes (P<.01). Superiority of 1/4Fx ewes for litter size at 60 d and 120 d per ewe exposed was .24 and .22 lambs, respectively (P<.01). Lambs from mature 1/4Fx dams were lighter at birth (P<.01), but not significantly different at 60 d and 120 d than those from mature WF dams. Survival to 60 d was 4.6% higher for lambs from 1/4Fx dams (P<.05), and 7.6$ higher for lambs from 1/4Fx dams when adjusted for birth weight (P<.01). Litter weights at 60 d and 120 d per ewe exposed were 3.9 kg and 4.3 kg heavier for 1/4Fx than WF ewes (P<.01). Fleeces from 1/4Fx ewes ages 1 to 6 were lighter and coarser than those from WF ewes (P<.01), but had similar staple lengths. COMPARISON OF FINNISH LANDRACE CROSSBRED EWES WITH COLUMBIA, RAMBQUILLET, AND TARGHEE ON WESTERN RANGE by Robert David Lewis A t h e s i s sumbmitted i n p a r t i a l f u l f i l l m e n t of th e re q u ire m e n ts f o r th e degree of M aster of Science in Animal Science MONTANA STATE UNIVERSITY Bozeman, Montana June 1984 ii APPROVAL o f a t h e s i s s u b m it te d by R o b ert David Lewis T h is t h e s i s h a s been r e a d by each member o f t h e t h e s i s c o m m itte e a n d h a s b e e n f o u n d t o be s a t i s f a c t o r y r e g a r d i n g c o n t e n t , E n g l i s h u sa g e , f o r m a t , c i t a t i o n s , b i b l i o g r a p h i c s t y l e , and c o n s i s t e n c y , and i s r e a d y f o r s u b m i s s i o n t o th e C o lle g e o f G ra d u a te S t u d i e s . A pproval f o r t h e M ajor D epartm ent (o ^ D ate -Q-H Head, Major D epartm ent Approved f o r th e C o lle g e o f G ra d u a te S t u d i e s ^ D a te G ra d u a te D ean _ iii STATEMENT OF PERMISSION TO USE In p re se n tin g th is th e sis in p a rtia l f u l f i l l m e n t of th e r e q u i r e m e n t s f o r a m a s t e r ’s d e g r e e a t M ontana S t a t e U n i v e r s i t y , I a g re e t h a t th e L ib r a r y s h a l l make i t a v a i l a b l e to b o rro w ers under th e r u l e s of th e L ib ra ry . B r i e f q u o t a t i o n s from t h i s t h e s i s a r e a llo w a b le w ith o u t s p e c i a l p e rm is s io n , provided t h a t a c c u r a te acknowledgement Of so u rce i s made. P e rm iss io n f o r e x te n s iv e q u o t a t i o n from or r e p r o d u c t io n o f t h i s t h e s i s may be g ra n te d by ,my major p r o f e s s o r , or i n h i s absence, by the D ir e c t o r of L i b r a r i e s when, i n th e o p in io n of e i t h e r , th e proposed use of th e m a t e r i a l i s f o r s c h o l a r l y purposes. m a te ria l Any copying or use of the i n t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l n o t be a l l o w e d w ith o u t my w r i t t e n p erm issio n . iv ACKNOWLEDGEMENTS I w ish to e x p r e s s my s i n c e r e thanks t o Dr. P. J. B urfening and Dr. D. D. K ress f o r t h e i r guidance, encouragem ent, and p a tie n c e throughout my g ra d u a te program. A p p re c ia tio n i s extended t o Dr. M. D. Huffman and Dr. W. W. Kezar f o r t h e i r counsel and i n s t r u c t i o n . S p e c i a l t h a n k s a r e e x p r e s s e d t o Ms. D. M. Doede f o r h e r s u p p o r t and f r i e n d s h i p which made the s u c c e s s of my g ra d u a te program p o s s i b le , and to Mr. D. E. T r o w b r id g e fo r h is a ssista n c e w ith c o m p u te r programming. Thanks a r e e x t e n d e d t o Dr. J. L. Van Horn, Mr. K. L. C olem an, and th e sta ff of th e Red B l u f f R esearch Ranch f o r m an ag em en t and c o l l e c t i o n of d ata. I w ish to o f f e r my s i n c e r e g r a t i t u d e t o my f a m i l i e s , f a r and near, w ith o u t whose d e d i c a t i o n t o my e d u c a t i o n my g r a d u a t e p ro g ra m w ould n e i t h e r have been a tte m p te d nor completed. L a s t ly , I e x p r e s s my most s p e c i a l thanks and a d m ir a ti o n t o my w if e . Sarah. V TABLE OF CONTENTS Page LIST OF ........................................................................................................................... v ii ABSTRACT............................................................................................................................ ix 1. INTRODUCTION......................... ........................................................... 1 2. LITERATURE REVIEW.................................................................................................. 6 F e r t i l i t y . . . . . . .................................................................................................. P r o l i f i c a c y . ........................................................................................................ S u r v i v a b i l i t y .................................... ................................................................. I n d iv i d u a l Progeny Weaning Weight Performance.................................. Composite R eproductive P e rfo rm a n c e ........................................................ Wool......................................................................................................................... 6 3. MATERIALS AND METHODS........................................................... 15 19 23 26 29 P o p u la t io n s .......................................................................................................... Management.................................................................................. .......................... T r a i t s S tu d ie d ...................................................................................... S t a t i s t i c a l Methods........................................................... 29 31 32 33 4 . RESULTS AND DISCUSSION....................................................................................... 36 Mature Ewe and Progeny Performance i n Phase I ............................... Breed Group o f S i r e E f f e c t s ................................................................. Breed Group of Ewe E f f e c t s ........................ Other Main E f f e c t s ............................................ Ewe Lamb and Progeny Performance i n Phase I I .................................. F e r t i l i t y and P r o l i f i c a c y ..................................................................... Progeny Perform ance.................................................................................. ■Composite T r a i t s . ....................................................... Mature Ewe and Progeny Performance i n Phase I I .......................... Breed Group o f Ewe E f f e c t s ................................... Age of Ewe and I n t e r a c t i o n E f f e c t s ................................................. Breed Group o f S i r e E f f e c t s ....................................................... Other Main E f f e c t s .................................................................................. 36 36 42 42 36 43 44 46 4? 4? 58 60 62 vi TABLE OF CONTENTS-C o n tin u e d Page Wool P r o d u c tio n ................................................................................................. Grease F leece W e ig h t........................................ F le e c e Grade...................... '.......................................................................... S ta p l e Length............................................ D is c u s s i o n ..................................................................................................... REFERENCES C IT E D ............................................. 62 62 64 65 66 68 v ii LIST OF TABLES Table 1 2 3 4 5 6 7 8 9 10 11 12 Page Comparison o f 1/2 and 1/4 Finn C rosses w ith S tandard Breeds f o r F e r t i l i t y i n Ewe Lambs................................................... 10 T o ta l Number of Phase I Ewes Lambing by Year and Age a t Lambing...................................... 30 T o ta l Number o f Phase I I Ewes Exposed i n P revious Breeding Season by Year and Age a t Lambing................................. 30 Mean Squares from Analyses o f V ariance f o r Phase I Ewe R eproductive T r a i t s per Ewe Lambing (1975 - 1 9 7 8 ) . . . . 37 Mean Squares from Analyses o f V ariance f o r Phase I I n d iv i d u a l Progeny T r a i t s (1975 - 1978)...................................... 38 L e a s t-S q u a re s Means and S tandard E r r o r s f o r Phase I Ewe R eproductive T r a i t s p er Ewe Lambing (1975 - 1 9 7 8 ) . . . . 39 L e a s t-S q u a re s Means and S tandard E r r o r s f o r Phase I Progeny T r a i t s (1975 - 1978)......... .................................................... 40 Mean Squares from Analyses o f V ariance f o r Phase I I Ewe P ro d u c tio n T r a i t s (1976 - 19 8 2 )............................................... 41 L e a st-S q u a re s Means and S tandard E r r o r s f o r Phase I I Fx Ewe Lamb R eproductive T r a i t s (1976 - 1979)......................... 44 Mean Squares from Analyses o f V ariance f o r Phase I I I n d iv i d u a l Progeny T r a i t s from Fx Ewe Lambs (1976 - 1.979)............................................................................................... 45 L e a st-S q u a re s Means and S tan d a rd E r r o r s f o r Phase I I I n d i v i d u a l Progeny T r a i t s from Fx Ewe Lambs . (1976 T 1979).................... ........................................................................... 45 L e a s t-S q u a re s Means and S tan d a rd E r r o r s f o r Phase I I Fx Ewe Lamb Composite R eproductive T r a i t s p er Ewe Exposed (1976 - 1979)..............'................................................... 46 I v iii LIST OF TABLES-Continued T ab le 13 14 15 16 17 18 19 20 21 22 23 Page L e a s t-S q u a r e s Means and S tan d ard E rro r s f o r F e r t i l i t y of Phase I I Mature Ewes (1976 - 19 8 2 )...................................... 48 L e a s t-S q u a re s Means and S tan d a rd E r r o r s f o r P r o l i f i c a c y of Phase I I Mature Ewes (1976 - 1982).......................................... 49 Mean S q u ares from A n a ly se s o f V a r ia n c e f o r Phase I I . Mature Ewe Progeny T r a i t s (1977 - 19 8 2 )...................................... 51 L e a s t-S q u a re s Means and S tandard E r r o r s f o r I n d i v i d u a l Lamb Performance from Dams Ages 2 through 6 (1977 - 1982) .......... ..................................... .............................................. 52 L e a st-S q u a re s Means.and S tan d a rd E r r o r s f o r L i t t e r S ize a t 60 Days p e r Ewe Exposed (LS60, 1977 - 1982)....................... 54 L e a st-S q u a re s Means and S tan d a rd E r r o r s f o r L i t t e r Weight a t 60 Days per Ewe Exposed (LW60, 1977 - 1 9 8 2 ) . . .................. 55 L e a st-S q u a re s Means and S tan d a rd E r r o r s f o r Phase I I L i t t e r S ize a t 120 Days per Ewe Exposed (LSI20, 1977 - 1982)................................................................................. 56 L e a s t-S q u a re s Means and S tan d a rd E r r o r s f o r Phase I I L i t t e r Weight a t 120 Days per Ewe Lambing ( LWI20, 1977 - 1.982).......................................................... 57 L e a st-S q u a re s Means and S tan d a rd E r r o r s f o r Phase I I Mature Ewe P ro d u c tio n and I n d i v i d u a l Lamb Performance by B re e d Group o f S i r e (1976 - 19 8 2 )............................... 61 Mean Squares from V ariance A nalyses f o r Ewe Wool P ro d u c tio n T r a i t s (1976 - 19 8 2 )........................................................ 63 L e a s t-S q u a r e s Means and S tan d ard E rro r s f o r Phase I I Wool C h a r a c te r s , Ages I to 6 (1976 - 1982)........................... 63 ix ABSTRACT E f f e c t s o f c r o s s in g 1/2 F in n is h Landrace and 1/2 R am b o u illet (FxR) r a m s w i t h C o lu m b ia (C), R a m b o u i l l e t (R), and T a rg h e e (T) ew es on p r o d u c t i v i t y t r a i t s under w e s te r n range s p r in g lam bing management were e v a lu a te d . From 1975 t o 1978, 563 w h ite f a c e (WF) ewes composed of C, R, and T, b r e d t o e i t h e r FxR o r WF ( o f t h e same b r e e d ) r a m s p r o d u c e d 821 la m b s . Ewes b r e d t o FxR had s i m i l a r l e v e l s o f p r o l i f i c a c y and i n d i v i d u a l lam b w e i g h t s a t b i r t h , 60 d, and 120 d ( w e a n in g ) a s t h o s e b r e d t o WF ra m s . Lambs s i r e d by FxR had 7.3% h i g h e r s u r v i v a b i l i t y t o 60 d t h a n t h o s e s i r e d by WF (P < .01), r e s u l t i n g i n 4.1 kg m ore lam b weaned per ewe lam bing f o r ewes bred to FxR rams (P<.01). From 1976 t o 1982, 146 F x R - s i r e d ew es ( 1 /4 F x ) and 143 W F - s ir e d ew es w e r e e x p o s e d t o S u f f o l k r a m s t o la m b b e g i n n i n g a t I y r o f age through to 6 y r of age, f o r a t o t a l o f 1264 lam bing o p p o r t u n i t i e s . At I y r o f a g e 1 / 4 F x ew es h ad h i g h e r f e r t i l i t y (P<.01) t h a n WF ew es (37.7% v s 1.3%). P r o l i f i c a c y ( la m b s p e r ewe l a m b in g ) f o r 1 /4 F x ew es was 1.10 lam bs, and t o t a l kg o f lamb weaned p er 1/4Fx ewe exposed was 14.0 kg. No d i f f e r e n c e s i n f e r t i l i t y were found between m ature (ages 2 t o 6 y r) 1/4Fx and WF ewes. P r o l i f i c a c y f o r m ature 1/4Fx was .36 lambs h i g h e r t h a n t h a t f o r WF ew es (P<.01). S u p e r i o r i t y o f 1 /4 F x ew es f o r l i t t e r s i z e a t 60 d and 120 d p e r ewe e x p o s e d w as .24 and .22 la m b s , r e s p e c t i v e l y (P<.01). Lambs fro m m a t u r e 1 /4 F x dam s w e r e l i g h t e r a t b i r t h (P < .01), b u t not s i g n i f i c a n t l y d i f f e r e n t a t 60 d and 120 d than th o s e from m ature WF dams. S u r v i v a l t o 60 d w as 4.6% h i g h e r f o r la m b s fro m 1 /4 F x dams (P<.05), and 7.6% h i g h e r f o r la m b s f r o m 1 /4 F x dams when a d j u s t e d f o r b i r t h w eig h t (P<.01). L i t t e r w e ig h ts a t 60 d and 120 d p e r ewe exposed w ere 3.9 kg and 4.3 kg h e a v i e r f o r 1/4Fx th a n WF ewes (P<.01). F le e c e s from 1/4Fx ewes ages I to 6 were l i g h t e r and c o a r s e r th a n th o se from WF ewes (PC.01), but had s i m i l a r s t a p l e le n g th s . I CHAPTER I . INTRODUCTION R eproductive e f f i c i e n c y o f th e ewe, t o t a l k ilo g ra m s o f lamb weaned p e r ewe i n im p o rta n t th e flo c k fa c to r ( S i d w e l l and M i l l e r , a ffe c tin g p ro fita b ility 19 7 1 a), is t h e m o st i n c o m m e r c i a l sh e e p p r o d u c t i o n s y s t e m s i n t h e U n i t e d S t a t e s ( D ic k e r s o n and G lim p , 1975; P arker and Pope, 1983; S id w e ll and M i l l e r , 1971a). L i t t e r s i z e , number o f la m b s p roduced p er p a rtu ritio n , r e p r o d u c t iv e e f f i c i e n c y (Bradford, r e g io n (Arizona, Colorado, Idaho, is a m a jo r com ponent of 1972a). L i t t e r s i z e i n th e Mountain Montana, Nevada, New Mexico, Utah, and Wyoming), which co m p rises 36% o f th e t o t a l sheep p o p u la tio n in, th e U.S., h a s i n c r e a s e d by o n ly .057 lam bs per. p a r t u r i t i o n (ap proxim ately 6 % ) from 1958 t o 1981 ( P a r k e r , 1981). Such l i m i t e d g r o w th i n any i n d u s t r y 's most im p o r ta n t economic t r a i t demands a t t e n t i o n from the r e s e a r c h community. R e p ro d u c tiv e e f f i c i e n c y may be i n c r e a s e d by im p r o v e m e n t s i n m an ag em e n t, n u t r i t i o n , and g e n e t i c s . C o n s i d e r a t i o n o f a l l o f t h e s e a r e a s i n t h e w e s te r n ra n g e la n d environm ent i s beyond th e scope of t h i s paper; e f f o r t s w i l l be c o n c e n tr a te d on improvement through th e use of g e n e tic s . New te c h n o lo g ie s loom on th e h o riz o n ; y e t , a t th e p re s e n t, two methods a r e a v a i l a b l e t o the com m ercial range sheep manager f o r g e n e t i c im p r o v e m e n t : (I). s e l e c t i o n among b r e e d i n g a n i m a l s b a s e d on in d iv id u a ls a n d /o r r e l a t i v e p erfo rm an ce and, (2) u t i l i z a t i o n of 2 g e n e t i c d i v e r s i t y between breeds i n c r o s s b r e e d in g schemes (Dickerson, 1969; T urner, 1969). Both o f t h e s e t e c h n i q u e s h a v e b e e n u n d e r s t u d y and i n p r a c t i c e u n d e r d i f f e r e n t m anagem ent s y s t e m s a r o u n d t h e w o r l d f o r many y e a r s ( N i t t e r , 1978). Improvements i n r e p r o d u c t iv e e f f i c i e n c y from s e l e c t i o n s t r a t e g i e s dep en d on t h e r e p e a t a b i l i t y and h e r i t a b i l i t y a s s o c i a t e d w ith th e components under s e l e c t i o n (Turner* 1969; F a lc o n e r, p a r t i a l l i s t o f components provided by B radford (1972) a t pu b erty , 1981). A i n c l u d e s : age frequency of p a r t u r i t i o n ( f e r t i l i t y and i n t e r v a l to f i r s t p o st-p artu m (p ro lific a c y ), e stru s), num ber o f la m b s produced per p a rtu ritio n s u r v iv a l of th e lambs, and le n g t h of r e p r o d u c t iv e l i f e . As t r a d i t i o n a l management in v o lv e