Physiological and behavioral responses of cows on Montana foothill range to winter and supplement by Roger William Dunn A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science in Range Science Montana State University © Copyright by Roger William Dunn (1986) Abstract: Supplementation of gestating beef cows spending the winter on range has shown benefits to cow productivity. In order to understand how these benefits operate, detailed studies of the nutritional costs for cows on winter range, and the effects of supplements, are necessary. As part of that research goal, a study was conducted in 1984 and 1985 that recorded the grazing and travel behavior, and estimated the forage intake, of gestating crossbred cows onwinter range receiving protein supplement. The rangeland available to the cows contained cool season grasses dominated by bluebunch wheatgrass (Agropvron snicatum (Pursh) Scribn. & Smith). The study pasture contained 324 hectares in the northwest foothills of the Madison Range of southwest Montana. The cows wore vibracorders and pedometers for 48 continuous days in January and February. Daily intake was estimated using the chromic oxide dilution technique. Correction factors for these- estimates were derived by determining chromic oxide-recovery and by comparing the estimates to total fecal collection. Cow age had a significant (p<.05) effect on daily grazing time, but not on daily intake. Cows receiving supplement showed significantly higher intake (p<.05) than unsupplemented cows, but spent the same or less time grazing; Protein supplementation seems to have lowered the nutritional cost of obtaining forage. Daily temperature swings brought about adjustments in the cows' daily grazing schedule. Severe temperature changes also brought short term reductions in daily grazing time. During these periods the cows may have been experiencing acute cold stress. No consistent correlation between grazing time, past grazing time, present temperature, and past temperature was found. This led to the conclusion that in 1 984 and 1985 the cows did not experience chronic cold stress, and were not forced to re-acclimate, even though temperature varied between 8 and -26 degrees Celcius. PHYSIOLOGICAL AND BEHAVIORAL RESPONSES OF COWS ON MONTANA FOOTHILL RANGE TO WINTER AND SUPPLEMENT by Roger W i ll i a m Dunn A th e s is submitted in p a r t i a l f u lf i llm e n t o f t h e r e q u i r e m e n t s f o r t h e d eg r ee of Master of S c ie n c e ! in Range S c i e n c e MONTANA STATE UNIVERSITY Bozeman, Montana March, 1986 /V37£ ii APPROVAL o f a t h e s i s s u b m i t t e d by Roger W i ll i am Dunn T h i s t h e s i s h a s been r e a d by each member of t h e t h e s i s c o m m it te e and ha s been fo und t o 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 usage, 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 ll eg e o f G r a d u a te S t u d i e s . Date C h a i r p e r s o n , G r ad u at e Committee Approval f o r t h e Major Department ___ V Date w V? Vc V % G> Head, Major Department Approved f o r th e Co ll ege o f G rad u ate S t u d i e s Date G r ad ua te Dean iii STATEMENT OF PERMISSION TO COPY In p resen tin g th is th e sis in p artial fu lfillm en 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 o n t a n a S t a t e U n i v e r s i t y , I a g r e e t h a t t h e L i b r a r y s h a l l make i t a v a i l a b l e t o b o r r o w e r s under t h e r u l e s of t h e L i b r a r y . 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 l l o w a b l e w ithout sp ecia l perm ission, p r o v i d e d t h a t a c c u r a t e acknowledgement o f s o u r c e i s made. P e r m i s s i o n f o r e x t e n s i v e q u o t a t i o n from or r e p r o d u c t i o n o f t h i s t h e s i s may be g r a n t e d by my m a jo r p r o f e s s o r , o r i n h i s a b s e n c e , by th e D i r e c t o r o f t h e L i b r a r i e s when, i n t h e o p i n i o n of e i t h e r , t h e p r op os ed use of t h e m a t e r i a l i s f o r s c h o l a r l y p u r p o s e s . Any c o p y in g or use of t h e m a t e r i a l 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 i t h o u t my w r i t t e n p e r m i s s i o n . Signature Date iv ACKNOWLEDGEMENTS I wo uld l i k e t o e x p r e s s my a p p r e c i a t i o n t o t h e f a c u l t y and s t a f f o f t h e D e p a r t m e n t o f A n im a l a n d Range S c i e n c e s , and th e M o n ta n a A g r i c u l t u r a l Experiment S t a t i o n . The s t a f f a t t h e A n i m a l N u t r i t i o n L a b o r a t o r y , e s p e c i a l l y Nancy Roth and Kent W i l l i a m s , gave a s s i s t a n c e and p a t i e n c e when b o th were v e r y much needed, and v e r y much a p p r e c i a t e d . I My t h a n k s t o them. would l i k e t o t h a n k J e s s Miner f o r b e i n g o u t t h e r e w i t h me and r e m a i n i n g my f r i e n d t h r o u g h i t a l l . He w i l l n ev er be f o r g o t t e n . My t h a n k s t o S h a n n o n Q u i n s e y f o r h e r s i g n i f i c a n t ( p < . 0 I ) h e l p w i t h s t a t i s t i c s and SAS. Without h e r h e l p P d s t i l l be h e r e . T h e r e a r e tw o men I h a v e t h a n k e d o f t e n a n d w i t h -the u t m o s t s i n c e r i t y . I do so a g a i n now, b u t I ’m s u r e t h i s w i l l n o t be t h e l a s t tim e. S pecial thanks to K ris H avstad an d Eldon e n c ou r a ge , h e l p , and a l l o w me t o be and do my b e s t . Ayers. Such a g i f t means m o r e t o me t h a n I c a n s a y , a n d I am h o n o r e d t o be i t s r e c i p i e n t . friends, you’r e welcome by my f i r e a n y t i m e , y o u r s warm me. They My and t h a n k you f o r l e t t i n g , And f i n a l l y , s i n c e r e t h a n k s t o my f a m i l y , e s p e c i a l l y my m o t h e r a n d f a t h e r , S a r a h a n d L a r r y Dunn, a n d my b r o t h e r a n d s i s t e r - i n - l a w , Barry a nd J a n e Dunn, remember w here home i s . for th eir fa ith i n me, and f o r h e l p i n g me V TABLE OF CONTENTS LIST OF TABLES............................................................................................ .................... Page vi LIST OF FIGURES............................................................................................................. .. viii ABSTRACT............................................................................................................................ ix INTRODUCTION............................... I LITERATURE REVIEW........................................................................................................... 3 I n t a k e o f W in te r F o r a g e ................................................ W in te r a s i t A f f e c t s Metabolism and N u t r i t i o n ................................. Pregnancy as i t A f f e c t s D i e t a r y Requi rem ent s and I n t a k e . . . . S u p p l e m e n t a t i o n a s i f A f f e c t s W in te r Forage I n t a k e ................... I n t a k e B e h a v i o r ............................................................................................ 3 8 14 16 23 MATERIALS AND METHODS................................................................................................. 28 Study S i t e D e s c r i p t i o n .................................................................................... G r a z in g Management................ I n t a k e Measurement............................................................................................. B eh a v i o r E v a l u a t i o n .......................................................................................... Data A n a l y s i s .................................................................................. 28 29 29 34 35 RESULTS..................................................................... ........................................................ 38 1984 T r i a l .................................................................................. Body Weight and C o n d i t i o n S c o r e ........................................................ G r az in g B e h a v i o r .......................................................................................... I n t a k e .................. 1985 T r i a l ......................................................................................................... Body Weight and C o n d i t i o n S c o r e .................. G r az in g B e h a v i o r .......................................................................................... I n t a k e ...............................................................■................................................. 38 38 38 42 44 44 45 50 DISCUSSION........................................................................................................................... 53 . Forage U t i l i z a t i o n and F o r a g in g P a t t e r n s ............................................ E n v i r o n m e n ta l S t r e s s . . . . . ............................................................................ Forage I n t a k e ........................................................................................................ 53 54 57 SUMMARY................................................................................................................. 61 RECOMMENDATIONS AND QUALIFICATIONS................................................................... 63 LITERATURE CITED............................................................................................................. 66 APPENDIX...............................................................................................................................' 78 vi LIST OF TABLES Ta b le 1 2 3 4 5 6 7 8 9 10 11 12 Page S e a s o n a l c o m p o s i ti o n o f r an ge g r a s s e s i n e a s t e r n Montana and n o r t h e r n U ta h ................................... ............................... ............. ......... 4 A ppa ren t d i g e s t i b i l i t y of g r a s s c o n s t i t u e n t s d u r i n g . summer and w i n t e r i n U tah ............ .................... ...................... ............... g P r e d i c t e d g r a z i n g time o f cows a t v a r i o u s t e m p e r a t u r e s and wind v e l o c i t i e s ..................................................................................... 25 S i g n i f i c a n t c o r r e l a t i o n c o e f f i c i e n t s between c a t t l e a c t i v i t i e s and w e a th e r p a r a m e t e r s ..................................................... 26 Supplement c o m p o s i ti o n f o r w i n t e r , 1985 a t Red B l u f f R es ear ch Ranch, N o r r i s , Montana.................................................. E s t i m a t i o n o f f e c a l o u t p u t and f o r a g e i n t a k e f r o m f e c a l g r a b s a m p l e s ..................................................................... ...................... .. R e g r e s s i o n model f o r e x p l a i n i n g v a r i a t i o n i n th e d a i l y time s p e n t g r a z i n g d u r i n g w i n t e r .......................................................... Time s e r i e s a n a l y s i s of d a i l y g r a z i n g time bas ed on p a s t g r a z i n g t i m e s and p r e s e n t and p a s t t e m p e r a t u r e s ............ .. D a ily g r a z i n g t i m e s and d i s t a n c e s t r a v e l e d f o r 3 cow age groups., I 9 8 4 . .................................................. , . . . ............. Stepwise l i n e a r r e g r e s s i o n of d a i ly grazing time, 1984 ... 31 33 36 37 39 43 Time s e r i e s a n a l y s i s o f t o t a l d a i l y g r a z i n g t i m e and av e r a g e d a i l y t e m p e r a t u r e , 1984..................................... T o t a l d a i l y i n t a k e (0MB %BW) e s t i m a t e d from g r a b samp les and a d j u s t e d f o r Cr2 Og r e c o v e r y , 1984.................................. .. 43 44 13 Linear r e g r e s s i o n of d a i ly t o t a l in ta k e as r e l a t e d to d a i l y g r a z i n g t i m e , 1984. . . . . ....................................................................44 14 D a i l y g r a z i n g t i m e s and t r a v e l o f each t r e a t m e n t group, 1985........................................................................................................................... 15 S te p w ise l i n e a r r e g r e s s i o n o f d a i l y g r a z i n g t i m e , 1 9 8 5 . . . 16 Time s e r i e s a n a l y s i s of t o t a l d a i l y g r a z i n g time and av e r a g e d a i l y t e m p e r a t u r e , I 985.......................................... 46 49 50 v ii L IS T OF TABLES - C o n tin u e d Tablfe Page 17 D a i l y f o r a g e o r g a n i c m a t t e r i n t a k e (%BW), 1985........................ 51 18 L i n e a r r e g r e s s i o n o f d a i l y t o t a l i n t a k e , 1985.............. .... 52 APPENDIX 19 Body w e i g h t s and c o n d i t i o n s c o r e s f o r 5 cow age g r o u p s . 1984..................... ........................ ...................................... .................................... yg 20 Body w e i g h t s and c o n d i t i o n s c o r e s f o r 4 cow age g r o u p s , 1 9 8 5 . ............................................................................ ..................................... .. 79 Body w e i g h t s and c o n d i t i o n s c o r e s f o r each t r e a t m e n t g ro u p , 1985............................................................................ ...................... .. 80 21 22 D a i l y g r a z i n g t i m e s and d i s t a n c e s t r a v e l e d f o r f o u r cow age g r o u p s , 1985................................................ ........................... ............... 80 23 Chromic o x id e r e c o v e r y (% f e d ) , 19 85 ....................... ......................... 81 24 D a i l y f e c a l o u t p u t (0MB %BW) e s t i m a t e d u s i n g t o t a l f e c a l c o l l e c t i o n and Cr2 Og, I 985.......................................................... Si 25 T o t a l d a i l y i n t a k e (0MB %BW), 1985..................................................... 82 v iii L IS T OF FIGURES Figure 1 Page Heat p r o d u c t i o n a s r e l a t e d t o t h e th e rm al e n v ir o n m en t of b e e f c a t t l e ................................................................................................. 2 Heat p r o d u c t i o n and co l d a d a p t a t i o n i n b e e f c a t t l e ............ 3 A normal w i n t e r t e m p e r a t u r e p a t t e r n and t h e a s s o c i a t e d g r a z i n g a c t i v i t y . ............................................................ 4 i 11 4 G raz in g a c t i v i t y a s s o c i a t e d w i t h t h e 1984 t r i a l ' s c o l d e s t t e m p e r a t u r e s ......................................................... 5 A warming t r e n d and t h e a s s o c i a t e d g r a z i n g a c t i v i t y . . . . . ' 6 A t e m p e r a t u r e d e c l i n e and t h e a s s o c i a t e d g r a z i n g a c t i v i t y ....................................................................... 41 47 ' APPENDIX 7 Chromic o x i d e r e c o v e r y p a t t e r n d u r i n g t o t a l f e c a l c o l l e c t i o n , 1985.............................................................................................. o 83 ix ABSTRACT S u p p l e m e n t a t i o n o f g e s t a t i n g b e e f cow s s p e n d i n g t h e w i n t e r on r a n g e h a s shown b e n e f i t s t o cow p r o d u c t i v i t y . In order to understand how t h e s e b e n e f i t s o p e r a t e , d e t a i l e d s t u d i e s o f t h e n u t r i t i o n a l c o s t s f o r co w s on w i n t e r r a n g e , a n d t h e e f f e c t s o f s u p p l e m e n t s , a r e n e c e s s a r y . As p a r t o f t h a t r e s e a r c h g o a l , a s t u d y was c o n d u c t e d i n 1984 a n d 1985 t h a t r e c o r d e d t h e g r a z i n g a n d t r a v e l b e h a v i o r , a n d e s t i m a t e d t h e f o r a g e i n t a k e , o f g e s t a t i n g c r o s s b r e d co w s o n w i n t e r r a n g e r e c e i v i n g p r o t e i n s u p p l e m e n t . The r a n g e l a n d a v a i l a b l e t o t h e cows c o n t a i n e d cool s e a s o n g r a s s e s d o m i n a te d by bluebunch w h e a t g r a s s ( Agropyron s p ic a tu m (Pursh) S c rib n . & Smith). The s t u d y p a s t u r e c o n t a i n e d 324 h e c t a r e s i n t h e n o r t h w e s t f o o t h i l l s o f t h e Madison Range o f s o u t h w e s t Montana. The cows wore v i b r a c o r d e r s and p e d o m e te r s f o r 48 c o n t i n u o u s d a y s i n J a n u a r y and F e b r u a r y . D a i l y i n t a k e w as e s t i m a t e d u s in g the chrom ic oxide d i l u t i o n te c h n iq u e . C orrection f a c t o r s f o r these- e s t i m a t e s w ere d e r i v e d by d e t e r m i n i n g c h r o m i c oxider e c o v e r y a n d by c o m p a r i n g t h e e s t i m a t e s t o t o t a l f e c a l c o l l e c t i o n . Cow a g e h a d a s i g n i f i c a n t (p <.05) e f f e c t on d a i l y g r a z i n g t i m e , b u t n o t on d a i l y i n t a k e . Cows r e c e i v i n g s u p p le m e n t showed s i g n i f i c a n t l y h i g h e r i n t a k e (p<.05) t h a n u ns u p p le m en te d cows, but. s p e n t t h e same or l e s s time grazing. P r o t e i n s u p p l e m e n t a t i o n seems t o h av e l o w e r e d t h e n u t r i t i o n a l co st of o b ta in in g fo ra g e . ■D aily te m p e r a tu r e swings b r o u g h t a b o u t a d j u s t m e n t s i n th e cows’ d a i l y g r a z i n g s c h e d u l e . Severe te m p e r a t u r e changes a l s o brought s h o r t term r e d u c t io n s i n d a i l y g ra z in g tim e. D u r i n g t h e s e p e r i o d s t h e c o w s may h a v e b e e n e x p e rie n c in g acu te cold s t r e s s . No c o n s i s t e n t c o r r e l a t i o n b e t w e e n g r a z i n g tim e , p a s t g r a z i n g tim e , p r e s e n t t e m p e r a t u r e , and p a s t t e m p e r a t u r e w a s f o u n d . T