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The response of axial growth gradients within the skeleton of the black-tailed deer to variations in… Addison, Ralor Blendle 1970

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The  Response o f A x i a l Growth G r a d i e n t s  W i t h i n the  S k e l e t o n o f the  Deer t o V a r i a t i o n s Pattern  o f Energy  i n the  Black-Tailed Level  and  Availability.  by  Ralor Blendle  Addison 1963 1966  B.Sc, U n i v e r s i t y o f B r i t i s h Columbia, M o S c . j U n i v e r s i t y o f B r i t i s h Columbia,  A Thesis the  Submitted i n P a r t i a l F u l f i l m e n t Requirements f o r the Doctor of i n the  of  Degree o f  Philosophy  Department of  Zoology  We  a c c e p t t h i s t h e s i s as conforming t o  required  The  standard  U n i v e r s i t y of B r i t i s h April,  1970  Columbia  the  In presenting  this thesis in partial  fulfilment of the requirements for  an advanced degree at the University of B r i t i s h Columbia, I agree that the Library shall make i t freely available for reference  and  study.  I further agree that permission for extensive copying of this thesis for scholarly purposes may by his representatives.  be granted by the Head of my  Department or  It is understood that copying or publication  of this thesis for financial gain shall not be allowed without my written permission.  Department of  Zoology  The University of B r i t i s h Columbia Vancouver 8, Canada  Date  June 8,  1970  ABSTRACT: A s t u d y was d e s i g n e d t o examine t h e i n t e r a c t i o n o f a x i a l g r o w t h g r a d i e n t s and t i m e s p e c i f i c e n e r g y r e s t r i c t i o n s upon, t h e g r o w t h r a t e s o f s k e l e t a l d i m e n s i o n s i n t h e B l a c k t a i l e d deer. A t o t a l o f 53 new-born fawns was c a p t u r e d and t h e n r e a r e d i n c a p t i v i t y on c o n t r o l l e d e n e r g y r e g i m e s . The t e s t p e r i o d o f 322 d a y s was d i v i d e d i n t o f i v e i n t e r v a l s w i t h t h e e n e r g y l e v e l i n e a c h i n t e r v a l b e i n g s e t as h i g h , medium, o r low p l a n e . R e p r e s e n t a t i v e a n i m a l s were k i l l e d a t t h e end o f e a c h i n t e r v a l t o e x a m i n e g r o w t h o v e r t h e p r e v i o u s i n t e r v a l s and t o e s t a b l i s h a b a s e f o r e v a l u a t i n g growth i n the next i n t e r v a l . F i v e w i l d fawns w e r e c o l l e c t e d a t 112 d a y s and f o u r a t 175 d a y s f o r c o m p a r i s o n with the l a b o r a t o r y standards. R e l a t i v e g r o w t h p r i o r i t i e s o f 23 s k e l e t a l d i m e n s i o n s were i n v e s t i g a t e d by c h a l l e n g i n g s k e l e t a l g r o w t h w i t h a s e r i e s of energy r e s t r i c t i o n s . The amount o f g r o w t h e x h i b i t e d by e a c h s k e l e t a l e l e m e n t i n r e s p o n s e t o t h e l e v e l o f a v a i l a b l e e n e r g y c o u l d be i n t e g r a t e d i n t o an i n t e r p r e t a t i o n o f r e l a t i v e growth p r i o r i t i e s . S k e l e t a l g r o w t h was examined f o r i t s a b i l i t y t o p r o v i d e an i n d i c a t i o n o f t o t a l amount and p a t t e r n o f e n e r g y i n t a k e o v e r two i n t e r v a l s a t t h e end o f 112 d a y s , o v e r t h r e e i n t e r v a l s a t t h e end o f 175 d a y s , and o v e r f i v e i n t e r v a l s a t t h e end o f 322 d a y s . The d e g r e