s b re e d in g ewes f i r s t a s y e a r l i n g s and s u b se q u e n tly on an annual b a s is , c h o ic e s f o r s e l e c t i o n s t r a t e g i e s have focused on f e r t i l i t y and p r o l i f i c a c y . I n a r e v i e w , T u r n e r (1969) r e p o r t e d r e p e a t a b i l i t y f o r f e r t i l i t y a t .08 t o .09, and h e r i t a b i l i t y a t 0 t o .03. More r e c e n t l y , C la r k e and Hohenboken (1983) fo u n d t h e h e r i t a b i l i t y t o be .02 i n a c r o s s b r e d p o p u la tio n . E s ti m a t e s of such low m agnitude would s u g g e s t g a in s from s e l e c t i o n t o be slow. S h e lto n (1962) r e p o r t e d g a in s from c u l l i n g open 2 - y r - o l d s o f .05 la m b s p e r ewe b r e d p e r y r . T u r n e r (1966) r e p o r t e d g a i n s o f o n ly .005 and .059 i n tw o f l o c k s , c o n c l u d i n g t h a t s e l e c t i o n f o r f e r t i l i t y (or a g a i n s t i t s converse, b a rre n n e s s ) i s i n e f f e c t u a l . H e r i t a b i l i t y e s t i m a t e s f o r p r o l i f i c a c y a r e a l s o q u i t e low i n magnitude. I n a re v ie w , T urner (1969) r e p o r t e d an average e s t i m a t e to be .1 5 ; o t h e r v a l u e s h a v e b e e n .06 t o .16 i n w e s t e r n f i n e w ool sh e e p ( S h e l t o n and M e n z ie s , 1970; B a s u t h a k u r e t a l . , 1973), and .12 i n 3 c r o s s b r e d s ( C la r k e and H ohenboken, 1983). R e p e a t i b i l i t y e s t i m a t e s rev ie w e d by T u r n e r f a l l b o th a b o v e and below th e h e r i t a b i l i t y e s t i m a t e s . B r a d f o r d ( 1972a) s u g g e s t s t h a t a n e g a t i v e e n v i r o n m e n t a l c o r r e l a t i o n m ight e x i s t between c o n s e c u tiv e r e c o r d s due t o ewes who r a i s e d t w i n s b e i n g i n p o o r e r c o n d i t i o n a t m a t i n g t h a n ew es r a i s i n g o n ly I lamb. Again w ith such low e s t i m a t e s re s p o n s e to s e l e c t i o n w i l l be slow. Turner (1969) and B urfening (unpublished d a ta ) both r e p o r te d an ann u al d iv e rg e n c e p ro lific a c y of b e tw e e n l i n e s se le c te d fo r h ig h and low .023 lambs per ewe exposed. I n th e l a t t e r experim ent no d i f f e r e n c e s w e re fo u n d b e t w e e n h i g h and c o n t r o l l i n e s . (1972) r e p o r t e d on a New Z e a la n d l o n g - t e r m C la r k e s e l e c t i o n e x p erim en t i n i t i a t e d i n 1948. The h ig h l i n e , s e l e c t e d f o r m u l t i p l e lam bings, had an a n n u a l r a t e o f im p r o v e m e n t o f .018 t o .019 la m b s p e r ewe p e r y r above the c o n t r o l l i n e , s e l e c t e d f o r g e n e ra l appearance. As a r e s u l t o f t h e slow g a i n s due t o s e l e c t i o n , focused on u t i l i z a t i o n of g e n e tic d iv e rsity re s e a rc h has b e tw e e n breeds in c r o s s b r e e d i n g s c h e m e s f o r i n c r e a s i n g r e p r o d u c t i v e e f f i c i e n c y . The F in n is h Landrace (Finn) and th e Romanov have s u r f a c e d a s seem ingly th e most p r o l i f i c breeds (Bradford, 1972a). R eports of d i r e c t com parisons b e tw e e n t h e two b r e e d s a r e fe w . Goot e t a l . , s ig n ific a n t d iffe re n c e s in f e r t i l i t y (1979) r e p o r t e d and p r o l i f i c a c y no b e tw e e n Fl p ro g e n y p r o d u c e d by F in n and Romanov ra m s c r o s s e d w i t h M erino and Awassi ewes. However, d i f f e r e n c e s did e x i s t between progeny f o r wool t r a i t s . Romanov c r o s s e s had s i g n i f i c a n t l y lo w e r wool g ra d e s and h ig h e r in c id e n c e of kemp f i b e r s th a n th e Finn c r o s s e s , r e s u l t i n g i n a poorer q u a lity fle e c e . 4 P u re F i n n s a r e n o te d f o r t h e i r e a r l y age a t p u b e r t y and l a r g e litte rs. I n F in la n d , Finn ewe lam bs have an average 96.1% co n ce p tio n r a t e and h a v e l i t t e r s i z e s o f 1.80, 2 . 3 9 , and 2.75 a t 1 , 2 and 3+ y r of age, r e s p e c t i v e l y (Maij a l a and O stepberg, 1977). Under d i f f e r e n t B rita in , m anagem ent and e n v i r o n m e n t a l co n d itio n s in p u r e F in n ewe la m b s had a s i m i l a r c o n c e p t i o n r a t e , 93%; h o w e v e r, l i t t e r s i z e s w e r e c o n s i d e r a b l y h i g h e r a t 2 . 0 , 3. 0 , and 3.4 f o r a g e s I , 2 and 3+ y r , r e s p e c t i v e l y (D onald and Read, I 967). T hese r e s u l t s may be due to th e i m p o r t a t i o n o f sheep more i n t e n s e l y s e l e c t e d f o r p r o l i f i c a c y r a t h e r t h a n s h e e p m ore r e p r e s e n t a t i v e o f th e b r e e d . C ro ssb red F in n s i n B r i t a i n have a l s o shown s u p e r i o r p r o l i f i c a c y com pared t o t h r e e t r a d i t i o n a l c r o s s b r e d g r o u p s . At a g e s I , 2 and 3+ y r , f e m a l e p ro g e n y from F in n s i r e and S c o t t i s h B l a c k f a c e ew es had l i t t e r s i z e s o f 1.5, 2.0 , and 2.3, r e s p e c t i v e l y , w h i l e t h e c r o s s b r e d a v e r a g e s fro m B o r d e r L e i c e s t e r , Clun F o r e s t , and D o r s e t Horn s i r e s w e r e 1.1, 1.6, and 1.6, r e s p e c t i v e l y (D onald e t a l . , 1968). E v a l u a t i o n o f F in n c r o s s e s (Fx) i n t h e U.S. b e g a n i n I 970 a t t h e U.S. M eat and A nim al R e s e a r c h C e n t e r , Clay C e n t e r , N e b r a s k a . Under p a s tu r e c o n d i tio n s c r o s s e s w ere made u s in g Finn and R am b o u ille t s i r e s w ith s e v e n b r e e d s o f ew es. At I y r o f age Fx ew es had h i g h e r co n ce p tio n r a t e s th a n R a m b o u illet c r o s s e s (Rx), 85% vs 66%, and h ig h e r t h a n p u r e b r e d a v e r a g e , 58%. At a g e s I , 2, 3, and 4 y r l i t t e r s i z e s w e r e h i g h e r f o r Fx t h a n Rx, 1.55, 1.94, 2 .0 3 , and 2.25 vs 1.06, 1.38, 1.68, 1.51, r e s p e c t i v e l y , and h i g h e r th a n p u r e b r e d a v e r a g e , 1.08, I .3 2 , 1.52, 1.73, r e s p e c t i v e l y . O v e r a l l r e p r o d u c t i v e p e r f o r m a n c e f o r the Fx was 28% g r e a t e r th a n th e R am bouillet c r o s s e s (D ickerson, 1977). 5 Under w e s te r n range c o n d i tio n s Fx s u p e r i o r i t y over th e average of p u reb red Columbia, R am b o u illet, and Targhees i n co n c e p tio n r a t e a t I y r o f age w as e v e n m ore d r a m a t i c , 90.6% vs. 24.0%. L i t t e r s i z e was a l s o s i g n i f i c a n t l y h i g h e r f o r Fx e w e s , 1. 2 3 , 1.53 a t a g e s I , Knight, ' 2 , and 3+ y r , 1.53, 1.90, re sp e c tiv e ly 2.25 v s . 1.06, ( E r c a n b r a c k and unp u b lish e d d ata). The p r e li m i n a r y r e s u l t s c i t e d above i n d i c a t e t h a t u t i l i z a t i o n of c r o s s b r e e d i n g s c h e m e s i n v o l v i n g F in n b r e e d i n g may be an a t t r a c t i v e a lte rn a tiv e for im p ro v in g c o n d itio n s. The p r i m a r y re p ro d u c tiv e o b je c tiv e of e ffic ie n c y th is stu d y is under to ran g e com pare r e p r o d u c t iv e e f f i c i e n c y and wool p ro d u c tio n o f ewes composed o f 1/4 F in n b r e e d i n g w i t h s t r a i g h t b r e d C o lu m b ia , R a m b o u i l l e t , and T a rg h e e ewes on w e s te r n range. The secondary o b j e c t i v e i s t o a n a ly z e breed of s i r e e f f e c t s (F in n x R a m b o u i l l e t v s . s t r a i g h t b r e d r a m s ) k ilo g ra m s o f lamb weaned p e r ewe lambing. on t o t a l 6 CHAPTER 2 LITERATURE REVIEW F e rtility In t h i s paper, fe rtility i s d e f i n e d a s t h e p r o p o r t i o n o f ew es l a m b i n g r e l a t i v e t o ew es e x p o s e d t o t h e ram ( N i t t e r , 1978). I n a review of Finn c r o s s b r e d perform ance i n numerous c o u n t r i e s , M a ija la and O s t e r b e r g ( 1977) fo u n d F in n b r e e d i n g c o n t r i b u t e d t o i n c r e a s e d f e r t i l i t y o n l y i n ewe la m b s (ew es l a m b i n g a t a p p r o x i m a t e l y I y r of age). I n B r i t a i n , B arker (1975) r e p o r t e d s l i g h t l y n e g a t i v e , a l t h o u g h non—s i g n i f i c a n t , d i f f e r e n c e s i n f e r t i l i t y f o r m ature ewes between Finn c r o s s e s and p u r e b r e d s . T hese r e s u l t s a r e i n a g r e e m e n t w i t h s t u d i e s c o n d u c t e d u n d e r s e m i - i n t e n s i v e m anagem ent i n t h e U.S. in v o lv in g 1/4Finn c r o s s e s (1/4Fx) compared w ith D orset x R am b o u illet (Thomas and W hitem an, 1979); and I / 2 F in n ( I / 2Fx) compared w ith Border L e i c e s t e r c r o s s e s ( O l t e n a c u and B o y la n , 1981a). H ow ever, u n d e r w e s t e r n r a n g e c o n d itio n s at H o p elan d , C a lifo rn ia , 1 /2 F x (x W h i t e f a c e ) have d em o n strated s i g n i f i c a n t l y h ig h e r f e r t i l i t y a t ages 2 th ro u g h 5 than 1 /2 S u ffo lk (x W h ite face ), 98$ vs. 88$, r e s p e c t i v e l y (D ickerson, 1977). E r c a n b r a c k and K n ig h t ( u n p u b l i s h e d d a t a ) r e p o r t s i m i l a r r e s u l t s , a lth o u g h not s i g n i f i c a n t , b e t w e e n 1/2 F x (x W h iteface) and W hiteface b reed s ag es 2 through 7 on w e s te r n range. F e r t i l i t y i n ewe la m b s i s l a r g e l y a f u n c t i o n o f age a t p u b e r t y (N itte r, 1978). Puberty may be d e f in e d a s t h a t p o in t when r e p r o d u c t io n 7 i s f i r s t p o s s i b le . Age a t p u berty i s u s u a l l y determ in e d by o b s e r v a tio n o f f i r s t b e h a v io r a l e s t r u s (D ickerson and L a s t e r , 1975; Drymundssonf 1981). A l i s t o f f a c t o r s a f f e c t i n g age a t p u b e r t y w ould a t l e a s t in c lu d e breed, n u tritio n , breed-cross, year, d a y lig h t, and d a te o f b i r t h , te m p e ra tu re w eig h t a t breeding, (L a ste r et a l., 1972; Drymundsson, 1981). Review of each of th e s e a r e a s i s beyond the scope o f t h i s paper, e f f o r t s w i l l be c o n c e n tr a te d on breed and b r e e d - c r o s s effects. Age a t p u b e r t y i n ewe la m b s i s n o t o n ly o f i n t e r e s t f o r t h e p o t e n t i a l o f r e d u c i n g t h e num ber o f n o n - p r o d u c t i v e ew es i n a f l o c k , b u t a l s o f o r i t s e f f e c t s on l i f e t i m e r e p r o d u c t i v e p e r f o r m a n c e . On w e s t e r n r a n g e ewe la m b s who d i s p l a y b e h a v i o r a l e s t r u s d u r i n g t h e i r f i r s t b re e d in g season, but bred to lamb f i r s t a t 2- y r - o l d , have been r e p o r te d to be more p ro d u c tiv e a t su b se q u en t ages th a n lam bs f a i l i n g t o c y c l e ( H u l e t e t a l . , I 969; B u r f e n i n g e t a l . , 1972). S e v e ra l s t u d i e s have r e p o r t e d s m a ll s i g n i f i c a n t breed d i f f e r e n c e s i n ewe lambs o b ta in in g p u b erty i n t h e i r f i r s t b re e d in g " s e a s o n (Hulet e t a l . , 1969; S o u th a n e t a l . , 1971; B u r f e n i n g e t a l . , 1974). Due t o w id e v a r i a t i o n s i n m anagem ent and e n v i r o n m e n t a t v a r i o u s r e s e a r c h s ite s, l a r g e d i f f e r e n c e s e x i s t b e t w e e n r e p o r t e d m eans among and b e tw e e n b r e e d s (D rym undsson, 1981). R e s u l t s i n t h e l i t e r a t u r e may r a n g e fr o m 0% i n e s t r u s d u r i n g f i r s t b r e e d i n g s e a s o n f o r M erin o ewe la m b s i n A u s t r a l i a (W atson and Gamble, 1961), and 24% f o r C olum bia, R am b o u illet, and Targhee ewe lambs on w e s te r n range (fiurfening e t a l., 1972), t o 90.0% and 51.4% ewe la m b s l a m b i n g a t 12 mo o f age f o r 8 S u f f o l k and T a r g h e e u n d e r s e m i - i n t e n s i v e m anagem ent ( O l t e n a c u and Boylan, 1981a). S tu d ie s in v o l v in g pure F inns i n d i c a t e c o n s i s t e n t l y high f e r t i l i t y i n ewe la m b s i n v a r i o u s e n v i r o n m e n t s . M a i j a l a (1977) r e p o r t s i n F i n l a n d t h a t o v e r 90% o f t h e F in n ewe la m b s a r e b r e d t o la m b a t I y r of age, and lam bs on a h ig h l e v e l of n u t r i t i o n breed a t 5 mo. S i m i l a r i n d i c a t i o n s o f f e r t i l i t y were found i n S c o tla n d where 97% o f th e pure F in n ewe l a m b s e x h i b i t e d e s t r u s a t 6 mo o f ag e (Land e t a l . , 1974). R ep o rts o f s u ch l a r g e breed d iffe re n c e s in age a t p u b e rty and f e r t i l i t y i n ewe lam bs h a s l e d r e s e a r c h e r s t o ask "how” and "to what degree" w i l l in c o rp o ra tio n o f F in n i n t o sta n d a rd breeds a f f e c t f e r t i l i t y i n c r o s s b r e d progeny. I n a s t u d y i n v o l v i n g F in n c r o s s e s (Fx) and R a m b o u i l l e t c r o s s e s (Rx) w i t h s e v e n s t a n d a r d b r e e d s ( S u f f o l k , H a m p s h ire , R a m b o u i l l e t , D o r s e t , T a r g h e e , C o r r i e d a l e , and C o a rs e W ool), D i c k e r s o n and L a s t e r (1975) r e p o r t e d s i g n i f i c a n t l y lo w er age a t p u b erty f o r Fx over Rx, 219 d and 238 d, r e s p e c t i v e l y . I n t h i s s t u d y la m b s w e r e b o r n e i t h e r i n F e b r u a r y o r A p r i l and e x p o s e d t o v a s e c t b m i z e d ra m s f o r a b o u t 10 wk p r i o r to November 10th. No breed by d a te i n t e r a c t i o n was r e p o r te d . Fx la m b s w e r e 4 kg l i g h t e r a t p u b e r t y , b u t d i d n o t w e ig h s i g n i f i c a n t l y l e s s th a n Rx a t 230 d o f age. The number of Fx lambs r e a c h in g puberty by November IO th g r e a t l y e x c e e d e d t h e Rx la m b s , resp e c tiv e ly . 70.2% v s 36.4%, While acknow ledging high r e p o r t s of f e r t i l i t y by Donald and Read (1 9 6 7 ), D i c k e r s o n and L a s t e r s u g g e s t e d h e t e r o s i s p l a y e d a l a r g e r r o l e th a n average gene e f f e c t s i n th e s u p e r i o r i t y of Fx lambs. T h is s u g g e s t i o n was b a s e d on d a t a i n d i c a t i n g 41% m ore 1 /2 F x r e a c h e d 9 p u b e rty by November IO th t h a n 3 /4 F x ( 3 /4 F x w e r e n o t m e n tio n e d e l s e w h e r e i n t h e s t u d y ) . Due t o a l a c k o f p u r e F i n n s , a h e t e r o s i s e s t i m a t e was a v a i l a b l e only f o r Rx: 8$, n o n - s i g n i f i c a n t . On the c o n tr a r y , C e d illo and Hohenboken (1977) found no d i f f e r e n c e i n p e rc e n ta g e r e a c h in g p u b e rty or average age a t f i r s t e s t r u s between S u f f o l k and C o lu m b ia ewe la m b s s i r e d by F in n ra m s v s t h o s e s i r e by D orset, Targhee, C o r r ie d a le, and Coarse Wool rams. But, i n t h i s same study a l a r g e d i f f e r e n c e i n f e r t i l i t y between Fx and s ta n d a r d c r o s s e s was observed, 72% and 38%, r e s p e c t i v e l y . While th e l i t t l e d a t a a v a i l a b l e on th e a f f e c t of Finn b re e d in g on age a t p u b e r t y i s c o n t r a d i c t o r y , much e v i d e n c e e x i s t s u p p o r t i n g Fx su p e rio rity fo r f e r t i l i t y i n ewe lambs over purebred s ta n d a rd breeds (see t a b l e I). Unweighted av e ra g e s o f f e r t i l i t y f o r Fx ( e i t h e r 1/2 or* 1 /4 ) and s t a n d a r d p u r e b r e d s , w i t h w h ic h t h e F i n n s w ere c r o s s e d , a r e l i s t e d w i t h i n th e r e s p e c t i v e s t u d ie s . Only i n one study, Magid e t a l. (1 9 8 1 b ), w e re Fx n o t r e p o r t e d s i g n i f i c a n t l y s u p e r i o r . T h is s t u d y co m p ared ewe la m b s s i r e d by e i t h e r B o r d e r L e i c e s t e r o r F in n r a m s ; purebred com parisons were n o t r e p o r te d . B ased on s i m i l a r l e v e l s f o r f e r t i l i t y f o r p u r e F i n n s (81%) and F in n x M erin o (76%), Land e t a l . (1974) c o n c lu d e d t h a t t h e t r a i t show ed h e t e r o s i s (87.7%) a t l e a s t " s u p e r f i c i a l l y " . I n t h e o n ly s tu d y i n v o l v i n g p u r e F i n n s i n t h e U.S., O l t e n a c u and B o y la n (1 981 a) a l s o r e p o r t e d s i m i l a r l e v e l s o f f e r t i l i t y b e t w e e n p u r e F i n n s (95.0%) and F in n s c r o s s e d w i t h T a r g h e e , M in n e s o ta 100, and S u f f o l k : 9 4 .7, 94.1, and 87.0,. r e s p e c t i v e l y . Reported l e v e l s o f f e r t i l i t y f o r th e Targhee, M in n e s o ta 100, and S u f f o l k w e r e 5 1 .4 , 7 4 .7, and 90.0, r e s p e c t i v e l y ; TABLE I . Finn Breeding 1/2 1/2 1/2 1/2 1/2 1/2 1/4 1/4 1/4 • COMPARISON OF 1 / 2 AND 1 / 4 FINN CROSSES WITH STANDARD BREEDS FOR F E R T IL IT Y IN EWE LAMBS S tandard Breeds Involved i n Crosses F e r t i l i t y (?) Fx S td. R eference Ml 00, S u f f o lk , Targhee Columbia, R am bouillet, Targhee S u f f o lk , R am bouillet, Targhee C o r r ie d a le , D orset, Hampshire Coarse Wool Tasmanian Merino Hampshire, R a m b o u ille t,Targhee D o rse t, Targhee, C o rrie d a le Coarse Wool D orsetxR am bouillet Columbia, R am bouillet, Targhee Ml 00, S u f f o lk , Targhee 91.9 90.6 84 71.7 24.0 51 O ltenacu and Boylan (1981 a) Ercanbrack and Knight (unpublished d a t a ) L a s t e r e t a l . , (1972) 81 78 72 5 78* 38 Land e t a l . (1974) Magid e t a l . (1981b) C e d illo and Hohenboken (1977) 80.5 76.1 79.2 56.5 24.0 71.7 Thomas and Whiteman (1979) Ercanbrack and Knight (unpublished d a ta ) O ltenacu and Boylan (1981 a) E s tim a te e q u a ls average of Finn and Border L e i c e s t e r c r o s s e s . Authors r e p o r te d no s i g n i f i c a n t d i f f e r e n c e between Finn and Border L e i c e s t e r c r o s s ewe lam bs; purebred e s ti m a t e s not r e p o r te d . 