h i s l e d t o t h e c o n c l u s i o n t h a t i n I 984 and 1985 t h e cows d i d n o t e x p e r i e n c e c h r o n i c c o l d s t r e s s , a n d w e r e n o t forced to re -a c c lim a te , even though t e m p e r a t u r e v a r i e d betwe en 8 and - 26 d e g r e e s C e l c iu s . I INTRODUCTION I n many a r e a s o f w e s t e r n Montana snow blows f r e e from r a n g e l a n d , and f o r a g e i s a c c e s s i b l e t o c a t t l e y e a r round. However, content of forage W inter range varies g reatly c o n ta in in g m ature, d e f i c i e n t i n some n u t r i e n t s , th e n u t r i e n t among a r e a s a n d p l a n t dormant g r a s s e s w i l l species. usually be and i s o f t e n i n a d e q u a t e t o m e e t t h e n u t r i e n t r e q u i r e m e n t s o f p r e g n a n t b e e f cows ( e s p e c i a l l y a s p r o d u c e r s t r y t o m a i n t a i n o r im pr o ve cow p r o d u c t i v i t y ) . N u t r i e n t s u p p l e m e n t a t i o n f o r b e e f cows w i n t e r i n g on r a n g e l a n d h a s b e e n s h o w n t o i m p r o v e cow p e r f o r m a n c e w i t h t h e a d d e d a n d d i r e c t l y related of b e n e f i t of im pr ove d c a l f p er f o r m a n c e . nutrien t w asteful. supplem entation can be However, p ro fitab le or th e p r a c t i c e econom ically The m o s t e f f e c t i v e s u p p l e m e n t f o r cows g r a z i n g w i n t e r f o r a g e i s one t h a t w i l l a i d t h e rum en m i c r o b i a l p o p u l a t i o n i n i t s d i g e s t i o n of t h e m a t u r e w i n t e r f o r a g e . The most, e f f i c i e n t s u p p le m en t i s one t h a t p r o v i d e s t h i s d i g e s t i v e a s s i s t a n c e w i t h o u t b e c o m i n o a d ie ta r y s u b s t i t u t e fo r w in te r forage. effectiv en ess, efficiency, In order to and p r o f i t a b i l i t y determ in e the of a supplem ent, a p r e c i s e d e t e r m i n a t i o n o f t h e b i o l o g i c a l b e n e n i t s g a i n e d w i t h each u n i t o f expense i s r e q u i r e d . Such d e t e r m i n a t i o n s w i l l p r o v i d e i n f o r m a t i o n f o r d e s i g n i n g o t h e r , more p r o f i t a b l e s u p p l e m e n t s . This study was a segment of an ongoing research project i n v e s t i g a t i n g several param eters of w in te r c a t t l e n u t r i t i o n including the b io lo g ic a l b e n e fits of supplem entation. This p o r t i o n of th e p r o j e c t h a s c o n c e n t r a t e d on t h e i n t e r a c t i o n o f d i e t a r y s u p p l e m e n t a t i o n and w i n t e r forage intake. The s p e c i f i c o b j e c t i v e s i n c l u d e d (I) 2 estim atio n of w in ter forage in tak e of b o t h s u p p l e m e n t e d and u n s u p p l e m e n t e d cow s i n t h e t h i r d t r o m e s t e r o f g e s t a t i o n , an d ( 2 ) me as ur e m en t o f t h e b e h a v i o r a s s o c i a t e d w i t h w i n t e r f o r a g e i n t a k e of both s u p p l e m e n t e d a n l u n s u pp le m en te d cows i n gestation. the t h i r d trim ester of The s t u d y used A n g u s - c r o s s cows g r a z i n g f o o t h i l l r a n g e l a n d t y p i c a l o f s o u t h w e s t Montana. Major g r a s s e s on t h e s t u d y s i t e i n c l u d e b l u e b u n ch u h e a t g r a s s ( A g r o p y r o n s p i c a t u m ( P u r s h ) S c r i b n . & S m i t h ) , n e e d l e a n d t h r e a d ( S t i p a , c o m a ta ’ T rin . & Rupr.), Id ah o f e s c u e ( F e s t u c a id a h o e n s is Elm er), a n d 'b a s in w ildrye (E l v m u s c i n e r e u s S cribn. & Mmrr. ) . Funding for the study was p r o v i d e d . j o i n t l y by t h e A g r i c u l t u r a l Expe rim ent S t a t i o n and Montana S t a t e U n i v e r s i t y . M o nt an a 3 LITERATURE REVIEW Any s t u d y of th e b e n e f i t s o f s u p p l e m e n t a t i o n must be l i m i t e d i n scope t o a l l o w s o l u t i o n s t o be a p p l i e d t o a s p e c i f i c s i t u a t i o n . This s t u d y was l i m i t e d t o c r o s s b r e d cows i n t h e l a s t t r i m e s t e r o f g e s t a t i o n (w in te r m onths), Montana, grazing n ativ e f o o th i ll rangeland in and r e c e i v i n g a p r o t e i n s u p p le m e n t. lim itatio n s behavior. h a s a n e f f e c t on t w o v a r i a b l e s , The f o l l o w i n g l i t e r a t u r e lim itatio n , th eir in teractio n s, review and th e ir southw est Each o f t h e a b o v e i n t a k e an d f o r a g i n g w ill deal effects w ith on each the tw o variables. I n t a k e o f Wi nte r Forage Forage av ailab le n u tr itio n a l q uality. shrubs for w in ter grazing is generally The n u t r i e n t c o n t e n t o f g r a s s e s , o f low forbs, and c h a n g e s s e a s o n a l l y , w i t h mi nim um l e v e l s o f s u c h i m p o r t a n t n u t r i e n t s a s p r o t e i n , phosp ho rou s, and c a r o t e n e o c c u r i n g i n t h e w i n t e r (Cook a nd H a r r i s 1 950b , M ars h e t a l . 1959). u s u a l l y low i n p r o t e i n d u r i n g t h e w i n t e r , energy to c a t t l e i f H a r r i s 1968b, th e y may p r o v i d e s u f f i c i e n t c o n s u m e d i n l a r g e e n o u g h q u a n t i t i e s (Cook a n d C la n to n 1981). ; A lthough g r a s s e s a r e ' S e a s o n a l l e v e l s o f i m p o r t a n t components ■ ! o f g r a s s e s h a v e b e e n m e a s u r e d i n U t a h an d M o n ta n a (Cook an d H a r r i s 19 5 0 a, 1 968b ; levels are M ar s h e t a l . in w inter Lignin w hile o th e r n u t r i e n t l e v e l s , e s p e c i a l l y crude p r o te i n , a re low er. Along than in and Thes e e a r l y summer, higher Table I . 1 965). levels generally in Van Dyne e t a l . cellulose- are summarized 1959; w i t h l o w n u t r i e n t c o n t e n t , m a t u r e d o r m a n t g r a s s e s a r e a l s o o f low 4 Tabl e I„ S ea s o n al c o m p o s i t i o n o f r a n g e g r a s s e s i n and n o r t h e r n ' Utah. eastern Montana Time Dec-Apr Dormant May-June Immature-bloom Component Species Years Crude Protein % Agropryon s p i ca turn 19551957 2.7-3.4 10-15 S tipa comata 19551960 1 . 9- 11 .2 11.3-14.7 Ii 19481953 3.5-6.0 9.0-17.0 grazing sample 19481953 3.0-5.0 8 . 0- 20.0 Agropyron s p ic a t u m 19551957 8.5-12.5 11. 0- 13.0 S tipa comata 19551960 12.0-21.5 10.7-16.9 ran ge grass mix 1968 12.7 (Sept) 9 .7 Cook and Harris 1968b 1950 12.5 ( S e p t) 10.0 Cook and Harris 1950a Lignin % U Citation VanDyne et al. 1965 it Marsh e t a l . 1959 ir VanDyne et al. 1965 Ir Cellulose % Bromus carinatus 1950 49.5-749.8 (Sept)■ 42.7-43.1 Cook and Harris 1950b Ca ro te n e m illigram % Agropyron sp ic at u m 19551960 6 . 2- 5 8 .0 (Sept) 118 .4-1 36 .1 VanDyne et al. 1965 Stipa comata 19551960 1 . 0- 8 .3 6 3 . 4 -1 3 9. 1 Il " 1948- 0-12.5 75.0-160.0 Marsh et al. 1959 5 T a b le I . c o n tin u e d Time Dec-Apr Dormant May-June Immature-bloom mS1953 0- 10.0 75.0-200.0 Marsh et al. 1959 Agropyron s p ic a t u m 19551957 0.0310.042 0.062-0.148 VanDyne et al. 1965 S tipa comata 19551960 0 . 0210.118 0.104-0.230 19481953 0.050.08 0.17-0.23 19481953 0.030.09 0.14-0.28 Component Species Years Car ot ene cont. grazing sample Phosphorous % IT grazing sample d ig estib ility cellu lo se due t o t h e higher (Cook a n d H a r r i s le v e ls of 1968 b). C itation It Marsh e t al. 1959 !I in d ig estib le l i g n i n and A com parison of the ap p are n t d i g e s t i b i l i t y o f g r a s s e s i n summ er a n d w i n t e r c a n be f o u n d i n T a b l e 2. Cook and H a r r i s (1968b) s t r e s s e d t h e f a c t t h a t t h e c h e m ic a l c o n t e n t s o f p l a n t s on w i n t e r r a n g e v a r i e d l i t t l e months. Van Dyne and Heady constituents te ste d for, season (1965) found th at over the w in te r of all chem ical l i g n i n v a r i e d l e a s t over t h e w i n t e r g r a z i n g 6 Tabl e 2. Apparent d i g e s t i b i l i t y o f g r a s s c o n s t i t u e n t s d u r i n g and w i n t e r i n Utah (from Cook and H a r r i s 1968b). Digestion C o effic ien t % Total Protein Gross ener gy summer Metabo­ lizable D i g e s t i b l e • ener gy protein (kcal/kg) Time Species Summer Agropyron in te r m ed iu m 66.2 71 .4 62.4 7.3 2253 A g ro p yr o n. in erm e 50.1 68.7 57.0 5. 2 1936 Agropyron sm ithii 7 3 .8 64.0 61 .2 11.1 2028 Agropyron inerme 0. 0 76.4 56.9 0. 0 1991 Agropyron sm ithii 6.4 70.1 64 .3 0.2 2469 27.6 69 .7 50.7 1.2 1647 69.5 45 .9 0.3 1616 W in te r Stipa comata O r y z o p s is ? hymenoides 6. 4 : Cellulose I f c a t t l e r e l y on t h i s l e s s d i g e s t i b l e , less nutritio u s more must be consumed t o o b t a i n n e c e s s a r y n u t r i e n t l e v e l s , t i m e and en er g y must be Under t h e s e c o n d i t i o n s , rumen and th e forage, and more expended t o d i g e s t what th e y have consumed. both the p h y sical cap acity of the digestive c a p a c i t y of t h e rumen beco me l i m i t i n g ( C r a m p t o n I 952 ). reticulo- m i cr o b e • p o p u l a t i o n This le a d s to d e c re a se d in t a k e , b e c a u se i n t a k e cannot ex ce ed o u tf l o w from th e rumen (p a s s a g e r a t e ) , and o u t f l o w i s dep endent on how f a s t t h e a n i m a l and i t s m i c r o b e s can b r e a k down t h e consumed p l a n t m a t e r i a l . For w i n t e r f o r a g e h i g h i n c e l l 7 w all c o n te n t d i g e s t i o n r a t e i s slow ed. This re d u c e s o u tflo w and l e a v e s l a r g e am ounts of fo o d r e s i d u e s i n th e rumen i n t h e form of a f l o a t i n g f i b e r ma t (Weston 1967, Jo n es 1972). and f u r t h e r i n t a k e i s l i m i t e d intake The rumen r e m a i n s f u l l ( C a m p l i n g e t a l . 196 2) . Dry m a t t e r c a n i n c r e a s e o n l y i f an i n c r e a s i n g a m o u n t o f d r y m a t t e r i s d i g e s t e d (Conrad e t a l . 1964). ', t I M o n t g o m e r y ' a n d B a u m g a r d t (1 96 5) s t a t e d t h a t c a t t l e w i l l a d j u s t t h e volume o f fo o d i n t a k e t o meet p h y s i o l o g i c a l en er g y demand i f rumen fill does not l i m i t consum ption. H ow e v er , f o r cow s g r a z i n g low q u a l i t y w i n t e r f o r a g e r u m e n f i l l d o e s a p p e a r t o be t h e m a i n f a c t o r l i m i t i n g i n t a k e ( B a l c h and C a m p l i n g 196 2, Grovum 1 96 9, J o u r n e t an d Bemond 1976 , E l l i s 197 8) . When t h e d i g e s t i b i l i t y o f f o r a g e i s l e s s t h a n a p p r o x i m a t e l y 65 p e r c e n t , r u m e n f i l l i s f a i r l y c o n t i n u o u s , and therefore c o n t i n u o u s l y l i m i t i n g ( J o n e s 1972). f o r a g e i n t a k e by c a t t l e , In a review co n c e r n in g C o r d o v a e t a l . ( 1 97 8 ) r e p o r t e d t h a t i n t a k e e s t i m a t e s i n t h e w e s t e r n U n i t e d S t a t e s r a n g e f r o m 1.0 t o 2.8 p e r c e n t o f body w e i g h t o r 40 t o 90 g dm/kg body w e ig h t . study at Montana S tate U n i v e r s i t y ’s In a previous w in te r Red B l u f f Research Ranch, e s t i m a t e d f o r a g e o r g a n i c m a t t e r i n t a k e r a n g e d from 0.8 t o 1.4 p e r c e n t o f body w e i g h t ( T u r n e r 1985). Und er t h e f i l l - l i m i t i n g conditions i m p o s e d by w i n t e r f o r a g e o f low q u a l i t y , d e c l i n i n g a v a i l a b i l i t y o f such f o r a g e does n o t a p p e a r to l i m i t i n t a k e . estim ated quality, the intake of heifers grazing but d e c l i n i n g a v a i l a b i l i t y . as th e q u a n t i t y of fo ra g e d ecre ase d , levels. H a v s t a d e t a l . (1 98 3) forage of c o n s t a n t low H e if e r s in c re a s e d grazing time and a c h i e v e d c o n s t a n t i n t a k e 8 Und er c i r c u m s t a n c e s w h e r e r u m e n f i l l i s n o t th e main f a c t o r l i m i t i n g i n t a k e , VFA ( v o l a t i l e f a t t y a c i d s ) , e s p e c i a l l y a c e t a t e a n d propionate, a p p e a r t o be i m p o r t a n t s i g n a l s u b s t a n c e s f o r t h e c o n t r o l of in ta k e . N e u r a l s e n s o r s f o r VFA a r e b e l i e v e d t o be l o c a t e d i n o r n e a r t h e r u m e n , ( B a i l e a nd May er 1 968, B i n e s 1971) a n d i n t h e p o r t a l v e i n ( B a i l e I 969 a s c i t e d i n B i n e s 1 9 7 1 ) . N e u r a l s i g n a l s a r e s e n t t o t h e hy p o th a lm u s which i s t h e c e n t e r of t h e i n t a k e (Conrad 1966). c o n t r o l mechanism As VFA l e v e l s go up, i n t a k e i s d e p r e s s e d (B h a t t a c h a r y a and Warner 1968, B a i l e and F o rb es 1974, Anil and F or b es 1977). K a r t c h n e r a n d Adams ( I 982) f o u n d l o w e r i n t a k e a n d h i g h e r r u m e n VFA l e v e l s i n cows s u p p le m e n te d e v e r y o t h e r day vs. cows sup p le m en te d e v e r y day i n t h e w i n t e r . T h i s s u g g e s t s t h e VFA i n t a k e m e c h a n i s m i s o p e r a t i n g d u r i n g t h e w i n t e r w i t h a lo w q u a l i t y r o u g h a g e d i e t . But J o n e s (1 972) p o i n t e d o u t t h a t l o w q u a l i t y r o u g h a g e d i e t s c o n t a i n i n g h i g h l e v e l s o f l i g n i n a r e s l o w l y d i g e s t e d , and rumen VFA l e v e l s may be to o low i n t h e s e intake. If so, c irc u m s ta n c e s to play a r o l e in the control of o t h e r m e c h a n i s m s may be i n v o l v e d i n t h e r e s u l t s r e p o r t e d by K a r t c h n e r and Adams (1982). These a u t h o r s s u g g e s t e d t h a t rumen c o n d i t i o n s i n t h e cows f e d g r a i n on a l t e r n a t e days may have been l e s s favorable f o r f ib e r digestion. Thi s c o n d i t i o n i s d i s c u s s e d i n a l a t e r s e c t i o n of t h i s review. W in te r a s i t A f f e c t s Metabolism and N u t r i t i o n W i n te r h a s some s t r i k i n g p h y s i o l o g i c a l and b e h a v i o r a l e f f e c t s on cattle. C li m a t e can a f f e c t a r an ge cow's r e p r o d u c t i v e p e r f o r m a n c e and m ilk p ro d u ctio n (B ellido e t al. 19 8 1 ). E ffe c ts of environm ental 9 s t r e s s , i n c l u d i n g t h e en er gy c o s t o f g r a z i n g and w a l k i n g , can r a i s e an a n i m a l s m a i n t a i n e n c e en er g y r e q u i r e m e n t 25 t o 50 p e r c e n t above t h a t of a n i m a l ’s k e p t i n d o o r s ( O s u j i 197*0. environment are s i g n i f i c a n t , A lthough th e e f f e c t s o f a cold th e y a r e n o t a l w a y s e x c e s s i v e and do n o t e x c l u d e p r o d u c t i v i t y ( e x p r e s s e d by cow w e i g h t , cow c o n d i t i o n , and f e t a l g r o w th ) d u r i n g t h e w i n t e r (W eb s te r 1970a). The most o b vi o u s e f f e c t o f t h e c o l d w i n t e r e n v i ro n m e n t i n v o l v e s t h e f a c t t h a t heat, e m i s s i o n from t h e a n i m a l r i s e s a s t h e t e m p e r a t u r e d r o p s ( B l a x t e r an d Wa in ma n 1 9 6 1 ) . Heat i s l o s t th ro u g h r a d i a t i o n , c o n d u c t i o n , a n d e m i s s i o n o f w a t e r v a p o r ( B l a x t e r a n d Wainman 196 1) . In fact, all in d irectly , animal physiologic responses to cold r e l a t e , directly or to an i n c r e a s e i n th e r a t e of h e a t t r a n s f e r betw een an and i t s e n v i r o n m e n t (C h r i s t o p h e r s o n and Young 1981). Webster (I 9 7 0 a , b) h a s sh o w n t h a t c a t t l e may g a i n r a d i a n t h e a t f r o m t h e s u n d u r i n g t h e day; how ever, r a d ia n t h e a t l o s s a t n ig h t outw eighs th e g a i n b et w ee n November and March i n s o u t h e r n A l b e r t a , Canada. This n e t r a d i a n t h e a t l o s s h a s an i m p o r t a n t and i m m e d i a t e e f f e c t . The an im al must produce h e a t to m ain tain counteract lo sses in order to homeothermy (Young and C h r i s t o p h e r s o n 1974). Cold s t r e s s , of exposure. and t h e a n i m a l ’s r e s p o n s e s t o i t , D uring acute (short conserves h e a t through p il o e r e c t io n , surface, low ered r e s p i r a t o r y r a te , a n d s e e k i n g s h e l t e r ( S l e e 1971, term ) cold vary w ith d u ratio n stress an a n i m a l v a s o c o n s t r i c t i o n a t the s k in d e c r e a s e d w a t e r and f oo d i n t a k e , Young 1 975a* Gonyou e t a l . 1979). S h o r t te rm c o l d s t r e s s i s a l s o acc ompanied by m a ss iv e h e a t p r o d u c t i o n ( t h e r m o g e n e s i s ) v i s i b l e a s s h i v e r i n g (Gonyou e t a l . 