e t o w h i c h t h e e f f e c t s o f e a r l i e r r e s t r i c t i o n s were removed by c o m p e n s a t o r y g r o w t h was a l s o examined i n t h e f i f t h i n t e r v a l . A s h o r t a g e o f a n i m a l s p r e c l u d e d t h e s a c r i f i c e o f any s t a n d a r d s a t 259 d a y s . When s k e l e t a l . s i z e was g r a p h e d a g a i n s t t o t a l e n e r g y i n t a k e , e x p r e s s e d i n C a l o r i e s o f apparent d i g e s t i b l e energy, a t a f i x e d age, t h e p a t t e r n o f e n e r g y i n t a k e c r e a t e d a d i s t r i b u t i o n o f p o i n t s w h i c h c o u l d by c o n v e n i e n t l y bounded by a t r i a n g l e . On d i f f e r e n t p a t t e r n s o f e n e r g y i n t a k e but e q u a l t o t a l energy i n t a k e s , the r e l a t i v e s i z e s o f t h e 23 s k e l e t a l d i m e n s i o n s change t h e i r p o s i t i o n s w i t h i n the b o u n d a r i e s s u r r o u n d i n g the d i s t r i b u t i o n of points. A c o m p a r i s o n o f a s e t o f measurements f r o m an a n i m a l o f unknown n u t r i t i o n a l h i s t o r y w i t h t h e r e f e r e n c e l i n e s i n each of the s t a n d a r d t r i a n g u l a r d i s t r i b u t i o n s leads t o a s e r i e s of energy i n t e r c e p t s that should u n i q u e l y c h a r a c t e r i z e t h e amount o f e n e r g y consumed by t h a t a n i m a l , and t h e p a t t e r n i n w h i c h i t was a v a i l a b l e .  Ill  A t 112 d a y s , t h e t o t a l e n e r g y i n t a k e and p a t t e r n o f r e s t r i c t i o n s c o u l d be d e r i v e d f r o m s k e l e t a l g r o w t h . A t 175 d a y s , a l t h o u g h e s t i m a t e s o f t o t a l e n e r g y i n t a k e s t i l l a p p e a r e d t o be good, t h e p a t t e r n o f e n e r g y i n t a k e c o u l d n o t be e s t a b l i s h e d e x a c t l y , b u t c o u l d be l i m i t e d t o a s m a l l number o f p a t t e r n s . By 322 d a y s , c o m p e n s a t o r y g r o w t h had r e d u c e d d i f f e r e n c e s i n s k e l e t a l s i z e t o t h e p o i n t where n e i t h e r amount n o r p a t t e r n o f e n e r g y a v a i l a b i l i t y c o u l d be deduced. F o u r m e a s u r e m e n t s o f t h e f o r e cannons f r o m w i l d fawns a t 150 d a y s w e r e u s e d t o t r y t o d e f i n e t h e n u t r i t i o n a l r e g i m e w i t h a p o r t i o n o f t h e s k e l e t o n w h i c h w o u l d be r e a d i l y a v a i l a b l e from h u n t e r - k i l l e d a n i m a l s . This did not a l l o w a p r e c i s e e v a l u a t i o n of the n u t r i t i o n a l regime f o r i n d i v i d u a l animals, but i t d i d p o i n t out that there c a n be t r e m e n d o u s v a r i a b i l i t y i n t h e e n e r g y i n t a k e o f w i l d fawns f r o m a r e l a t i v e l y s m a l l a r e a o f h a b i t a t . It is s u g g e s t e d t h a t a c o l l e c t i o n o f f o r e cannons s h o u l d be made a l o n g w i t h t h a t o f body w e i g h t s t o a l l o w an i m p r o v e d e v a l u a t i o n of range c o n d i t i o n s .  i i i exT a b l e o f Contents  Page  Abstract:  i i  T a b l e o f Contents: List  of Tables:  List  of Figures:  i i i V v i i  Acknowledgements:  x  G l o s s a r y o f Terms:  x i i  Introduction:  1  Methods:  6  I. II.  III.  Rations.  10  1.  Milk:  10  2.  Weaner r a t i o n  3.  Adult r a t i o n  V. VI.  VII.  (U.B.C. 3 6 - S - 6 3 ) :  D i g e s t i b l e Energy.  