11 e s t i m a t e s o f h e t e r o s i s w ere 29.5, 10.8, and -,5.9, r e s p e c t i v e l y . These r e s u l t s combined w ith th o se from Land e t a l . (1974) p o in t t o a s tr o n g l i n k between i n c r e a s i n g e s t i m a t e s o f h e t e r o s i s f o r Finn c r o s s e s and d e c r e a s in g f e r t i l i t y i n th e p u re b re d s i n c o r p o r a te d i n th e cro ss. P ro lific a c y In a d d i t i o n t o e a r l y age a t pu b erty , pure F inns a r e most noted f o r e x c e p t i o n a l l y h ig h p r o l i f i c a c y ( l i t t e r s i z e ) , num ber o f la m b s b o rn (dead or a l i v e ) p e r ewe lambing. The average l i t t e r s i z e f o r ewe lambs i n F i n l a n d a v e r a g e s fro m 1.6 t h r o u g h 1.90 (Goot and M a i j a l a , M a i j a l a and O s t e r b e r g , 1 977; 1977) i s w e l l above m a t u r e s h e e p o f any s t a n d a r d b r e e d i n t h e U.S. M a i j a l a and O s t e r b e r g (I 977) r e p o r t t h a t t h e l a r g e l i t t e r s i z e s c o n t i n u e t o i n c r e a s e w i t h age up t o 3 y r s o f age., w i t h a v e r a g e s o f 2.39 and 2.75 f o r a g e s 2 and 3 y r . From ag e 3 onto a t l e a s t 8 y r old th e l i t t e r s i z e re m a in s c o n s ta n t. U n w e ig h te d m eans o f p u r e F in n p e r f o r m a n c e i n o t h e r E u ro p e a n c o u n t r i e s (not i n c lu d in g B r i t a i n ) , under v a r io u s management system s, ag reed w ith th e r e s u l t s in F in la n d , a l t h o u g h t h e r e w as a s l i g h t d e c l i n e a t 6 y r o f age. L i t t e r s i z e s w e r e 1.84, 2 .4 5 , and 2.82 f o r a g e s I , 2 and 3+ y r , r e s p e c t i v e l y ( M a i j a l a and O s t e r b e r g , 1977). I n B rita in , Donald and Read (1967) r e p o r t e d s i m i l a r r e s u l t s o f in c r e a s in g l i t t e r s i z e t o 3 y r o f a g e , 2 .0 , 3 .0 , 3 .3 , 3 .4 , 3.4 f o r ew es a g e s I , 2, 3, 4 and 5 y r , r e s p e c t i v e l y . I n t h e U.S., r e s u l t s f o r p u r e F in n s a r e l i m i t e d t o one s t u d y r e p o r t i n g o n ly t h e f i r s t 3 y r . Yet th e d a t a i s i n c l o s e a g r e e m e n t w i t h t h o s e r e p o r t e d by M a i j a l a and O s t e r b e r g (1977); average l i t t e r s i z e was 156.8, 271.0, and 300.4 lambs born per 12 10O ew es e x p o s e d f o r a g e s 1 , 2 and 3 y r , r e s p e c t i v e l y ( O l t e n a c u and Boylan, 1981a). Of major concern t o com m ercial sheep managers i s how w e ll t h i s breed s u p e r i o r i t y i s t r a n s m i t t e d to progeny i n c r o s s b r e e d in g schemes. The f i r s t F in n c r o s s b r e e d i n g p r o j e c t i n B r i t a i n d e m o n s t r a t e d c l e a r s u p e r i o r i t y f o r l i t t e r s i z e o f 1 /2 F x o v e r s t a n d a r d b r e e d s . Fx ew es a v e r a g e d 1.52, 2. 0 2 , and 2.30 la m b s b o r n p e r ewe l a m b i n g a t I , 2 and 3 y r of age; and, d i f f e r e n c e s between Fx and s ta n d a rd b reed s in c r e a s e d w ith age: .4 8 , .5 5 , and .79 a t a g e s I , 2 and 3 y r , re sp e c tiv e ly (D onald e t a l . , 1968). A l a t e r s tu d y i n B r i t a i n i n v o l v i n g d i f f e r e n t b r e e d s fo u n d t h e same p a t t e r n . But t h e d i f f e r e n c e s w e r e s m a l l e r and d i d n o t i n c r e a s e a s d r a m a t i c a l l y w i t h ag e . Fx m inus s t a n d a r d b r e e d (Border L e i c e s t e r , S u ffo lk , and Clun F o r e s t) average was .30, .32, .39 f o r ages I , 2 and 3, r e s p e c t i v e l y (B arker, 1975). S e v e r a l e x p e r i m e n t s c o n d u c t e d i n t h e U.S. have fo u n d s i m i l a r r e s u l t s t o t h o s e i n B r i t a i n . L a s t e r e t a l . (1972) r e p o r t e d on. ewe la m b s from cro sses in v o lv in g seven b r e e d s o f dam and e i t h e r R a m b o u i l l e t o r F in n s i r e s . Fx ew es la m b s h ad .58 m ore la m b s p e r ewe l a m b i n g t h a n Rx ewe la m b s and .52 m ore t h a n p u r e b r e d . C e d i l l o e t a l . (1977) r e p o r t e d a d i f f e r e n c e o f .44 b e tw e e n Fx and s t a n d a r d c r o s s e s ( D o r s e t , C h e v i o t , and Romney c r o s s e d w i t h C o lu m b ia and S u f f o l k ) ewe la m b s . I n a more e x t e n s i v e s t u d y , O l t e n a c u and B o y la n (1 9 8 1 a ) fo u n d l a r g e and s h a r p ly i n c r e a s i n g d i f f e r e n c e s i n l i t t e r s i z e betw een Fx and th e average of M in n e so ta 100 (Ml 0 0 ) , S u ffo lk , and T arghee p u r e b r e d s : .5 1 , .7 5 , and .86 a t a g e s I , 2 and 3. y r , r e s p e c t i v e l y . 13 Two s t u d i e s under h arsher c o n d itio n s on t h e w e s te r n ran g e d e m o n stra te t h a t Fx s u p e r i o r i t y f o r p r o l i f i c a c y i s n o t r e a l i z e d only i n s e m i - i n t e n s i v e m anagem ent s c h e m e s. At H o p land, C a l i f o r n i a t h e average d i f f e r e n c e f o r ag es 2 through 5 between 1/2Fx and w h ite breeds w as .60 ( D i c k e r s o n , 1977). E r c a n b r a c k and K n ig h t ( u n p u b l i s h e d d a t a ) r e p o r t e d l a r g e d i f f e r e n c e s b e t w e e n 1 /2 F x and s t a n d a r d w h i t e - f a c e breeds (Columbia, Targhee, and R am b o u illet) on w e s te r n r a n g e . . Fx ewes e x c e e d e d th e s t a n d a r d b r e e d s by .47 a t I y r o f ag e. S u b s e q u e n t b r e e d group d i f f e r e n c e s were n e a r l y i d e n t i c a l , .71 f o r ages 2 through. 7 yr. S i m i l a r i t y b e tw e e n r e s u l t s u n d e r w e s t e r n r a n g e and s e m i - i n t e n s i v e m an ag em e n t, a r e i n a g r e e m e n t w i t h c o n c l u s i o n s d raw n by Meyer and Bradford (1973). Finn x Targhee and Targhees both on high and low feed l e v e l s d em o n strated no t r e a tm e n t x genotype i n t e r a c t i o n f o r f e r t i l i t y and p r o l i f i c a c y . The r e s e a r c h e r s c o n c l u d e d t h a t f e e d l e v e l s a r e o f l i t t l e im p o rtan ce f o r e v a l u a t i o n of r e p r o d u c tiv e t r a i t s . R e s u lts from s t u d i e s in v o lv in g 1/4Fx ewes a t v a r io u s ag es show a r e d u c t io n i n p r o l i f i c a c y from 1/2Fx; however, 1/4Fx w ere s t i l l found to be s u p e r i o r to the p u reb red s. The s m a l l e s t breed group d if f e r e n c e , was r e p o r t e d by Thomas and W h item a n (1979) u n d e r s e m i - i n t e n s i v e management f o r ewe lam bs. 1/4Fx average l i t t e r s i z e was 1.1.7 compared to 1.08 f o r R a m b o u i l l e t x D o r s e t s , a n o n -s ig n ific a n t d iffe re n c e o f .09. Also under s e m i - i n t e n s i v e management, 1/4Fx ewes, produced by b a c k c r o s s i n g 1 /2 F x w i t h t h e s t a n d a r d b r e e d s , w ere s u p e r i o r t o t h e s t a n d a r d b r e e d s . 1/4 F x ewe la m b s s i g n i f i c a n t l y e x c e e d e d s t a n d a r d s by .2 4 ; w h ile l i t t e r s i z e s from 2 y r o l d 1/4 F x w e r e .12 h i g h e r , alth o u g h n o n - s i g n i f i c a n t ( O ltenacu and Boylan, 1981), 14 Iro n ic a lly , under h a r s h e r w e s te r n range management, o v e r a l l breed means f o r both 1/4Fx and sta n d a rd b reed s w ere much h ig h e r than under s e m i - i n t e n s i v e management. I n a d d i t i o n , d i f f e r e n c e s betw een 1/4Fx and s t a n d a r d b r e e d s w e re g r e a t e r i n m a g n itu d e . A c r o s s a g e s 2 t o 5 y r a t Hopland, l i t t e r s i z e s were .32 h ig h e r f o r 1/4Fx (D ickerson, 1977). The d i f f e r e n c e r e p o r t e d by E r c a n b r a c k and K n ig h t ( u n p u b l i s h e d d a t a ) f o r ewe lambs was h a l f as g r e a t , .16, but s i g n i f i c a n t . The amount by which I / 4Fx e x c e e d e d t h e s t a n d a r d b r e e d s a t l a t e r a g e s (2 th o u g h 7 y r ) rem ained c o n s ta n t, Cl e a r l y , .38 average. t h e F in n h a s t h e a b i l i t y to t r a n s m i t i t s s u p e rio r p r o l i f i c a c y to c r o s s b r e d progeny under v a r y in g management system s and a t d e g r e e s o f b r e e d i n g a s .l o w a s 1 /4 . The q u e s t i o n now m ust be a s k e d a s t o what g e n e t i c mechanism i s t h i s d i f f e r e n c e a t t r i b u t e d - h e t e r o s i s or a d d i t i v e gene e f f e c t ? In an ex p erim en t in v o lv in g pure F in n s, M e r in o s , and t h e i r r e c i p r o c a l c r o s s e s , average p r o l i f i c a c y f o r ag es 2 through 4 was 2.68, 1. 0 0 , and 1.70, r e s p e c t i v e l y . E s t i m a t e d h e t e r o s i s , c r o s s b r e d mean m in u s t h e p a r e n t a l mean a s a p r o p o r t i o n o f th e p a r e n t a l mean, was ■7.6%. T h erefo re, be a t t r i b u t e d no h e t e r o s i s was p r e s e n t and th e s u p e r i o r i t y could t o a d d i t i v e g en e e f f e c t s (Land e t a l., 1974). r e s u l t s o f O ltenacu and Boylan (1981) a r e i n s tr o n g agreem ent. d ata d e m o n stra te d sig n ific a n t. h e te ro sis ( 10. 1%) a t I yr of age, but The Their non­ However, a t ages 2 and 3 y r h e t e r o s i s was z e ro . The mean o f t h e 1/4 F x a t I y r o f a g e f e l l a l m o s t e x a c t l y h a l f way b e tw e e n th e m id -p a re n t v a lu e and th e s ta n d a rd breed average, 1.01 (1/4Fx) and 0.97 15 (m id -p a re n t). However, a t age 2 yr, th e 1/4Fx were lo w e r th a n th e mid­ p a r e n t v a lu e , but n o n - s i g n i f i c a n t . There do e x i s t r e p o r t s i n th e l i t e r a t u r e of p o s i t i v e h e t e r o s i s f o r p r o l i f i c a c y (T urner, 1969; S id w e ll and M i l l e r , 1971a); however, most average about 3% and a r e n o n - s i g n i f i c a n t ( H i t t e r , e s t i m a t e t h i s a u t h o r r e v i e w e d , 30%, s ig n ific a n t 1978). The h ig h e s t i n v o l v e d Fx e w es, b u t was non­ (H ohen b o k en and C o c h ra n , 1 97 6 ). T h erefo re, w h ile d i f f e r e n c e s between Fx and s ta n d a rd b re e d s may v ary w ith environm ent and age, th e com m ercial producer may ex p ec t, i n g e n e r a l, l i t t e r s i z e s t o i n c r e a s e by t h e same p r o p o r t i o n o f F in n b r e e d i n g due t o a d d i t i v e gene e f f e c t s . S u rv iv a b ility U tiliz a tio n o f F in n b r e e d i n g t o i n c r e a s e p ro lific a c y may be unw arranted i f c o r r e s p o n d in g l a r g e d e c r e a s e s i n s u r v i v a b i l i t y fro m b irth t o w e a n in g e x i s t . F acto rs a f f e c tin g s u r v iv a b ility (o r th e converse, m o r t a l i t y ) i n lam bs a r e n u m e ro u s and o f t e n u n a c c o u n t a b l e . Any l i s t o f f a c t o r s w o u ld i n c l u d e b r e e d , b i r t h w e i g h t o f t h e la m b , n u t r i t i o n (p re-p artu m and p o s t-p a rtu m ), l i t t e r s i z e , age of ewe, sex, season, and management system (B radford, 1972b; Jakubee, 1977). A ll of th e s e f a c t o r s a r e im p o r ta n t to the com m ercial sheep p ro d u cer; however, t h i s r e v i e w w i l l c o n c e n t r a t e on t h e g e n e t i c f a c t o r s : b i r t h weigh];, l i t t e r size, and breed. S u b s t a n t i a l e f f e c t s o f l i t t e r s i z e on p o s t-p a rtu m lamb m o r t a l i t y have been r e p o r t e d i n th e l i t e r a t u r e under v a r io u s environm ents. Under adverse c o n d i tio n s tw in s have been found t o have 68% h ig h e r m o r t a l i t y r a t e s than s i n g l e s (Sm ith, 1974). But, under more i n t e n s i v e management 16 i n w h ic h m o r t a l i t y r a t e s a r e l o w e r , t w i n s may h a v e i n e x c e s s o f 80$ h ig h e r m o r t a l i t y th a n s i n g l e s ( S id w e ll e t a l . , 1962; S i d w e l l and M i l l e r , 1971a; O l t e n a c u and B o y la n , 1981). The e f f e c t i s e v e n m ore d r a m a t i c f o r la m b s b o r n a s t r i p l e t s , m o r t a l i t y th a n tw in s and s i n g l e s , o v e r 50$ and 2 00$ h i g h e r r e s p e c t i v e l y (O ltenacu and Boylan, 1981). I n a w e ll-p la n n e d study under two env iro n m en ts ( s e m i - i n t e n s i v e and open p a s tu r e ) , P u rse r and. Young (1964) concluded t h a t m u l t i p l e b i r t h s had no d i r e c t e f f e c t on s u r v i v a b i l i t y , but r a t h e r b i r t h w e ig h t was th e m a j o r f a c t o r . Lambs o f e q u a l b i r t h w e i g h t , r e g a r d l e s s o f ty p e of b irth , had equal s u r v i v a l . Numerous s t u d i e s have r e p o r t e d e f f e c t s of l i t t e r s i z e on i n d i v i d u a l b i r t h w e ig h ts ; tw in s w eighing on average 15 t o 20$ l e s s t h a n s i n g l e s ( P u r s e r and Young, 1964; B r a d f o r d , 1972b; S i d w e l l and M i l l e r , 1971b, Magid e t a l . , 1981a), and t r i p l e t s 30 to 35$ l e s s t h a n s i n g l e s ( R a s t o g i e t a l . , 1975; Magid e t a l . , 1981a; O ltenacu and Boylan, 1981b) As p u r e F i n n s h a v e much h i g h e r l i t t e r s i z e s t h a n m o s t s t a n d a r d breeds, b i r t h w e i g h t s and s u r v i v a b i l i t y w ould be e x p e c t e d t o be s u b s t a n t i a l l y lo w e r . M a i j a l a and O s t e r b e r g ( 1977) r e p o r t e d av erage, b i r t h w eig h t of pure F inns a c r o s s a l l ages, and a t v a r i o u s l o c a t i o n s , t o be 2.57 kg. A lth o u g h n o t an a c c u r a t e c o m p a r is o n , b i r t h w e i g h t s i n t h e U.S. a v e r a g e 4 t o 5 kg ( S i d w e l l and M i l l e r , 1971b; R a s t o g i , 1975; D ickerson e t a l . , 1975). D e s p ite th e much lo w er b i r t h w e ig h ts , M a ija la and O s t e r b e r g ( 1977) r e p o r t e d m o r t a l i t y a c r o s s a l l a g e s o f dams and litte r s i z e s , b e t w e e n b i r t h and 2 wk o f ag e , t o be 10 t o 15$. The e f f e c t s o f l i t t e r s i z e and age o f dam w e re s i g n i f i c a n t . M o rta lity 17 r a t e s o f lambs from y e a r l i n g s were tw ic e a s high as t h a t from m ature ewes a t every l i t t e r s iz e . Lambs born a s q u i n t u p l e t s t o y e a r l i n g s had m o r t a li ty - r a t e s i n e x c e ss o f 70/6. Donald and Read (1967) r e p o r t e d s i m i l a r e s t i m a t e s f o r b i r t h w eight o f p u r e F in n la m b s i n B r i t a i n . A verage b i r t h w e i g h t f o r s i n g l e s and t w i n s was 2.8 kg and 2.2 kg, r e s p e c t i v e l y . I n c l u d i n g la m b s dead a t b i r t h , F in n s had 38/6 m o r t a l i t y fro m b i r t h t o w e a n in g . D e s p i t e s e m i i n t e n s i v e m a n ag em e n t, m o r t a l i t y f o r la m b s b o rn a l i v e fro m b i r t h to w e a n in g w as 255», a v e r a g e d a c r o s s a l l a g e s o f dam and l i t t e r s i z e s . They a t t r i b u t e d th e high death l o s s e s t o low b i r t h w e ig h ts. B ir th w e ig h ts fo r p u r e F in n s h e e p i n t h e U.S. are in c lo se agreem ent w ith th e r e s u l t s r e p o r t e d i n F in la n d and B r i t a i n . I n d iv id u a l pure Finn lam bs from ewes had an average b i r t h w eig h t o f 2.60 kg, 1.00 (28/6) kg l e s s t h a n t h e s t a n d a r d b r e e d s (Ml 00, T a r g h e e , and S u f f o l k ) a c r o s s ages I , 2 and 3 y r (Oltenacu and Boylan, 1981b). As th e s e means a r e a d ju s te d f o r type of b i r t h and p u re Finns had much l a r g e r l i t t e r s i z e s , th e a c t u a l d i f f e r e n c e i n means would be l a r g e r . The d i f f e r e n c e r e p o r t e d i n t h i s s t u d y may r e f l e c t t h e s m a l l e r m a t u r e w e i g h t o f t h e F in n ewe, a l t h o u g h F in n la m b s a r e s m a l l r e l a t i v e t o m a t u r e ewe s i z e ( B r a d f o r d , 1972b). I n c o r p o r a t i o n o f F in n b r e e d i n g i n t o s t a n d a r d b r e e d s h a s a l s o served to reduce b i r t h w e ig h ts a t a l l ages. In B r i t a i n , 1 /2 F x ew es w ere found t o have c o n s i s t e n t l y lo w er b i r t h w e ig h ts th a n purebreds a t ag es I , 2 and 3 yr. U nadjusted f o r type o f b i r t h the d i f f e r e n c e was on a v e r a g e .5 kg (12%) l e s s t h a n p u r e b r e d a v e r a g e (D onald e t a l . , 1968; B ark er, 1975). 