1979) and powered 10 by f r e e f a t t y a c i d s (Young I 9 7 5 a ) o b t a i n e d t h r o u g h t h e o x i d a t i o n o f a d i p o s e t i s s u e ( B l a x t e r and Wainman 1961). Acute c o l d s t r e s s o c c u r s i f an a n i m a l i s removed from i t s th erm o ­ n e u t r a l zone. T h i s i s t h e r a n g e of t e m p e r a t u r e s i n which an a n i m a l ' s heat production i s constant. F i g u r e I shows t h e t h e r m a l n e u t r a l zone and t h e r e g i o n s of t h e r m o g e n e s i s a s t e m p e r a t u r e s drop below o r r i s e above t h i s l e v e l . Heat Production Cold \ Thermo-' genesis Thermo­ neutral Zone Cold Thermal Environment F i g u r e I. Heat p r o d u c t i o n a s r e l a t e d t o t h e t h e r m a l e n v i ro nm en t of b e e f c a t t l e ( a d a p t e d from Young 1980). As c o l d e x p o s u r e c o n t i n u e s ( c h r o n i c c o l d s t r e s s ) , r e s p o n s e s t o c o l d c h a n g e , b u t t h e i n c r e a s e d m e t a b o l i c r a t e c o n t i n u e s ( S l e e 1971, Young and C h r i s t o p h e r s o n 1974, Young 1980). Young and C h r i s t o p h e r s o n ( 1 9 7 4 ) r e p o r t e d a n 8 t o 40 p e r c e n t r i s e i n t h e r m o n e u t r a l m e t a b o l i c h e a t p r o d u c t i o n d u r i n g p r o lo n g e d c o l d ex p o s u r e . The new l e v e l of h e a t p r o d u c t i o n depended on t h e av er a ge t e m p e r a t u r e t o which a n i m a l s had previously been exposed. S pecifically, the average increase in t h e r m o n e u t r a l r e s t i n g h e a t p r o d u c t i o n f o r e a c h 1°C r i s e i n a v e r a g e t e m p e r a t u r e t o which a n i m a l s had been exp os ed was 0.6 k c a l p e r kg body w e ig h t'? ^ p e r day. D uring p ro lo n g ed cold exposure the in c re a se d 11 m etab o lic rate acclim atization. becom es th e b asis for cold ad ap tatio n or The t h e r m o n e u t r a l z o n e s h i f t s t o i n c l u d e t h e new l e v e l o f h e a t p r o d u c t i o n a s s e e n i n F i g u r e 2. Heat Production Cold adapted Warm adapted Tem perature F i g u r e 2. Heat p r o d u c t i o n and co l d a d a p t a t i o n i n b e e f c a t t l e from Young 1980). (adapted Each s h i f t i n t h e t h e r m o n e u t r a l zone b r i n g s a new l o w e r c r i t i c a l tem perature. critical Even W ebster (1970a) r e p o r t e d that the absolute low er t e m p e r a t u r e f o r p r e g n a n t b eef cows r a n g e s from -11 to -23°C. though tem peratures drop below -23°C in w inter, c a ttle can u s u a l l y s u r v iv e th e s e p e r io d s w ith a c u te cold re s p o n s e s . Young (1975b) s t u d i e d a group of cows a c c l i m a t i z e d t o J a n u a r y t e m p e r a t u r e s a p p r o a c h i n g -25°C. When exposed t o -30°C, metabolic r a t e s increased up t o 37%, b u t r e c t a l t e m p e r a t u r e s d e c r e a s e d s h o w i n g t h a t t h e cows w er e n o t m a i n t a i n i n g homeothermy. A cclim atization also involves oth er m echanism s a s d e fe n s e s a g a i n s t co ld such as an i n c r e a s e d h e a r t r a t e w hich increased m etabolic r a te (W ebster e t a l . 1970). c o n t r i b u t e s to R espiratory ra te d e c r e a s e s t o m i n i m i z e l o s s o f h e a t f r o m w a t e r v a p o r ; Young ( I 9 7 5 a ) reported re sp ira to ry r a t e s i n cows a s lo w a s 8 t o 11 b r e a t h e s p e r 12 m in u te . V asoconstriction a t the skin surface continues, v a s o d ila tio n occurs, prevent tis s u e follow ed by s l o w constriction, but p e r i o d i c in order to damage a t t h e e x t r e m i t i e s (B l a x t e r and Wainman 1961). H a i r s h e d d i n g i s r e d u c e d , which can l e a d t o a h a i r c o a t t w i c e a s heavy a s unexposed a n i m a l s (Webster e t a l . 1970). Cold e x p o s u r e b r i n g s on a r e l a t i v e l y dehydrated c o n d itio n i n exposed an im als as t h e i r w ater c o n s um p ti o n d r o p s (Young 1975a). As c a t t l e became a d a p t e d t o c o l d e r tem peratures, m etabolic c a p a b ili ty i n r e s p o n s e to a c u t e co ld s t r e s s i n c r e a s e d ( S l e e 1971). These mechanisms combine t o i m p l y a g r e a t e r e n e r g y consumption by a n i m a l s a c c l i m a t e d t o c o l d . The h e a t c o n s e r v a t i o n a t t e m p t e d d u r i n g a c u t e c o l d s t r e s s i s a c c o m p a n i e d by a d r o p i n f o o d i n t a k e ( W e b s t e r a n d Young 1 970, S l e e 1971)=. The g r e a t e r e n e r g y o u t f l o w t h a t f o l l o w s d u r i n g c h r o n i c c o l d s t r e s s i s a c c o m p a n i e d , by a n i n c r e a s e i n f o o d i n t a k e a b o v e n o r m a l levels (Webster and Young 1970, Webster e t a l . 1970, S l e e 1971, B a i l e and F o r b e s 1974, Ames and Br in k 1977). The pr o lo n g ed co l d changes t h e a n i m a l s p h y s i o l o g i c and e n v i r o n m e n t a l c i r c u m s t a n c e s . In o rd e r to m a in ta i n a c o n s ta n t energy b alan ce w ith th e se changes, c a t t l e must in crease t h e i r food i n t a k e ( B a i l e a n d F o r b e s 1 97 4) . H ow ev er, the i n c r e a s e d h e a t p r o d u c t i o n needed d u r i n g p r o lo n g e d co l d e x p o s u r e w i l l o c c u r no m a t t e r w h at t h e f o o d i n t a k e l e v e l o f t h e a n i m a l (B l a x t e r and Wianman 1961, Young 1975b). t i o n to th e animal, This shows t h e i m p o r t a n c e o f h e a t p r od u c­ and s u p p o r t s t h e f i n d i n g s t h a t a d i p o s e t i s s u e i s a l s o us ed t o f u e l h e a t p r o d u c t i o n . Thus f a r , te m p e r a tu r e alone has been d is c u s s e d to i n d i c a t e th e d e g r e e o f c o l d exp os ur e. Wind can a g g r a v a t e t h e e f f e c t o f t e m p e r a t u r e 13 w i t h p r o n o u n c e d a d d i t i o n s t o t h e e f f e c t i v e t h e r m a l .en vir onm en t, cold tem peratures, wind w i l l At i n v o k e h i g h e r h e a t p r o d u c t i o n (Webster 19 7 0 a) a n d a h i g h e r m e t a b o l i c r a t e (C hristopherson e t al. 19 7 9 ). I n s l e y and Ames (1972) d e t e r m i n e d t h a t wi nd v e l o c i t e s u n d e r 32 k m/ hr rem oved an i n s u l a t i n g l a y e r of a i r s u r r o u n d i n g t h e a n i m a l and t r a p p e d i n t h e h a i r co at . Above, 32 km/hr t h e wi nd a l s o d e s t r o y e d t h e s t r u c t u r e o f t h e i n s u l a t i n g h a i r c o a t c a u s i n g a second, accelerated r is e in heat flow . Cold e x p o s u r e n o t o n l y i n c r e a s e s an a n i m a l ’s m a i n t a i n e n c e ener gy requirem ents, i t also in flu e n c e s th e d ig e s 'tiv e system of ru m in a n ts . S e v e r a l s t u d i e s hav e shown a s i g n i f i c a n t d e c r e a s e i n DMD (dry m a t t e r d i g e s t i b i l i t y ) d u r i n g p r o l o n g e d c o l d e x p o s u r e ( B l a x t e r a n d Wainman 1961 , Young a n d C h r i s t o p h e r s o n 19 7 4 , Kennedy e t a l . 1 9 7 6 , Ames and B r in k 1977). The mechanisms of t h i s d e c r e a s e i n d i g e s t i b i l i t y i n v o l v e in c re a s e d d i g e s t s flow t r a c t (Kennedy e t a l . th r o ug h t h e rumen and l o w e r g a s t r o i n t e s t i n a l 1976, Kennedy and M i l l i g a n 1978) a s s o c i a t e d w i t h i n c r e a s e d r u m i n a t i o n t i m e (Gonyou e t a l . 1979) and i n c r e a s e d r e t i c u l a r c o n t r a c t i o n s ( W es tr a and C h r i s t o p h e r s o n 1976). The increased passage rate through the reticulorum en is p a r t i a l l y c o u n t e r a c t e d by t h r e e r e s p o n s e s : i n c r e a s e d i n t a k e d u r i n g p r o l o n g e d c o l d e x p o s u r e and t h u s a n i n c r e a s e i n t h e t o t a l am ou nt o f n u trien ts p resen ted for d ig estio n ( Young 1981, W estra and C h r i s t o p h e r s o n 1976), an i n c r e a s e i n t h e a p p a r e n t d i g e s t i b i l i t y i n t h e in testin e s (Kennedy e t al. 197 6) , and an i n c r e a s e in m icrobial e f f i c i e n c y a s s o c i a t e d w i t h t h e d e c r e a s e d t u r n o v e r t i m e (Kennedy e t a l . 1 976 ). An e f f e c t o f c o l d e x p o s u r e p o s s i b l y r e l a t e d t o c h a n g e s i n 14 d i g e s t i b i l i t y i s a d e c r e a s e i n r u m e n f l u i d v o l u m e (D ege n an d Young 1980, Kennedy e t a l . 1 976) f o l l o w e d by a d e c r e a s e i n t h e v o l u m e o f in terstitial f l u i d and p l a s m a (Degen and Young 1980). Young (1981) and Young and C h r i s t o p h e r s o n ( 1974) have su mmarized t h e e f f e c t s o f cold. • They found a 16% av er a ge i n c r e a s e i n a n nu al f e e d needs caused asso ciated in w ith equal proportions increased passage by rate (I) a reduction and d e c r e a s e d in DMD, d igestive e f f i c i e n c y , and ( 2 ) a h i g h e r m a i n t a i n e n c e r e q u i r e m e n t , a s s o c i a t e d w i t h i n c r e a s e d r e s t i n g m e t a b o l i c r a t e and an i n c r e a s e d en er g y r e q u i r e m e n t . These authors e f f e c t ", also discussed th e need to a "relativ ely produce h e a t minor but im portant during ac u te cold s t r e s s (i.e. s t o r m s ) i n o r d e r t o m a i n t a i n homeothermy. Pregnan cy as i t A f f e c t s D i e t a r y Re qu ir em ent s and I n t a k e Added to t h e e f f e c t s o f p r o lo n g e d c o l d ex p o su re a r e some im p o r ­ t a n t e f f e c t s of l a t e pregnancy. cavity i s lim ite d , volume, as the fe tu s S in c e t h e volum e of t h e a b d o m i n a l ' g r o w s a n d o c c u p i e s mo re o f t h a t rumen volume d e c r e a s e s (Grahm and W i l l i a m s 1962; F o r b e s 1969, 1970; B in e s 1971;). Although u t e r u s volume i n c r e a s e s s t e a d i l y , F or b es ( 1968) found t h a t rum en volum e d i d n o t d e c r e a s e u n t i l t h e l a s t f i v e weeks of pregnancy. I n an e x p e r i m e n t c o n d u c te d w i t h ewes (F o rb es 1969) i t was conc lu ded t h a t t h e volume o f t h e abdominal c a v i t y has an u p p e r . l i m i t and t h a t th e ex p a n d a b le o r g a n s ' w i t h i n th e c a v i t y must compete f o r t h a t space. The t h r e e t i s s u e s most i n c o m p e t i t i o n a r e t h e u t e r u s , t h e rumen, and d e p o s i t s of ab d o m in al f a t . A decline in intake i n th e l a s t weeks o f pregnancy has been a t t r i b u t e d to th e r a p i d l y 15 e x p a n d i n g u t e r u s c a u s i n g a r e s t r i c t i o n on t h e a m o u n t o f v o l u n t a r y i n t a k e p o s s i b l e (John son e t a l . 1966, Cu r r an e t a l . 1967, F o r be s 1970, B i n e s 1971). Pregnancy d i d n o t seem t o a f f e c t t h e volume of d i g e s t a i n t h e l o w e r i n t e s t i n a l t r a c t (For bes 1969). C a m p l i n g (1966) pregnancy. h a s shown ch a n g e s i n i n t a k e behavior during P r e g n a n t cows a t e more s l o w l y and r u m i n a t e d l o n g e r th a n t h e i r n o n - p r e g n a n t mo no z y go tic t w i n s . Thi s b e h a v i o r r e s u l t e d i n a 13 p e r c e n t l o w e r hay i n t a k e f o r th e p r e g n a n t cows. It has also been shown t h a t in creased passage rate is a p h y s i o l o g i c a l s i d e e f f e c t o f l a t e p r e g n a n c y (Grahm and W i l l i a m s 1962). A lthough i n c r e a s e d p a s s a g e r a t e would te n d to o f f s e t t h e d e c re a s e d i n t a k e caused by s m a l l e r rumen volume, b o th f a c t o r s would c o n t r i b u t e t o un de r n u t r i t i o n i n t h e l a s t weeks o f pregnancy. Forb es (1970) p o s t u l a t e d t h a t a change i n e n d o c r i n e b a l a n c e cou ld a l s o cause a d e p r e s s i o n i n fo o d i n t a k e . p r o g e s t e r o n e w a s sh o w n t o d e p r e s s Estrogen i n t h e p r e s e n c e of v o l u n t a r y i n t a k e , and e s t r o g e n s e c r e t i o n by t h e p l a c e n t a i n c r e a s e s d u r i n g t h e l a s t t h i r d o f pregnancy (Robin so n 1957 a s c i t e d i n F o rb es 1970). R o b i n s o n a nd F o r b e s (1 967) s t u d i e d t h e p r o t e i n r e q u i r e m e n t s o f m a t u r e b r e e d i n g ew es . A d e c l i n e i n i n t a k e was s e e n a s p r e g n a n c y advanced; w as m o r e p r o n o u n c e d on l o w - p r o t e i n d i e t s the decline (5.5 p e r c e n t v s . H o w e v er , the 11 p e r c e n t efficien cy pregna nc y advanced. of crude p ro tein in digested the e n t i r e N u tilizatio n ration). im proved as The i n t a k e of a p p a r e n t l y d i g e s t e d N r e q u i r e d f o r maximum r e t e n t i o n e f f i c i e n c y d e c r e a s e d from 0.623 g/kgW^'^^/day a t 10-12 weeks p r e g n a n t t o 0.567 g / k g W0,7 3 / d ay a t 18-20 weeks p re g n an t . 16 T he fetu s has undernutrition. p rio rity for am ino acids in co n d itio n s of The d e c r e a s e i n e q u i l i b r i u m N r e q u i r e m e n t s s e e n i n t h i s s t u d y s u g g e s t s some t r a n s f e r o f N from m a i n t a i n c e t o p r o d u c t i o n d u r i n g pr egn anc y and u n d e r n u t r i t i o n . B esid es usin g N for feta l developm ent, the anim al also e x p e r i e n c e s an i n c r e a s e d e n e r g y r e q u i r e m e n t d u r i n g p r e g n a n c y . This h a s been shown w i t h m e a s u r e s o f h e a t l o s s from p r e g n a n t and non­ p r e g n a n t ewes. During t h e 145 days o f pregnancy, p r e g n a n t ewes l o s t 39.5 m e a l m o r e h e a t t h a n - n o n - p r e g n a n t e w e s ( B r o c k w a y e t a l . 19 6 3 ). During pr egn anc y t h e a n i m a l e x p e r i e n c e s g r e a t e r n u t r i t i o n a l n eed s w i t h a d e c r e a s i n g a b i l i t y t o o b t a i n them. S u p p l e m e n t a t i o n a s i t A f f e c t s W in te r F or ag e I n t a k e The most i m p o r t a n t and most v a r i e d s u p p l e m e n t s f o r cows g r a z i n g w i n t e r r a n g e a r e p r o t e i n and energ y. As d i s c u s s e d p r e v i o u s l y , a r e o f te n l i m i t i n g i n a w in te r range s e t t i n g . these A compounding f a c t o r i s t h e f a c t t h a t e n e r g y n eed s a r e met f i r s t and p r o t e i n w i l l be used f o r e n e r g y u n t i l i t i s no l o n g e r l i m i t i n g ( C la n to n 1981). When f e e d i n g a d i e t a r y s u p p l e m e n t t o cows on w i n t e r r a n g e , i t i s im portant to feed ju s t th a t, Therefore, a supplem ent, and n o t a r e p l a c e m e n t . th e s u p p l e m e n t ' s e f f e c t s on t h e i n t a k e and d i g e s t i b i l i t y o f t h e f o r a g e a r e m o s t i m p o r t a n t a n d w i l l be d i s c u s s e d h e r e . S e v e r a l e x p e r i m e n t s have shown t h a t p r o te in supplem ent can i n c r e a s e t h e i n t a k e of low p r o t e i n f e e d (Cla nto n and Zimmerman 1965, Cook a n d H a r r i s 1 9 6 8 a, To pp s 1972, B r a n i n e a n d G a l y e a n ( 1 985) C l a n t o n 1981, Turner rep o rte d increased passage r a te 1985). (a n d 17 d e c r e a s e d t u r n o v e r t i m e ) a nd i n c r e a s e d i n t a k e w he n s t e e r s g r a z i n g m a t u r e b l u e g r a m a r a n g e l a n d w e r e f e d .5 k g o f c o r n / head / day c o m p a r e d t o s t e e r s f e d no s u p p l e m e n t a n d I kg o f c o r n / h e a d / da y . However, th is e f f e c t i s n o t alw ays seen (R itte n h o u se 1 9 6 9 ). The i n f l u e n c e s seem t o be dep endent on t h e c o n c e n t r a t i o n o f crude p r o t e i n i n the forage. A gen era lly accepted r u l e i s w ill i n t a k e w h en p r o t e i n l e v e l s i n f o r a g e a r e b el o w 8% increase th a t p r o t e i n supplements ( A l l i s o n 19 8 5 ). As h a s a l r e a d y been d i s c u s s e d , of p assag e i n pr egn anc y, the re tic u lo ru m e n , cold weather s t i m u l a t e s th e r a t e rum en volume during and l i g n i n and c e l l w a l l c o n s t i t u e n t c o n t e n t s a r e h ig h i n w i n t e r f o r a g e w h i l e p r o t e i n c o n t e n t i s low. p article decreases G iven t h e s e f a c t o r s , s i z e r e d u c t i o n of i n d i g e s t i b l e f i b e r i n t h e rumen would a p p e a r t o be t h e l i m i t i n g p r o c e s s i n r e m o v i n g m a t e r i a l from t h e rumen and allow ing achieving fu rth er particle ferm entation. intake size (We lc h reduction 1982). are The t w o r u m i n a t i o n and methods of m icrobial R u m i n a t i o n i s a p r o c e s s d i r e c t l y c o n t r o l l e d by t h e a n im a l i n r e s p o n s e t o . t h e f i b e r f r a c t i o n of th e f o r a g e . The t i m e s p e n t r u m i n a t i n g h a s shown a s t r o n g p o s i t i v e c o r r e l a t i o n t o t h e - c e l l w a l l c o n s t i t u e n t c o n t e n t o f f e e d ( W e l c h a n d S m i t h I 9 6 9 ). M icrobial f e r m e n t a t i o n r e l i e s on t h e a c t i v i t y and numbers o f rumen m i c r o f l o r a . Any f a c t o r , i n c l u d i n g a l l t h o s e m e n t i o n e d above, t h a t d e c r e a s e s m i c r o f l o r a a c t i v i t y o r n u m b e r s c a n be e x p e c t e d t o d e c r e a s e DMD ( C r a m p t o n 1952) and, therefore, decrease intake. The p o s i t i v e e f f e c t of p r o t e i n s u p p l e m e n t a t i o n on d i g e s t i b i l i t y a n d i n t a k e may be due t o a p o s i t i v e ' e f f e c t on m i c r o b i a l dig estio n (Cam pling e t a l . 1 962, Eg an 19 6 5 )^ 18 P r o t e i n s u p p l e m e n t a l l o w s t h e o p t i m i z a t i o n of m i c r o b i a l s y n t h e s i s , num bers, and a c t i v i t y a s th e a v a i l a b l e n i t r o g e n c o n t e n t of th e d i e t in c r e a s e s a b o v e ,th a t a v a i l a b l e i n fo rag e alone (A rias e t a l. 1951, Moir and H a r r i s 1962, Zin n and Owens 1981). -1 ' , These c o n c l u s i o n s a r e n o t s u p p o r t e d by r e c e n t s t u d i e s t h a t have a t t e m p t e d t o show i n more d e t a i l t h e e f f e c t s of p r o t e i n s u p p le m e n t on N d i g e s t i o n and rumen m i c r o b i a l e f f i c i e n c y . G o e t s c h and Owens ( a s c i t e d i n G o e t s c h e t a l . I 984) a n d G o e t s c h e t a l . ( 1 98 4 ) h a v e r e p o r t e d c o n f l i c t i n g r e s u l t s on t h e im p a c t o f p r o t e i n s u p p le m e n t. The f i r s t s t u d y showed im p ro v ed m i c r o b i a l e f f i c i e n c y (g m i c r o b i a l N / kg o r g a n i c m a t t e r f e r m e n t e d ) when a d i e t c o n t a i n i n g 12 p e r c e n t CP (crude p r o t e i n ) was s u p p l e m e n t e d w i t h soybean or c o t t o n s e e d meal t o a l e v e l p r o v i d i n g 17 p e r c e n t CP. The s e c o n d s t u d y s h o w e d no s u c h i m p r o v e d m i c r o b i a l e f f i c i e n c y a s CP l e v e l s w e r e i n c r e a s e d f r o m 9.2 t o I 5.1 p e r c e n t . addition, In b a c t e r i a l numbers and t h e amount o f m i c r o b i a l N p a s s i n g t o th e duodenum r e m a i n e d t h e same a s CP i n th e d i e t r o s e . However, rumen p r o t o z o a numbers and ADF ( a c i d d e t e r g e n t f i b e r ) d i g e s t i o n i n c r e a s e d i n t h e se cond s t u d y when t h e b a s a l d i e t was s up p le m en te d . Grummer e t a l . (1984) co n cl u d ed t h a t changes i n rumen ammonia c o n c e n t r a t i o n "do n o t r e g u l a t e p r o t e i n or dry m a t t e r degradation by r u m i n a l m i c r o b e s " . There r e m a i n s a need f o r c l a r i f i c a t i o n o f t h i s t o p i c . There i s s t r o n g e v i d e n c e s h o w i n g i n c r e a s e d i n t a k e o f l o w q u a l i t y f o r a g e when s m a l l amounts o f p r o t e i n su p p le m e n t a r e p r o v id e d . S t u d i e s sh owing e v i d en ce t o t h e c o n t r a r y may ,have f e d d i e t s d i f f e r i n g t o s u c h a n e x t e n t t h a t ' I r e s u l t s s h o u l d h o t be compared. 