II.  13 Moisture, 17  E x p e r i m e n t a l Treatments.  20  C o l l e c t i o n of F i e l d  21  Specimens.  Measurement o f Growth.  23  1.  Body weight:  23  2.  S k e l e t a l measurements:  27  Treatment o f Data.  Cumulative  30 31  Results: I.  12  (U.B.C. 3 6 - 5 7 ) :  Determination of Ration D i g e s t i b i l i t y , and Apparent  IV.  6  R e a r i n g the E x p e r i m e n t a l Animals.  Energy I n t a k e .  V a r i a t i o n s i n N u t r i t i o n a l Plane.  31 36  iv  Page III.  V a r i a t i o n s o f Body Weight w i t h C u m u l a t i v e 48  Energy I n t a k e . IV.  S k e l e t a l Dimensions o f L a b o r a t o r y  Reared Deer.  54  Discussion: I. II.  III.  E s t i m a t i o n o f C u m u l a t i v e Energy I n t a k e . Determination  o f P a t t e r n o f Energy I n t a k e  (Energy Regime).  169  Regimes o f N u t r i t i o n t o 112 Days.  180  Regime *HH*  184  Regime 'MM'  16*5  Regime 'ML'  187  Regime •MH»  188  Regime «LH»  189  Regime «LL»  190  Regime HM  f  191  Regime HL»  191  f  T  IV. V. VI.  VII. VIII.  161  Energy I n t a k e s o f W i l d Fawns t o 112 Days.  192  Regimes o f N u t r i t i o n t o 175 Days.  194  P a t t e r n s o f Energy I n t a k e s o f W i l d Fawns t o 175 Days.  196  The S k e l e t a l Growth o f Fawns t o 322 Days.  203  The Dimensions o f Fore Cannons C o l l e c t e d from H u n t e r - K i l l e d Favms i n 1966 and 1967.  204  Summary and C o n c l u s i o n s :  214  Bibliography:  219  Appendix:  224  L i s t o f Tables  Table  1.  The pre-weaning  and weaning d a i l y  allotments of milk. Table  2.  The f o r m u l a t i o n o f weaner r a t i o n ,  U.B.C  36-S-63.  Table  3-  The f o r m u l a t i o n o f a d u l t r a t i o n ,  U.B.C  36-57.  Table  4.  N u t r i e n t c o m p o s i t i o n o f r a t i o n s U.B.C. 36-S-63  and 3 6 - 5 7 compared w i t h t h e  r e q u i r e m e n t s f o r g r o w i n g sheep (N.A.S. 1963).  Table  5.  E s t i m a t e d c u m u l a t i v e energy i n t a k e s o f w i l d fawns based on body w e i g h t .  Table  6.  Fore cannons c o l l e c t e d from h u n t e r killed  Table  7.  fawns.  C u m u l a t i v e energy i n t a k e ( C a l o r i e s ) o f l a b o r a t o r y males, by i n t e r v a l .  Table  S.  C u m u l a t i v e energy i n t a k e ( C a l o r i e s ) o f l a b o r a t o r y f e m a l e s , by i n t e r v a l .  Table  9.  R e - e v a l u a t i o n o f energy i n d i c e s and regimes o f l a b o r a t o r y - r e a r e d  fawns.  Tables 10  The d i m e n s i o n s and energy i n t e r c e p t s o  to 5 6 .  t h e s k e l e t a l components o f l a b o r a t o r y fawns.  VI  Page T a b l e s 57  The d i m e n s i o n s and energy i n t e r c e p t s o f  to  the  65.  Table  66.  s k e l e t a l components o f w i l d fawns.  The e s t i m a t e d t o t a l energy i n t a k e s o f w i l d fawns.  Table  67.  152-160  165  The d e v i a t i o n s o f the energy i n t e r c e p t s o f t h e s k e l e t a l d i m e n s i o n s from the e s t i m a t e s o f a c t u a l energy i n t a k e f o r l a b o r a t o r y and w i l d fawns.  Table  68.  Fore cannon d i m e n s i o n s o f h u n t e r k i l l e d fawns.  Table  69.  197  206  T o t a l energy i n t a k e s o f fawns based on measurements o f t h e f o r e cannon.  210  vii Page List Figure  1.  of Figures  The proposed energy regimes f o r male 22  fawns (females d u p l i c a t e «HHH» and L L L » ) . f  Figure  2  V a r i a t i o n s i n t h e exponent y » r e q u i r e d ,  0  to the  f i t heat p r o d u c t i o n slope, 39.65,  data o f deer when  i s held  constant; y i . e . , B.H.P. = 39.65 W  41  lb. Figure  3.  