18 Most o f t h e e s t i m a t e s o f d i f f e r e n c e s i n t h e U.S. f a l l b e tw e e n .3 and .9 kg l e s s f o r t h e 1/2 F x t h a n s t a n d a r d b r e e d s ( D i c k e r s o n e t a l . , 1 975; Magid e t a l . , I 981; O l t e n a c u and B o y la n , 1981, C o c h ra n e t a l . , 1982, E r c a n b r a c k and K n ig h t, u n p u b lish e d d a ta ). B ut one r e p o r t i n v o l v i n g F i n n x Panama (FP) and Panama (P) p u r e b r e d s f o u n d S u f f o l k s i r e d la m b s fro m FP ew es t o be s i g n i f i c a n t l y h e a v i e r a t b i r t h th a n la m b s fro m P ew es by .89 kg (Dahmen e t a l . , 1 979). The s t u d y d i d n o t r e p o r t i f type of b i r t h was in c lu d e d i n th e model. E s ti m a t e s f o r IA F x a r e i n t e r m e d i a t e between 1/2Fx and p u re b re d s , ran g in g from .3 to .6 kg l e s s t h a n t h e p u r e b r e d s ( O lte n a c u and B o y la n , 1981; C o c h ra n e t a l . , 1992; Ercanbrack and Knight, u npublished d a ta ). B ased on t h e l o w e r e s t i m a t e s o f b i r t h w e i g h t f o r l a m b s fro m Fx dams, one would ex p ec t s u r v i v a b i l i t y to be lo w er as w e ll. But, d e s p i t e sig n ific a n t d iffe re n c e s in b irth w e ig h t, th is au th o r fo u n d no e s t i m a t e s o f s i g n i f i c a n t l y lo w er s u r v i v a b i l i t y i n lambs from 1/2Fx or I M F x ew es a g e 2 y r o r o l d e r when ty p e o f b i r t h was n o t i n c l u d e d i n th e model (Donald e t a l . , 1968; B arker, 1975; D ickerson, 1977; Thomas and W h item a n , 1979; C o ch ran e t a l . , I 982). Only two s t u d i e s r e p o r t e d s i g n i f i c a n t l y l o w e r s u r v i v a b i l i t y f o r la m b s fro m Fx y e a r l i n g dams. D onald e t a l . (1969) and B a r k e r (1975) b o th r e p o r t e d a b o u t 9% l o w e r s u r v i v a b i l i t y f o r Fx th a n p urebreds when weaned a t about 90 d of age. Although s t a t i s t i c a l l y n o n - s i g n i f i c a n t , s i m i l a r r e s u l t s w ere r e p o r te d by D ickerson (1977) and Thomas and Whiteman (1979). C o n f li c ti n g , but s t a t i s t i c a l l y n o n - s i g n i f i c a n t r e s u l t s w e r e fo u n d by E r c a n b r a c k and Knight (unpublished d a ta ) when lambs w ere r e a r e d on w e s te r n range and weaned a t a p p ro x im a te ly 120 d of age . Lambs from 1/2Fx y e a r l i n g ewes 19 had 3/5 h ig h e r s u r v i v a b i l i t y th a n purebreds. These r e s u l t s may b ia s Fx s u r v i v a b i l i t y upwards due t o th e management procedure which c r e d i t s a ewe w i t h h a l f a la m b when s u c c e s s f u l l y r e a r e d by a n o t h e r ewe o r a rtific ia lly . E s t i m a t e s o f s u r v i v a b i l i t y u s i n g m o d e ls w h ic h i n c l u d e ty p e o f b i r t h show i n c r e a s e d s u r v i v a b i l i t y w i t h i n c r e a s i n g a m o u n ts o f F in n breeding. O ltenacu and Boylan (1981a) found s ta n d a r d breeds and 1/4Fx t o have s i m i l a r s u r v i v a l r a t e s f o r p e r c e n t of t o t a l born t o weaning; however, 1/2Fx and p u re F inns exceeded s ta n d a r d breeds by 9% and 25%, r e s p e c t i v e l y . H ow ever, D i c k e r s o n e t a l . (1975) r e p o r t e d ev e n more d r a m a t i c d i f f e r e n c e s ; 1 /2 F x e x c e e d e d b o th Rx and p u r e b r e d s by more t h a n 30%. I n c o n t r a s t , H ohenboken and C l a r k e (1981) f o u n d l o w e r s u r v i v a l r a t e s f o r la m b s fro m 1 /2 F x dam s; a l t h o u g h , due t o t h e i r s u p e r i o r p r o l i f i c a c y , . the Finn s u r v i v a b i l i t y was b ia sed downwards as a l l la m b s i n e x c e s s o f 2 w e re rem o v e d fro m t h e dam and c o u n t e d a s dead. The l i t e r a t u r e r e v i e w e d s u g g e s t , t h a t no l a r g e breed d i f f e r e n c e s e x i s t b e tw e e n F in n c r o s s b r e d s and s t a n d a r d breeds, or t h a t any m a t e r n a l h e t e r o s i s i s p r e s e n t . These c o n c l u s i o n s a g r e e w i t h o t h e r r e p o r t s i n th e l i t e r a t u r e t h a t l i t t l e t o no h e t e r o s i s has been found in v o l v in g o t h e r s ta n d a rd breed c r o s s e s (Sidwe l I e t a l . , 1962; S id w e ll and M i l l e r , 1971a; R asto g i e t a l . , 1975; Hohenboken and Cochran, 1976; N itte r, 1978). J 20 I n d i v i d u a l Progeny Weaning Weight Perfnrmannm The l a s t com ponent o f r e p r o d u c t i v e e f f i c i e n c y , i n d i v i d u a l lam b weaning w eight perform ance, o f t e n r e c e i v e s th e most a t t e n t i o n from the p ro d u c e r; how ever, t h e t o t a l k i l o g r a m s o f lam b w eaned i s m ore a f u n c t i o n of th e t o t a l number o f lam bs weanec? than th e i n d i v i d u a l lamb w e ig h ts (Sidwe l I and M i l l e r , 1971a). A com posite view o f th e p revious s e c t i o n s of t h i s re v ie w would c l e a r l y s u g g e s t th e Finn c o n t r i b u t i o n to t o t a l number of lambs weaned i s im p o r ta n t. Now th e q u e s t i o n becomes w hether the g a in s r e a l i z e d i n number weaned from c r o s s b r e d Finns a re negated or dampened by poor weaning w e ig h t performance. Any l i s t o f t h e f a c t o r s a f f e c t i n g t h e w e a n in g w e i g h t o f la m b s would a t l e a s t in c lu d e , breed, sex, type of b i r t h / r e a r i n g , age of dam, age o f lamb, n u t r i t i o n , season, and management. Of th e en v iro n m e n ta l f a c t o r s , ■ty p e o f b i r t h / r e a r i n g h a s t h e l a r g e s t e f f e c t on w e a n in g w eight. A ll s t u d i e s have shown a n e g a tiv e e f f e c t on w eaning w eight due to in c r e a s in g re p o rts th a t ty p e of b i r t h / r e a r i n g w e a n in g w e ig h ts of (B radford, tw in s reared 1972b). as B radford s in g le s fa ll i n t e r m e d i a t e to w e ig h ts o f tw in s r e a r e d a s tw in s and s i n g l e s r e a r e d as sin g le s, s u g g e s tin g a permanent en v iro n m e n ta l e f f e c t due to type of b i r t h . B ut, e v i d e n c e p u t f o r t h by Doney and Munro (1962) i n d i c a t e s d i f f e r e n c e s i n w e a n in g w e i g h t a r e n o t due d i r e c t l y t o t y p e o f b i r t h but ra th e r b irth w e i g h t w hich i s a fu n c tio n of l i t t e r siz e as d i s c u s s e d e a r l i e r . The r e s e a r c h e r s f o u n d no d i f f e r e n c e i p w e a p in g w e ig h t a t 90 d o f age between s i n g l e s r e a r e d as s i n g l e s and tw in s when r e a r e d a s s i n g l e s from I d of age when a d j u s t e d f o r b i r t h w eight. 21 Due t o . a la rg e am ount of v a r i a t i o n in m anagem ent no v a l i d e s t i m a t e s a r e a v a i l a b l e f o r pure Finn i n d i v i d u a l w eaning w e ig h ts i n F in la n d ; however, i n a review of e x p e rim e n ts in v o lv in g p u re Finn sheep i n E u ro p e , M a i j a l a and O s t e r b e r g (1977) r e p o r t e d an a v e r a g e o f 18 kg a t 90 d o f age. A ll s t u d i e s r e v i e w e d by t h i s a u t h o r h a v e shown r e d u c e d lam b weaning w e ig h ts due to th e i n c o r p o r a t i o n o f Finn b re e d in g when means a r e u n a d ju ste d f o r type of b i r t h / r e a r i n g . Barker (1975) found s i n g l e and tw in lam bs from 1/2Fx dams l i g h t e r a t 70 d of age than purebreds. The p u r e b r e d s u p e r i o r i t y i n c r e a s e d w i t h age r a n g i n g f r o m 1.1 kg f o r t w i n s fr o m y e a r l i n g s t o 3.4 f o r s i n g l e s a t a g e s 3 and 4 y r . S i m i l a r r e s u l t s h a v e b e e n r e p o r t e d i n t h e U.S. u n d e r b o th s e m i - i n t e n s i v e management and w e s te r n range. Lambs from Fx ewes were 1.3 t o 2.7 kg l e s s t h a n p u r e b r e d s a t 70 t o 105 d o f age ( C e d i l l o e t a l . , 1977; D i c k e r s o n , 1977). B u t i n c o n t r a d i c t i o n t o B a r k e r ( 1 9 7 5 ), when o n ly s i n g l e s w e r e co m p ared D i c k e r s o n f o u n d no breed d i f f e r e n c e s e x is te d . A lso i n c o n tra d ic tio n , Magid et al. (1981) re p o rte d no breed d i f f e r e n c e s between u n a d ju ste d weaning w e ig h ts o f lambs a t 50 d o f age fro m Fx and B o r d e r L e i c e s t e r s i r e d ew es. T hese r e s u l t s may i n d i c a t e th a t th e l i m i t a t i o n s o f th e m a te rn a l e n v i r o n m e n t h a v e n o t been exceeded a t such an e a r l y weaning d ate. Thomas and W h item an (1979) a l s o fo u n d u n a d j u s t e d w e i g h t s t o be s i g n i f i c a n t l y lo w e r f o r lam bs from Fx ewes when compared w i t h Dorset x R a m b o u i l l e t . B ut, once t h e m odel i n c l u d e d ty p e o f b i r t h / r e a r i n g t h e w e i g h t s w e re n o t s i g n i f i c a n t l y d i f f e r e n t . I n a g r e e m e n t w i t h t h e s e r e s u l t s , O l t e n a c u and B o y l a n (1981b) r e p o r t e d no d i f f e r e n c e b e tw e e n 22 la m b s fro m p u r e F i n n s , I / 2Fx, and t h e t h r e e s t a n d a r d b r e e d s (M100, Targhee, and S u ffo lk ) when a d j u s t e d f o r type o f b i r t h / r e a r i n g . from I A F x w ere s i g n i f i c a n t l y h e a v ie r th a n s ta n d a r d b reed s, Lambs but only by 1.0 kg. E x clu sio n o f th e S u f f o lk breed from the s ta n d a r d breed and Fx m eans r e s u l t s i n la m b s fro m p u r e F i n n s , e x c e e d in g t h o s e fro m 1 /2 F x , and I A F x dams T a rg h e e and M100 by 2 .0 , 1.4, and 2.3 kg, re sp e c tiv e ly . T h i s a u t h o r fo u n d no u n a d j u s t e d e s t i m a t e s o f m a te r n a l h e t e r o s i s f o r i n d i v i d u a l la m b w e a n in g w e i g h t fro m b re e d in g i n th e l i t e r a t u r e . s t u d i e s i n v o l v i n g F in n E s tim a te s i n t h e l i t e r a t u r e f o r m a tern al h e t e r o s i s range from 0 t o 8% i n s t u d i e s in v o l v in g s ta n d a rd breeds, and a r e c o n s i d e r e d o f m i n im a l i m p o r t a n c e ( S i d w e l l and M i l l e r , 1971b; Hohenboken and Cochran, 1976; Jakubec, 1977; N i t t e r , 1978). Most o f th e s t u d i e s review ed above have u t i l i z e d a "meat" breed as th e sire of th e p ro g e n y fro m t h e Fx and p u r e b r e d r e s u l t i n g p r o g e n y a r e o n ly I A F x o r 1 /8 F x , dams. As th e d e p e n d i n g on t h e F in n b r e e d i n g i n t h e dam, o n ly s m a l l d i f f e r e n c e s i n i n d i v i d u a l w e a n in g w e ig h ts a d j u s t e d f o r type of b i r t h a r e expected. The m ajor cause f o r r e d u c t i o n s i n la m b w e i g h t s r e m a i n s l i t t e r s i z e , and t h e a s s o c i a t e d l e s s th a n optimum environm ent f o r lamb grow th (Thomas and Whiteman, 1979). T herefore, perform ance are r e d u c t i o n s i n i n d i v i d u a l lam b w e a n in g , w e i g h t to be e x p ec ted under any s y s te m of b reed in g , n u t r i t i o n , o r management which s e r v e s t o in c r e a s e p r o l i f i c a c y . ) 23 Composite R eproductive Performance I n c r e a s i n g p r o l i f i c a c y and f e r t i l i t y f o r the com mercial sheep producer. are in te rm e d ia te o b je c tiv e s I n th e f i n a l a n a l y s i s , e v a lu a tio n of th e c o n t r i b u t i o n t o r e p r o d u c t iv e perform ance of v a r y in g l e v e l s of F in n b r e e d i n g w i t h s t a n d a r d b r e e d s m u s t c e n t e r on a c o m p o s i te o f r e p r o d u c t iv e perform ance, th e t o t a l k ilo g ra m s o f lamb weaned per ewe exposed a t a l l ages. U n f o r tu n a te ly , n o t a l l th e components o f th e th e above com posite measurement a r e a v a i l a b l e i n each study, no r i s th e r e agreem ent among sy stem s of a c c re d ita tio n for rearin g . E stim a te s of pure Finn, perform ance a r e l i m i t e d to an average o f l i t t e r w e ig h ts a c r o s s F inland u n d e r v a r i o u s m anagem ent s y s t e m s . M a i j a l a and O s t e r b e r g (1977) r e p o r t e d t o t a l k ilo g ra m s per ewe lam bing f o r 2- y r - o l d and o ld e r ewes o f 2 9 .1 , 4 5 .9 , 6 2 .8 , 8 7 . 8 , and 95.1 f o r l i t t e r s i z e s o f 1 , 2 , 3 , 4 and 5, r e s p e c t i v e l y . A v erag e l i t t e r w e i g h t a t 150 d o f a g e wa$ 7 1 kg. As re p o rte d e a rlie r, fe rtility fo r p u r e F in n s h e e p i n F in la n d is e x c e p t i o n a l l y h i g h ; t h e r e f o r e , e s t i m a t e s b a s e d on a p e r ewe e x p o s e d b a s i s would n o t be c o n s id e r a b ly low er. These r e s u l t s a r e g a th e re d from s m a ll system s of i n t e n s i v e management where s u p p le m e n ta tio n o f lambs from l a r g e l i t t e r s i s r e g u l a r l y p r a c t ic e d . On a p e r ewe l a m b i n g b a s i s , D onald e t a l . (1968) r e p o r t e d o n ly s l i g h t d i f f e r e n c e s i n t o t a l l i t t e r w e i g h t b e tw e e n p u r e F i n n s and s t a n d a r d b r e e d s ; h o w e v e r , a t a g e s 2 and 3 y r p u r e F i n n s r a i s e d 5.9 kg and 9.5 kg m ore la m b , r e s p e c t i v e l y . T hese r e s u l t s r e f l e c t s m a l l e r d i f f e r e n c e s i n p r o lif ic a c y - and h ig h e r lamb m o r t a l i t y r a t e s f o r Finn 24 y e a r l i n g s th a n s ta n d a r d breeds. L arger d i f f e r e n c e s i n p r o l i f i c a c y a t l a t e r ag es r e s u l t e d i n l a r g e d i f f e r e n c e s i n l i t t e r w eight. Under s e m i - i n t e n s i v e m anagem ent i n t h e U.S. p u r e F i n n s have g r e a t l y e x c e e d e d s t a n d a r d b r e e d s ' (M l00, S u f f o l k , and T a rg h e e ) f o r com posite r e p r o d u c t iv e perform ance. Y e a rlin g F inns weaned a t 70 d 10.7 kg (117%) m ore la m b t h a n t h e s t a n d a r d b r e e d s p e r ewe m a te d . Much o f th is d iffe re n c e c o u ld be a t t i b u t e d to su p e rio rity f e r t i l i t y a t I y r of age. But, a t 2 y r of age breeds o f F in n s f o r Finns exceeded sta n d a rd by 19.6 kg (104%); F inn s u p e r i o r i t y f o r p r o l i f i c a c y being th e m a jo r f a c t o r . ( O l t e n a c u and B o y la n , 1981b). From a c o m m e r c ia l p ro d u cers v ie w p o in t th e s e r e s u l t s may be b ia se d i n f a v o r of th e Finns, a s th e w e ig h ts of a l l lam bs r e a r e d a r t i f i c i a l l y were a c c r e d i t e d to th e ewe. C ro ssb red p erfo rm a n ce i n B r i t a i n w as r e p o r te d by B ark er (1975) on a p er ewe ex p o sed b a s is . Fx a d v a n ta g e o v er p u reb red s i n t o t a l w e ig h t w e a n e d w a s s i g n i f i c a n t o n l y a t I y r o f a g e (+3.3 k g ) . A t a g e s 2 and 3 y r th e Fx a d v a n ta g e w as s l i g h t l y p o s i t i v e , but n o n - s i g n i f i c a n t . B arker c o n clu d e d i n c r e a s i n g t o t a l k ilo g r a m s th a t th e F in n ’s c o n t r ib u t io n in o f lamb weaned per ewe exposed i s p r i m a r i l y by i n c r e a s i n g f e r t i l i t y i n ewe lambs. T h is c o n c l u s i o n i s i n a g r e e m e n t w i t h t h e r e s u l t s r e p o r t e d by Thomas and W hitem an (1979) f o r management, 1 /4 F x ewes u n d e r s e m i - i n t e n s i v e and by Hohenboken and Cochran (1981) f o r 1/2Fx ewes under h i l l c o n d itio n s . Thomas and Whiteman found t o t a l w eig h t weaned a t 70 d o f age f o r 1/4Fx y e a r l i n g ewes t o exceeded D orset x R a m b o q ille t by 3.5 kg, w h ile d i f f e r e n c e s a t 2 and 3 y r of age were only s l i g h t l y p o s i t i v e 25 and n o n - s i g n i f i c a n t . These r e s u l t s l e d t o t h e c o n c l u s i o n t h a t t h e m a j o r a d v a n t a g e o f 1/4 F in n b r e e d i n g i s i n c r e a s e d f e r t i l i t y i n ewe lambs. Hohenboken an< Clarke (1981) fount} a 10 kg s u p e r i o r i t y f o r lamb p r o d u c t i o n fr o m y e a r l i n g 1/2F x ew es o v e r s t a n d a r d b r e e d s , and no d i f f e r e n c e s a t l a t e r a g e s . B u t, u n d e r m ore f a v o r i b l e c o n d i t i o n s on i r r i g a t e d p a s tu r e 1/2Fx exceeded s ta n d a r d s a t a l l ag es r e s u l t i n g i n a c u m m u la t iv e d i f f e r e n c e o f +36kg (27%) by age 5 y r . T hese r e s u l t s a r e r e p o r t e d on a per ewe e n t e r i n g the e x p erim en t b a s i s , and would r e f l e c t h a rd in e ss of t h e ewe a s w e l l th e re p r o d u c tiv e t r a i t s d is c u s s e d e a r l i e r . I n a d d i t i o n , ewes r e c e iv e d c r e d i t i n t h i s study only f o r the w e ig h t o f lamb they a c t u a l l y r e a r e d th e m selv es. Under w e s te r n range c o n d itio n s , E rcanbrack and Knight (unpublished d a t a ) a l s o fo u n d 1/2 F x and 1/4F x t o e x c e e d p u r e b r e d s a t I y r o f age f o r c o m p o s i t e r e p r o d u c t i v e p e r f o r m a n c e by 23.5 kg (452%) 13.8 kg (265%), resp ectiv ely . The magnitude o f th e d i f f e r e n c e d ecre ase d from a g e s 2 to 6 y r w ith a s l i g h t in c r e a s e a t age 7 y r. D if f e r e n c e s a t a l l ages between Finn c r o s s e s and p u re b re d s were h ig h ly s i g n i f i c a n t . The cum