1-9 W h i l e s u p p l e m e n t s may i n c r e a s e t h e d i g e s t i b l e p r o t e i n r e c e i v e d from ra n g e f o r a g e , Cook a n d H a r r i s (1 96 8a ) sh o we d t h a t t h e r e i s a n a c c o m p a n y i n g d e c r e a s e i n t h e d i g e s t i b i l i t y o f c e l l u l o s e and g r o s s en erg y. The cows r e c e i v e d l e s s m e t a b o l i z a b l e ener gy when r a n g e f o r a g e was s u p p le m en te d. A p o r t i o n o f t h e p r o t e i n i n any f e e d s t u f f i s u n a v a i l a b l e i n t h e r u m e n , an d b y p a s s e s o r e s c a p e s t o t h e l o w e r g a s t r o i n t e s t i n a l t r a c t . The amount o f bypass o r es ca p e p r o t e i n v a r i e s n o t o n l y b et we en f e e d s , b u t a l s o w i t h i n f e e d s due t o f e e d p r o c e s s i n g and "a n im al , d i e t a r y , and m i c r o b i a l v a r i a b l e s " (NEC 19 8 4 ). The NEC ( I 984) h a s s u g g e s t e d t h r e e c a t e g o r i s e s o f f e e d s t u f f s based on t h e i r p e r c e n t a g e o f bypas s p r o t e i n : ( I ) low b y p a s s ( l e s s t h a n 40 p e r c e n t ) , i n c l u d i n g s o y b e a n m e a l and peanut to m eal; ( 2 ) medium bypass (40 60 percent), including c o t t o n s e e d m e a l , d e h y d r a t e d a l f a l f a m e a l , c o r n g r a i n , and b r e w e r s d r i e d g r a i n s ; and (3) h i g h bypass ( g r e a t e r th a n 60 p e r c e n t ) , i n c l u d i n g me at me al, c o r n g l u t e n me al, blood m e al , f e a t h e r me al, and f i s h meal. The s u p p l e m e n t s u se d i n t h i s s t u d y had a so ybean meal base. H arris ( 196 8 a) u s e d t h e n y l o n b ag t e c h n i q u e to Cook and determ ine th a t a soybean meal s u p p le m e n t c o n t a i n i n g 48.2 p e r c e n t t o t a l p r o t e i n was 83.4 percent d i g e s t i b l e i n t h e rumen. Hume (1 974) r e p o r t e d a s i m i l a r a p p a r e n t DMD f o r s o y b e a n m e a l o f 83.1 p e r c e n t . bypass p r o te in , In h i s review of C h a l u p a ( 197 5) r e p o r t e d t h a t 40 t o 59 p e r c e n t o f so ybean meal p r o t e i n w i l l bypas s or e s c a p e t h e rumen, and t h e r e f o r e , w i l l be u n a v a i l a b l e t o rumen mi cr o b es . The benefits of bypass protein resu lt from a increase in. e s s e n t i a l amino a c i d s a v a i l a b l e t o th e a n i m a l i n t h e s m a l l i n t e s t i n e 20 and g r e a t e r t r u e d i g e s t i b i l i t y o f q u a l i t y p r o t e i n s c o m p a r e d t o t h e digestibility However, of crude p r o t e i n from rumen m i c r o b e s as p r o t e i n b yp as s o r e s ca p e i n c r e a s e s , (Van S o e s t 1983). t h e o p t i m i z a t i o n of m i c r o b i a l s y n t h e s i s , n u m b e rs , and a c t i v i t y i s n o t p o s s i b l e , and t h e p o s i t i v e fo ra g e in ta k e e f f e c t seen w ith p r o te i n supplem ent i s l e s s likely. E x p e r i m e n t s w her e s u p p l e m e n t a l en erg y ( c a r b o h y d r a t e ) was f e d have g e n e r a l l y shown a d e c r e a s e i n f o r a g e i n t a k e due t o a s u b s t i t u t i o n of t h e s u p p l e m e n t f o r f o r a g e ( H o l d e r I 9 6 2 , C a m p l i n g a n d Mu rdoch 1 966, E llio tt 1 96?, R ittenhouse e t al. 1 970, B e l l o w s a n d Thomas M e r r i l l and K l o p h e n s t e i n 1984, A l l i s o n 1985). been shown to decrease the d ig estio n 1 976, Energy s u p p le m e n ts have of cellu lo se and o th e r c a r b o h y d r a t e s (Cook and H a r r i s 1968a ) „ S t u d i e s w here both p r o t e i n and e n e r g y a r e f e d a r e i n c o n c l u s i v e . C la nt on (1981) f e d c o m b i n a t i o n s o f p r o t e i n and en er g y t o cows, but saw no s i g n i f i c a n t d i f f e r e n c e i n dry m a t t e r i n t a k e or d i g e s t i b i l i t y when averages for supplem ented i groups were compared to the control i ( un s up p le m en te d ) group. R al e ig h and W a l la c e (1963) found an i n c r e a s e i n f o r a g e i n t a k e whe n l o w l e v e l s o f p r o t e i n w e r e f e d r e g a r d l e s s o f energy le v e l. When h i g h l e v e l s o f p r o t e i n w e r e f e d , d ecre ase d , ag ain , r e g a r d l e s s of energy le v e l. forage in tak e T u r n e r (1 985) f o u n d i n c r e a s e d i n t a k e o f r a n g e f o r a g e w i t h 1.8 k g / d a y o f a 1 5 % p r o t e i n supplem ent. B u t w hen h a l f t h a t same s u p p l e m e n t a n d 0.9 k g / d a y o f a 30% p r o t e i n s u p p le m e n t w ere f e d , was seen. no i n c r e a s e i n i n t a k e o v er c o n t r o l s 21 K a r t c h n e r (1981) s u p p l i e d a p r o t e i n and an en er g y su pp le m en t t o g r o u p s of cows d u r i n g two s u c c e s s i v e w i n t e r s . The f i r s t w i n t e r was m i l d w i t h r e a d i l y a v a i l a b l e f o r a g e , no i n t a k e a d v a n t a g e w a s g a i n e d fr om e i t h e r s u p p le m e n t. The second w i n t e r was more s e v e r e and i n t a k e and DMD w ere enhanced by p r o t e i n s u p p l e m e n t , bu t d e p r e s s e d w i t h energy su p p le m en t. The e f f e c t o f s u p p l e m e n t c a n a l s o be a f f e c t e d by t h e t i m i n g o f supplem entation. Adams (1984b) and K artchner and Adams (1982) r e p o r t e d t h a t cows f e d e v e r y day g a i n e d w e i g h t a n d c o n d i t i o n w h i l e t h o s e f e d e v e r y o t h e r day g a i n e d l e s s w e i g h t a n d l o s t These f i n d i n g s w ere a t t r i b u t e d to the condition. e f f e c t s w i t h i n t h e rumen. A l t e r n a t e day f e e d i n g b r o u g h t r e d u c e d ph and i n c r e a s e d VFA c o n c e n t r a ­ tio n s. Lower r u m e n ph b r i n g s a b o u t r e d u c e d f i b e r d i g e s t i o n w h i c h c o u l d a c c o u n t f o r t h e l o w e r w e i g h t g ai n . Adams ( 1 9 8 4 a , b) a l s o s t u d i e d t h e e f f e c t o f t h e t i m e o f day o f supplement feeding. S te e rs fe d i n the e a r l y afternoon spent l e s s time g r a z i n g , but consumed more f o r a g e , and had a h i g h e r a v e r a g e d a i l y g a i n t h a n s t e e r s f e d i n t h e morning. supplem ent i n The a u t h o r postu lated th a t feeding th e morning d is r u p te d normal g r a z in g a c t i v i t y and a f f e c t e d b o th b e h a v i o r and pe r f or m a n ce . Many stu d ies have d ealt w ith th e c u r r e n t and s u b s e q u e n t p e r f o r m a n c e o f co w s r e c e i v i n g w i n t e r s u p p l e m e n t . Cows h a v e l o s t s u b s t a n t i a l a m o u n t s o f body f a t o v e r t h e w i n t e r w i t h o u t a d v e r s e l y a f f e c t i n g t h e i r s u b s e q u e n t p e r f o r m a n c e or th e p e r f o r m a n c e of t h e i r c a l v e s (Thompson e t a l . 1983, Turner 1985). But s u p p le m en t l e v e l has b e e n d i r e c t l y r e l a t e d t o a cow’s w i n t e r w e i g h t l o s s a n d i n v e r s e l y 22 r e l a t e d t o h e r co ns um pt ion o f s p r i n g , summer f o r a g e and summer w e i g h t g a i n ( R i g h t 1 966, J o r d a n e t a l . 1 968, L u s b y e t a l . I 976, Thompson e t a l . 1 9 8 3 , T u r n e r I 985 ). Cows r e c e i v i n g l i t t l e o r no w i n t e r s u p p l e m e n t h a v e sh own a reduced reproductive cap a b ility . S e v e r a l s t u d i e s have shown a d el a y e d r e t u r n t o e s t e r u s i f p r e - c a l v i n g n u t r i t i o n i s low ( W i ltb a n k e t a l . I 962, J o r d a n e t a l . I 968, Dunn e t a l . I 9 6 9 , W h i t m a n 1 975, B e l l o w s a n d Short 1 978). calving, For cow s i n good, m oderate, and t h i n condition a t t h e i r l i k e l i h o o d of e s t r u s 60 days p o s t p a r t u m was 0.91, 0.61, a n d 0.46 r e s p e c t i v e l y ( W h i t m a n 1 9 7 5 ) . C o n c e p t i o n r a t e may a l s o be l o w e r f o r cows on a low p la n e o f w i n t e r n u t r i t i o n ( H a r r i s e t a l . 1956, W i l t b a n k e t a l . 1 962 , C l a n t o n an d Z i m m e r m a n 1 9 6 5) . When cows on a h ig h p la n e of n u t r i t i o n b e fo re and a f t e r c a l v i n g w ere bred, percent did not conceive, only 5 c o m p a r e d t o 33 p e r c e n t o f t h e cows n o t c o n c e i v i n g h a v i n g b e e n on a l o w p l a n e o f n u t r i t i o n ( W i l t b a n k e t a l . 1962) . I t i s i m p o r t a n t t o r e m e m b e r t h a t a cow c a n o b t a i n t h e e x c e s s e n e r g y needed d u r i n g t h e w i n t e r th r o u g h e x t r a f e e d o r m o b i l i z a t i o n of fat. I f cows a r e n o t r e c e i v i n g e x t r a p r o t e i n a n d e n e r g y p r i o r t o calving, t h e y w i l l l i k e l y be l o s i n g w e i g h t a s t h e y u s e body t i s s u e reserves. The f e r t i l i t y o f a cow 70 t o 90 d ay s p o s t p a r t u m i s d i r e c t l y r e l a t e d t o t h e amount of body en er g y r e s e r v e s she ha s (Whitman 1975). Tho mp son e t a l . ( 1983) d i s c u s s e d t h e t o t a l e n e r g y r e q u i r e m e n t f o r t h r e e me thods o f w i n t e r i n g c a t t l e : I) f a l l f a t t e n i n g and l i t t l e w i n t e r s u p p l e m e n t , 2 ) k e e p i n g co w s i n a t h i n c o n d i t i o n y e a r r o u n d , and 3 ) k e e p i n g cows f a t y e a r round. Al l cows were k e p t a t s i m i l a r p r o d u c t i o n 23 lev els. energy, Covzs t h a t w e r e f a t y e a r r o u n d r e q u i r e d t h e l e a s t t o t a l and f a l l fa tte n in g req u ired t h e most. The body e n e r g y r e s e r v e s o f t h e f a t oows a l l o w e d them t o o v e r w i n t e r w i t h l e s s s u p p l e ­ ment t h a n t h e t h i n cows w h i l e s t i l l r e m a i n i n g p r o d u c t i v e . A l i m i t e d amount o f p r e - c a l v i n g w e i g h t l o s s i s n o t d e t r i m e n t a l . Jones e t al. ( 19 79 ) f o u n d t h a t cows c a n l o s e up t o 60 kg (+3 6. 2) w i t h o u t a f f e c t i n g p r e - weaning d a i l y g a i n o r weaning w e i g h t . Weight l o s s be y o n d t h i s l e v e l h a s b e e n s h o w n t o l o w e r c a l f b i r t h w e i g h t s , c a l f gain, c a l f weanin g w e i g h t , and t h e number of c a l v e s weaned (Black e t a l . 1 938, S p e t h e t a l . 1 963» C l a n t o n a n d Zimmerman. 1 9 6 5 , R i g h t 19 6 6 , H i r o n a k a a n d P e t e r s 1969 , 1979 , B e l l i d o e t a l . 1981). Corah e t a l . 19 7 5 , R ussell et al. . I n t a k e B eh av io r B e h a v io r i s an i m p o r t a n t , tion. u n d e r l y i n g f e a t u r e o f a n i m a l p r odu c­ I t i s i m p o r t a n t - t o produce b e e f e f f i c i e n t l y ; therefore, i t is i m p o r t a n t t o u n d e r s t a n d b e e f c a t t l e b e h a v i o r ( S t r i c k l i n an d K a u t z Scanavy 1984). But t h e r e s u l t s of b e h a v i o r s t u d i e s a r e o f t e n v a r i b l e an d d i f f i c u l t to i n t e r p r e t . (H a n c o c k 1953, cattle (H a n c o c k V ariation S t o b b s 1970) due t o 1950 as cited in among anim als i s high the in h e re n t in d i v id u a l ity Hancock 1953). The f a c t of th at o b s e r v e r s have a te n d e n c y t o av o id i n c l e m e n t w e a t h e r and d a r k n e s s adds i n c o n s i s t e n c i e s w i t h i n an d b e t w e e n b e h a v i o r s t u d i e s 1970). When s t u d y i n g i n t a k e b e h a v i o r , . (Gary e t a l . . animal i n d i v i d u a l i t y i s expressed t o th e g r e a t e s t e x t e n t a t n i g h t w h i l e d a y t im e g r a z i n g i s more l i k e l y 24 to be a group a c t i v i t y ( B a i l e and F o rb e s 1974, S t r i c k l i n e t a l . 1976). Many s t u d i e s h a v e i d e n t i f i e d t h e h o u r s j u s t a f t e r s u n r i s e an d a t s u n s e t a s t h e m o s t a c t i v e g r a z i n g p e r i o d s ( S t o b b s 1 9 7 0 , A r n o ld an d D u d z i n s k i I 978, Bueno a n d B u c k e b u s c h 1 979, R o a th a n d K r u e g e r 1982, S t r i c k l i n a n d K a u t z - S c a n a v y 1 9 8 4 ). c o n s is te n t g ra z in g a c t iv it y around m i d n ig h t (S to b b s S tu d ie s have a ls o d u rin g th e 1970, sh o w n l e s s day a n d a t n i g h t , S tric k lin e t a l. D u d z in s k i 1978, S t r i c k l i n and K au tz-S can av y 1984). 1976, u su a lly A r n o ld and Gary e t a l . (1970) r e p o r t e d t h a t 17 p e r c e n t o f b e e f c a t t l e g r a z i n g o c c u r ed a t n i g h t w h i l e 83 p e r c e n t o c c u re d d u r i n g d a y l i g h t . G iv e n t h e su n se t, b a sic p a tte rn of d a y lig h t g ra z in g a t su n rise and w e a th e r and tim e o f y e a r have been shown t o i n f l u e n c e g r a z i n g p a t t e r n s (A rnold and D u d z in sk i 1978, Ruckebusch and Bueno 1978, Low e t al. 1981, Roath and K rueger 1982). L ength o f d a y l i g h t h a s been shown t o a f f e c t t h e t i m e o f day w hen g r a z i n g b e g i n s a n d e n d s ; a s d a y s g e t s h o r t e r more d a y l i g h t h o u r s w ere s p e n t g r a z i n g ( T a y l e r 1951 a s c i t e d i n Hancock 1953, A rnold and D u d z in sk i 1978, Bueno and Ruckebusch 1979, W ils o n and F ly n n 1979, Low e t a l . 1981). However, d a y le n g th does n o t d e t e r m i n e t h e t o t a l , am o u n t o f t i m e s p e n t g r a z i n g e a c h d a y ; cow s co m p en sa te f o r s h o r t d a y l i g h t h o u r s i n w i n t e r by g r a z i n g more a t n i g h t (S tric k lin e t al. 1976, W ilson and F ly n n 1979). A lth o u g h t e m p e r a t u r e h a s n o t b e e n show n t o i n f l u e n c e w hen a g r a z i n g a c t i v i t y o c c u r s (A rnold and D u d z in sk i 1978, Roath and K rueger I 9 8 2 ), i t d o e s a f f e c t t h e a m o u n t o f d a i l y g r a z i n g t i m e . Cows s p e n d l e s s t i m e g r a z i n g on c o ld days th a n on warm days ( S m o liak and P e t e r s 1 955, M a le c h e k a n d S m i t h I 976, Adams 1 9 8 4 a ). T ab le 3 sh o w s t h e 25 e x te n t to w h ic h g ra z in g tim e c a n be e x p e c t e d t e m p e r a t u r e f a l l s o r w ind v e l o c i t y r i s e s . to d ecrease w hen This d e c r e a s e d g r a z i n g tim e on c o ld days h a s been a s s o c i a t e d w i t h d e c r e a s e d en e rg y e x p e n d i t u r e i n o r d e r t o c u t m a i n t a i n e n c e en e rg y r e q u i r e m e n t s . ' R i t te n h o u s e and S e n f t (1982) showed t h a t g r a z i n g t i m e d e c r e a s e s w i t h any d e v i a t i o n from th e th e r m a l n e u t r a l zo n e t o w h ich c a t t l e a r e a c c l i m a t e d . been shown t o a f f e c t g razin g a c t i v i t i e s b u t m o s tl y i n warm o r t r o p i c a l s e t t i n g s . g ra z in g tim e H u m id ity h a s (A rnold and D u d z in sk i 1978), Adams (1984a) h a s shown t h a t d e c r e a s e s a s w in d v e l o c i t y i n c r e a s e s . T a b l e 4 sh o w s s i g n i f i c a n t c o r r e l a t i o n c o e f f i c i e n t s b etw ee n v a r i o u s a n im a l a c t i v i t i e s and t h r e e w e a th e r f a c t o r s . T ab le 3 . P r e d i c t e d g r a z i n g tim e o f cows a t v a r i o u s wind v e l o c i t i e s (from Adams 1984a). Mimimum d a i l y a i r t e m p e r a t u r e °F Average d a i l y wind v e l o c i t y (MPH) O 5 10 15 30 9.1 8.8 8 .4 8.0 15 8.1 7 .8 7 .4 7 .0 0 7.1 6 .7 6 .4 6 .0 -15 6.0 5.6 5.3 4 .9 -30 5 .0 - 4.6 4.3 3 .9 D ra stic b e h a v io r. t e m p e r a t u r e s and changes i n w e a th e r Bueno and d iso rg a n iz a tio n of th e b rin g d r a s ti c R ucke bus ch (1 9 7 9 ) r e p o r t e d d a y lig h t r a t i o ex tre m e changes i n w e a th e r. ch an g es i n an im al a " c o m p le te of g ra z in g " d u rin g q u ic k , S t r i c k l i n e t a l . (1 976) r e p o r t e d t h a t 26 c a t t l e d id n o t g r a z e a t a l l d u r in g a th u n d e r s to r m ; th e y t u r n e d t h e i r b ack s t o t h e w ind and s to o d o r w alk ed w i t h t h e wind. F orage a v a i l a b i l i t y f o r a g e becomes s c a r c e , h as been shown t o a f f e c t g r a z i n g ti m e s . As th e number o f d a i l y g r a z i n g h o u r s i n c r e a s e s a s a n i m a l s t r y to o b t a i n a c o n s t a n t i n t a k e l e v e l (A lld e n and W h i tta k e r 1970, H av stad e t a l . 1983, L a th ro p e t a l . 1985). T ab le 4 . S i g n i f i c a n t c o r r e l a t i o n c o e f f i c i e n t s betw een c a t t l e a c t i v i ­ t i e s and w e a th e r p a r a m e te r s ( a d a p te d from Malechek and Smith 1 9 7 5 ). A c tiv itie s D is ta n c e T r a v e le d A ir Temp Baro P re s change Wind Speed G ra z in g ns ns .65 .56 VO O Weather F acto rs ns Time s p e n t R um inating S ta n d i n g ns .63 ns . 6 8 .70 ns' ns - n o n s i g n i f i c a n t c o r r e l a t i o n c o e f f i c i e n t s D a ily t r a v e l by c a t t l e h a s shown a c l o s e c o r r e l a t i o n t o th e tim e spent g razin g , w ith in itia tio n of tra v e l o ccu rin g alo n g w ith i n i t i a t i o n o f g r a z i n g a t s u n r i s e a n d s u n s e t ( R u c k e b u s c h a n d Bueno 1978). The t i m e s p e n t w a lk in g and t h e d i s t a n c e t r a v e l e d p e r day a r e a ffe c te d by w e a th e r a v a ila b ility , and (esp. w a te r Ruckebusch and Bueno 1978). as re p o rte d w in d ), fo rag e lo c a tio n a v a ila b ility , sh e lte r ( A r n o l d a n d D udzi n s k i . 1978, The r a n g e o f d i s t a n c e s t r a v e l e d by c a t t l e i n th e l i t e r a t u r e ( A r n o l d a n d D u d z i n s k i 1 9 7 8 ). is q u ite broad, 1.3 t o 12.6 k m /d a y Some s p e c i f i c e x a m p l e s i n c l u d e : M o n ta n a sum m er m o u n t a i n r a n g e c a ttle tra v e le d On 3.7 t o 5.7 k m /d a y 27 (L ath ro p e t a l . tra v e le d 19 8 5 ). On o p e n r a n g e i n c e n t r a l A u s t r a l i a c a t t l e 9.3 k m / d a y (S.D.=2„6) (Low e t al. 1981). Texas r a n g e l a n d w alk ed 5.3 km/day (Hancock 1953). B eef c a ttle on 28 METHODS AND MATERIALS S tudy S i t e D e s c r i p t i o n The S t u d y w as c o n d u c t e d i n a 324 h e c t a r e p a s t u r e l o c a t e d on Red B l u f f R e s e a r c h R anch o p e r a t e d by t h e D e p a r t m e n t o f A n im a l a n d Range S c i e n c e s , M o n ta n a S t a t e U n i v e r s i t y , fo o th ills of th e M a d is o n R ange i n in c lu d e d in th e le g a l I 8, T .3S ., R.1 E. The r a n c h i s i n t h e n o r t h w e s t so u th w e st d e sc rip tio n of th e M o n ta n a , s tu d y S e c tio n s p a stu re are P a s t u r e e l e v a t i o n s r a n g e f r o m 1 ,400 t o 1 ,9 0 0 m e t e r s ( 4 ,0 0 0 t o 6 ,2 0 0 f e e t ) w i t h l o n g , s l i g h t t o m o d e r a t e s l o p e s an d a r e a s 1 i o f s t e e p s l o p e s and r o c k o u tc r o p s . Annual p r e c i p i t a t i o n a v e r a g e s from 350 to 406 mm (14 t o 16 i n c h e s ) ( USDA-SCS 1 976). A 1980 S o i l C o n s e r v a t i o n S e r v i c e s u r v e y o f t h e r a n c h d e s c r i b e d t h e p a s t u r e a s a s i l t y r a n g e s i t e i n good c o n d i t i o n w i t h v e g e t a t i o n composed o f 6 5 p e r c e n t g r a s s e s and 35 p e r c e n t f o r b s and weedy s p e c i e s . Dominant g r a s s e s i n c l u d e bluebunch w h e a t g r a s s , fescue, and b a s in w ild r y e . n e e d l e a n d t h r e a d , Idaho O th e r m a jo r f o r a g e s p e c i e s l i s t e d by M ueggler and S t e w a r t (1980) and Payne (1 973) in c l u d e p r a i r i e J u n e g r a s s ( K o e l e r i a o v r a m i d a ta (Lam.) Beav,), t h r e a d l e a f s e d g e ( C a r e x f i l i f o l i a N u tt.), green n e e d le g ra ss (S tio a v ir id u la ( Bromus m a r g i n a tu s P i p e r (H itc h c .) ) , T rin .), m o u n t a i n brom e Kentucky blu e g r a s s (Poa o r a t e n s i s L .) , w e s t e r n y a r r o w ( A c h i l l e a m i l l e f o l i u m L .), and f r i n g e d s a g e w o r t ( A r t e m i s i a f r i g i d a W illd .) . S h allo w s i t e s s u r r o u n d i n g r o c k o u t c r o p s i n t h e p a s t u r e c o n t a i n po n d ero sa p in e (P in u s p o n d e ro sa D ougl.), l i m b e r p i n e ( P i n u s f l e x u l i s J a m e s ) , a n d Rocky M o u n t a i n j u n i p e r ( J u n i o e r u s s c o D u lo r u m S a r g.) A Red B l u f f Ranch p l a n t s p e c i e s l i s t c o m p i l e d by P a t P la n t e n b u r g can be fo u n d i n T u rn e r ( 1 9 8 5 ). 29 Payne (1973), i n r e v i e w i n g w e s t e r n M ontana's m a jo r r a n g e s i t e s , s t a t e d t h a t f o o t h i l l r a n g e l a n d d o m i n a t e d by b l u e b u n c h w h e a t g r a s s i s one of th e m ost common f o r a g e ty p e s in th a t p a rt of th e s t a t e . E s t i m a t e d c a r r y i n g c a p a c i t y f o r t h i s f o r a g e ty p e was t h r e e a c r e s p e r AUM (a n im a l u n i t month). G ra z in g Management I n 1984, 48 cows w e re p la c e d i n t h e s tu d y p a s t u r e on December 20 a n d r e m o v e d M arch 2. S t o c k i n g d e n s i t y w a s 0.15 AU ( a n i m a l u n i t s ) p e r • h e c t a r e (0 .0 6 A U / a c r e ) . The s t o c k i n g r a t e i n 1984 w as 2 .7 8 h e c t a r e s p e r AUM (7 a c r e s / AUM). I n 1 9 8 5 , 6 0 cow s w e r e i n t h e p a s t u r e f r o m D e c e m b e r 14 t o M arch I . ( 0 .0 7 AU/a c r e ) . S t o c k i n g d e n s i t y w as 0 .1 9 AU p e r h e c t a r e The s t o c k i n g r a t e w a s 2.08 h e c t a r e s p e r AUM (5.6 acres/A U M ). O b s e r v a t i o n s o f g r a z i n g d i s t r i b u t i o n w e r e made e a c h w i n t e r ; how ever, g r a z i n g d i s t r u b u t i o n was n o t q u a n t i f i e d . was e s t i m a t e d w i t h a u t i l i z a t i o n gauge (U. S. F orage u t i l i z a t i o n F o re st S erv ice ). g a u g e w as d e v e l o p e d u s i n g p l a n t h e i g h t / w e i g h t r a t i o s g r a s s e s and s e d g e s . The o f m a jo r M easurem ents i n c l u d e d u n g ra z e d and g ra z e d p l a n t h e i g h t a v e r a g e s o f b lu e b u n ch w h e a t g r a s s o r n e e d l e a n d t h r e a d , and number o f g r a z e d p l a n t s oh s i x s t e p t r a n s e c t s . T ra n s e c ts w ere d i s t r i b u t e d th r o u g h o u t th e p a s t u r e . I n t a k e Measurment The e x p e r i m e n t a l cows w ere ra n d o m ly s e l e c t e d from t h e Red B l u f f R e s e a r c h Ranch h e r d b a s e d on a g e , e x p e c t e d c a l v i n g d a t e s , • and c a l f sire s. Herd s i z e was chosen based on t h e q u a n t i t y o f f o r a g e a v a i l a b l e 30 i n t h e s tu d y p a s t u r e , se le c tio n c r ite r ia . and t h e number o f cows on t h e r a n c h t h a t f i t t h e The cows w ere 3 to 7 y e a r s of age. The cows w ere a r t i f i c i a l l y in s e m in a te d th e p re v io u s June u sin g angus or h e re fo rd semen. C a lv in g was s c h e d u le d to s t a r t March 4 e a c h y e a r ; S u p p l e m e n t w a s f e d i n d i v i d u a l l y on a l t e r n a t e d a y s i n t h e e a r l y a fte rn o o n . I n 1984, cow s w e r e f e d t h e sam e s u p p l e m e n t : 0.9 kg / day o f a soybean meal su p p le m e n t p r o v i d i n g 250 g CF (cru d e p r o t e i n ) / day. I n 1985 t h e cow s w e r e s p l i t i n t o f i v e t r e a t m e n t g r o u p s . Cow a g e g r o u p s , e x p e c t e d c a l v i n g d a t e s , and c a l f s i r e s w e r e d i s t r i b u t e d a s e v e n ly a s p o s s i b l e th ro u g h a l l t r e a t m e n t s . The t r e a t m e n t s , and t h e i r I n t e r n a t i o n a l Feed Numbers (IFN#) w ere a s f o l l o w s : TRT I - Forage o n ly (CON) TRT 2 - F orage p lu s so y b ean m eal (SBM, IFN# 5-04-600) TRT 3 - F orage p l u s soybean m eal and a n im a l f a t (FAT, IFN# 4-00-409) TRT 4 - F orage p l u s so y b ean m eal and b lood meal (SM, IFN# 5 -0 0 -3 8 0 ) TRT 5 - F o r a g e p l u s s o y b e a n m e a l and c o r n g l u t e n meal(CGM, IFN# 5 -2 8 -2 4 1 ) Blood m eal and c o rn g l u t e n meal w ere f e d a s s o u r c e s o f b y p a s s p r o t e i n , fro m a n a n im a l and p l a n t s o u r c e r e s p e c t i v e l y . Animal f a t was f e d a s a s o u r c e o f b y p ass en e rg y i n o r d e r to d e t e r m in e i f any e f f e c t s e e n w i t h b y p ass p r o t e i n was due t o 1e x t r a e n e r g y o r e x t r a p r o t e i n . A d e ta ile d d e s c r i p t i o n o f t h e s u p p le m e n ts f e d i n 1985 i s shown i n T a b le 5. C h r o m ic o x i d e w as u s e d a s a n e x t e r n a l m a r k e r t o e s t i m a t e f e c a l o u t p u t ( R a le ig h e t a l . sm all 1980). am ount of ground corn; Chromic o x id e powder was m ixed w i t h a th is p r e m i x w as t h e n a d d e d t o th e 31 T ab le 5 . Supplem ent c o m p o s itio n f o r w i n t e r 1985 a t Red B l u f f R esearch Ranch, N o r r i s , Montana (Miner e t a l . 1986). TRT I Supplem ent C o n stitu e n ts g fed 2 SBM 570 250 430 50 200 3 FAT SBM 210 51SL 0 252. 250 410 430 840 0 52. 50 0 2m 200 370 100 100 50 52. 200 40 162. 200 g TDN Bypass (g) 780 290 60 SSL 0. T o ta l 450 140 Ii 606 400 470 190 10 2. ■ 200 T o tal 230 4B0 680 200 222. 400 340 IM 480 160 M 200 BM SBM 5 g CP T o ta l CGM SBM Urea 4 P ro te in A v a ila b ility Rumen (g) 1P o ta s s iu m c h l o r i d e , d e calciu m p h o s p h a te , V itam in A, and a m o la s s e s b o o s t e r w ere added t o a l l s u p p le m e n ts . s u p p le m e n ts . i n 1985. The s u p p le m e n t was p e l l e t e d i n 1984, F e c a l g ra b s a m p le s from t h e e n t i r e h e rd w e re t a k e n i n 1984 and b iw e e k ly i n 1985. also b u t was f e d l o o s e made on I n 1985, t o t a l f e c a l f o u r cows from each group u s i n g c o lle c tio n s w ere f e c a l bags and u rin e / f e c e s s e p e r a t o r f l a p s (K a r tc h n e r and R it te n h o u s e 1979). w ere made t w i c e d u r i n g t h e s tu d y p e r io d . h o u rs f o r 4 c o n s e c u tiv e days. m ixed, and sam pled. w eekly C o lle c tio n s Bags w ere changed e v e ry 24 E ach d a y 's c o l l e c t i o n w a s w e i g h e d , Sam ples w ere f r o z e n f o r l a t e r a n a l y s i s . A f t e r th a w in g , dry m a t t e r and o r g a n i c m a t t e r w ere d e te r m in e d f o r e ac h f e c a l sam ple f o l l o w i n g AOAC (1975) p r o c e d u r e s . th e s am p le was ground th ro u g h a 2mm s c r e e n . tio n in The r e m a in d e r o f Chromic o x id e c o n c e n tr a ­ e a c h -sam p le w a s d e t e r m i n e d u s i n g t h e s p e c t r o p h o t o m e t r y te c h n i q u e ( W il li a m s e t a l . 1962). a to m ic a b s o r p tio n F e c a l dry m a t t e r 32 o u t p u t p e r day was e s t i m a t e d from g r a b s a m p le s based on t h e f o l l o w i n g eq u atio n : g chrom ic o x id e f e d / day . g f e c a l o u t p u t / day = ---------------------------------------------------- % ioo $ chrom ic o x id e i n dry f e c a l sam ple Two a d j u s t m e n t s w ere made t o t h i s o r i g i n a l e s t i m a t e from g r a b sam p les. C h r o m ic o x i d e r e c o v e r y w as c a l c u l a t e d a n d t h e q u a n t i t y o f c h r o m i c o x id e fe d per day a d j u s t e d a c c o rd in g ly . A lso , fecal o u tp u t m e a su re m e n ts from t o t a l f e c a l c o l l e c t i o n s w e re used t o a d j u s t o u tp u t c a l c u l a t e d f r o m g r a b s a m p l e s ( R i t t e n h o u s e 1969, R i t t e n h o u s e e t a l . 1970). S am p les o f g r a z e d f o r a g e w e re c o l l e c t e d th r o u g h o u t t h e s tu d y p e rio d s u sin g 4 e s o p h a g e a lly f i s t u l a t e d r e v ie w e d by Van Dyne and T o r r e l l (1964). freeze d rie d d ig e stib ility ) and of g round. each IVOMD sam p le w as cow s. The t e c h n i q u e w a s E x tr u s a was f r o z e n and l a t e r (in v itro d e te rm in e d o rg a n ic u sin g m o d i f i c a t i o n o f th e , T i l l e y 1arid T e rry te c h n i q u e ( H a r r i s m a tte r th e B arnes 1970). Rumen f l u i d f o r t h e IVOMD t r i a l s was c o l l e c t e d from rumen f i s t u l a t e d cows on a 60% d i g e s t i b l e (IVOMD) a l f a l f a h a y d i e t . The h a y w a s u s e d a s t h e s t a n d a r d f o r t h e IVOMD c o m p ariso n s. F e c a l o u tp u t and f o r a g e d i g e s t i b i l i t y e s ti m a t e s w ere used to e s ti m a t e in ta k e u sin g th e f o llo w in g eq u atio n : g D a i l y f e c a l o u tp u t (OMB) g o r g a n i c m a t t e r i n t a k e = —------------------------------------------I - IVOMD o f d i e t T ab le 6 g iv e s a c o m p le te l i s t of th e c a l c u l a t i o n s u sed to e s t i m a t e t o t a l o rg a n ic m a tte r in ta k e . 33 T ab le 6 . E s t i m a t i o n o f f e c a l ouput and f o r a g e i n t a k e from f e c a l g ra b s am p les. Item D e sc rip tio n A PPM Chromium i n f e c a l sample B O rganic m a t t e r o f sample C Weight o f sample 0.6842 C o n v ersio n o f PPM Cr t o Cr2 Og 0.1 D i l u t i o n f a c t o r f o r a c i d ' d i g e s t i o n t e c h n iq u e D g Cr2 Og f e d p e r day E Cr2 Og r e c o v e r y ($ o f f e d ) DxE=F A d ju s te d Cr2 Og f e d p e r day 0.6842xCxBxF ---------------------=G AxO. I • kg f e c a l o u tp u t p e r day H A djustm ent f a c t o r f o r each TRT group comparing t o t a l c o l l e c t i o n w ith g ra b sam ples HxG=I A d ju s te d f e c a l o u p u t J Body w e ig h t i n kg (I/J)x 1 0 0 = K L F e c a l o u t p u t a s a % o f body w e ig h t •. . IZI-L=M (MZJ)x100=N Forage d i g e s t i b i l i t y (IVOMD) T o t a l i n t a k e (kgZday) T o ta l i n t a k e (% BW) Cows w e r e w e i g h e d a n d c o n d i t i o n s c o r e d i m m e d i a t e l y b e f o r e and a f t e r th e s tu d y p e r io d s . I n t e r m e d i a t e w e i g h t s an d c o n d i t i o n s c o r e s w e re t a k e n once d u r i n g t h e 1985 s tu d y p e r io d . C o n d itio n s c o r e s w ere a 34 v i s u a l and p a l p a b l e s c o r e f r o m one t o t e n ( t h i n t o f a t ) (L aM o n ta g n e 1981). B e h a v io r E v a l u a t i o n ■ V i b r a c o r d e r s w e re u sed t o r e c o r d t h e t i m e s p e n t g r a z i n g . Use of v i b r a c o r d e r s i s d e s c r i b e d by S t o b b s (I 9 7 0 ), S t r i c k l i n e t a l . (1 97 6 ), Bueno and Ruckebusch (1979), and Adams (1984a). each 5, o f 3 age groups, 3, and 7 y e a r s v i b r a c o r d e r s and g r a z i n g a c t i v i t y > F e b r u a r y 27. I I n 1984, 4 cows from o ld , w ere f i t t e d w ith was r e c o r d e d betw ee n J a n u a r y 9 and 'I I n 19 8 5 , 4 cow s f r o m e a c h t r e a t m e n t g r o u p an d t h e c o n t r o l group w ore v i b r a c o r d e r s from J a n u a r y 3 to F e b ru a ry 28. b o th y e a r s , cow s w e a r i n g v i b r a c o r d e r s p e d o m e te r s . In d iv id u a l w ere a lso fitte d During w ith cows w ore t h e same v i b r a c o r d e r and pedom eter th r o u g h o u t t h e r e c o r d i n g p e r io d . P e d o m e te rs were s t r a p p e d t o e i t h e r f r o n t f o o t i n 1984, b u t r e m a in e d on t h e l e f t f r o n t f o o t th r o u g h o u t t h e 1985 t r i a l . V ib r a c o r d e r c h a r t s w ere r e a d to in c lu d e o n ly g r a z i n g o r n o t g r a z i n g , no o t h e r l e v e l s o f a c t i v i t y w e r e a s s i g n e d . w e re rounded t o 15 m in u te i n t e r v a l s . G ra z in g tim e s Each day was d i v i d e d i n t o t h r e e p e r i o d s : 7AM t o 1PM, IPM to 7PM, and 7PM t o 7AM. The number of h o u r s s p e n t g r a z i n g d u r i n g e a c h p e r i o d a n d e a c h day w e r e r e c o r d e d a s a. p e r c e n ta g e of th e t o t a l num ber o f h o u r s i n t h a t p e r io d . w e re r e a d and z e r o e d e v e r y 6 to 8 days. P e d o m e te rs C a l i b r a t i o n was a c c o m p lish e d by z e r o i n g and r e i n s t a l l i n g th e p e d o m e te r s , th e n moving th e cows w h ile f o l l o w i n g w i t h a c a l i b r a t i o n w h eel. The w heel i s 2 m i n c ir c u m f e r e n c e and r e c o r d s th e number o f w heel r e v o l u t i o n s . Cows w ere th e n p u t back 35 i n t h e c h u te and p e d o m e te r s removed, read, and compared t o t h e a c t u a l d i s t a n c e r e c o r d e d by th e c a l i b r a t i o n w heel. Ambient a i r t e m p e r a t u r e and r e l a t i v e h u m i d ity w ere r e c o r d e d each y e a r u s i n g a h y g r o t h e r m o g r a p h p l a c e d on a n o r t h f a c i n g s l o p e i n t h e a p p ro x im a te c e n te r of th e p a s tu re . T e m p e r a t u r e an d h u m i d i t y w e r e a v e ra g e d f o r eac h p e r i o d and each day u s i n g t e m p e r a t u r e s r e c o r d e d a t tw o h o u r i n t e r v a l s . D ata A n a ly s is Data w e re a n a ly z e d f o r s i g n i f i c a n t d i f f e r e n c e s (p<.05) i n g r a z i n g b e h a v i o r and t o . ta l in t a k e , among age g r o u p s and t r e a t m e n t groups. w as a c c o m p lish e d u s in g This S t a t i s t i c a l A n a l y s i s S y stem ’s (SAS) General L i n e a r Model (GLM) p r o c e d u r e , and th e a n a l y s i s o f v a r i a n c e p r o c e d u r e s a v a i l a b l e i n MSUSTAT (Lund 1983). The SAS GLM p r o c e d u r e was a l s o used t o t e s t a l i n e a r r e g r e s s i o n model u s i n g tim e s p e n t g r a z i n g each day t o e x p l a i n v a r i a t i o n i n t o t a l o r g a n ic m a t t e r i n t a k e : I n t a k e = f ( D a ily G r a z in g Time) A m o r e e x t e n s i v e m o d e l w as t e s t e d u s i n g t h e SAS S t e p w i s e p r e c e d u r e . The model was u sed t o e x p l a i n ■ v a r i a t i o n i n d a i l y g r a z i n g ti m e ; i t s com ponents a r e fo u n d i n T a b le 7. And f i n a l l y , A u to re g re ssiv e t h e SAS A rim a p r o c e d u r e w as u s e d t o d e v e l o p a n M oving A v e ra g e (ARMA) m o d e l t h a t d e sc rib e d d a ily g r a z i n g t i m e on any g iv e n day based on p a s t g r a z i n g t i m e s a lo n g w i t h p r e s e n t and p a s t te m p e ra tu re s. T h is m o d e lin g p r e v io u s ly been a p p lie d to a n im a ls , p r o c e d u r e had n o t b u t w as u s e d i n t h i s s t u d y t o d e t e r m i n e t h e e f f e c t o f p r e v i o u s e x p e r i e n c e s on p r e s e n t g r a z i n g 36 T ab le 7• R e g r e s s io n model f o r e x p l a i n i n g v a r i a t i o n i n t h e d a i l y tim e sp en t g ra z in g d u rin g w in te r. T— ; ---------------------------------------— Item D e sc rip tio n b0-7 C o n s ta n ts I T otal d a ily in ta k e A Age IW I n i t i a l W eight a t b e g in n in g o f s tu d y T Average d a i l y te m p e r a t u r e H Average d a i l y h u m id ity DG Day o f g e s t a t i o n DT D is ta n c e t r a v e l e d p e r day GT Time s p e n t g r a z i n g each day R e g r e s s io n M odel: GT=b0+ b 1( I ) + b 2 (A)+b3 (IW)+b4 (T )+b5 (H)+b6 (DG)+b7 (DT) b eh av io r. S uch a r e l a t i o n s h i p h a s b e e n s u g g e s t e d a s t h e b a s i s f o r t h e r m a l a c c l i m a t i o n i n b e e f c a t t l e ( S e n f t and R it te n h o u s e 1985). The f i r s t s t e p i n t h i s p r o c e d u r e was an a u t o r e g r e s s i o n o f t e m p e r a t u r e and d a i l y g r a z i n g ti m e . The e f f e c t s o f a u t o r e g r e s s i o n w e re t h e n removed w i t h a " p r e w h ite n i n g " p r o c e s s which t r a n s f o r m s an a u t o c o r r e l a t e d i n p u t s e r i e s t o an u n c o r r e l a t e d , w h i t e n o i s e ( p u r e ly random) s e r i e s (Box and J e n k i n s 1 976, C h a t f i e l d 1 980). The tw o v a r i a b l e s w e r e t h e n c r o s s - c o r r e l a t e d to e s t a b l i s h a tim e s e r i e s r e l a t i o n s h i p . t h i s model a r e fo u n d i n T able 8. C o m p o n e n ts o f 37 T ab le 8. Time s e r i e s a n a l y s i s o f d a i l y g r a z i n g tim e b ased g r a z i n g tim e s and p r e s e n t and p a s t te m p e r a t u r e s . Item past D e sc rip tio n GTd G r a z in g tim e on day d GT^_p G ra z in g tim e on p t h day p r e v io u s t o d Td Average t e m p e r a t u r e on day d Tp - q Average t e m p e r a t u r e on q t h day p r e v io u s t o d a ,b on C o n s ta n ts A u t o r e g r e s s i v e Moving Average .