mulative advantage o f 1/2Fx and 1/4Fx over ages I th ro u g h 7 y r was 107.8 kg (50%) and 63.5 kg (30%), r e s p e c t i v e l y . As m e n t i o n e d e a r l i e r i n t h i s re v ie w , ewes were c r e d i t e d w ith h a l f th e w eig h t of a l l t h e i r lambs r e a r e d a r t i f i c i a l l y or by a n o th e r ewe. Due to v a r y in g system s of a c c r e d i t a t i o n and a n a l y s i s th e v e r d i c t on th e F inns c o n t r i b u t i o n t o c r o s s b r e e d in g schemes in v o l v in g s ta n d a rd b r e e d s i s n o t c l e a r . B u t, a l l t h e l i t e r a t u r e d o es p o i n t t o a c l e a r s u p e r i o r i t y of F inns and Fx f o r f e r t i l i t y i n ewe lambs which r e s u l t s i n in c r e a s e d k ilo g r a m s o f lamb weaned per ewe lamb exposed. I p term s 26 o f c o m m e r c i a l s h e e p o p e r a t i o n s , u t i l i z a t i o n o f F in n b r e e d i n g w i l l red u ce the number o f u n p ro d u ctiv e ewes i n th e flo c k . Wool This rev ie w has been p r i m a r i l y aimed a t th e p o t e n t i a l c o n t r i b u t i o n of the Finn breed to w ard s i n c r e a s i n g r e p r o d u c t i v e e f f i c i e n c y due to t h e l a r g e p r o p o r t i o n o f g r o s s in c o m e d e r i v e d fro m t h e s a l e o f lam b com pared t o w o o l. N e v e r t h e l e s s , t h e q u a l i t y and q u a n t i t y o f wool produced re m a in s e c o n o m ic a lly im p o r ta n t t o th e w e s te r n range producer, and should n o t bo overlooked i n e v a l u a t i n g any in tro d u c e d breed. I n F i n l a n d t h e q u a n t i t y and q u a l i t y o f wool a r e c o n s i d e r e d o f secondary im p o rta n c e , and b r e e d s o t h e r t h a n t h e F in n a r e b e t t e r adapted f o r th e p ro d u c tio n of wool. Pure F inns i n F in la n d produce on a v e r a g e 2,7 kg o f g r e a s e w ool p e r y r , h a v e l o n g s t a p l e l e n g t h s , and h a v e a g r a d e a v e r a g e a ro u n d 3 / 8 b lo o d ( M a i j a l a and O s t e r b e r g , 1977). S i m i l a r r e s u l t s were r e p o r t e d from B r i t a i n where pure F in n s had f l e e c e w e i g h t s o f 2.2 kg w i t h 3 / 8 t o 1 /2 b lo o d g r a d e s (D onald and Read, 1967). I n t h e U.S. O l t e n a c u and B o y la n (1981b) r e p o r t e d a v e r a g e g r e a s e f l e e c e w e i g h t s f o r p u r e F i n n s o f 2.0 kg and 2.2 kg f o r a g e s I. and 2 y r , r e s p e c t i v e l y , compared to s ta n d a r d breed av era g es (Ml 00, S uffolk, and T a r g h e e ) o f 3.2 kg and 3.4 kg. From t h e s e e s t i m a t e s p u r e F in n s have ap p ro x im a te ly 35% l i g h t e r g re a s e f l e e c e w eight. Wool g ra d e s f o r the pure Finn f e l l between 1/4 and 3/8 blood; grades f o r th e purebred v a r i e d w id e ly due t o t h e breeds involved. 27 I n th e same study, 1/2Fx $wes had f l e e c e w e ig h ts s l i g h t l y h ig h e r th a n th e m id - p a r e n t v a lu e , tra it. d e m o n s tr a tin g about 8% h e t e r o s i s f o r th e E s t i m a t e s o f h e t e r o s i s f o r g r e a s e f l e e c e w e i g h t h a v e been r e p o r t e d b e f o r e r a n g i n g fr o m 0 t o 27%; h o w e v e r , S i d w e l l and M i l l e r (1971c) concluded t h a t more s t u d i e s w ere r e q u i r e d b e f o r e e s t a b l i s h i n g the e x i s t e n c e of h e t e r o s i s i n wool t r a i t s . The s u p e r i o r i t y o v e r t h e m i d - p a r e n t mean was m a i n t a i n e d i n t h e 1/4 F x a s t h e i r a v e r a g e w e i g h t s w e re t h e same a s t h e 1 /2 F x a t b o th I and 2 y r o f age. 1 /2 F x w ool g r a d e s w e r e s i m i l a r t o p u r e F i n n s , w h i l e th e 1/4Fx were no c o a r s e r th a n the s ta n d a rd breeds. Breed d i f f e r e n c e s i n q u a l i t y and q u a n t i t y w e re c o n s i s t e n t a c r o s s a g e s I , 2 , and 3 y r ( l i m i t e d d a ta r e p o r t e d f o r age 3). I n a study in v o lv in g ewes from S u f f o lk and Columbia dams crossed w i t h e i t h e r F in n , D o r s e t , C h e v i o t, o r Romney s i r e s , no c r o s s b r e d d i f f e r e n c e s e x i s t e d f o r wool g r a d e . D e s p i t e h a v i n g s i m i l a r s t a p l e l e n g t h s t o Romenys, Fx ew es had s i g n i f i c a n t l y l i g h t e r w e i g h t s ( - .3 kg) t h a n t h e c r o s s b r e d a v e r a g e . S i m i l a r i t y i n w ool g r a d e s i s t o be expected a s a l l the o th e r s i r e b re e d s t y p i c a l l y have g ra d e s averaging from 1/4 to 3 /8 . I n com parison w ith Columbia, R am b o u illet, and Targhee breeds on w e s te r n ra n g e , 1/2Fx and 1/4Fx had c o n s i s t e n t l y l i g h t e r f l e e c e w e ig h ts a t a g e s I t h r o u g h 7 y r t h a n t h e w h i t e f a c e b r e e d s by 1.10 kg and .48 k g, r e s p e c t i v e l y . D i f f e r e n c e s i n g r a d e s w e re c o n s i s t e n t a c r o s s a l l ages, 1/2Fx w ere of lo w er grade w ith 1/4Fx i n t e r m e d i a t e (Ercanbrack and Knight, u npublished d ata). But, i n a p re v io u s study in v o l v in g the 28 same g r o u p s o f s h e e p Drpmmond e t a l . (1982) fo u n d no d i f f e r e n c e s i n q u a l i t y o f y a r n s o r f a b r i c s produced by th e th r e e breed groups. ' A s t u d y i n v o l v i n g Panama and F i n n x Panama ew es Dahmen e t a l . (1978) r e p o r t e d average Panama g r e a s e f l e e c e w eight t o exceed the Fx by 1.1 kg and 1.5 kg a t I and 2 y r o f age. At 2 y r o f age Fx ew es w a re fo u n d t o h a v e s i g n i f i c a n t l y s m a l l e r a v e r a g e f i b e r d i a m e t e r ( f i n e r f l e e c e ) , ev e n th o u g h v i s u a l e s t i m a t e s had p r e v i o u s l y r a n k e d t h e Fx lo w e r i n wool grade th a n th e Panama. 29 CHAPTER 2 MATERIALS AND METHODS P o p u la tio n s From I 974 t h r o u g h 1977 a t t h e M ontana A g r i c u l t u r a l E x p e r im e n t S t a t i o n , Red B l u f f R e s e a r c h Ranch, N o r r i s , Mt., a p p r o x i m a t e l y 240 w e s t e r n r a n g e ew es o f a g e s 2 t h r o u g h 6 y r , com posed o f c o m p a r a b le n u m b e rs o f C o lu m b ia (C), R a m b o u i l l e t (R), and T a rg h e e (T), w e re exposed t o F in n is h Landrace x R a m b o u ille t rams (FxR). The (FxR) rams were o b ta in e d from th e U. S. Sheep Experim ent S ta t io n , Dubois, Idaho. I n a d d i t i o n , a p p ro x im a te ly 360 s t r a i g h t b r e d , w h ite fa c e ewes (WF) ages o f 2 t h r o u g h 6 y r , cpm posed o f C, R, and T i n n e a r l y e q u ^ l n u m b e rs , w ere m a in ta in e d a s c o n t r o l s , ^o c r o s s e s w ere made between t h e t h r e e WF g r o u p s . P e r f o r m a n c e o f t h e 563 ew es w h ic h la m b e d and o f t h e i r 821 progeny was d e s ig n a te d "Phase I" o f t h i s study ( t a b l e 2). All h e a l th y fe m a le 1/4Fx lam bs from Phase I were r e t a i n e d i n the f l o q k e a c h y e a r , t o t a l l i n g 146 f o r 4 y r . Each y e a r a p p r o x i m a t e l y t h e same number o f WF lam bs a s 1/4Fx lam bs, r e p r e s e n t i n g a l l t h r e e breeds comparably, were s e l e c t e d randomly from th e c o n tr o l group, 143 f o r 4 y r . to ta llin g P h ase I I o f t h i s s t u d y i n v o l v e d t h e la m b and wool p r o d u c t i o n o f 289 ew es fro m t h e s i x b r e e d g r o u p s , C, R, T, (FxR)xC, (FxR)xR, and (FxR)xT, f o r a t o t a l o f 1264 s p r in g lam bing o p p o r t u n i t i e s from I 976 t o 1982 a c r o s s a g e s I t o 6 y r ( T a b l e 3). B e g i n n in g a s ewe lambs, a t a p p r o x i m a t e l y 7 mo o f a g e , b o t h 1/4F x and WF ew es w e re 30 TABLE 2. TOTAL NUMBER OF PHASE I EWES LAMBING LAMBING BY YEAR AND AGE AT Age a t lambing Year 2 3 4 5 6 1975 18 26 23 20 —— 87 1976 51 46 47 27 22 193 1977 51 57 33 32 17 190 1978 18 34 32 9 138 163 135 88 T otal Total 93 39 TABLE 3. TOTAL NUMBER OF PHASE I I EWES EXPOSED IN PREVIOUS SEASON BY YEAR AND AGE IN YEARS AT LAMBING 563 BREEDING Age a t lambing Year I 2 3 4 5 6 1976 42 — —— — — — 42 1977 88 38 — —— —— — 126 1978 80 84 35 — — — 199 1979 79 84 73 42 — — 278 1980 —— 73 77 65 39 — 254 1981 —— — 58 67 55 33 213 1982 —— —— — 51 56 45 152 243 225 150 78 1264 TOTAL 289 279 Total 31 exposed to S u f f o lk rams th roughout th e b re e d in g season f o r y e a r s 1976 t o 1979. From 1980 t o 1982 ewes were randomly d iv id e d i n t o two groups and exposed t o e i t h e r FxT o r S u f f o lk rams. Management A ll P h ase I and I I m a t u r e ew es ( a g e s 2 th r o u g h 6 y r ) from each breed group were herded a s one band on th e range th roughout th e y ear w i t h t h e e x c e p t i o n o f b r e e d i n g and l a m b i n g s e a s o n s . Ewe la m b s w ere m anaged s e p a r a t e l y fro m w e a n in g u n t i l m i d - A p r i l on a f t e r m a t h hay meadows r e c e i v i n g an a d d i t i o n a l 150 g. of supplem ent (ap p ro x im ate ly 1556 p r o t e i n , 7356 TDN) composed of b a r le y (62.556), w h e a tm ill run (30$), cane m o lasse s (6.25$), u re a (1$), t r a c e m i n e r a ls (.25$), and D icalcium Phosphate (.25$) w ith v it a m i n s A and D added (.25$). From th e second week i n November to th e end o f December, a l l ewes w e re b re d i n s m a l l p a s t u r e s . Common "sh e d l a m b in g " p r a c t i c e s w ere f o l l o w e d a t l a m b i n g , m i d - A p r i l t h r o u g h May, e x c e p t i n 1976 and 1977 when 65 and 80 P hase I ew es la m b ed on open p a s t u r e . At t h i s t i m e l a m b 's i d e n t i f i c a t i o n , dam 's i d e n t i f i c a t i o n , s e x , b i r t h d a t e , b i r t h w eight (BWT), and type of b i r t h were re c o r d e d ; a l l lam bs were docked and male lambs were c a s t r a t e d . B ir th d a te and w eight f o r th e 205 Phase I la m b s b o rn on p a s t u r e w ere n o t r e c o r d e d . Lambs w e re w e ig h e d t w i c e s u b se q u e n tly , once i n th e m iddle of June (ap p ro x im ate ly 60 d of age, WT60) and a g a in th e l a s t week i n August a t weaning (ap p ro x im a te ly 120 d o f age, WTI20). Ewes were shorn d u rin g th e f i r s t week o f A pril each y e a r e x c e p t i n I 981 when s h e a r i n g o c c u r r e d i n t h e l a s t week o f May. Ewes were removed from Phase I I p r i o r to b reed in g a t s i x y e a r s of age, 32 o r a t e a r l i e r a g e s due t o u n s o u n d n e s s su c h a s m a s t i t i s , e t c . . . ; no ewes w ere removed based on perform ance. The r a n g e s i t e i s c h a r a c t e r i z e d by a 25 t o 35 cm p r e c i p i t a t i o n zone, w i t h san d y and s i l t y s o ils, d o m i n a te d by I d a h o f e s c u e and b lu e b u n c h w h e a t^ g ra s s , aqd c l a s s i f i e d i n good c o n d i t i o n . D u r in g t h e b reed in g seaso n th e ewes were f lu s h e d w i t h f i r s t c u t t i n g g r a s s hay and g r a i n . D u r in g g e s t a t i o n ew es r e c e i v e d .2 kg p e r h e a d p e r day o f t h e same supplem ent a s th e ewe lambs. Only under s e v e re w in te r c o n d itio n s when snow a c c u m u l a t i o n p r e v e n t e d g r a z i n g d i d t h e ew es r e c e i v e hay ( a p p r o x i m a t e l y 5 d p e r y e a r ) . S t a r t i n g A p r i l I e a c h y e a r t h e ew es r e c e iv e d an a d d i t i o n a l 1/2 kg of b a r le y p er sheep. In th e lam bing pens f i r s t c u t t i n g hay was provided f r e e choice. A fte r th e lam bing perio d ewes were r e tu r n e d t o th e range. T r a i t s S tudied T r a i t s a s s o c i a t e d w i t h b r e e d g ro u p d i f f e r e n c e s i n o v e r a l l lamb p ro d u c tio n , t o t a l k ilo g ra m s o f lamb weaned per ewe exposed, and wool p r o d u c t i o n w e r e o f p r i m a r y i n t e r e s t . C om ponents o f la m b p r o d u c t i o n measured ;Ln t h i s study c o n s is te d of ewe f e r t i l i t y , lamb s u r v i v a b i l i t y , ewe p r o l i f i c a c y , and i n d i v i d u a l lamb w eig h ts. For t,he purposes of t h i s ex p erim en t f e r t i l i t y was d e f i n e d a s t h e p e r c e n t a g e o f ew es lam bing o f those exposed f o r breeding. Ewe p r o l i f i c a c y was d e fin e d as the number o f lambs born per ewe lam bing. S u r v i v a b i l i t y was measured a t m i d - l a c t a t i o n (S60), a t a p p ro x im a te ly 60 d, and a t w eaning (S120), a p p ro x im a te ly 120 d o f age. S u rv iv a b ility was d e fin e d p e rc e n ta g e of lambs a t th e r e s p e c t i v e ages o f those born, as th e and coded 0 33 and I f o r dead and a l i v e , resp e c tiv e ly , f o s s e s in c lu d e d s t i l l b i r t h s , lam bs removed from th e ewe to be r e a r e d a r t i f i c i a l l y or by a n o th e r ewe n o t i n t h e s t u d y , and la m b s t h a t d i e d p r i o r t o w e i g h i n g . C o m p o s ite t r a i t s f o r re p ro d u c tiv e perform ance c o n s is te d of l i t t e r s i z e and l i t t e r w e i g h t a t m i d - l a c t a t i o n (LS60 and LW60), and l i t t e r s i z e and l i t t e r w e i g h t a t y e a n i n g (L S I20 and LWI? p ). C o m p o s ite t r a i t s yere m e a su re d on a p e r ewe l a m b i n g b a s i s i n P h ase I and on a p e r ewe exposed b a s i s i n Phase I I . Wool c h a r a c t e r i s t i c s s tu d ie d in c lu d e d k ilo g ra m s o f g re a s e f l e e c e w e i g h t , g r a d e , and f i b e r l e n g t h . F l e e c e s w e re g r a d e d b a s e d on th e " b lo o d " s c a l e and t h e n coded I t h r o u g h 5: F in e ( I ) , 1 /2 ( 2 ) , 3 / 8 (3 ), 1 /4 (4 ), and lo w 1 /^ (5). L e n g t h s w ere coded a s S t a p l e ( I ) , F r e n c h ( 2 ) , and C l o t h i n g (3). S t a t i s t i c a l Methods A ll d a t a w e re a n a l y z e d a c c o r d i n g t o t h e p r o c e d u r e d e s c r i b e d by Harvey (1975) f o r l e a s t s q u a r e s a n a l y s i s of d a t a w ith unequal s u b c la s s n u m bers. The f i x e d - e f f e c t s m odel f o r P h ase I and I I m a t u r e ewe r e p r o d u c tiv e perform ance in c lu d e d th e f a c t o r s breed group of ewe (B), breed group q f lam b's s i r e (S), age o f ewe a t lam bing (A), and y e a r of lam bing (Y). A ll p o s s i b le two-way i n t e r a c t i o n s b e tw e e n m a in e f f e c t s were t e s t e d (P<.10) i n p r e li m i n a r y a n a ly se s. R e s u lts i n d i c a t e d BxA to be th e o n ly i n t e r a c t i o n s i g n i f i c a n t f o r any o f t h e t r a i t s ; RxA was in c lu d e d i n a l l a n a ly s e s , except LW60 and LWI20 i n Phase L Composite t r a i t s f o r t o t a l k i l o g r a m s r e a r e d t o 60 and 120 d o f a g e p e r ewe 34 exposed w ere not a d j u s t e d f o r sex, type of b i r t h / r e a r i n g , and d a te of b irth . The b a s ip f i x e d - e f f e c t s model f o r Phase I and I I i n d i v i d u a l lamb s u r v iv a l, and w eig h t t r a i t s f o r m ature ewe progeny in c lu d e d th e f a c t o r s b r e e d g ro u p o f dam (B ), b r e e d g ro u p o f s i r e ( S ) , age o f dam (A), y e a r (Y), sex (X), and type of b i r t h / r e a r i n g (BR). P r e lim in a ry a n a l y s i s f o r s i g n i f i c a n c e (P<.10) o f a l l p o s s i b l e tw o-w ay i n t e r a c t i o n s i n d i c a t e d t h a t BxY end SxY s h o u ld be i n c l u d e d i n t h e f i x e d - e f f e c t s m o d e ls f o r P hase I , and t h a t AxBR be i n c l u d e d i n t h e m o d e ls f o r P hase I I . Q u a d ra tic r e g r e s s i o n on b i r t h w e ig h t was in c lu d e d i n th e models f o r e ac h t r a i t o t h e r t h a n b i r t h w e i g h t and s u r v i v a l from 60 t o 120 d. I n Phase I , 205 lam bs, born on p a s tu r e , had no b i r t h w eig h t re c o rd e d , and con seq u en tly were not in c lu d e d i n th e a n a l y s i s o f i n d i v i d u a l progeny p e r f o r m a n c e . ( A n a l y s i s o f c o m p o s i t e t r a i t s , l e t t e r s i g e and w e i g h t , f o r Phase I ewes in c lu d e d a l l 8 2 1 lambs born.) Q u ad ratic r e g r e s s i o n on b i r t h d a te was in c lu d e d i n model f o r b i r t h w eig h t; l i n e a r r e g r e s s i o n on b i r t h d a t e was in c lu d e d i n th e f i x e d - e f f e c t s models f o r 60 and 120 d w e ig h ts. A d d itio n a l a n a ly s e s of a l l t r a i t s was conducted w ith type o f b i r t h / r e a r i n g main e f f e c t and r e g r e s s i o n on b i r t h w eig h t removed from th e model. As a r e s u l t o f o n l y tw o ewe la m b s o u t o f 143 c o n c e i v i n g fro m t h e WF g r o u p , a s e p a r a t e a n a l y s i s o f v a r i a n c e f o r ewe la m b r e p r o d u c t i v e perform ance was c o n d u c t e d . The f i x e d - e f f e c t s m odel f o r ewe lam b r e p r o d u c t iv e perform ance in c lu d e d breed group (B), y e a r (Y), and BxY i n t e r a c t i o n . F i x e d —e f f e c t s m odel f o r ewe la m b i n d i v i d u a l p ro g en y 35 t r a i t s i n c l u d e d B, Y, X, ty p e o f b i r t h / r e a r i n g , and r e g r e s s i o n s on b i r t b d a t e ( l i n e a r ) and b ir t h w e i g h t ( q u a d r a tic ) . The f i x e d - e f f e c t s model f o r wool p ro d u c tio n t r a i t s in c lu d e d breed g ro u p o f ew e’s dam (D), b r e e d g r o u p o f ew e’s s i r e (S)t age p o ssib le of ewe (A). P re lim in a ry a n a ly sis of a ll y e a r (Y), and tw o-w py i n t e r a c t i o n s i n d i c a t e d t h a t DxS, DxY, SxY, and SxA should be in c lu d e d i n th e f i x e d e f f e c t s model. T e s t o f s i g n i f i c a n c e p r o c e d u r e s d e s c r i b e d by H arvey (1975) f o r a n a ly sis v a ria b le s. of v a ria n c e a ssu m e norm al d istrib u tio n s D i s t r i b u i t o n s f o r ewe f e r t i l i t y of dependent and la m b s u r v i v a l a r e b in o m ia l; and a l s o , p r o l i f i c a c y does n o t f o llo w a normal d i s t r i b u t i o n . C o n seq u en tly , te sts of sig n ific a n c e a c c u r a t e l y d e s c r ib e th e p o p u la tio n s . fo r th e se tra its may n o t 36 CHAPTER 3 RESULTS AND DISCUSSION M ature Ewe and P rogeny P eform an oe i n P hase I B reed Group o f S ir e E f f e c t s Mean s q u a re s from a n a l y s i s o f v a r ia n c e f o r Phase I a r e p re s e n te d in t a b l e s 4 and 5. in d iv id u a l p ro g e n y L ea st-sq u a res perform ance m eans f o r by breed ewe p r o d u c t i o n and g ro u p o f sire (S) a r e p r e s e n t e d i n t a b l e s 6 and ?• Ewes b r e d t o FxR ra m s e x h i b i t e d n e a r l y id e n tic a l le v e ls of p r o lif ic a c y , had l a r g e r , but not s i g n i f i c a n t , l i t t e r s i z e s and w e i g h t s a t 60 d, and w eaned more lambs. (P<.1 0) and more t o t a l k ilo g ra m s of lamb p e r ewe lam bing (P<.05) th a n ewes bred to WF ra m s . Land e t a l . (I 974) r e p o r t e d no d i f f e r e n c e s i p f e r t i l i t y or p r o l i f i c a c y between ewes bred t o e i t h e r F inn or Tasmanian Merino rams, e v e n th o u g h ew es o f t h e two b r e e d s d i f f e r g r e a t l y i n p r o l i f i c a c y . F u rth e rm o re , B radford (1972b) i n d i c a t e s th a t d if f e r e n c e s in p r o l i f i c a c y a r e more a f u n c t i o n o f g e n o t y p e o f t h e dam r a t h e r t h a n genotype of th e o f f s p r in g . B irth lig h te r. w e i g h t s o f la m b s s i r e d Magid e t al. by FxR w e re n o t s i g n i f i c a n t l y (1981) fo u n d pure F in n sire s to produce c r o s s b r e d lam bs w ith l i g h t e r BWT th a n Border L e i c e s t e r rams. D espite l i g h t e r BWT ( n o n - s i g n i f i c a n t ) , 1 /4 F x la m b s had b e t t e r s u r v i v a b i l i t y (P < .0 1 ) from b irth to 60 d t h a n WF l a m b s ; how ever, in c re a se d TABLE 4 . MEAN SQUARES FROM LAMBING ( 1 9 7 5 - ANALYSES OF VARIANCE FOR PHASE I EWE REPRODUCTIVE TRAITS PER EWE 1978)a Source df P ro lificacy LS60 LW60 LSI 20 LWI20 Breed o f dam (B) Breed of s i r e (S) Year (Y) Age of dam (A) BxA E r r o r MS E rro r df 2 I 3 4 8 .992* .008 '1 6 6 , , 3.606 .421 + .221 544 1.680** .7011 .5 3 8 ,, 4.355 .439 .283 544 693** 1.664** 1.012+ 2444** 1718 4.125 .269 .295 544 5257 ——— 319 552 a LS60, l i t t e r s i z e a t 60 d ; w eight a t 120 d. t p < . io % 1915 ——— 95 552 LW60, l i t t e r w eight a t 60 d; LSI20, l i t t e r s i z e a t 120 d; LWI20, l i t t e r FROM ANALYSES OF VARIANCE TABLE 5. MEAN SQUARES - 1978)a FOR PHASE I INDIVIDUAL PROGENY TRAITS TRAITS (1975 Source df BWT S60 WT60 SI 20 WTI20 Breed o f dam (B) Breed of s i r e (S) Year (Y) Age of dam (A) Sex (X) Type of b i r t h / r e a r i n g (BR)" BxY SxY R eg ressio n s BDT ( l i n e a r ) BWT ( l i n e a r ) BWT ( q u a d .) E r r o r MS E rro r df 2 I 3 4 I 6.66** '1 2 5 ,, .793** •330 •476“ 1.214 45.85** •0 1 ,, 6 5.31% 57.62** 40.99 .025 .035 .067 .035 .033 171.8** 1 .3 ,, 244.1 * 92.6 218.2 .419+ ——— — —— 455.30** ——— — —— .095 ——— ——— 732.6** ——— ——— ## 1 2 6 2 .0 9 ,, 524.99 4.65 6.18 499 ——— ——— —— .056 502 713.6% 933.3** 19.5 12.5 489 1(2) 6 3 I I I 4 .0 4 “ 6 . 09; , 9.54** 90.80** .84 1.27 »» 23.626* ——— ——— .38 592 —— 2.897 ——— .110 601 a BDT, b i r t h d a t e ; BWT, b i r t h w e i g h t ; S60, s u r v i v a l t o 60; WT60, w e i g h t a t 60 d ; SI 20, s u r v i v a l to 120; WTI20, w eig h t a t 120 d. ^ d e g r e e s o f fre e d o m i n p a r e n t h e s i s f o r WT60, SI 20, and WTI20 m o d e ls ; r e a r i n g w as coded 11 ( s i n g l e r e a r e d s i n g l e ) , 21 (tw in r e a r e d a s s i n g l e ) , and 22 (tw in r e a r e d a s twin). T r i p l e t s were coded as tw in s due to s m a ll numbers. * PC.10 PC.05 P<.01 UJ CD TABLE 6 . LEAST-SQUARES MEANS AND STANDARD ERRORS FOR PHASE I EWE REPRODUCTIVE TRAITS PER EWE LAMBING (1975 - 1978)a E ffect Breed Group o f S i r e b FxR WF Breed Group of Dam Columbia (C) R am bouillet (R) Targhee (T) N 220 343 176 194 193 P ro lific a c y LS60 LW60, kg LSI 20 LWI20, kg NS NS NS 26.8 ±. 0.9 25.4 ± 0.8 + * 1.31 ± .04 1.21 + .04 1.50 + .04 1.49 ±. .03 • 1.35 1.27 ** •• •• 1.41 1.50 1.58 1.24 ± .05 1.25 + .05 1.44 ± .05 24.8 ± 1.1 24.9 ± 1.0 28.5 ± 1.0 1.19 ± .05 1.21 ± .05 1.39 ± .05 ± ± ±. .04 .04 .04 ± ± .04 .04 48.1 ± 1.7 44.0 ± 1.6 43.8 ±. 1.9 43-7 + 1.8 50.6 ± 1 .8 a LS60, l i t t e r s i z e a t 60 d; LW60, l i t t e r w eight a t 60 d; LSI 20, l i t t e r s i z e a t 120 d ; LWI20, l i t t e r w eight a t 120 d. b FxR, Finn x R a m b o u ille t; WF, com posite of Columbia, R am bouillet, and Targhee. NS, n o n - s i g n i f i c a n t (PXIO) t PC.10 TABLE 7. LEAST-SQUARES MEANS AND STANDARD ERRORS FOR PHASE I PROGENY TRAITS (1975 - 1978)a BWT Mean, kg E ffect N Breed Group of Ramb FxR WF NS 290 4.56 ± .06 326 4.64 ± .05 Breed Group of Dam Columbia (C) R am bouillet (R) Targhee (T) *• 188 4.39 + .06 203 4.53 + .06 225 4.88 ± .06 WT60 S60 N Mean, % •* 290 91.7 ± 2.6 326 84.4 ±. 2.6 NS 188 91.1 + 2 .9 203 86.5 ± 3.0 225 86.6 + 3.0 N Mean, kg NS 251 21.6 ± .3 265 21.6 + .3 •« 159 21.4 + .3 170 21.0 ± .3 187 22.3 ± .3 SI 20 N Mean, WTI20 % NS 251 96.3 + .1 265 94.8 ± .1 NS 159 95.0 ± 1.6 170 94.9 ± 1.5 187 96.8 + 1.5 N Mean, kg NS 246 37.1 ± .4 259 37.0 ± .3 •• 154 37.5 ± .4 166 35.8 ± .4 185 37.8 ± .4 a BWT, b i r t h w e ig h t; S60, s u r v i v a l from b i r t h t o 60 d; WT60, w eig h t a t 60 d of age; S120, s u r v iv a l to I 20 d ; WTI 20, w e i g h t a t 120 d o f age. b FxRf Finn x R a m b o u ille t; WFf composite of Columbia, R am b o u illetf and Targhee. NS, non-s i g n i f i c a n t (P > .10) t P<.10 TABLE 8. MEAN SQUARES Source Fx Ewe lambs: Breed group (B) Years (Y) BxY E r r o r MS E rro r df Mature ewes: Breed group (B) Years (Y) Age (A) S i r e (S) BxA E r r o r MS E rro r df df FROM ANALYSES OF VARIANCE F e rtility 2 3 6 •044 1.305 .103 .222 134 5 5 4 I 20 .086 .218 .144 .084 939 a LS60, l i t t e r s i z e a t 60 d; w eig h t a t 120 d. PC. 10 FOR PHASE I I EWE PRODUCTION TRAITS (1976 - 1982)a P ro lific a c y LS60 LW60 .036 .063+ .183+ .090 43 •088 1.495 .312 .236 134 3.066*" 2.359 .625+ .198 .249 845 1.345+ .634+ .419 939 LSI 20 LWI20 36.2 732.9 135.4 109.8 134 . 0 3 9 .. 1.335 .241 .204 134 37 1929 296 255 134 1066.2** 3633.5** 938.9” 3277.0 250.4+ 162.3 939 2.597” 4.075* 1.291* 2.040 .885 .412 939 1597” 8 0 2 7 .; 1878 321 365 347 939 LW60, l i t t e r w eight a t 60 d; LSI20, l i t t e r s i z e a t 120 d; LWI20, l i t t e r s u r v i v a b i l i t y due t o i n d i v i d u a l h e t e r o s i s o f c r o s s b r e d la m b s o v e r p urebred lam bs i s w e ll documented i n th e l i t e r a t u r e (D ickerson e t a l., 1975; D ickerson and Glimp, 1975; W i tte r , 1978). Lambs s i r e d by FxR had v i r t u a l l y the same average w e ig h ts a s lam bs s i r e d by WF ram s a t botl) 60 d and 120 d; and, s u r v i v a b i l i t y from 60 to 120 d was only s l i g h t l y h i g h e r f p r 1/ftFx la m b s . T h e r e f o r e , t h e 4 k g a v e r a g e s u p e r i o r i t y i n l i t t e r w eight of ewes s i r e d by FxR rams (P<.05) was due p r i m a r i l y to improved s u r v i v a b i l i t y r e s u l t i n g from i n d i v i d u a l h e t e r o s i s . Breed Group o f Ewe E f f e c t s V a ria tio n due t o B w as l a r g e l y due to h i g h e r l e v e l s of perform ance from T ewes f o r a l l t r a i t s e x c e p t S60 and SI 20 ( t a b l e s 6 and 7). Lambs from T ewes had h e a v i e r WT60 and WTI20 th a n lam bs from R (PC.0 1), b u t w e re s i m i l a r t o la m b s fro m C ew es. B u t, t h e s u p e r i o r i t y i n LW12Q d p e r ewe l a m b i n g (6 k^, PC.01) o v e r b o th G pnd R was c}]u^ m o stly to h ig h e r p r o l i f i c a c y . Other Main E f f e c t s Y ear (Y) a f f e c t e d LW60 (PC.01) and a l l p ro g e n y t r a i t s (PC.05) e x c e p t S I 20 ( t a b l e 4). Age o f ewe (A) a f f e c t e d (PC.01) r e p r o d u c t i v e and w e i g h t t r a i t s a s w e l l a s S60 (PC.0 1 ). P r o l i f i c a c y and c o m p o s i te t r a i t s ( l i t t e r s i z e and w eig h t) in c r e a s e d t o 5 y r of age fo llo w e d by a s l i g h t d e c lin e a t age 6 yr. L a rg e st d i f f e r e n c e s occured between age 2 and s u b s e q u e n t a g e s . Ewes l a m b i n g a t 5 y r o f ag e w eaned 20 kg more la m b a t 120 d t h a n 2 - y r - o l d ew es (PC.01) due m o s t l y to h ig h e r p r o l i f i c a c y (.47 lambs, PC.01). B i r t h w e ig h t In c r e a s e d s t e a d i l y to age 43 6 y r ; how ever, WT60 and WTI20 w ere h i g h e s t f o r la m b s fro m 4 - y r - o l d ewes and d e c l in e d t h e r e a f t e r . The o n ly i n t e r a c t i o n s a f f e c t i n g BWT w e r e BxY and SxY (P<.05) and no tr e n d was e v id e n t. R eg ressio n on b i r t h d a te was h ig h ly s i g n i f i c a n t fo r BWT (.03 k g / d ) , r e s u l t i n g p e r h a p s fro m b o th d i f f e r e n c e s i n g e s t a t i o n l e n g t h s and a l o n g e r t i m e on h i g h p la n e o f n u t r i t i o n f o r ewes lam bing l a t e i n t h e season. Ewe Lamb and P rogeny P eform an ce i n Phase I I F e r t i l i t y and P r o l i f i c a c y E arly sex u al developem ent i s r e f l e c t e d i n f e r t i l i t y a t 7 months of a g e w i t h l a m b i n g a t I y r o f ag e . Out o f 143 e x p o s u r e s o v e r 4 y r o n ly tw o ew es fro m WF b r e e d g r o u p s la m b e d a t I y r o f ag e , co m p ared t o 55 o u t o f 146 f r p m 1/4 F x b r e e d gypups ovep t h e same p e r i o d (1.3? v s 3 7 . 7 ? p PC.OI). S u p e r i o r i t y o f 1 /4 F x e w e s o v e r WF b r e e d s and b r e e d c r o s s e s f o r e a r l y sex u al development h as been r e p o r t e d p r e v io u s ly i n t h e l i t e r a t u r e r a n g i n g fro m 7? t o 24? f e r t i l i t y f o r ewe la m b s u n d e r s e m i - i n t e n s i v e management (Thomas and W h item a n , 1979; O l t e n a c u and Boylan, 1981 a}, and up t o 52? pn w e s te r n range (Ercanbrack and Knight, u n p u b lish e d d ata). Y ear a f f e c t e d fe rtility (PC.OI), b u t was n o t s i g n i f i c a n t f o r p r o l i f i c a c y ( t a b l e 8). L ev els o f f e r t i l i t y ranged from 16? i n 1977 t o 57? i n 1976. A l t h o u g h B w as n o t sig n ific a n t, R a m b o u i l l e t c r o s s e s e x c e e d e d t h e p t h e r tw o b r e e d s f o r b o t h t r a i t s ( t a b l e 9). BxY i n t e r a c t i o n a p p r o a c h e d s i g n i f i c a n c e f o r p r o l i f i c a c y . E r c a n b r a c k and K n ig h t ( u n p u b l i s h e d d a t a ) fo u n d F in n x C o lu m b ia ewe la m b s t o be i n f e r i o r i n f e r t i l i t y (PC.0 5 ) , 77.7? v s . 85.9? and 86.5?, 44 f o r F in n x RambouilJLet and F in n x T a r g h e e , r e s p e c t i v e l y ; b u t th e y found no s i g n i f i c a n t d i f f e r e n c e s i n p r o l i f i c a c y . TABLE 9. LEAST-SQUARES MEANS AND STANDARD ERRORS FOR PHASE I I Fx EWE LAMB REPRODUCTIVE TRAITS (1976 - 1979) F e rtility Brqed o f dam Columbia (C) R am bouillet (R) Targpee (T) Mean N Mean, 42 50 54 146 NS 41.7 45.5 39 .? 42.1 P ro lific a c y % ± ± + + 8 .2 6.9 7.1 4 .3 N Mean 16 21 18 55 NS 1.04 + .10 1.14 ±. .07 1.13 + .OJ 1.10 ± .05 T NS, n o n - s i g n i f i c a n t (PX IO ) Progeny Performance Mean s q u a re s from a n a ly s e s o f v a r ia n c e and l e a s t - s q u a r e s means by brepd' of ewe f o r i n d i v i d u a l progeny t r q i t s a re p re s e n te d i n t a b l e s 10 and 11, r e s p e c t i v e l y . Breed of ewe approached s i g n i f i c a n c e f o r BWf due t o l a r g e r lam bs from (FxR)xT ewes. These r e s u l t s a r e i n agreem ent w ith r e s u l t s fo u n d i n P h ase I o f t h i s s t u d y , w h e re s t r a i g h t b r e d T ewes p r o d u c e d l a r g e r la m b s a t b i r t h t h a n e i t h e r R o r C (P < .01, t a b l e 7). F or b o th WT60 and WTl 20 B was n o t s i g n i f i c a n t , c o n t r a r y t o P hase I where lambp from T ewes were h e a v ie r a t both ages (P<.01). D ickerson (1977) r e p o r t e d t h a t th e s u p e r i o r i t y of purebred T lambs over R lambs was t r a n s m i t t e d t o c r o s s b r e d la m b s fr o m T and R dam s, a l t h o u g h i n n e i t h e r cpse d ifferen ces sig n ific a n t. Type o f b i r t h / r e a r i n g a f f e c t e d BWT (P<.01) and WT60 (P <.01), b u t d i d n o t s i g n i f i c a n t l y a f f e c t WTI20. N e v e r t h e l e s s , la m b s r e a r e d a s s i n g l e s w e r e £ Jcg h e a v i e r t h a n t w i n e . Q u a d r a t i c r e g r e s s i o n on b i r t h 45 w e i g h t i n f l u e n c e d WTl 20 (P < .0 1), i n d i c a t i n g perhaps t h a t lambs w ith h e a v i e r BWT had poorer subsequent perform ance to weaning due perhaps to more d i f f i c u l t b i r t h s . TABLE 10. MEAN SQUARES FROM ANALYSES OF VARIANCE FOR PHASE I I INDIVIDUAL PROGENY TRAITS FROM Fx EWE LAMBS (1976 - 1979)a Source df BWT, kg WT60, kg Breed of dam (B) Year (Y) Sex (X) Type of b i r t h / r e a r i n g (BR)b R eg ressio n s BDT ( l i n e a r ) BWT ( l i n e a r ) BWT (q u ad .) E rro r MS E rro r df 2 3 I .853+ .455 1.496* .048 3.591 .481 10 3 73.4 19.0+ 11.635** 13.045* 10.9 .215 ——— ——” .359 52 62.437** 21.746* 7.375 3.745 41 44.1* 48.5 101.0** 6.4 38 1(2) I I I WTI20, kg '„ a BWT, b i r t h w eig h t; WT60, w eight a t 60 d; WTI20, w eight a t 120 d. D d e g re e s of freedom i n p a r e n t h e s i s belong w ith models f o r WT60 and WT120. + PC.10 » ::: TABLE 11. LEAST-SQUARES MEANS AND STANDARD ERRORS FOR PHASE I I INDIVIDUAL PROGENY TRAITS FROM Fx EWE LAMBS (1976 - I 979)a BWT Breed Of Isweb N Mean, kg WT60 N Mean, kg N Mean, kg 16 21 16 53 NS 21.8 ± 21.8 ± 21.9 ± 21.8 ± 15 20 15 50 NS 36.1 ± 38.0 ± 37.5 ± 37.2 ± + (FxR) x C (FxR) x R (FxR) x T Mean 17 24 20 61 3.73 ± .17 4.10 ± .14 4.15 ± .15 3.99 ±. .10 WTI20 .5 .6 .5 .3 a BWT, b i r t h w eig h t; WT60, weight a t 60 d; WT120, w eight a t 120 d. b C = Columbia, F = F inn, R = R am bouillet, T = Targhee. NS, n o n - s i g n i f i c a n t (P>.10) + PC.10 .8 .9 .8 .6 46 A l i m i t e d am ount o f d a t a p r e v e n t e d t h e u t i l i z a t i o n o f l e a s t s q u a re s methods f o r a n a l y s i s of s u r v i v a b i l i t y . Out o f 61 lambs born, 53 s u rv iv e d t o ap p ro x im a te ly 60 d and 50 s u rv iv e d to ap p ro x im a te ly 120 d o f ag e . Lambs fro m C ew es had t h e h i g h e s t S60 r a t e s (94$) f o l l o w e d by R (88$) and T (80$). One lamb from each breed group d ie d between 60 and I 20 d. Composite T r a i t s Composite t r a i t s f o r 1/4Fx ewe lamb r e p r o d u c t iv e perform ance a r e r e p o r t e d i n t a b l e s 8 and 12. B reed o f ewe was n o n - s i g n i f i c a n t f o r litte r siz e and w e i g h t at ( w e a n in g ) p e r ewe e x p o s e d , fe rtility , b o th 60 d ( m i d - l a c t a t i o n ) re fle c tin g and 120 d th e la c k of B e f f e c t s f o r p r o l i f i c a c y , and i n d i v i d u a l lamb w eights. Year a f f e c t e d a l l f o u r com posite t r a i t s (P<.01). I n 1977 l i t t e r w eight per ewe exposed was 17 kg l i g h t e r than th e p rev io u s year. LW60 LSI 20 LWI20 42 50 54 146 NS .42 ± .08 .45 ± .07 .36 ± .07 .41 ± .04 NS 9 .5 ± 1.8 9.4 ± 1.5 7 .8 ± 1.6 8.9 ± .9 NS 14.6 ± 2.8 14.4 ± 2.4 12.9 ± 2.4 14.0 ± 1.5 CO LS60 O Columbia (C) R am bouillet (R) Targhee (T) Mean N -Cr O Breed of dam 1+ o5 TABLE 12. LEAST-SQUARES MEANS AND STANDARD ERRORS FOR PHASE I I Fx EWE LAMB COMPOSITE REPRODUCTIVE TRAITS PER EWE EXPOSED (1976 - 1979)* .40 ± .07 .35 ± .07 .38 ± .04 a LS60, l i t t e r s i z e a t 60 d ; LW60, l i t t e r w eight a t 60 d ; l i t t e r s i z e a t 120 d ; LWI20, l i t t e r w eig h t a t 120 d. NS, n o n - s i g n i f i c a n t ( P > .10) LS120, 47 M ature Ewe and Progeny P erfo rm a n ce i n P hase I I B reed Group o f Ewe E f f e c t s M eans' s q u a r e s fr o m a n a l y s i s o f v a r i a n c e f o r r e p r o d u c t i v e and c o m p o s ite t r a i t s o f m atu re ew es (a g e s 2 th ro u g h 6 y r ) a re p r e s e n te d i n T a b le 8 . Mean s q u a r e s f o r w e i g h t and s u r v i v a b i l i t y o f p r o g e n y from m ature ewes i s p r e s e n te d i n Table 15. While l a r g e d i f f e r e n c e s i n f e r t i l i t y e x i s t e d between 1/4Fx and WF ewe la m b s , B was n o t a s i g n i f i c a n t s o u r c e o f v a r i a t i o n i n t h e model for f e r t i l i t y i p m a t u r e ew es ( t a b l e 13). B reed g r o u p s w i t h I /4 F x b r e e d i n g had h i g h e r l e v e l s (P < .0 1) o f p r o l i f i c a c y t h a n WF b r e e d groups, a v e ra g in g a t .36 more lambs p e r ewe lam bing f o r ages 2 through 6 y r ( t a b l e 14). S u p e r i o r i t y o f 1/4Fx f o r p r o l i f i c a c y was pronounced a t a l l a^ee w ith no e v id e n t d e c l in e i n s u p e r i o r i t y w ith i n c r e a s i n g age of ewe (ta ttle 14). D ick erso n ( 1 977) apd E rcan b rack and K n ig h t (unpublished d a ta ) r e p o r t e d average 1/4Fx s u p e r i o r i t y over WF breeds on w e s t e r n r a n g e t o be .24 ( a g e s 2 t h r o u g h 5 y r ) and .38 ( a g e s 2 through 7 yr) lambs p er ewe lam bing, r e s p e c t i v e l y ; and t h e magnitude o f th e d i f f e r e n c e r e m a i n e d c o n s t a n t a t a l l a g e s . R e s e a r c h i n v o l v i n g 1/2Fx ewes h as shown g r e a t e r s u p e r i o r i t y i n p r o l i f i c a c y th a n r e s u l t s f o r 1/4Fx, but po tr e n d of i n c r e a s i n g d i f f e r e n c e s w ith i n c r e a s i n g age (D onald e t a l „ , 1968; Land e t a l . , 1974; B a r k e r , 1975; O l t e n a c u and Boyl an, 1981). B i r t h w eig h t f o r lambs from 1/4Fx m a tu re dams were .63 kg l i g h t e r (P<.01) than lam bs from WF dams ( t a b l e s . 