(ARMA) Model: GTd=E1 GTd e ^ a 2 GTde2+ . . . + a ^ GTd_p+ T ^ b 1Tde1+b2Tde2+ . . .+bq Td_q These m o d e lin g p r o c e d u r e s w e re u sed t o p ro v id e i n f o r m a t i o n and c l a r i f i c a t i o n o f th e p a r a m e t e r s m e a su re d i n d ev elo p ed a r e not s t a t is tic a lly c o m p le te , in c o m p l e t e a n a l y s i s o f th e o v e r a l l d a t a s e t . th is s tu d y . The m odels and t h e r e f o r e show an As t h i s s tu d y c o n tin u e s t h e m o d e l s a n d t h e i r v a r i a b l e s s h o u l d be r e - f o r m e d , and d i f f e r e n t p ro c e d u re s u sed , i n an a t te m p t to i n c r e a s e th e s i g n i f i c a n c e of th e m o d e ls and t h e outcome. 38 RESULTS 1Q84 T r i a l Body Weight and C o n d it io n S co re The t h r e e y e a r o ld cows e n t e r e d t h e s tu d y a t s i g n i f i c a n t l y lo w e r w e i g h t t h a n t h e 7 y e a r o l d cow s. The o t h e r a g e g r o u p s ( 4 , 5, and 6) f e l l i n betw ee n t h e 3 and 7 y e a r o ld s . T h is w e ig h t p a t t e r n p e r s i s t e d t h r o u g h o u t t h e t r i a l w i t h w e i g h t c h a n g e s r a n g i n g f r o m - 1 0 . 8 t o +1.2 lb . Mean w e i g h t changes among t h e age g ro u p s w ere n o t s i g n i f i c a n t l y d iffe re n t. C o n d itio n s c o r e s fo llo w e d th e w e ig h t p a tte rn ^ and, a g a in , 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 c o n d i t i o n s c o r e c h a n g e am ong a g e g r o u p s w e re seen. A ll g ro u p s l o s t c o n d itio n over th e w in te r. The f i n d i n g s a r e s u m m a rized i n A ppendix T ab le 19. G ra z in g B e h a v io r The o v e r a l l a v e r a g e o f d a i l y g r a z i n g t i m e w a s 8.8 h o u r s p e r day o r 36.8 p e r c e n t o f t h e 24 h o u r p e r i o d b e g i n n i n g a n d e n d i n g a t 7AM. T a b l e 9 g i v e s a c o m p a r i s o n o f g r a z i n g t i m e s f o r 3 age g r o u p s . The t h r e e y e a r o l d cow s s p e n t m o re t i m e g r a z i n g e a c h day t h a n t h e 5 o r 7 y e a r o ld cows. The p ed o m e te r r e c o r d i n g s showed t h a t t h e 3 y e a r o l d cows t r a v e l e d s i g n i f i c a n t l y f a r t h e r t h a n t h e 5 and 7 y e a r o ld cows. Age group means o f d i s t a n c e t r a v e l e d p e r day can be s e e n i n T able 9. For a l l th e g ro u p s, t h e a f t e r n o o n p e r i o d ( 1PM-7PM) w as m o s t i m p o r t a n t f o r g r a z i n g w i t h an a v e r a g e o f 4.3 (72.0%) o f t h e 6 h o u r s sp en t g razin g . A s u b s t a n t i a l p o r t i o n o f t h e m orning p e r i o d (7AM-1PM) 39 T ab le 9. D a ily g r a z i n g tim e s and d i s t a n c e s t r a v e l e d f o r 3 g r o u p s , 1984. cow age G ra z in g Time ( % ) D is ta n c e T r a v e le d (km) Age 7AM-IPM I PM-7PM 7PM-AM 3 5 4 .3a 7 5. I a 1 2 .7a 3 8 .7a 5 .6 a . 5 4 8 . Ob 6 9 .2b 1 2 .3a 3 5 . 4b 4 .8 b 7 50. I b 7 2 . Oab 11.3 a 36 *2b 4 .7 b D aily a >'-’Means i n t h e same column w ith d i f f e r e n t s u p e r s c r i p t s a r e d i f f e r e n t (p < .0 5 ). was s p e n t g r a z i n g ; t y p i c a l l y , o f th e p e r io d . g r a z i n g o c c u p ie d j u s t o v e r 3 h o u rs (51%) N ig h ttim e g r a z i n g (7PM-7AM) was f a i r l y c o n s i s t e n t , but r e p r e s e n t e d a r e l a t i v e l y s m a l l p o r t i o n o f each d a y 's g r a z in g . G razing d u r i n g t h i s p e r i o d was a d i s t i n c t l y i n d i v i d u a l a c t i v i t y w h i l e g r a z i n g d u r in g t h e day u s u a l l y in v o lv e d groups. F ig u r e 3 show s a t y p i c a l w i n t e r t e m p e r a t u r e r e s u l t i n g g ra z in g b eh av io r. s e c t i o n of th e g rap h , p a t t e r n and th e The t e m p e r a t u r e s r e c o r d e d on t h e t o p s h o u ld be: w i t h i n th e cow's t h e r m o n e u t r a l zone; th e re fo re , th e g ra z in g p a t t e r n situ a tio n s. The m i d - s e c t i o n show s t h e g r a z i n g t i m e p e r c e n t a g e s f o r e a c h d a y 's 3 p e r i o d s . tim e m eans. sh o u ld be t y p i c a l of u n s tre s s fu l The b o t t o m p o r t i o n s h o w s t h e d a i l y g razin g D u rin g t h e s e s i x d a y s, d a i l y g r a z i n g tim e a v e ra g e d 4 1 p e r c e n t o f t h e day. F i g u r e 4 sh o w s a much c o l d e r 6 d a y p e r i o d i n J a n u a r y . The a v e ra g e d a i l y g r a z i n g ti m e was lo w e r th a n d u r in g t h e w arm er p e r io d , 36 p e rc e n t. On J a n u a r y I 6 t h e t e m p e r a t u r e d r o p p e d f r o m - 1 0 ° t o - 2 4 ° C . 40 TEMP TIME SPENT 40 GRAZING 24 HRS 30 SPENT 20 GRAZING in F ig u re 3. A n o r m a l w i n t e r t e m p e r a t u r e p a t t e r n an d t h e a s s o c i a t e d g razin g a c t i v i t y . a , b, c, d, e, f , g M eans r e pr e s e n t e d by s i m i l a r b a r s c o n t a i n i n g d i f f e r e n t l e t t e r s are d if f e r e n t (p < .0 5 ). 41 TEMP 0C %0F TIME SPENT GRAZING %0F 2 4 HRS SPEN T GRAZING J a n . 14 F ig u re 4. J a n . 15 G ra z in g a c t i v i t y te m p e ra tu re s. i J a n . 16 J a n . 17 J a n . 18 J a n . 19 a s s o c i a t e d w i t h th e 1984 t r i a l ' s c o l d e s t g> h, i , j Me a n g r e p r e s e n t e d by s i m i l a r b a r s c o n t a i n i n g d i f ­ f e r e n t l e t t e r s a r e d i f f e r e n t (p<.05). 42 The f o l l o w i n g d ay t h e h ig h t e m p e r a t u r e w as - 2 2 ° and g r a z i n g tim e d r o p p e d t o 30.0 p e r c e n t o f t h e d a y , p e rio d s. H o w e v er, a ll th re e on t h e 18 t h and 1 9 t h , g r a z i n g t i m e r o s e t o t h e l e v e l i t h a d b e e n b e f o r e t h e 1 7 th . p e rio d in c re a s e d w ith a d ec re a se in slig h tly G r a z i n g t i m e i n t h e IPM t o 7PM d u rin g t h i s co ld p e rio d w h ile g ra z in g d e c r e a s e d i n t h e m o rn in g and a t n i g h t . T h i r t y - s e v e n p e r c e n t o f th e v a r i a t i o n i n d a i l y g r a z i n g ti m e was e x p l a i n e d by th e s e v e n f a c t o r s i n c l u d e d i n t h e r e g r e s s i o n model. o rd er of e n tra n c e in to sta tistic s are The t h e s t e p w i s e r e g r e s s i o n and t h e r e s u l t i n g show n i n T ab le 10. O nly tw o o f th e in d e p en d en t v a r i a b l e s had a s i g n i f i c a n t e f f e c t on d a i l y g r a z i n g t i m e : h u m id ity and ( day o f g e s t a t i o n . I n c r e a s i n g h u m i d ity h ad a n e g a t i v e im p a c t on d a i l y g r a z i n g t i m e w h i l e g e s t a t i o n had a s l i g h t l y p o s i t i v e e f f e c t . T a b l e 11 s h o w s t h e r e s u l t o f t h e t i m e s e r i e s a n a l y s i s o f d a i l y g r a z i n g t i m e and t e m p e r a t u r e . G razin g t i m e on any g iv e n day was n o t s i g n i f i c a n t l y e x p l a in e d by t h e a u t o r e g r e s s i v e m o v in g a v e r a g e m o d e l u sin g past g razin g tim e s and p r e s e n t and p a s t te m p e ra tu re s as co v ariab les. In ta k e As s h o w n i n T ab le 12, age d if f e r e n c e s i n t o t a l d a ily in ta k e . o f body w e ig h t p e r day. gro u p s e x h ib ite d sig n ific a n t The o v e r a l l mean i n t a k e was 0.87% The l i n e a r r e g r e s s i o n model u s i n g i n t a k e and d a i l y g r a z i n g tim e showed no s i g n i f i c a n t r e l a t i o n s h i p . model a p p e a r i n T a b le 13» no D e t a i l s of th e 43 T ab le 10. S te p w is e l i n e a r r e g r e s s i o n of 1 d a i l y g r a z i n g tim e , 1984. E n tr a n c e in to Model Source S ig n ific a n t P aram e ter V alues Mean Alpha=. I d f S quare f Prob>f Model r^ a t each step R e g r e s s io n I Humidity^ 2 Day o f G e s t a t i o n 3 -0 .2 4 I 86.9 2 .8 9 0 .1 0 0.26 0 .1 4 I 119 3.96 0 .0 5 0 .3 2 I n i t i a l Weight I 27.6 0.92 0 .3 5 0.34 4 Tem perature^ I 31.6 1 .05 0.31 0.37 5 D a il y I n ta k e I 2.71 0 .0 9 0 .7 7 0.37 6 Age I 1 .10 0 .0 4 0 .8 5 0.37 7 D a il y T ra v e l I 0.11 0.00 0 .9 5 0 .3 7 27 30.1 R e s id u a l I D a ily a v e r a g e T ab le Lag (Days) 11. Time s e r i e s a n a l y s i s o f t o t a l d a i l y a v e ra g e d a i l y t e m p e r a t u r e , 1984. g razin g tim e and C ross C o r r e l a t i o n s -------——— —---------------------------------------------------------------------------- :— C o e ffic ie n t -I - .8 -.6 - .4 - .2 0 .2 .4 .6 .8 I Marks two s t a n d a r d e r r o r s o r a l p h a = . 05 44 T ab le 12. T o ta l d a i l y i n t a k e (OMB %BW) e s t i m a t e d from g ra b sam ples and a d j u s t e d f o r Cr2 Og r e c o v e r y ( T u r n e r 1 9 8 5 ), 1984. Age T otal In ta k e (0MB %BW) 3 0 . 9 1 + 0 . 05a 4 0 .8 9 ± P .0 4 a 5 0 .8 4 + 0 .0 4 a 6 0 .9 0 ± 0 .0 3 a 7 0 . 8 3 + 0 .0 3 a a Means i n th e same column w ith th e same s u p e r s c r i p t a r e n o t d i f f e r e n t (PC.0 5 ) . T able 13. L in ear r e g r e s s io n o f d a ily t o t a l in ta k e as r e l a t e d to d a ily g r a z i n g tim e , 1984. S ource df Mean S quare I 0 .0 2 48 0 .1 2 f prob> f Model r^ 0 .7 2 0 .0 0 R e g r e s s io n D a ily g r a z i n g tim e R e s id u a l 0 .1 3 1085 T r i a l Body Weight and C o n d it io n S c o re As i n 1 984, 3 year o ld cow s e n t e r e d and l e f t th e t r i a l s i g n i f i c a n t l y lo w e r w e i g h t s th a n some o f th e o l d e r age g r o u p s ; at th e re was no s i g n i f i c a n t d i f f e r e n c e i n t h e w e i g h t change o r c o n d i t i o n s c o r e change e x p e r i e n c e d by th e age groups. Appendix T a b le 20. These r e s u l t s a r e su m m arized i n 45 T h e r e w e r e no w e i g h t o r c o n d i t i o n s c o r e d i f f e r e n c e s am ong t h e t r e a t m e n t g ro u p s a t th e b e g in n in g o f t h e t r i a l . group lo s t w e ig h t (I kg) d u rin g s u p p le m e n te d gro u p s g a in e d w e i g h t . s c o r e d u r in g t h e t r i a l , , c o n t r o l group. th e However, th e c o n t r o l tria l w h ile a ll of th e W h i le a l l g r o u p s l o s t c o n d i t i o n t h e f a t group l o s t s i g n i f i c a n t l y l e s s t h a n t h e The f a t group f i n i s h e d t h e t r i a l w i t h t h e h i g h e s t mean c o n d itio n s c o re w ith o u t h av in g g ain ed th e m ost w e ig h t. c o n d itio n s sc o re s fo r W e i g h t s and t r e a t m e n t g r o u p s a r e f o u n d i n A p p e n d ix T a b l e 21. G ra z in g B e h a v io r Appendix T a b le 22 shows t i m e s p e n t g r a z i n g by age g ro u p s i n 1985. T h r e e y e a r o l d cow s g r a z e d s i g n i f i c a n t l y l o n g e r e a c h day t h a n t h e o l d e r age g ro u p s. D istan ce s tr a v e le d , a l s o i n T ab le 22, d id n o t show th e same age p a t t e r n a s i n 1984. Of t h e t r e a t m e n t g ro u p s, cows r e c e i v i n g a n im a l f a t su p p lem en t s p e n t l e s s tim e g r a z in g th a n d id th e r e c e i v i n g b lo o d m eal s u p p le m e n t. c o n t r o l g ro u p and th e group The b lo o d meal group and t h e c o n t r o l group d id n o t d i f f e r s i g n i f i c a n t l y i n g r a z i n g tim e , g r o u p t r a v e l e d s i g n i f i c a n t l y l e s s e a c h day . b u t t h e blood meal The c o r n g l u t e n m e a l g r o u p t r a v e l e d m o re t h a n t h e c o n t r o l , s o y b e a n m e a l , a n d b l o o d m e a l g ro u p s. Mean g r a z i n g t i m e s and d i s t a n c e s t r a v e l e d f o r each t r e a t m e n t group can be s e e n i n T ab le 14. F i g u r e s 5 and 6 show t e m p e r a t u r e and g r a z i n g p a t t e r n s f o r tw o p e rio d s in Jan u ary . D u rin g th e t e m p e r a t u r e r o s e from - 1 8 ° t o O0C. p e rio d show n i n F ig u re 5 th e The a f t e r n o o n p e r i o d l o s t r e l a t i v e 46 T ab le 14. D a ily g r a z i n g tim e s and t r a v e l o f each 1985. tre a tm e n t group, G r a z in g Time (%) D is ta n c e T ra v e le d (km) TRT 7AM-IPM CON 5 8 . 6a 6 7 .8a 1 5 . Oa 3 9 .3a 4 .8 b SBM 5 6 . I ab 6 5 . 7 ab 1 4 .5 a 3 7 . 6 bc 4 .5 b FAT 5 4 . 5b 6 3 . 9b . 1 3 . 9a 3 6 .6 ° 5 . Oab CGM 5 4 . 2b 6 4 . 8 ab 1 3 .9 a 3 6 . 9 bc 5 .4 a BM 5 6 .3 ab 6 5 . Oab 1 6 .2 a 3 8 .2 ab 3 .9 ° I PM-7PM a , b , c l e a n s i n th e same d if f e r e n t (p < .0 5 ). column 7PM-7AM w ith D a ily d iffe re n t su p e rsc rip ts are im p o rta n c e i n th e d a ily g ra z in g sc h e d u le as th e te m p e ra tu re ro s e . T h i s w a s a c c o m p a n i e d by a n i n c r e a s e i n m o r n i n g g r a z i n g a c t i v i t y . T o t a l g r a z i n g t i m e d e c l i n e d on J a n u a r y 14, t h e day t h e t e m p e r a t u r e peaked a t 0°. The a v e r a g e o f d a i l y g r a z i n g t i m e s f o r t h e s e s i x d a y s was 38.6 p e r c e n t , v e ry c l o s e t o t h e tw o 1984 p e r i o d s shown i n F ig u r e s 3 a n d 4. F i g u r e 6 show s a t e m p e r a t u r e d e c l i n e fro m - 2 ° to -2 4 °C . On J a n u a r y 28 and 29 t h e te m p e r a t u r e d e c l i n e d s t e a d i l y , and d a i l y g r a z i n g t i m e f e l l t o 28.4 p e r c e n t . a fte rn o o n in c re a se d As t e m p e r a t u r e d e c l i n e d , g r a z i n g i n t h e sig n ific a n tly and n ig h t g ra z in g decreased s i g n i f i c a n t l y . B e fo re and a f t e r t h e 2 9 th , d a i l y g r a z i n g t i m e was c l o s e t o 40 p e r c e n t . The 6 day av era g e d a i l y g r a z i n g ti m e was 38.2 p e r c e n t , v e r y c l o s e t o t h e p e r i o d s shown i n F i g u r e s 3» 4 and 5. 47 -2 TEMP -6 °C - I O -14 -18 7 am %0F TIME SPENT GRAZING % OF 2 4 HRS SPENT GRAZING 4U 30 20 m J a n . 12 J a n . 13 J a n . 14 J a n . 15 J a n . 16 J a n . 17 1985 F i g u r e 5. A warming t r e n d and th e a s s o c i a t e d g r a z i n g a c t i v i t y . a ,b ,c ,d ,e ,f,g ,h ,i Means rep resen ted by s i m i l a r bars d i f f e r e n t l e t t e r s a re d i f f e r e n t (p < .0 5 ). c o n ta in in g 48 TIME SPENT GRAZING 2 4 HRS 40 SPENT 30 GRAZING 20 Jan. 2 6 Jan. 2 7 Jan. 2 8 Jan. 2 9 Jan. 3 0 Ja n . 31 1985 F ig u re 6 . A t e m p e r a t u r e d e c l i n e and t h e a s s o c i a t e d g r a z i n g a c t i v i t y . a , b , c , d , e , f , g , h , i Means r e p r e s e n t e d by s i m i l a r bars d if f e r e n t l e t t e r s a re d if f e r e n t (p < .0 5 ). c o n ta in in g 49 F o r t h e 1985 t r i a l , t r e a t m e n t g ro u p w as added t o t h e l i s t o f v a r i a b l e s i n t h e l i n e a r r e g r e s s i o n m o d e l. T h i s m o d e l e x p l a i n e d ' 51 p e r c e n t o f t h e v a r i a t i o n i n d a i l y g r a z i n g tim e . Once a g a i n , h u m id ity was t h e f i r s t v a r i a b l e t o e n t e r t h e m odel, b u t w i t h a p o s i t i v e i n s t e a d of n e g a tiv e e f f e c t. T e m p e ra tu re e n t e r e d seco n d w i t h a n e g a t i v e e f f e c t on d a i l y g r a z i n g tim e . S t a t i s t i c a l d e t a i l s of t h e model, can be found i n T a b l e 15. T able 15. S tep w ise l i n e a r r e g r e s s i o n o f d a i l y g r a z i n g tim e , 1985. S ig n ific a n t P a ra m e te r V alues A lpha=. I df E n tr a n c e in to Model S ource Mean S quare f ]Prob>f Model r ^ a t each ste p R e g r e s s io n I Humidity^ 2 Tem perature^ 3 1 .4 I 302 17.5 0 .0 0 0 .3 7 -0 .5 I 6 7 .6 3 .9 2 0 .0 5 0 .4 2 D a ily I n ta k e I 3 7 .7 2 .1 9 0 .1 5 0 .4 5 4 D a ily T r a v e l I 4 4 .8 2 .6 0 0 .1 2 0 .4 7 5 Age I 35.6 2 .0 7 0.16 0 .4 8 I n i t i a l Weight I 2 7 .5 1.59 0 .2 2 0.50 7 Day o f G e s t a t i o n I 12.3 ' 0 .7 2 0 .4 0 0.51 8 T re a tm e n t Group I 5 .7 5 0 .5 7 0.51 32 17.2 ■6 R e s id u a l *1 0 .3 3 D a il y a v e ra g e The t i m e s e r i e s a n a l y s i s o f 1985 shown i n T ab le 16. g r a z i n g and t e m p e r a t u r e d a t a i s G ra z in g tim e and t e m p e r a t u r e on t h e f i f t h p r e v io u s day h ad a s i g n i f i c a n t ( a l p h a = . 