15 and 16). L i g h t e r BWT would be e x p e c t e d due t o l a r g e r l i t t e r s i z e s ; h o w e v e r , when ty p e o f b i r t h TABLE 1 3 . LEAST-SQUARES MEANS AND STANDARD ERRORS FOR F E R T IL IT Y OF PHASE I I MATURE EWES ( 1 9 7 6 - 1982) Age of ewe a t time o f lambing 2 yr 4 yr 3 yr 5 yr Breed Groupa N C R T WF mean 44 43 51 138 94.9 ± 5.0 89.0 ± 5.1 91.2 ± 4 .7 91.7 ± 2.8 37 38 41 116 90.7 ±. 5.6 102.6 ± 5.5 108.4 ± 5.1 100.6 ± 3.1 33 38 41 112 103.6 ± 102.3 ± 98.9 ± 101.6 + 43 46 52 141 99.7 ± 5.3 89.5 + 5.2 101.1 ± 4 .8 96.8 ± 2.9 34 42 51 127 96.5 + 5 .8 105.8 ± 5 .2 83.8 + 4 .9 95.4 ± 3.0 30 36 47 113 91-3 108.8 90.3 96.8 + ±. ± ±. 279 94.5 ± 2 .3 243 (FxR) (FxR) (FxR) IA Fx Mean x C x R x T mean a C1 C o lu m b ia ; R, Mean,, N % R a m b o u i l l e t ; T, Mean, 97.8 N % ± T a rg h e e ; F, 2.4 F in n ; N Mean, 6 .2 5.7 5.3 3.3 20 22 29 71 96.8 98.3 99.3 98.1 6.5 5.6 5.3 3.3 18 29 32 79 99.3 + 2.5 150 Mean, 225 breed % of s ire lis te d 6 yr N Mean, % 7.7 7.2 6.6 4.1 10 13 15 38 107.4 112.2 94.4 104.7 ±10.5 ± 9.5 ± 8 .9 ± 5.5 95.8 + 8.4 87.5 ± 6.7 92.1 ± 6.6 91.8 ± 4.1 8 16 16 40 77.8 76.7 96.5 83.7 ±11.9 ± 9.4 ± 9.1 ± 5.7 94.8 78 95.4 ± 4.1 firs t. % ± ± ± ± ± 3.1 T ab le 14. LEAST-SQUARES MEANS AND STANDARD ERRORS FOR PROLIFICACY OF PHASE I I MATURE EWES ( 1 9 7 6 - 1982). Age of ewe iat time o f lambing 2 yi N Mean C R T WF mean 40 37 43 120 .99 1.12 1.14 1.08 ± + + zt (FxR) (FxR) (FxR) 1/4Fx 41 38 50 129 1.48 1.44 1.43 1.45 ± zt zt ± 249 Mean a C, x C x R x T mean C o lu m b ia ; R, 4 yr 3 yir Breed Group3 N Mean N Mean .08 .09 .08 .05 31 36 41 108 1.24 1.16 1.34 1.25 .10 .09 .09 .05 32 35 39 106 1.39 1.40 1.45 1.41 .09 .09 .08 .05 31 39 41 111 1.65 zt .10 1.58 zt .09 1.72 ± .09 1.65 zt .05 26 35 42 103 1.67 1.79 1.67 1.71 1.27 zt .04 219 1.45 zt .04 209 1.56 R a m b o u i l l e t ; T, zt zt zt + T a rg h e e ; F, 5 yr N Mean .10 .09 .09 .05 18 19 27 64 1.34 1.50 1.76 1.54 ± .10 .09 .09 .05 Zt .04 ± ± ± ± Zt Zt Zt 6 yr N Mean .13 .12 .11 .07 10 13 14 37 1.50 1.43 1.75 1.56 ± + zt ± .17 .15 .14 .09 16 23 30 69 1.81 ;t .13 1.81 ± .11 1.73 zt .09 1.78 ± .07 6 13 15 34 2.18 1.80 1.83 1.94 ± zt ± zt .21 .14 .14 .09 133 1.66 ± .06 71 1.75 zt .08 F in n ; breed o f s i r e l i s t e d ± zt zt zt firs t. 50 was i n c l u d e d i n t h e m odel BWT f o r la m b s fro m I /4 F x dam s r e m a in e d s m a l l e r by .34 kg (P < .0 1). R e s u l t s w e r e i n a g r e e m e n t w i t h B r a d f o r d (1972) who found BWT f o r Finn lambs t o be s m a ll r e l a t i v e t o m ature ewe s i z e a f t e r a d ju s tm e n t f o r type o f b i r t h . But r e s u l t s do n o t a g r e e w ith E r c a n b r a c k and K n ig h t ( u n p u b l i s h e d d a t a ) who fo u n d no s i g n i f i c a n t d i f f e r e n c e s i n BWT betw een 1/4Fx and and WF b reed s when u n a d ju s te d f o r type of b i r t h . D e s p i t e l i g h t e r BWT, la m b s fro m 1 /4 F x dams had .6 and .7 kg h e a v i e r ( n o n - s i g n i f i c a n t ) WT60 and WTI20, r e s p e c t i v e l y , t h a n la m b s from WF breeds ( t a b l e 16). When type of b i r t h / r e a r i n g was in c lu d e d i n t h e m odel t h e d i f f e r e n c e s i n c r e a s e d t o 1.3 and 1.5 kg f o r WT60 and WT l 20, re sp e c tiv e ly ( P < .0 1). O l t e n a c u and B o y la n (1981) fo u n d a s i g n i f i c a n t a d j u s t e d w eig h t s u p e r i o r i t y o f 1.6 kg f o r lam bs from 1/4Fx over WF breeds a t a p p ro x im a te ly 70 d o f age. D ipkerson (1977) r e p o r te d u n a d j u s t e d la m b w e i g h t s f r o m 1/4F x dams t o be .4 kg h e a v i e r a t 105 d o f age t h a n t h o s e fro m WF b r e e d s ( n o n - s i g n i f i c a n t ) . I n d i s a g r e e m e n t w ith th e s e f i n d i n g s , Thomas and Whiteman (1979) found u n a d ju s te d lamb w e a n in g w e i g h t s fro m 1 /4 F x dams t o be 1.5 kg l e s s t h a n D o r s e t x R am b o u illet, but when type o f b i r t h / r e a r i n g was in c lu d e d i n th e model no s i g n i f i c a n t d i f f e r e n c e s were e v id e n t. Amounts o f F inn b reed in g i n e x c e s s o f 1/4 h a v e l o w e r e d u n a d j u s t e d w e a n in g w e i g h t s by 1.1 t o 3.5 kg, and h a v e had no s i g n i f i c a n t e f f e c t s when a d j u s t e d f o r ty p e o f b i r t h / r e a r i n g (Donald e t a l . , 1968; B ark er, 1975; D ickerson, 1977). Lambs from 1/4Fx dams had 7.6% (P<.01) g r e a t e r S60 when q u a d r a t ic r e g r e s s i o n on BWT w as i n c l u d e d i n t h e m o d e l ( t a b l e 16). Even though BWT w ere s m a l l e r f o r lambs w ith F inn b re e d in g and s m a l l e r BWT cause TABLE 1 5 . MEAN SQUARES FROM ANALYSES OF VARIANCE FOR PHASE I I MATURE EWE PROGENY TRAITS ( 1 9 7 7 - 1982)a Source df BWT, kg S60 WT60, kg Breed of dam (B) Breed o f s i r e (S) Year (Y) Age of dam (A) Sex (X) Type of b i r t h / r e a r i n g (BR) A x BR R eg ressio n s BDT ( l i n e a r ) BDT ( q u a d .) BWT ( l i n e a r ) BWT (quad.) E rro r MS E rror d f 5 I 5 4 I 3 .8 S ” 1 9 .3 5 ., 5.41%. 2 .1 5 ,, 39.20* .384** .2 0 4 .. 1.267 .0 2 0 . .515 4 5 .2 % 5 5 .8 % 407.4* 18.2 3.8 .051 . 105._ .280** .033 .036 63.7** 6.4 690.4% 5 0 .5 ,, 88.3 2(4) 7 95.49** 1.02 .273+ .170 250.5** 11.1 + .028 .063 746.5% 27.5** 5 .6 6 ” 2.30 —— 1631.6** — — 991.5** 5 .265% 2.338** .104 1268 1400.6% 72.4 7.5 1069 — .047 1072 1946.8** 35.1 + 13.7 952 a BWT, b i r t h w eig h t; w eight a t 120 d. t PC.10 I I I I S60, —— .48 1268 s u r v i v a l to 60 d ; WT60, SI 20 WTI20, kg w eight a t 60 d; SI20, s u r v i v a l to 120 d; WTI20, TABLE 1 6 . LEAST-SQUARES MEANS AND STANDARD THROUGH 6 ( 1 9 7 7 - 1 9 8 2 ) a BWT Breed o f damb N Mean, kg ERRORS FOR INDIVIDUAL LAMB PERFORMANCE FROM DAMS AGES WT60 S60 N Mean, ? *# *# SI 20 Mean, kg #* 4.14 + .14 4.53 ± .13 4.37 + .08 4.35 ± .08 86 • 0 +. 3.6 86.6 + 3.4 92.2 ± 3 .2 88.3 ± 2.0 125 143 199 467 20.2 + .5 20.7 ± .6 21.5 + .4 20.8 ± .3 (FxR) (FxR) (FxR) 1/4Fx 195 249 290 734 3.84 ± .08 4.08 ± .07 4.12 ± .07 4.01 ± .04 97.6 ± 3.3 97.1 ± 3.0 92.9 + 3.1 95.9 ±. 1.8 166 224 241 631 22.3 ± .4 22.1 ± .4 21.9 ± .4 4.18 o> x C x R x T mean O v erall mean 1296 ± .03 -H 158 175 229 562 CU Columbia (C) R am bouillet (R) Targhee (T) WF mean 1.5 1098 Mean, 2 Wl 20 % N Mean, kg ** NS 95.3 + 87.4 ± 93.9 ± 92.5 ± 3.7 3.4 5.6 2.6 117 123 180 420 33.1 ± .6 33.0 ± .8 34.7 + .6 33.6 ± .4 4.9 3.2 4.0 2.3 145 199 219 563 35.4 ± .6 34.8 ±. .5 35.1 ± .5 35.1 ±. .3 983 34.4 + .2 2 2 .1 ±. .2 92.6 94.2 87.5 91.8 ± ± ± ± 21.5 ± .2 92.2 ± 2.2 a BWT, b i r t h w e ig h t; S60, s u r v i v a l t o 60 d ; WT60, w eight a t 60 d ; SI20, s u r v iv a l to 120 d; WT120, w eight a t 120 d. b C rosses d e s ig n a te d w ith l e t t e r s , breed of s i r e f i r s t ; F = F inn. NS, n o n - s i g n i f i c a n t (P > .10) PC.05 53 d e c r e a s e d s u r v i v a b i l i t y ( P u r s e r and Young, 1964), la m b s fro m 1/4 F x dams had 4.6% (P<.05) g r e a t e r S60 when th e q u a d r a t i c r e g r e s s i o n on BWT was removed from th e model. R eg ressio n s on BWT were n o t s i g n i f i c a n t and w e re n o t i n c l u d e d i n t h e m odel f o r S120. B reed o f ewe e f f e c t s w e r e n o t s i g n i f i c a n t ; la m b s fro m WF b r e e d s h a v i n g s l i g h t l y b e t t e r S I20. R e s u lts i n t h i s study f o r S60 a r e i n d is a g re e m e n t s tu d ie s w h ic h fo u n d la m b s from 1 /4 F x dam s t o w ith p re v io u s have s im ila r s u r v i v a b i l i t y w ith s ta n d a r d p u re b re d s and c r o s s b r e d s (D ickerson, O l t e n a c u and B o y la n , Knight, 1981; C o ch ran e t a l . , 1977; 1982; E r c a n b r a c k and u npublished d ata). Average LS60 ( m i d - l a c t a t i o n ) p e r ewe exposed o f 1/4Fx b reed groups exceeded t h e WF b r e e d a v e r a g e f o r a g e s 2 t h r o u g h 6 ( t a b l e r e s u l t i n g i n an o v e r a l l d i f f e r e n c e 17), o f .24 la m b s p e r ewe e x p o se d (PC.01). L a r g e s t d i f f e r e n c e s o c c u r r e d a t a g e 2 y r w h e r e 1 /4 F x ewes r e a r e d .46 m ore I ^mbs. S u p e r i o r i t y o f 1/4F% d e c l i n e d f o r 5 and 6 - y r old ewes p r i m a r i l y due to number o f lam bs r e a r e d by T ewes in c r e a s i n g w ith age. Targhee ewes exceeded R and C a t a l l ages and 1/4Fx average a t 5 and 6 y r o f a g e , a l t h o u g h n o t s i g n i f i c a n t l y . LS60 r a t h e r t h a n i n d i v i d u a l lamb w e ig h ts was ,mostly r e s p o n s i b l e f o r 1/4Fx s u p e r i o r i t y f o r LW60 p e r ewe e x p o s e d ( t a b l e 18). W e i g h ts w ere n o t a d j u s t e d f o r s e x , age o f la m b , o r t y p e o f b i r t h / r e a r i n g . A veraged a c r o s s a g e s 2 t h r o u g h 6 y r , 1/4 F x e x c e e d e d WF b r e e d s by 3.9 kg (P<.01). The l a r g e s t d i f f e r e n c e o ccu rred a t age 2 y r (8.2 kg), and th e s m a l l e s t a t age 6 yr (1.4 kg). TABLE 1 7 . LEAST-SQUARES MEANS AND STANDARD ERRORS FOR L ITTER S IZ E AT 6 0 DAYS 1977 - 1982) PER EWE EXPOSED (L S 60, Age of ewe a t time of lambing 2 yr Breed Groupa N Mean 4 yr 3 yr N N Mean 6 yr 5 yr Mean N Mean N Mean Columbia (C) Ram bouillet (R) Targhee (T) WF mean 44 43 51 138 .68 ± .10 .79 ± .10 • 86 ±_ *10 .77 ± .06 37 38 41 116 .97 ± .11 .94 ± .11 1.19 ± .11 1.03 ± .06 33 38 41 112 1.14 ± .12 1.09 ± .11 1.17 ± .11 1.13 ± .06 20 22 29 71 1 .09 ± .15 1.12 + .14 I .47 ± .13 I .23 ± .08 10 13 15 38 1.20 ± .21 1.03 ± .19 1.50 ± .17 1.24 + .11 (FxR) (FxR) (FxR) 1/4Fx 43 46 52 141 1.25 ± .11 1.17 ± .10 1.30 ±. .10 1.24 + .06 34 42 51 127 1.32 ± .12 1.44 ± .11 1.25 ± .10 1.34 ± .06 30 36 47 113 1.12 1.58 1.29 1.33 + + ±. ±. .13 .11 .10 .06 18 29 32 79 1.58 ± .16 1.24 ± .13 1.33 ± .12 1.38 ± .08 8 16 16 40 1.46 1.30 1.25 1.34 279 1.01 dt .05 243 1.19 .05 225 1.23 ± .05 150 I .31 ± .06 78 1.29 ± .09 x C x R x T mean O v e ra ll mean a C ro sses d e s ig n a te d w ith l e t t e r s , ± breed o f s i r e firs t, F = F in n . ± + ± ± .24 .17 .17 .11 TABLE 1 8 . LEAST-SQUARES MEANS 1977 - 1982) AND STANDARD ERRORS FOR LITTER WEIGHT AT 6 0 DAYS PER EWE EXPOSED (LW60, Age of ewe a t time of lambing 2 yr Breed Groupa N Mean, kg 3 yr N 4 yr N Mean,, k:g 5 yr 6 yr Mean, kg N Mean,» kg N Mean,, kg 44 43 51 138 13.7 ± 2.0 18.5 ± 2.0 18.9 ± 1.9 17.0 ± 1.1 37 38 41 116 20.8 21.8 29.0 23.9 + + + + 2 .2 2 .2 2.1 1.3 33 38 41 112 25.4 26.4 27.7 26.5 + + ± + 2 .3 2 .2 2.1 1.3 20 22 29 71 24.0 ± 3.0 21.3 ± 2 .8 30.2 ± 2.5 25.2 + .1 .6 10 13 15 38 25.0 21.7 31.6 26.1 (FxR) (FxR) (FxR) 1/4Fx 43 46 52 141 25.5 ± 2.0 23.1 ± 2.0 26.9 dL 1.9 25.2 ± 1.1 34 42 51 127 27.3 29.5 25.8 27.5 ± ± ± ± 2.3 2.1 1.9 1.2 30 36 47 113 24.8 32.5 28.1 28.5 ± ± ± ± 2.5 2.2 2 .0 1.3 18 29 32 79 29.9 ±. 3.1 27.2 ± 2.5 30.8 ± 2.4 29.3 ± 1.5 8 16 16 40 30.9 ± 4.6 27.1 ± 3.4 24.6 ± 3.4 27.5 ± 2.1 279 21.1 ± 1.0 243 25.7 ± 1.1 225 27.5 ± 1.2 150 27.0 78 26.8 O verall mean a C ro sses d e s ig n a te d w ith l e t t e r s , breed o f sire firs t; F = F in n . + ± ± ± 4.1 3.7 3.4 2.1 Hru O x C x R x T mean ■H Columbia (C) Ram bouillet (R) Targhee (T) WF mean Ul Ul TABLE 1 9 . LEAST-SQUARES MEANS EXPOSED ( L S I 2 0 , 1 9 7 7 - AND STANDARD 1982) ERRORS FOR PHASE I I L ITTER S I Z E AT 1 20 DAYS PER EWE Age of‘ ewe a t time of lambing 2 yr Breed Groupa N Mean 4 yr 3 yj N N Mean 5 yr 6 yr Mean N Mean N Mean Columbia (C) R am bouillet (R) Targhee (T) WF mean 44 43 51 138 .69 ±. .10 .73 ± .10 .84 ± .10 .75 ± .06 37 38 41 116 .91 ± .11 .88 ± .11 1.11 ± .11 .97 ± .06 33 38 41 112 1.03 ± .12 1.05 + .11 1.11 ± .11 1.06 ± .06 20 22 29 71 1.03 ± .15 .93 ± .14 1.35 ± .13 1.10 ± .07 10 13 15 38 1.15 + .21 •91 ± .19 1.32 ± .17 1.13 ± .11 (FxR) (FxR) (FxR) 1/4Fx 43 46 52 141 1.11 + .10 1.06 ±. .10 1.16 ± .10 1.11 ± .06 34 42 51 127 1.22 ± .11 1.36 ± .11 1.17 ± .10 1.25 ± .06 30 36 47 113 1.07 ± .12 1.44 ± .11 1.18 ± .10 1.23 ± .06 18 29 32 79 1.33 + .16 1.18 + .13 1.28 ± .12 1.26 + .07 8 16 16 40 1.30 ± .23 1.20 ± .17 1.21 ± .17 1.24 + .11 279 .93 ± .05 243 1.11 225 1.15 150 1.18 ±. .06 78 1.18 x C x R x T mean O v e ra ll mean a C r o sse s d e s ig n a t e d w ith l e t t e r s , ± .05 ± breed o f s i r e f i r s t ; F = F in n .05 ± .09 TABLE 2 0 . LEAST-SQUARES MEANS AND STANDARD LAMBING ( LWI2 0 , 1 9 7 7 - 1 9 8 2 ) ERRORS FOR PHASE I I L ITTER WEIGHT AT 1 2 0 DAYS PER EWE Age of ewe a t time of lambing 2 yr Breed Group3 N 4 yr 3 yr Mean, kg N Mean, kg N 5 yr 6 yr Mean, kg N Mean, kg N Mean, kg Columbia (C) R am bouillet (R) Targhee (T) WF mean 44 43 51 138 24.3 + 2 .8 26.8 ±_ 2 .9 28.5 ± 2 .7 26.5 ± 1.6 37 38 41 116 30.5 ± 3.1 31.9 ± 3.1 41.0 ± 2 .9 34.5 ± 1.7 33 38 41 112 35.9 ± 3.3 38.5 ± 3.1 39.0 ± 3.0 37.8 + 1.8 20 22 29 71 37.2 34.1 46.7 39.3 ± ± ± ± 4 .2 4.0 3.5 2 .2 10 13 15 38 40.5 34.3 44.3 39.7 ± ± ± ± 5.8 5.1 4.8 3.0 (FxR) (FxR) (FxR) 1/4Fx 43 46 52 141 36.7 ± 2 .9 33.3 + 2 .8 38.0 + 2 .7 36.0 ± 1.6 34 42 51 127 40.2 ± 3.2 42.9 ± 2 .9 37.7 ± 2 .7 40.3 + 1.7 30 36 47 113 37.2 44.5 39.1 40.3 3.5 3.1 2.8 I .8 18 29 32 79 42.9 39.8 44.4 42.4 ± ± ± + 4.4 3.6 3.3 2.1 8 16 16 40 41.2 40.1 39.5 40.3 ± ± ± ± 6.4 4.7 4.8 3.0 279 31.1 1.4 243 37.4 jl 1.5 225 39.1 ± 1.6 150 40.9 ± 2.0 78 40.0 ± 2.8 x C x R x T mean O v e ra ll mean ± a C r o sse s d e s ig n a t e d w ith l e t t e r s , ± ± ± ± b reed o f s i r e f i r s t ; F = F in n . 