0 5) p o s i t i v e e f f e c t on c u r r e n t g r a z i n g 50 T able Lag (Days) 16. Time s e r i e s a n a l y s i s o f t o t a l d a i l y a v e ra g e d a i l y t e m p e r a t u r e , 1985. g razin g tim e and C ro ssc o rre la tio n s ------------------------------------------- — —— ------------------------- --------------------C o e ffic ie n t -I - .8 -.6 - .4 - .2 0 .2 .4 .6 .8 I - 0.26 Marks two s t a n d a r d e r r o r s o r a l p h a = .05. tim es. However, none o f t h e days b e tw e e n t h e f i f t h p r e v io u s day and t h e c u r r e n t day h a d a n e f f e c t , a n d n o n e o f t h e d a y s p r e v i o u s t o t h e fifth day had an e f f e c t . In ta k e Grab s a m p le s t a k e n once d u r in g t h e t o t a l f e c a l c o l l e c t i o n p e r io d a l l o w e d f o r t h e r e c o v e r y o f 6 0 .3 p e r c e n t o f t h e s u p p le m e n t. -fed w ith th e Mean r e c o v e r y f o r each t r e a t m e n t group r a n g e d from 44 to 7 1 p e r c e n t , a lth o u g h t h e s e m eans w ere 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 ( A p p e n d ix T ab le 23). Chromic o x id e r e c o v e r y from s a m p le s ta k e n from each day’s t o t a l c o l l e c t i o n s h o w e d a d i s t i n c t p a t t e r n ( A p p e n d ix F i g u r e 7 ). H o w e v er, LSD m u l t i p l e c o m p a r i s o n o f r e c o v e r y m e a n s s h o w e d f e w s i g n i f i c a n t d iffe re n c e s - 51 The CrgO^ r e c o v e r y f o r 1985 w as l o w e r t h a n p r e v i o u s s t u d i e s a t Red B l u f f R e s e a r c h R anch ( T u r n e r 1 9 8 5 ). The r e c o v e r y le d to an o v e r e s t i m a t i o n o f f e c a l o u tp u t com pared t o t o t a l f e c a l c o l l e c t i o n . These co m p ariso n s a r e l i s t e d i n A ppendix T ab le 24. A ppendix T ab le 25 shows t h e t o t a l o r g a n i c m a t t e r i n t a k e f o r each t r e a t m e n t g r o u p a n d e a c h c o l l e c t i o n day . These f i g u r e s have been a d j u s t e d f o r CrgOg r e c o v e r y and g r a b s a m p le o v e r e s t i m a t i o n . I n T ab le 17» th e amount o f s u p p le m e n t f e d each day h a s been rem oved i n o r d e r t o a l l o w t h e c o m p a r i s o n o f f o r a g e i n t a k e am ong a l l g r o u p s . The b lo o d m eal and f a t g ro u p s e x h i b i t e d th e h i g h e s t d a i ly f o r a g e in t a k e . The c o n t r o l and c o r n g l u t e n m e a l g r o u p s c o n s u m e d t h e l e a s t f o r a g e e a c h day. T a b le 17. D a ily f o r a g e o r g a n i c m a t t e r i n t a k e (%BW), 1985. Day T rea t­ ment J a n 17 J a n 23 Feb 6 Feb 20 A ll Days C CON 0 .9 4 + 0 .1 0 I .01+0.07 0 .9 2 + 0 .0 8 1 .04+ 0.08 0 .9 8 ± 0 .0 4 SBM 1 .5 1 + 0 .0 9 I .0 7 ± 0 .0 5 1 .0 3 ± 0 .0 7 1.26±P .10 I .2 3 ± 0 .0 5 ab FAT I .13+ 0.14 0 .9 1 + 0 .1 4 1 .5 6 + 0 .1 7 2 .1 3 + 0 .4 3 1 . 4 4 + 0 .13a COM I .01+ 0.13 1 .0 2+ 0.79 I .0 0 ± 0 .1 0 1.08+ 0.13 1 .0 8 ± 0 .0 5 bc BM I .89+ 0.22 I .08+0.11 I .3 5 + 0 .1 3 1.94+ 0.33 1 .5 6 ± 0 .1 1 a Mean of a ll TRT’ s 1 .3 0+ 0.07 de f I .02+ 0.04 ef I • 17±0.06 d 1 .49±P.13 1.25 a »b>cMeans i n th e same column w ith d i f f e r e n t s u p e r s c r i p t s a r e d if f e r e n t (p < .0 5 ), d»e >^Means i n t h e same row w ith d i f f e r e n t s u p e r s c r i p t s a r e d i f f e r e n t ( p < .0 5 ) . 52 The l i n e a r r e g r e s s i o n model u s i n g i n t a k e and d a i l y g r a z i n g t i m e a s v a r i a b l e s s h o w e d no s i g n i f i c a n t r e l a t i o n s h i p . T a b l e 18 l i s t s d e t a i l s o f t h e model. T ab le 18. L in ear r e g r e s s io n o f d a ily t o t a l in ta k e as r e l a t e d to d a ily g r a z i n g ti m e , 1985. S ource df Mean S quare f prob> f Model r 2 I I .29 2 .1 3 0 .1 5 0 .0 5 40 0.60 R e g r e s s io n D a ily G ra z in g Time R e s id u a l •X 53 DISCUSSION F o rag e U t i l i z a t i o n and F o r a g in g P a t t e r n s The, s t o c k i n g r a t e d u r in g both 1984 and 1985 was w e l l below t h a t s u g g e s t e d by P ayne ( I 9 7 3 ). H o w e v e r, o b s e r v a t i o n s a n d u t i l i z a t i o n m e a s u r e m e n t s d i d n o t show e v e n g r a z i n g d i s t r i b u t i o n t h r o u g h o u t t h e p a stu re , so a n o v e r a l l sto c k in g ra te fig u re d isc u ssin g fo rag e u t i l i z a t i o n p a tte rn s . is m i s l e a d i n g w hen The p a s t u r e c o n t a i n s l a r g e a r e a s w i t h a c o n s t a n t , s l i g h t s l o p e ; t h e s e a r e a s r e c e i v e d h e a v i e s t u se w ith a p p r o x im a te ly 75 p e r c e n t u t i l i z a t i o n o f th e key f o r a g e s p e c i e s . A re a s w i t h c o n s t a n t , m o d e ra te s lo p e showed l i g h t g r a z i n g p r e s s u r e , a p p r o x i m a t e l y 30 p e r c e n t u t i l i z a t i o n . The a r e a s c o n t a i n i n g s t e e p s l o p e s w e re l e a s t u t i l i z e d show ing o n l y 15 p e r c e n t u t i l i z a t i o n of key sp e c ie s. T h i s u t i l i z a t i o n p a t t e r n w a s m o re p r o n o u n c e d i n I 984 t h a n 1 9 8 5 ; p e r h a p s t h e h i g h e r s t o c k i n g r a t e i n 1985, a n d d i f f e r e n c e s i n f o r a g e c h a r a c t e r i s t i c s , f o r c e d more ev e n f o r a g e u t i l i z a t i o n . T h r e e a r e a s i n t h e p a s t u r e w e r e u s e d by t h e co w s f o r s h e l t e r : ro ck o u tc ro p s, draw s s h a llo w s i t e s w i t h an o v e r s t o r y o f c o n i f e r s , c o n t a i n i n g s h r u b s and b a sin w ild ry e . and deep J u d g in g from fecal d ep o sits, t h e cows s p e n t a c o n s i d e r a b l e amount o f ti m e i n t h e s e a r e a s ; how ever, t h e a r e a s d i d n o t c o n t a i n l a r g e q u a n t i t i e s o f key f o r a g e s p e c i e s ( e x c e p t b a s i n w i l d r y e ) and so w e re p r o b a b ly n o t i m p o r t a n t fo ra g in g a re a s. The b e h a v i o r d a t a r e a d i l y show t h a t d a y l i g h t h o u r s w e r e p r e f e r r e d o v e r n i g h t t i m e f o r g r a z i n g , and t h e h o u r s betw ee n IPM and 7PM a r e m ost preferred , e s p e c i a l l y a t c o l d e r te m p e r a tu r e s * grazin g a c t i v i t y As t e m p e r a t u r e s warm, spread out th ro u g h th e day; m o rn in g and a f te r n o o n 54 became e q u a l l y i m p o r t a n t and n i g h t a c t i v i t y 5). al. These f i n d i n g s a r e (19 7 0 ), i n c r e a s e d ( F i g u r e s 3 and c o n s i s t e n t w i t h f i n d i n g s r e p o r t e d by Gary e t A rn o ld and D u d z in s k i (1 9 7 8 ), and S t r i c k l i n a n d K a u t z - S canavy ( 1 9 8 4 ) . C o n tr a ry t o t h e f i n d i n g s o f Hancock (1953) and S to b b s (1970), t h e v a r i a t i o n among cows f o r d a i l y g r a z i n g t i m e i n t h i s s tu d y was low. In 1984 t h e c o e f f i c i e n t o f v a r i a t i o n f o r d a i l y g r a z i n g t i m e r a n g e d from 13.7 t o 20.6 ( a v g . 1 8 .6 ) ; i n 1 985 t h e c o e f f i c i e n t s r a n g e d f r o m 12.8 t o 19.7 (avg. 15.6). W in te r may im pose s t r i c t e r l i m i t s on b e h a v io r th a n o th e r seasons, r e s u l t i n g i n l e s s v a r ia tio n . E n v iro n m e n ta l S t r e s s C o m p a riso n s of th e b eh av io r and w e a th e r i m p o r t a n c e o f w e a t h e r a s a n e f f e c t on d ata in d ic a te g ra z in g b eh av io r. th e F ig u re s 3 t h r o u g h 6 show r e a c t i o n s a n d a d j u s t m e n t s made t o d a i l y f o r a g i n g p a tte r n s as te m p e ra tu re f lu c tu a te s . H o w e v e r, th e se d a ta f a i l to e x p l a i n a s i g n i f i c a n t am ount o f th e v a r i a t i o n s e e n i n tim e s p e n t g razin g . I n tw o s t u d i e s ( M a le c h e k a n d S m ith 1 976, Adams 1 9 8 4 a ) t e m p e r a t u r e w as r e p o r t e d a s b e in g c l o s e l y c o r r e l a t e d to t i m e s p e n t g r a z i n g . Adams (1984a) d e v e lo p e d a p r e d i c t i v e t a b l e g ra z in g tim e te m p e ra tu re s as a ir te m p e ra tu re d u rin g th a t stu d y decreased ra n g e from show ing d e c r e a s e d (see 8 to T ab le -3 4 °C . 3). The G ra z in g b e h a v i o r was r e c o r d e d w i t h v i b r a c o r d e r s from December 9 t o J a n u a r y 14. The a u t h o r c a l c u l a t e d t h a t , b e g in n in g a t 0°C, f o r e v e r y 10° r e d u c t i o n i n t e m p e r a t u r e , a one h o u r r e d u c t i o n i n d a i l y g r a z i n g t i m e c o u l d be e x p e c te d . I n d ic a tio n s of t h i s r e l a t i o n s h i p w ere a l s o s e e n i n t h i s 55 s tu d y . The d i f f e r e n c e i n a v e ra g e t e m p e r a t u r e betw ee n t h e two p e r i o d s s h o w n i n F i g u r e s 3 a n d 4 w a s 16.6°C . T h i s w a s a c c o m p a n i e d by a 1.2 h o u r r e d u c t i o n i n ' a v e r a g e d a i l y g r a z i n g ti m e . d id n o t e x p la in a s u b s t a n t i a l p o rtio n However, te m p eratu re of th e v a r i a t i o n in d a ily g r a z i n g t i m e d u r in g t h i s s tu d y . A lso, a s s e e n i n F i g u r e 6, r e d u c t i o n s i n g r a z i n g t i m e w i t h c o l d e r t e m p e r a t u r e s may be o n ly te m p o ra ry . Adams and F i g u r e s 3 and 4 s u g g e s t a s t r a i g h t f o r w a r d r e l a t i o n s h i p b e t w e e n t e m p e r a t u r e a n d g r a z i n g t i m e , t h e r e g r e s s i o n m o d e l s a n d F i g u r e 6 do n o t. T e m p e ra tu re and h u m i d ity by t h e m s e lv e s d e s c r i b e a p o r t i o n o f t h e th e r m a l e n v iro n m e n t, and w ere somewhat r e l a t e d t o d a i l y g r a z i n g ti m e a s s e e n by t h e i r e a r l y e n t r a n c e i n t h e r e g r e s s i o n m o d e l s . may n e e d t o be c o m b in e d w i t h w i n d s p e e d , But t h e y b a ro m e tric p re ssu re , and r a d i a n t h e a t l o s s o r g a i n t o d e v e lo p a t h e r m a l in d e x t h a t would more c o m p le te ly d e s c r ib e th e th e rm a l e n v iro n m e n t as i t is re la te d to g r a z i n g b e h a v io r . The r e l a t i v e c o n sta n c y of d a ily g ra z in g in d e p e n d e n c e o f g r a z i n g tim e and t e m p e r a t u r e s e e n i n in d i c a t e a la c k o f s e v e re e n v iro n m e n ta l s t r e s s . tim e s and t h e th is s tu d y may When d a i l y g r a z i n g d e c r e a s e d i t d id so i n c o n j u n c t i o n w i t h s e v e r e t e m p e r a t u r e changes (up or d o w n ), but re m a in e d lo w fo r one i n d i c a t i o n of a c u te th erm al s t r e s s ; day o n ly . T h is may be a n o t h e r s t u d i e s (W e b ste r and Young 1 9 7 0 , S l e e 1971) h a v e s h o w n t h a t a c u t e s t r e s s i s a c c o m p a i'n e d by a d e c lin e in in tak e. a p p ro x im a te ly However, a t no tim e d id d a i l y g r a z i n g r e m a in below 40 p e r c e n t f o r m ore th a n one d a y , even i f th e t e m p e r a t u r e change p e r s i s t e d , and e v e n t h o u g h r e c o r d e d t e m p e r a t u r e s 56 r a n g e d f r o m 8 ° t o - 2 6 °C. Change i n t h e d a i l y g r a z i n g s c h e d u l e s h o w s d e f in i te behavioral adjustm ent as tem peratures f lu c a tu a te , prolonged change i n t o t a l daily b u t l a c k of g r a z in g tim e as d a i l y sc h e d u le s c h a n g e d may i n d i c a t e a l a c k o f p h y s i o l o g i c a d j u s t m e n t . w ords, behavioral adjustm ents physiologic adjustments. w ere made w ithout In o th e r the need of Perhaps b e h a v i o r a l a d j u s t m e n t s a l l o w th e cow to r e m a in p h y s i o l o g i c a l l y s t a b l e d u rin g ti m e s o f a c u te s t r e s s . This e o n s l u s i o n i s - s i m i l a r t o o n e made by S e n f t a n d E i t t e n h b u s e ( 1 9 8 5 ) . They pr op os e " b e h a v i o r a l a c c l i m a t i o n " a s a "mechanism f o r o f f s e t t i n g STTS ( s h o r t t e r m t h e r m a l s t r e s s ) " . In t h i s study, such b e h a v io ra l a c c l i m a t i o n i n v o l v e d a d j u s t m e n t s o f w he n d u r i n g t h e day t h e cows grazed, but d id n o t n e c e s s a r i l y in v o l v e a d ju s tm e n t of t o t a l d a i ly g razin g tim e. I f t h e cow s w e r e r e m o v e d f r o m t h e i r TNZ f o r b r i e f p e r i o d s , a d j u s t m e n t s w e r e made q u i t e r a p i d l y a n d t h e co w s q u i c k l y r e t u r n e d t o nor ma l p h y s i o l o g i c s t a t u s . Young (1981) e s t i m a t e d t h e l o w e r c r i t i c a l t e m p e r a t u r e of t h e dry p r e g n a n t cow t o be =20o C, t o be - 2 3 0C. W eb s te r (1970a) r e p o r t e d t h i s t e m p e r a t u r e These two e s t i m a t e s o f t h e l o w e r boundary o f t h e TNZ a r e s u p p o r t e d by t h i s s tu d y. t h e i r TNZ down t o , I t would a p p e a r t h a t t h e s e cows r e m a i n e d i n and p o s s i b l y below, - 2 3 ° . R e s u l t s of t h e t i m e s e r i e s a n a l y s i s a l s o i n d i c a t e t h a t cows w er e not fo rced to r e - a c c l i m a t e them selves during e i t h e r w in te r. Grazing t i m e on an y g i v e n day w a s i n d e p e n d e n t o f p r e v i o u s g r a z i n g t i m e a n d present and past E i t t e n h o u s e (1985). tem perature. This is supported by Senft and T h e i r " t h e r m a l a c c l i m a t i o n model" i s based on th e c o n c e p t t h a t p h y s i o l o g i c a l a c c l i m a t i o n d e p e n d s on a t i m e l a g t h a t 57 a llo w s th e i n t e g r a t i o n of e x p e rie n c e over tim e. The t i m e s e r i e s a n a l y s i s would i n d i c a t e t h a t s uch i n t e g r a t i o n and a c c l i m a t i o n was n o t o c c u r r i n g ; t h e r e f o r e , t h e cows w e r e w i t h i n t h e i r TNZ. F or age I n t a k e I n t a k e e s t i m a t e s i n 1985 w e r e s i m i l a r t o t h o s e o f t h e p r e v i o u s s t u d y a t Bed B l u f f Ranch (T ur n er 1985). E s t i m a t e s i n 1984 were l o w e r t h a n t h e I 985 e s t i m a t e s a n d s l i g h t l y l o w e r t h a n t h e p r e v i o u s s t u d y . The o v e r a l l i n t a k e a v e r a g e f o r 1984 was 0.87 p e r c e n t of body w e i g h t , f o r 1985 i t was 1.25 p e r c e n t , and i n t h e s tu d y by Turn er ( 1985) i n t a k e r a n g e d f r o m 0.8 t o 1.4 p e r c e n t w i t h a n a v e r a g e o f a b o u t 1.1 p e r c e n t . These i n t a k e d i f f e r e n c e s may s t e m f r o m t h e f o r a g e p o r t i o n of t h e i n t a k e e q u a t i o n . vitro d ig estib ility used i n t h i s study) Holechek e t a l . d ig e stib ility (1986) s t a t e d t h a t i n t e c h n i q u e s u s i n g a 48 hour i n c u b a t i o n p e r i o d (as poorly e s tim a te fo rag e d ig e stib ility . -True d i f f e r e n c e s i n d i g e s t i b i l i t y among y e a r s , e r r o r s i n e s o p h a g e a l e x t r u s a c o l l e c t i o n s , and e r r o r s i n i n v i t r o te c h n i q u e co u ld a l l a c c o u n t f o r i n t a k e d i f f e r e n c e s s e e n b et w ee n 1981 and 1985. The model r e l a t i n g d a i l y i n t a k e t o d a i l y g r a z i n g t i m e f a i l e d t o show a s i g n i f i c a n t r e l a t i o n s h i p . th is failure. F irst, intake i s Again t h e r e may be tw o r e a s o n s f o r a f u n c t i o n of t h r e e f a c t o r s : grazing t i m e , b i t e r a t e , and b i t e s i z e ( S p e d d i n g e t a l . a s c i t e d i n Chacon e t al. 1976). need to Second, These t h r e e f a c t o r s can o p e r a t e i n d e p e n d e n t l y and. a l l may be q u a n t i f i e d before a re la tio n s h ip c a n be e s t a b l i s h e d . e s t i m a t e s of f o r a g e i n t a k e w e r e made f o r o n l y 6 days i n 1984 58 and 4 days i n 1985. E s t i m a t e s f o r more days may be r e q u i r e d b e f o r e a r e l a t i o n s h i p can be e s t a b l i s h e d . S u p p le m e n ts used i n 1985 had d e f i n i t e e f f e c t s i n 3 r e l a t e d a r e a s . F i r s t , i t w o u l d seem t h a t s u p p l e m e n t l o w e r e d t h e c o s t o f o b t a i n i n g forage. Supplem ented grazing, b ut consumed more f o r a g e th a n th e u n s u p p le m e n te d group. Second, groups trav eled less an d s p e n t less tim e t h e h i g h e r i n t a k e o f s u p p l e m e n t e d co w s may i n d i c a t e a s u p p l e m e n t e f f e c t on d i g e s t i o n , p o s s i b l y on p a s s a g e r a t e th r o u g h t h e r e t i c u l o - r u m e n . F a s t e r p a s s a g e r a t e may h a v e b e e n a p r e q u i s i t e f o r h i g h e r i n t a k e u n d e r t h e b u l k - l i m i t i n g c o n d i t i o n s i m p o s e d by m a t u r e w in te r forage. This i s s u p p o r t e d by f i n d i n g s o f C a m p l i n g e t a l . (1 9 6 2 ) , a n d B r a n i n e a n d G a l y e a n ( 1 9 8 5 ) . I n a d d i t i o n , t h e b l o o d m e a l supplem ent w ith i t s bypass p r o t e i n had th e l a r g e s t in t a k e e f f e c t . P a rt of the d ig e s t iv e e f f e c t i n t h i s indirect, since i t was assumed t h a t b y pa ss p r o t e i n d i d n o t d i r e c t l y a f f e c t rumen f e r m e n t a t i o n . study may have c a s e would have t o have been Two o t h e r mechanisms n o t me as u re d i n t h i s contributed to the increased intake seen w ith s u p p l e m e n t : an i n c r e a s e i n rumen dry m a t t e r c o n t e n t and a n i n c r e a s e i n ru m in a tio n tim e. Welsh (1982) c o n cl u d ed t h a t r u m i n a t i o n was t h e most i m p o r t a n t method of a c h i e v i n g p a r t i c l e s i z e r e d u c t i o n and p a r t i c l e p a s s a g e (more i m p o r t a n t t h a n rum en m ic ro b ia l a c t i v i t y ) . tim e showed c l o s e a s s o c ia tio n w ith intake; anim als Rum ination deprived of r u m i n a t i o n showed d e c r e a s e d i n t a k e and s p e n t f e e d i n g t i m e s r u m i n a t i n g in s t e a d of eatin g . And t h i r d , t h e s u p p l e m e n t s a p p e a r t o h a v e h e l p e d t h e cow s m e e t th eir m aintainence requirem ent. K leiber (I 96 I ) . e x p r e s s e d th is 59 r e q u i r e m e n t a s t h e s t a t e i n which a n i m a l s n e i t h e r l o s e nor g a i n body substance. is Expanding on t h i s d e f i n i t i o n , t h e m a i n t a i n e n c e r e q u i r e m e n t the le v e l condition, o f n u t r i t i o n r e q u i r e d t o m a i n t a i n body w e i g h t , c o n s t a n t body t e m p e r a t u r e , and a g r o w in g f e t u s . body I n 1985, a l l s u p p le m e n te d cows g a i n e d w e i g h t , a p p r o x i m a t e l y t h e w e i g h t o f t h e i r calf. Also, alm ost t h e a n i m a l f a t s u p p l e m e n t h e l p e d t h a t group m a i n t a i n a n co n stan t body cond itio n . This in co n trast to the u n s u p p le m e n te d cows t h a t g e n e r a l l y l o s t w e i g h t and l o s t s i g n i f i c a n t l y more c o n d i t i o n t h a n t h e f a t group. to m ain tain in g The su p p le m en te d