58 T re n d s e s t a b l i s h e d f o r LS60 and LW60 co n tin u e d through weaning due p r i m a r i l y t o no s i g n i f i c a n t e f f e c t on S I 20 due t o b r e e d o f ewe. Average m ature 1/4Fx s u p e r i o r i t y f o r LSI20 and LWI20 p e r ewe exposed w as .2 2 , la m b s (P<.01) and 4.3 kg (P <.01), r e s p e c t i v e l y . T hese r e s u l t s r e p r e s e n t an a v e r a g e i n c r e a s e , due t o 1 /4 F x b r e e d i n g , i n LSI 20 and LW120 o f 22% and 12%, r e s p e c t i v e l y . Ercanbrack and Knight (unpublished d a t a ) fo u n d i n c o r p o r a t i o n o f 1 /4 F x b r e e d i n g i n t o t h e same WF b r e e d groups as in v o lv e d i n t h i s study t o i n c r e a s e l i t t e r w e ig h t a t weaning by 8.3 kg (24%) on a v e r a g e f o r ew es 2 t o 7 y r o f age. O th e r s t u d i e s h a v e r e p o r t e d o n ly s l i g h t , . n o n - s i g n i f i c a n t 1/4F x s u p e r i o r i t y f o r m ature ewes (D ickerson, 1977; Thomas and Whiteman, 1979; Magid e t a l . , 1981b ) . Age o f Ewe and I n t e r a c t i o n E f f e c t s . Although age of ewe (A) was n o t a s i g n i f i c a n t source of v a r i a t i o n i n th e model f o r f e r t i l i t y , th e BxA i n t e r a c t i o n was im p o r ta n t (P<.01). At 2 y r o f age 1 /4 F x b r e e d g r o u p s had s l i g h t l y h i g h e r f e r t i l i t y t h a n WF b r e e d s , but th is d i f f e r e n c e w as r e v e r s e d at 3 y r of age. S u p e r i o r i t y o f WF ewes continued t o i n c r e a s e a t subsequent ages ( t a b l e 13). Ijn c o n t r a s t , p r e v i o u s r e s e a r c h h a s i n d i c a t e d no s i g n i f i c a n t d e c l i n e s i n Fx f e r t i l i t y a t l a t e r ages, r a t h e r Fx f e r t i l i t y i s n e a r ly i d e n t i c a l to s ta n d a r d b re e d s a t a l l ag es p a s t I y r (B a rk e r, 1975; M a ija la and O stq rberg, 1977; Thomas and Whiteman, 1979; Magid e t a l . , 1981; E rcanbrack and Knight, u npublished d ata). Age of ewe s i g n i f i c a n t l y a f f e c t e d p r o l i f i c a c y and com posite t r a i t s ( t a b l e 8). Average l i t t e r s i z e s a t b i r t h in c r e a s e d s t e a d i l y from 2 to 59 6 y r o f a g e , r e s u l t i n g i n a d i f f e r e n c e o f .5 la m b s p e r ewe la m b in g between th e two ages (P<..01, t a b l e 13). E f f e c t s due to A on com posite t r a i t s f o r LS60 and LSI20 p er exposed r e s u l t e d p rim ary from A e f f e c t s on p r o l i f i c a c y a s A had no e f f e c t on f e r t i l i t y and S60 and S I20. Breed o f ewe x ag e o f ewe i n t e r a c t i o n a p p r o a c h e d s i g n i f i c a n c e f o r LS60 (PC.IO) due to combining f l u c t u a t i o n s i n p r o l i f i c a c y and §60 o f 4 - y r ° l d (FxR)xC and (FxR)xR, as w e ll as a s te a d y o v e r a l l improvement of T ewe perform ance from age 2 t o 6 yr. Breed of ewe x age o f ewe e f f e c t s on LW60 was due t o LS60 r a t h e r t h a n BxA e f f e c t s on i n d i v i d u a l lam b w e ig h ts. A n a ly s is o f combined I A F x breed groups vs. WF breed groups i n d i c a t e s no BxA i n t e r a c t i o n . B ir t h w eig h t, a d j u s t e d f o r type of b i r t h , a l s o in c r e a s e d w ith each y r i n age r e s u l t i n g i n lambe from 6- y r - o l d ewes w eighing on average .6 kg h e a v i e r th a n th o s e fro m R - y r - p l d s . Age o f ewe a l s o a f f e c t e d i n d i v i d u a l WT60 and WT120 (PC.OI), C o n t r a r y t o BWT, la m b w e i g h t s i n c r e a s e d w ith age of ewe up to 3 y r old th e n d e c lin e d w ith i n c r e a s in g age. T hese r e s u l t s a r e i n a g r e e m e n t w i t h D i c k e r s o n e t a l . (1975) who found a s tr o n g c u r v i l i n e a r r e l a t i o n s h i p between age of ewe w ith 4 and IQ wk la m b w e i g h t s . Age o f ewe a f f e c t e d LW60 and LWI20 p e r ewe e x p o s e d ( P ^ .0 1 ), and l a r g e l y r e f l e c t e d d i f f e r e n c e s b e t w e e n 2 - y r - o l d and o ld e r ewes due to the c o n f l i c t i n g e f f e c t s of age on th e components o f l i t t e r w e i g h t i n o l d e r ew es. A verage LW60 and LWI20 o f 3 t o 6- y r old ewes exceeded' 2- y r - o l d s by 6 ^nd 8 kg, r e s p e c t i v e l y ( t a b l e s 18 and 20). Age o f ewe x ty p e o f b i r t h / r e a r i n g a f f e c t e d WTI20 (P<,01) and approached s i g n i f i c a n c e f o r WT60 (P<.10). I n t e r a c t i o n e f f e c t s were due 60 p r i m a r i l y to w eig h t f l u c t u a t i o n s f o r lambs born a s t r i p l e t s and r e a r e d a s s i n g l e s o r t w i n s . Lamb n u m b e rs i n t h e s e c a t e g o r i e s w e re s m a l l . E f f e c t of A on s i n g l e s r e a r e d a s s i n g l e s and tw in s r e a r e d a s tw in s was n e a r ly i d e n t i c a l . B reed Group o f S ir e E f f e c t s Due t o u t i l i z a t i o n o f FxT ra m s i n a d d i t i o n t o S u f f o l k ra m s commencing I n th e f a l l o f 1980, breed o f s i r e e f f e c t s were in c lu d e d i n a l l models f o r m a tu re ewe r e p r o d u c t iv e and i n d i v i d u a l progeny t r a i t s ( t a b l e 21). I n P hase I , F x R - s i r e d la m b s d i d n o t show s i g n i f i c a n t l y h e a v ie r BWT, WT60, or WTI20 th a n W F-sired lambs. But i n com parison to S u f f o i k s , F x T - s i r e d la m b s had l i g h t e r BWT (P < .0 1) and WT60 (P < .0 1), but only .5 kg l i g h t e r WTI20. Oltenacu apd Boylan (1981b) found Finn x ^ u f f 9^ d^®? bred t o S u f f p lk rpms produced h e a v i e r BWT (+.3 kg, P<.05) but not s i g n i f i c a n t l y h e a v ie r weaning w e ig h ts a t 70 d o f age th a n when b r e d t o F in n ra m s . D i c k e r s o n e t a l . (1975) fo u n d Fx la m b s t o h av e l i g h t e r BWT (.8 kg) and l i g h t e r w e a n in g w e i g h t a t 70 d (1,4 Kg.) t h a n S u f f o lk c r o s s lambs.. Ewes e x p o s e d t o . FxT r a m s had h i g h e r f e r t i l i t y (P<.01) and h i g h e r p r o l i f i c a c y (P <.10) t h a n ew es e x p o s e d t o S u f f o l k rams,, i n d i c a t i n g s u p e r i o r f e r t i l i z a t i o n r a t e s a n d / o r l o w e r embyro m o r t a l i t y i n ewes exposed to FxT rams. fe rtility Due t o confounding o f SxY a d d i t i o n a l a n a ly s e s of and p r o l i f i c a c y w e re c o n d u c te d in c lu d in g o n ly th e y r 1981 and 1982. Ewes s i r e d by FxT ra m s i n t h e s e tw o y e a r s had 13/6 h i g h e r fe rtility th a n S u f f o l k - s i r e d ewes (P<.01), but w ere 9% l e s s p r o l i f i c (PC.10). P r e v i o u s r e s e a r c h by Land e t a l . (1974) fo u n d no d i f f e r e n c e s 61 in f e r t i l i t y or p r o li f ic a c y b e tw e e n ew es b r e d t o e i t h e r F in n o r Tasmanian Merino rams. L i t t e r s s i r e d by FxT were h e a v i e r a t 60 d per ewe exposed (PC.OI) th a n th o se s i r e d by S u ffo lk p r i m a r i l y as r e s u l t of su p e rio r f e r t i l i t y . Due t o l o w e r S60 ( n o n - s i g n i f i c a n t ) and SI 20 TABLE 21. LEAST-SQUARES MEANS AND STANDARD ERRORS FOR PHASE I I MATURE EWE PRODUCTION AND INDIVIDUAL LAMB PERFORMANCE BY BREED GROUP OF SIRE (1976 - 1982) S u f f o lk Finn x Targhee T rait .03 .06 .07 804 712 804 89.8 ±. .01 1 .58 ± .04 1.14 ± .03 171 171 28.7 ± 1.3 1.16 ±. .07 804 804 22.6 ± .7 1.06 + .03 171 37.2 1.9 804 35.3 234 234 214 214 196 3.87 ± .08 94.1 ±. .3 .4 20.9 97.2 ± .3 33.5 ± .6 1062 1062 884 884 787 171 169 171 103.0 1.49 1.26 ± ±. ± O Lamb performance B ir t h weight S u rv iv a l t o 60 NS Weight a t 60 ** S u rv iv a l t o 120 NS Weight a t 120 NS Mean Mean -H Ewe p r o d u c t i o n , _ F e rtility , % P ro lific a c y + L i t t e r s i z e a t 60 + L i t t e r w eight a t 60, kg ** L i t t e r s i z e a t 120 L i t t e r w eight a t 120, kg NS N N 4.28 ±. .06 97.3 ±. .2 21.7 ± .3 99.7 ± .3 33.8 ± .5 NS, n o n - s i g n i f i c a n t (PX IO ) + P4.10 P<.05 ** PC.OI ( n o n - s i g n i f i c a n t ) o f FxT s i r e d lam bs, average FxT s i r e d l i t t e r w eights at 120 d was o n l y 1.9 kg h e a v i e r (n o n -sig n ific a n t). Low er lam b v i a b i l i t y due to in c r e a s e d Finn b re e d in g found i n t h i s study i s not i n agreem ent w ith p re v io u s re s e a rc h . D ick erso n e t a l . (1975) fo u n d S u f f o l k s i r e d c r o s s b r e d la m b s t o h a v e 5$ l o w e r s u r v i v a b i l i t y (non­ s i g n i f i c a n t ) t h a n F in n s i r e d c r o s s b r e d la m b s . Magid e t a l . (1981) 62 r e p o r te d Finn s i r e d t o exceed Border L e i c e s t e r s i r e d c ro s s b r e d lambs by 12% i p s u r v i v a b i l i t y . O ther Main E f f e c t s Y ear e f f e c t s (Y) f o r f e r t i l i t y (P<.0 5 ) , p r o l i f i c a c y (P < .0 1 ), and la m b w e i g h t s (P<.01) c o n t r a d i c t p r e v i o u s r e s e a r c h w h ic h fo u n d Y e f f e c t s f o r p r o l i f i c a c y and i n d i v i d u a l lamb perform ance t r a i t s , but no s i g n i f i c a n t Y e f f e c t s f o r f e r t i l i t y (M agid e t a l . , . 1 9 8 1 ; E r c a n b r a c k and JCnight, u n p u b l i s h e d d a t a ) . Y ear e f f e c t w e r e a l s o fo u n d f o r LW60 (P<,01) and a p p r o a c h e d s i g n i f i c a n c e f o r LW120. Sex e f f e c t s w e re s i g n i f i c a n t f o r BWT, S60, and Wfl 20 ( t a b l e 14). M ale la m b s had .3 kg and .6 kg h e a v i e r BWT and WT120, r e s p e c t i v e l y , t h a n ,fezqales; however, S60 was 4% l o w e r f o r m a le s . S i m i l a r r e s u l t s f o r b i r t h and w e a n in g w eig h t have been r e p o r t e d i n s t u d i e s i n v o l v in g lambs from Fx ewes, but d i f f e r e n c e s i n su rv iv aty ijlity have been n o p - s i g n i f i c a n t (Magid e t a l ., 1981b; O ltenacu and Boylan, 1981a; Ercanbrack anb Knight, unpublished d ata). Wool P ro d u ctio n Grease F le e c e Weight S i g n i f i c a n t so u rces o f v a r i a t i o n f o r g re a s e f l e e c e w e ig h t (FLWT) o f P hase I l ew es a g e s I t h r o u g h 6 y r i n c l u d e d b r e e d o f dam (B), b r e e d g ro y p o f s i r e (S ), y e a r (Y), age (A), and SxA i n t e r a c t i o n ( t a b l e 22). Ewes s i r e d by WF r a m s h ad .4 kg h e a v i e r FLWT t h a n F x R -s i r e d ew es (P<.01, t a b l e 23). These r e s u l t s a r e i n c lo s e agreement w ith th e study by E r c a n b r a c k and K n ig h t ( u n p u b l i s h e d <iata) w h ich i n v o l y e d t h e same 63 dam breed groups and s i m i l a r s i r e breed groups. In t h a t study FLWTs of I A F x ewes were .5 kg l i g h t e r (P<.05) than WF ewes f o r ag es I through 7 yr. L arger d i f f e r e n c e were r e p o r te d by D ickerson (1977) w ith s i m i l a r TABLE 22. MEAN SQUARES FROM VARIANCE ANALYSES FOR EWE WOOL PRODUCTION TRAITS (1976 - 1982) df Source 2 Breed of dam (B) Breed o f s i r e (S) I 6 Years (Y) Age (A) 5 2 BxS 12 BxY 6 SxY SxA 5 Remainder 1224 + ** Grease weight F leece grade 3.84* 21. 68** 66.29** 28.63** 2.39 .83 2 . 10+ 10.05** 1.10 51.95** 15.99** 9 . 10** 12.87** 2 2 . 16** 1.44* 5.03** 1.13 .75 S ta p le le n g th 1.674* .5 9 5 .. 21.971 2.361** 1.487 .742* 1 .241 .684 .383 P<.10 PC.05 PC.01 TABLE 23. LEAST-SQUARES MEANS AND STANDARD ERRORS FOR PHASE I I CHARACTERS, AGES I TO 6 (1976 - 1982) WOOL Lengthb Grease w eight Gradea Mean Mean Breed group0 N Mean, kg Columbia (C) Ram bouillet (R) Targhee (T) WF mean 188 199 231 618 4.07 3.80 3.75 3.87 ± + ± ± .12 .10 .09 .07 2.75 ± .10 1.18 + .09 1.74 ± .08 1.89 ±. .06 1.42 ± .07 1.16 + .06 1.10 ± .05 1.23 ± .04 (FxR) (FxR) (FxR) IA Fx 175 219 252 646 3.63 3.27 3.51 3.47 ± ± ± ± .12 .09 .10 .07 2.54 ± .10 1.79 ± .08 2.37 ± .08 2.23 ± .06 1.35 ± .07 1.25 + .06 1.28 ± .06 1.29 ± .04 x C x R x T mean a F leece grades coded I to 5: Fine ( I ) , 1/2 blood ( 2 ) , 3/8 blood ( 3 ) , 1/4 blood ( 4 ) , low 1/4 blood (5) b S ta p l e le n g th coded I t o 3: S ta p le ( I ) , French ( 2 ) , C lo th in g (3) c C rosses d e s ig n a te d by l e t t e r s , breed o f s i r e f i r s t , Finn (F) 64 b r e e d g r o u p s . A verage FLWT f o r WF e w e s e x c e e d e d 1/4 F x e w e s by 1.1 kg f o r ag es I and 2 y r. S tu d ie s in v o l v in g com parisons o f 1/2Fx and 1/4Fx ewes w ith o th e r breeds have shown Fx ewes t o have FLWT ra n g in g from 0 t o 1.5 kg ( a v e r a g e .5 kg) l e s s t h a n s t a n d a r d b r e e d s ( C e d i l l o e t a l . , I 977; Dahmen e t a l . , 1978; M agid e t a l . , 1981b; O l t e n a c u and B o y la n , 1981b ). No SxA e f f e c t s w ere r e p o r te d i n th e l i t e r a t u r e c i t e d above; h o w e v e r , i n t h i s s t u d y , SxA w as a n i m p o r t a n t e f f e c t (P<.01). At I y r o f age 1 /4 F x ew e s had .7 kg h e a v i e r FLWTs t h a n WF e w e s , and a t 2 y r FLWTs were s i m i l a r . WF s u p e r i o r i t y f o r FLWT in c r e a s e d t o 5 y r of age, th e n showed a s l i g h t d e c l in e f o r 6- y r - o l d ewes. FLWT f o r 1/4Fx 2 - y r old ewes was s i m i l a r t o WF ewes ages 2 through 6 yr. S u p e r i o r i t y o f WF ewes f o r ages 2 through. 6 y r l a r g l y r e f l e c t e d a d e c l in e i n 1/4Fx FLWTs w ith age. E f f e c t s due to breed o f dam f o llo w e d normal breed c h a r a c t e r i s t i c s w i t h C ew es h a v i n g t h e h e a v i e s t FLWT and T and R b e i n g c o m p a r a b le . BxS, BxY, and SxY i n t e r a c t i o n s w ere not s i g n i f i c a n t (PX05). F le e c e G r a d e ' F l e e c e s fro m 1 /4 F x w e r e a b o u t 1 /3 o f a b lo o d c o u n t c o a r s e r t h a n th o se from WF ewes (P<.01, t a b l e 23). B a l s o a f f e c t e d f l e e c e grade i n th e expected manner w ith C ewes a v e ra g in g i n th e 3 /8 blood range, T i n th e 1/2 blood ran g e, and R g ra d in g c lo s e to Fine. F le e c e s from (FxR)xC ew es g r a d e d s i m i l a r i l y t o s t r a i g h t b r e d C, b u t w e r e a h a l f a b lo o d c o u n t c o a r s e r t h a n t h e f i n e r w o o le d R and T. R e s u l t s a r e i n c l o s e agreem ent th e study by Ercanbrack and Knight (unpublished d a ta ) which 65 i n v o l v e d t h e same dam b r e e d g r o u p s . The r e s u l t i n g BxS i n t e r a c t i o n (P C .O I) in d ic a te s c ro s s b re e d in g w ith ram s p o sse ssin g c h a r a c t e r i s t i c a l l y c o a r s e r f l e e c e s th a n dam breeds w i l l lo w er f l e e c e g ra d e s; however, c r o s s b r e e d in g w ith ewes of s i m i l a r f l e e c e g ra d e s h as no e f f e c t . C e d illo e t a l . (1977) found no d if f e r e n c e i n f l e e c e grades b e tw e e n ew es fro m C o lu m b ia and S u f f o l k dams c r o s s e d w i t h e i t h e r D orset, Cheviot, Romney or Finn, a l l o f which have ap p ro x im a te ly 3/8 blood f l e e c e grades. BxY and SxY w ere im p o r ta n t (P<.05) i n t h e -model f o r f l e e c e grades, b u t no t r e n d s w e r e e v i d e n t . U n lik e FLWT, SxA was n o t a s i g n i f i c a n t i n t e r a c t i o n f o r g r a d e s ( t a b l e 2 2 ). B u t, (P<.01), age e f f e c t s w e r e p r e s e n t and due l a r g e l y to d i f f e r e n c e s between y e a r l i n g s and m ature ewes. These r e s u l t s a r e i n agreem ent w ith p re v io u s r e s e a r c h i n d i c a t i n g a f f e c t s due t o age b u t no b r e e d x a g e i n t e r a c t i o n s (Dahmen e t a l . , 1978; E rcanbrack and Knight, u npublished d a ta ). ! S ta p le Length In ag reem en t w ith p re v io u s r e p o r t s i n th e l i t e r a t u r e in v o lv in g s i m i l a r s i r e and dam b r e e d s (Dahmen e t a l . , 1978; Drummond e t a l . , I 9 8 2 ), S d i d n o t s i g n i f i c a n t l y a f f e c t s t a p l e ( f i b e r ) l e n g t h i n t h i s s t u d y . B ut i n c o m p a r is o n o f o n ly R and T ew es s i r e d by e i t h e r FxR o r WF rams, s t a p l e le n g th was s l i g h t l y lo n g e r f o r 1/4Fx ewes ( t a b l e 23). C and (FxR)xC ew es had s i m i l a r s t a p l e l e n g t h s r e s u l t i n g i n a BxS in te ra c tio n (P<.05). S t a p l e l e n g t h i n c r e a s e d w i t h age (P < .0 1 ), b u t showed no SxA i n t e r a c t i o n . 66 D is c u s s io n Im proved r e p r o d u c t iv e e f f i c i e n c y L and race in te n s iv e b r e e d in g w ith m a n a g em en t lite r a tu r e . sta n d a r d sy ste m s Few r e p o r t s , th rou gh in c o r p o r a t io n o f F in n is h b reeds has h ow ever, b een i,n t h e w e ll U.S. under d o c u m e n te d s e m iin th e have b een p u b lis h e d e v a l u a t i n g th e p e r f o r m a n c e o f F in n c r o s s e s u n d e r m o re h a r s h c o n d i t i o n s o n w e s t e r n range. R e s u lts p r e s e n te d i n t h i s study i n d i c a t e g a in s i n t o t a l k ilo g ra m s o f la m b w eaned may be r e a l i z e d t h r o u g h t h e a d d i t i o n o f 1 /4 F in n b re e d in g i n t o t y p i c a l w e s te r n w h ite fa c e ewes d e s p i t e l e s s th a n optimum n u t r i t i o n and management. Based on unw eighted average o f LWI20 from I t h r o u g h 6 y r o f ag e , I /4 F x ew es w eaned 6 kg p e r ewe p e r y r m ore t h a n WF ewes under the same c o n d itio n s . Lamb p r o d u c tio n s u p e r i o r i t y of 1/4Fx ewes wa$ most e v id e n t a t age5 I , 2 and 3 y r . D i f f e r e n c e s a s ewe la m b s , l a m b i n g a t I y r o f a g e , re su lte d e n tire ly from in c re a se s in fe rtility (37.7# vs. 1.3#). I n c r e a s e d f e r t i l i t y a s ewe la m b s p r o v i d e d 1 /4 F x ew es w i t h a 14 kg p ro d u c tio n ad v an tag e b e f o r e WF ew es la m b e d f o r t h e f i r s t ti m e . A n a l y s i s by b r e e d o f dam show ed i n c r e a s e d l e v e l s o f p r o d u c t i o n i n 1 /4 F x ewe la m b s t o be i n d e p e n d e n t o f t h e WF b r e e d i n v o l v e d i n t h e cross. S u p e rio r LWI20 a t a g e s 2 and 3 y r r e s u l t e d p r o l i f i c a c y r a t h e r th a n f e r t i l i t y s ig n ific a n t, fro m in c re a s e d a s i n ewe la m b s . A lth o u g h non­ e s t i m a t e s o f f e r t i l i t y i n m ature ewes d em o n stra te d 1/4Fx ew es t o be a c t u a l l y i n f e r i o r t o WF ew es. D e c r e a s e s i n f e r t i l i t y i n 67 m ature 1/4Fx ewes may be a r e s u l t o f in c r e a s e d s t r e s s due t o lam bing a t I y r o f ag e ; how ever, t h i s stu d y does n o t a d d r e s s t h i s q u e s tio n . F or a g e s I and 2 y r I / 4 F x e w e s e x c e e d e d WF ew e s by an a v e r a g e o f .4 lam bs per ewe lam bing, r e s u l t i n g i n an 8 kg s u p e r i o r i t y i n LWI20. A n a l y s i s o f P hase I w as i n c l u d e d i n t h i s s t u d y t o e s t i m a t e any change i n t o t a l k ilo g ra m s o f lamb weaned due t o th e use of FxR rams t o o b t a i n 1/4Fx ewes f o r f u t u r e p ro d u ctio n . The s u b s t i t u t i o n o f FxR f o r WF ra m s show ed no e f f e c t s on i n d i v i d u a l la m b w e a n in g w e i g h t s , and t o t a l k ilo g r a m s o f lamb weaned i n c r e a s e d by 4 kg. S u p e r i o r i t y i n LW120 fo r ew es s i r e d by FxR. r a m s r e s u l t e d p rim a rily fro m h e te ro sis i n c r e a s i n g S60 f o r c r o s s b r e d lambs. . F i n n i s h L a n d r a c e a r e known f o r l i g h t e r c o a r s e r f l e e c e s t h a n WF e w es, and r e d u c t i o n s i n w ool p r o d u c t i o n and q u a l i t y due t o 1 /4 F in n b re e d in g w ere s i g n i f i c a n t i n t h i s study. 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