cows w e r e c l o s e r body s u b s t a n c e , and th erefo re m eeting m a i n t a i nance r e q u ir e m e n t, t h a n t h e u n s u p p l e m e n t e d cows. surface (i.e. body in dications. tem perature in the ear) would th e ir Measuring v erify these I t w ould g i v e a c o n t i n u o u s and a c c u r a t e i n d i c a t i o n of t h e h e a t p r o d u c t i o n , h e a t l o s s , and m e t a b o l i c s t a t u s of th e a n im a l. The p e r i o d i c v a s o d i l a t i o n and wa rming a t t h e s k i n s u r f a c e r e p o r t e d by B l a x t e r a n d Wainman ( 1 9 6 1 ) s h o u l d i n d i c a t e n o r m a l w i n t e r m e t a b o l i c status. Other t e m p e r a t u r e p a t t e r n s may i n d i c a t e an i n a b i l i t y t o meet t h e m a i n t a i nance r e q u i r e m e n t and r e m o v a l from t h e TNZ. The s u p p le m e n t e f f e c t s r e p o r t e d h e r e a r e open t o q u e s t i o n i n two areas, d i g e s t i b i l i t y and n u t r i e n t b a la n c e . When f o r a g e i n t a k e i s e s t i m a t e d u s i n g e s t i m a t e s o f f e c a l o u t p u t and f o r a g e d i g e s t i b i l i t y , an i n c r e a s e i n f e c a l o u t p u t may i n d i c a t e a n i n c r e a s e i n i n t a k e o r a decrease however, in d ig e stib ility . digestibility This study has assumed the form er; changes w i t h s u p p l e m e n t were n o t me asu re d and may r e p r e s e n t a p o r t i o n o f t h e e f f e c t s seen. For i n s t a n c e , t h e an im al f a t s u p p l e m e n t (which r e s u l t e d i n t h e h i g h e s t f e c a l o u t p u t ) may have 60 decreased forage d i g e s t i b i l i t y by c o a t i n g t h e f i b e r p a r t i c l e s w i t h a l a y e r of f a t and p r e v e n t in g or r e d u c i n g m i c r o b i a l a tta c h m e n t. This same mechanism may have d e c r e a s e d rumen d i g e s t i o n o f t h e soybean meal i n the supplement, t h u s t r a n s f o r m i n g t h i s s u p p le m e n t so t h a t i n s t e a d o f p r o v i d i n g r u m e n d e g r a d a b l e p r o t e i n a n d b y p a s s e n e r g y i t may h a v e p r o v i d e d b y p as s e n e r g y and p r o t e i n , protein. and l i t t l e or no rumen d e g r a d a b l e Such a m e c h a n i s m may h a v e p r o d u c e d n i t r o g e n a n d e n e r g y b a l a n c e s o t h e r t h a n t h o s e e x p e c t e d and assumed. t h e rumen, i n t h i s c a s e , Nitrogen balance i n may have been s i m i l a r t o t h a t e x p e c t e d i n t h e u n s u p p le m e n te d gro u p ; a p o s i t i v e n i t r o g e n and e n e r g y b a l a n c e i n th e lower g a s t r o i n t e s t i n a l t r a c t may have o f f s e t rumen d e f i c i e n c i e s and a l l o w e d f o r t h e p o s i t i v e e f f e c t s s e e n w i t h - f at s u p p le m en t. I n 1984 and 1985 t h e r e s u l t s o f age group a n a l y s i s b r o u g h t o u t an i m p o r t a n t f a c t o r i n a cow’s a b i l i t y range. I n b o t h y e a r s , t h e 3 y e a r o l d co w s w e r e s p e n d i n g t h e i r f i r s t w i n t e r on ran ge . mo re f o r a g e . This group s p e n t more t i m e g r a z i n g w i t h o u t consuming This s u g g e s ts t h a t g r a z in g e f f i c i e n c y in c r e a s e d w ith rangeland experience. able t o m a i n t a i n i t s e l f on w i n t e r to low er the G r a z i n g may be a l e a r n e d b e h a v i o r w i t h cow s n u tritio n al cost f a m i l i a r i z e them selves w ith w i n t e r range. ( of th is activ ity as they 61 SUMMARY The two main n u t r i t i o n a l c o s t s f o r t h e r a n g e b eef cow a r e (I) t h e m a i n t a i n e n c e r e q u i r e m e n t a n d (2) t h e l e v e l o f a c t i v i t y , g r a z i n g and t r a v e l . especially Bo th o f t h e s e c o s t s a r e h i g h i n w i n t e r due t o e n v i r o n m e n t a l s t r e s s and t h e a s s o c i a t e d i n c r e a s e i n h e a t production, an d due t o t h e r e l a t i v e l y low n u t r i t i o n a l v a l u e o f m a t u r e d o r m a n t forage. these Cows r e c i e v i n g p r o t e i n s u p p l e m e n t a r e b e t t e r a b l e t o m e e t costs, and a r e able to rem ain p ro d u ctiv e under the harsh c o n d i t i o n s imposed by t h e w i n t e r r a n g e e n v i r o n m e n t. In the w in ters of 1 984 and 198 5, a study w as c o n d u c t e d on f o o t h i l l r a n g e l a n d i n s o u t h w e s t Montana t o e s t i m a t e f o r a g e i n t a k e , and m e a su r e f o r a g i n g b e h a v i o r , o f s u p p le m e n te d and unsupplemented, cows i n t h e t h i r d t r i m e s t e r of g e s t a t i o n . In t h i s intake s tu d y , cows r e c e i v i n g p r o t e i n s u p pl em en t showed h i g h e r t h a n u n s u pp le m en te d cows w i t h o u t s p e n d i n g more t i m e g r a z i n g . T h i s s u g g e s t s a p o s i t i v e d i g e s t i v e i n f l u e n c e of 250g o f CP su pp le m en t p e r day. Of p a r t i c u l a r i n t e r e s t i s t h e b lo o d meal s u p p le m e n t (200g of r u m e n d e g r a d a b l e p r o t e i n a n d 200g o f b y p a s s p r o t e i n ) w h i c h b r o u g h t about the g r e a t e s t i n c re a s e i n d a ily fo rag e intake. It is d i f f i c u l t t o make c o n c l u s i o n s c o n c e r n i n g t h e r e l a t i o n s h i p of f o r a g e i n t a k e and t h e t h e r m a l e n v i r o n m e n t . intake, D aily e s t i m a t e s of though d i f f i c u l t t o o b t a i n and p r o c e s s , a r e n e c e s s a r y i n o r d e r t o show how a c u t e and c h r o n i c co l d s t r e s s a f f e c t f o r a g e i n t a k e . G r a z in g t i m e h a s shown an i m p o r t a n t b ut som ew hat am biguous relatio n sh ip to tem perature in th is study. As t h e tem perature f l u c t u a t e d b et w e e n 8 and - 2 6 °C t h e d a i l y g r a z i n g s c h e d u l e s h i f t e d t o 62 i n c l u d e m o r e a f t e r n o o n g r a z i n g a n d l e s s m o r n i n g an d n i g h t g r a z i n g . Also, severe tem perature daily g razin g tim e. changes caus ed a s h o r t (one day) d e c l i n e i n T h e s e a d j u s t m e n t s may h a v e b e e n b e h a v i o r a l r e a c t i o n s to acu te cold s t r e s s . T h is s tu d y a l s o showed t h a t p a s t grazing tim e, and p a s t t e m p e r a t u r e d i d n o t have a present tem perature, s i g n i f i c a n t e f f e c t on p r e s e n t g r a z i n g ti m e. Thi s may i n d i c a t e a l a c k o f any c h r o n i c c o l d s t r e s s d u r i n g t h e t r i a l s . U n f o r t u n a t e l y , t e m p e r a t u r e a l o n e i s n o t i n d i c a t i v e of t h e t h e r m a l e n v i r o n m e n t. An i n d e x t h a t c o n t a i n s t e m p e r a t u r e , h u m i d i t y , windspeed, and b a r o m e t r i c p r e s s u r e may e x p l a i n a g r e a t e r p o r t i o n o f t h e v a r i a t i o n i n b e h a v i o r s e e n i n cows on w i n t e r r a n g e . 63 RECOMMENDATIONS AND QUALIFICATIONS Four of th e d a t a s e t s c o l l e c t e d d u r i n g t h i s s t u d y may n o t p r e s e n t a c o m p l e t e o r c o m p l e t e l y a c c u r a t e i n d i c a t i o n o f t h e cow's s i t u a t i o n on w i n t e r range. Three o f t h e s e w ere m e n t i o n e d i n o t h e r p o r t i o n s of t h i s t e x t , b u t w i l l be d i s c u s s e d h e r e , a l s o . M o d ific a tio n of th e se fo u r d a t a s e t s w o u l d p r o v i d e a m or e c o m p l e t e p i c t u r e o f w i n t e r f o r a g i n g behavior. P e d o m e t e r s may n o t g i v e a n a c c u r a t e i n d i c a t i o n o f t h e d ai ly d i s t a n c e t r a v e l e d by c a t t l e . movement o f t h a t l e g , d irectly Pedom eters a tta c h e d to th e l e g r e c o r d but i t i s n o t alw ay s p o s s i b l e to t r a n s l a t e fro m l e g movement to d istance trav eled since movement p r o d u c e s a d i f f e r e n t q u a n t i t y o f f o r w a r d m ot ion . each le g Given t h i s i n a d e q u a c y , p e d o m e te r i n f o r m a t i o n i s n o t a c c e p t a b l e t o a l a r g e p a r t of the s c i e n t i f i c trav el com munity, rep resen ts expenditure. and i t i s an im p o rta n t b a s ic a lly ignored. part of a cow 's d aily Howe ver , energy F i n d i n g a n a c c u r a t e m e a su r e of d i s t a n c e s t r a v e l e d would h e lp a g r e a t deal i n e x p la in in g the n u t r i t i o n a l w i n t e r r an g e. I t may be p o s s i b l e , c o s t of s u r v i v i n g on and w o r t h w h i l e , t o d e v e l o p a more s e n s i t i v e r e c o r d i n g d e v i c e w i t h t h e g o a l o f more a c c u r a t e l y r e l a t i n g movement t o t r a v e l . A second a r e a of in a d e q u a te i n f o r m a tio n has been d a i ly in ta k e . As m e n t i o n e d p r e v i o u s l y , d a i l y f e c a l g r a b s a m p l e s f r o m a p o r t i o n o f th e herd, previous adjusted studies, using w ould to tal g reatly fecal co llection enhance our in th is ab ility c o n c l u s i o n s a n d show r e l a t i o n s h i p s b e t w e e n i n t a k e , environm ental fa c to r s . as to behavior, and make and 64 W eather tem perature, im p o rtan t in fo rm atio n hum idity, part of should w indspeed, th is also and in fo rm atio n be expanded to in c lu d e barom etric pressure. An w o u l d be d e t e r m i n i n g t h e c o n t r i b u ti o n of each catag o ry to the o v e r a l l th e rm al environm ent. T h i s may be a s t u d y i n i t s e l f , the understanding of the but would c o n t r i b u t e s u b s t a n t i a l l y t o environm ental pressures placed on t h e animals. Finally, t h e e s t i m a t e s o f f o r a g e d i g e s t i b i l i t y have shown a g r e a t deal of v a r ia tio n . I f t h i s v a r i a t i o n i s a c t u a l , i t would be i m p o r t a n t t o i n s u r e t h a t g r a z e d f o r a g e s a m p l e s be o b t a i n e d i n a l l a r e a s o f t h e p a s tu re throughout the t r i a l . R e c o r d i n g s a m p l e l o c a t i o n may h e l p e x p l a i n some o f t h e v a r i a t i o n seen. Also, t h e i n v i t r o te c h n i q u e may need t o be m o d i f i e d t o more c l o s e l y i m i t a t e a c t u a l rumen f e r m e n t a t i o n conditions. A r e c e n t s t u d y by H o l e c h e k e t a l . ( 1986) may p r o v i d e a b a s i s f o r modifying t h i s technique. Once a b a s e l i n e o f w i n t e r f o r a g i n g b e h a v i o r i s e s t a b l i s h e d , i t may be e n l i g h t e n i n g t o s t u d y b e h a v i o r a t o t h e r t i m e s of t h e y e a r . s p e c i a l i n t e r e s t m i g h t be O c t o b e r a n d N o v em b er , Of t h e m o n t h s when c h r o n i c c o l d s t r e s s and a c c l i m a t i o n t o c o l d a r e most l i k e l y t o oc cu r. The o t h e r m a j o r a r e a of i n t e r e s t i n t h i s s tu d y , i s t h e s up ple m ent and i t s e f f e c t on cow p erf or ma nce . shown d e f i n i t e b e n e f i t s , Providing p r o te i n supplement has but t h e r e r e m a i n two a r e a s o f concern. m echanism s of p r o t e i n b e n e f i t a r e n o t u n d e rs to o d , bypas s p r o t e i n e f f e c t especially s e e n w i t h blood meal i n t h i s s tu d y . Also, The the the e c o n o m i c s o f s u p p l e m e n t a t i o n , i n c l u d i n g w hen a n d how much t o f e e d , 65 have n o t been s a t i s f a c t o r i l y d e t e r m i n e d . Study of t h e s e two q u e s t i o n s s h o u ld c o n t i n u e and expand i f p o s s i b l e . So f a r t h e end p o i n t o f t h e s t u d y each y e a r h a s been body w e i g h t and c o n d i t i o n changes, i f t h i s e n d p o i n t w e r e r e - f o c u s e d i t may be m o r e h e l p f u l i n e x p l a i n i n g a n d d e s c r i b i n g so me o f t h e e f f e c t s s e e n . Body w e i g h t and c o n d i t i o n changes a r e one i n d i c a t i o n , inadequate indication, of w hether or not th e m a i n t a i n e n c e r e q u i r e m e n t over t h e w i n t e r . bu t p e r h a p s an co w s h a v e m e t t h e Me etin g t h e m a i n t a i n e n c e r e q u i r e m e n t i s t h e o v e r a l l g oal and t h e r e may be o t h e r m e a s u r e s t h a t would b e t t e r i n d i c a t e w h e th e r or n o t th e g o al i s b e i n g met. a c c u r a t e m e a su r e of t h e amount o f a d i p o s e t i s s u e , show the extent maintainence. to Also, w hich that tissue measuring s u r f a c e was A more for instance, being would u tilized body t e m p e r a t u r e (i . e . for in the e a r ) w o u l d i n d i c a t e i f t h e cows w e r e w i t h i n t h e i r TNZ a n d a b l e t o m a i n t a i n t h e h e a t p r o d u c t i o n r e q u i r e d i n t h e w i n t e r r a n g e en v i ro n m en t. 66 LITERATURE CITED 67 LITERATURE CITED Adams, D. 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E f f e c t o f p r o l o n g e d c o l d e x p os ur e on d i g e s t i o n and m e t a b o l i s m i n r u m i n a n t s . I n t . L i v e s t o c k Env. Symp., Am. Soc. Agr., Eng., S t . J o e s e p h . pp. 7 5 - 8 1 . Z i n n , R. A., a n d F. N. Owens. 1981. I n f l u e n c e o f l e v e l o f f e e d i n t a k e on d i g e s t i v e f u n c t i o n . I. N i t r o g e n m e ta b o li s m . Oklahoma Ag. Exp. S t a . Oklahoma S t a t e U n i v e r s i t y . USDA-SEA-AR, APPENDIX 79 Table 19 . Body w e i g h t s 1984. and c o n d i t i o n s c o r e s f o r 5 cow age g ro u p s , Initial Age , (yrs) Ending Weight C o nd it ion i (kg) Score Change■ Weight C o n d it io n Sco re 5.7° 503b 5.3° -1 0 .5a -0 .5 a 5.6° • 478b 5 . I bc +1.2a -0 .6 a - 1 0 . 8a -0 .4 a C o n d it io n Score Weight 7 518a 6 477a b 5 482ab 5 . 5 bc 4 7 I ab 5 . 2 bc 4 470ab 5 . I ab 470ab 4.8b -6 .4 a . -0.4a 3 43 5b 4.8a 435a 4.3a -3 .6 a a , ^»0Means i n t h e same column d i f f e r e n t (p<.05). T ab l e 20. Body w e i g h t s 1985. In itial Age (yrs) Weight C o n d it io n (kg) Score and with different -0 .4 a superscripts are c o n d i t i o n s c o r e s f o r 4 cow age gr oups, Ending Weight Change C o n d it io n Sc o re Weight Condition S co re 6 502ab 6 .Ib 534b 5 .Ib 32a -0 .9 a 5 537b 5.8b 56lb 4 . 7 ab 24a -1.2a 4 495ab 5 .Ia 542b 4 . 3 ab 41a -0 .7 a 3 461a 4.9a 490a 3.9a 29a - I . Oa a >^Means i n th e same column w i t h d i f f e r e n t s u p e r s c r i p t s a r e d i f f e r e n t (p<.05). 80 Tabl e 21 . Body w e i g h t s and c o n d i t i o n s c o r e s o f each t r e a t m e n t group, 1985. In itial TRT Ending Weight C o n d it io n (kg) Score Weight Change C o n d it io n S co re Weight Condition Sco re CON 501a 5.4a 500a 3 .9 a -Ia -1 .4 a SBM 523a 5.6a 556a 4 . 5 ab 33b - I . I ab FAT 520a 5 . Ta 547a 5 . 3b 27b - 0 . 4b CGM 502a 5.6a 546a 4 . 5 ab 44b - K I ab BM 499a 5.3a 538a 4 . 6 ab 37b - 0 . 7 ab 3. b ’ Means i n t h e same column w i t h d i f f e r e n t s u p e r s c r i p t s a r e d i f f e r e n t (p<.05). Ta b le 22,. D a il y g r a z i n g gr ou ps , 1985. times and d i s t a n c e s t r a v e l e d f o r 4 cow age G r a z in g Time ( %) Age 7AM-1PM ■ I PM-7PM 7PM-7AM Dail y D is ta n c e T r a v e l e d (km) 3 6 0 . 9a 68. 7a 14.I a 39.4a 4 . Ob 4 5 5 .8 ^ 6 6 .Sab 11. 2b 3 6 . 3b 3.8b 5 53.9^ 6 4 . 8b 1 4 .8a 37.3b 7 .9 a 6 55.Ib 6 4 .2b 1 5 . 9a 37.7b 3 . 7b ' i n t h e same column w i t h d i f f e r e n t s u p e r s c r i p t s a r e d i f f e r e n t (p<.05). a ’ ^M eans 81 Ta b le 23. Chromic o x i d e r e c o v e r y (% f e d ) , 1985. T r ea tm e nt Group Source SBM Samples from to ta l collection 95 .7 Grab samples 71 . I a FAT a CGM a BM a A ll TRTf s a 60.9 7 0 .2 102 43.7a 6 2 .4a 64.I a 82.3 • a Means i n th e same row w i t h th e same s u p e r s c r i p t a r e n o t (p <. 05) T a b l e 24. 60 .3 different D a i l y f e c a l o u t p u t (0MB % BW) e s t i m a t e d u s i n g t o t a l c o l l e c t i o n and CrgOg, 1985. fecal T rea tm en t Method Tot al^ C ollection CON SBM FAT ab a CGM b BM ab ab C 0.56 0 .6 3 0.71 Samples^ from t o t a l collection ----- a 0 .6 8 ' 0 .7 0 0 .8 9 1.10 0.90 Grab ---- 0.94a 1.00a I .IOa 1.40a I .IOe s a m p le st > 3 0 .6 0 Al l TRT 0.60 ab a 0.64 b d cL b ’ Means i n t h e same row w i t h d i f f e r e n t s u p e r s c r i p t s a r e d i f f e r e n t (p<.05). c > ’ eMeans i n t h e same column w i t h d i f f e r e n t s u p e r s c r i p t s a r e d i f f e r e n t (p<.05). ■‘Average f o r 24 h o u r s c a l c u l a t e d from a 96 hour c o l l e c t i o n p e r i o d . ^ E s t i m a t e d w i t h chromic o x id e t e c h n i q u e . ^Grab s am pl es t a k e n once d u r i n g t h e 96 hour t o t a l c o l l e c t i o n p e r i o d . 82 Ta b le 25. T o t a l d a i l y i n t a k e (0MB % BW), 1985. Day Treat­ ment J a n 17 J a n 23 Feb 6 Feb 20 Mean o f Al l Days CON 0 . 9 4 + 0 .1 0 ° I . 0 1 + 0 . OTa 0.92+0.08° 1 . 04 +0 .08 ° 0 . 9 8 + 0 .0 4 d SBM I . 6 2 + 0 . 09ab I . I 8 ±0. 05 a 1 . 14 ±p .0 7 bc 1 . 3 7 + 0 . 10bc 1.3 4+0 .05 b° FAT I . 2 8 + 0 . I 4bc I . 0 6 + 0 . 14a I . 7 1 + 0 . 17a 2.28+0.43a I . 5 9 + 0 . 14ab CGM I . 2 3 + 0 . I 3bc I . 2 4 + 0 . 09a 1 . 1 2 + 0 . 1 Obc 1. 20 +0. 13 ° 1 . 20+0.06° d BM 2.02+p.22a I . 4 8 + 0 . 13ab 2 . 0 7 + 0 . 3 3 ab I . 6 9 + 0 . 12a Mean a l l of TRT’ s I .21+0. I l a ef I .45+0 .08 S 1,1 5+0 .04 e fg 1. 31 +0. 07 I .62+0.14 1.37 a > b , c , d Means i n th e same GOlumn w i t h d i f f e r e n t s u p e r s c r i p t s a r e d iffe re n t (p<.05). e > >^Means i n t h e same row w i t h d i f f e r e n t s u p e r s c r i p t s a r e d i f f e r e n t (p<.05). 83 120 - - 110- - Cr O recovery IOO- 9 0 -- 80- 70-- 60 -■ Day I Day Feed SBM F i g u r e 7. 2 Day 3 day BM Day Feed FAT 4 day CGM Chromic o x i d e r e c o v e r y p a t t e r n d u r i n g t o t a l f e c a l c o l l e c t i o n , I 985. MONTANA STATE UNIVERSITY LIBRARIES 3 762 100 3603 3 Main Jf37Q D922 c o p .2 Dunn, Roger W i l l i am Physiological and ben a v io ra l responses o f . . . Main N378 D922 co p . 2 4