R e s t i n g heat p r o d u c t i o n o f deer d e s c r i b e d as a m u l t i p l e o f b a s a l heat  production;  i . e . , R.H.P. = 2.2645 B.H.P. = 2.2645  y (39.65) W  42  lb. Figure  4.  The r e l a t i o n s h i p between body weight and t o t a l energy i n t a k e o f l a b o r a t o r y r e a r e d fawns a t 49 days o f age.  Figure  5.  49  The r e l a t i o n s h i p between body weight and t o t a l energy i n t a k e o f l a b o r a t o r y r e a r e d fawns a t 112 days o f age.  Figure  6.  50  The r e l a t i o n s h i p between body weight and t o t a l energy i n t a k e o f l a b o r a t o r y r e a r e d fawns a t 175 days o f age.  Figure  7.  51  The r e l a t i o n s h i p between body weight and t o t a l energy i n t a k e o f l a b o r a t o r y r e a r e d fawns a t 322 days o f age.  Figure  8.  52 55  viii Page Figure  8a.  Changes w i t h age i n the l i v e r weights o f laboratory  Figure  8b.  fawns.  Changes w i t h age i n the head weights o f l a b o r a t o r y and w i l d  Figures to  9 50.  55  fawns.  55  The magnitude o f s e l e c t e d s k e l e t a l dimensions i n r e l a t i o n t o t o t a l  energy  i n t a k e i n C a l o r i e s o f apparent  digestible  energy:  Figure  51.  9 t o 22  —  at 112  days;  F i g u r e s 23 t o 36  —  at 175  days;  F i g u r e s 37 t o 50  --  a t 322 days;  Figures  57-99  The l i m i t s o f the range o f p o s s i b l e t o t a l energy i n t a k e s f o r l a b o r a t o r y fawn V26 (HML  regime) as d e f i n e d by i t s s k e l e t a l 163  dimensions. Figure  52.  The l i m i t s o f the range o f p o s s i b l e t o t a l energy i n t a k e s f o r l a b o r a t o r y fawn U33 (LMH regime) as d e f i n e d by i t s s k e l e t a l 164  dimensions. Figure  53.  The l i v e body weights o f w i l d fawns c o l l e c t e d between 49 and 270 days o f age.  Figure  54.  A diagram o f the growth o f the f o r e cannon r e l a t e d t o t o t a l energy i n t a k e .  Figure  55.  168  171  A diagram o f the growth o f the s c a p u l a r e l a t e d t o t o t a l energy  intake.  172  The l i m i t s o f t o t a l energy  intakes  p o s s i b l e w i t h i n each energy regime a t 112 days. The l i m i t s o f t o t a l energy  intakes  p o s s i b l e w i t h i n each energy regime a t 175 days. A diagram d e p i c t i n g the p o s s i b l e  relative  peaks o f growth p r i o r i t y t h r o u g h t h e growth g r a d i e n t s t h a t have been examined. The f r e q u e n c y d i s t r i b u t i o n the  by i n t e r v a l  of  f o r e cannon measurements f r o m h u n t e r -  k i l l e d male fawns i n 1966 and  1967.  X  Acknowledgements  I w i s h t o e x p r e s s my s i n c e r e g r a t i t u d e t o a l l o f t h e persons and a g e n c i e s whose i n t e r e s t and c o - o p e r a t i o n made p o s s i b l e the conducting o f t h i s r e s e a r c h p r o j e c t . Dr. H.C, Nordan, A s s o c i a t e P r o f e s s o r o f Zoology a t t h e U n i v e r s i t y o f B r i t i s h Columbia,  headed t h e a d v i s o r y  committee and was c l o s e l y i n v o l v e d i n a l l phases o f t h e work. Each o f t h e committee members as w e l l as some o t h e r members of f a c u l t y provided u s e f u l c r i t i c i s m o f experimental p r o c e d u r e s and a s s i s t e d i n e d i t i n g t h e f i n a l  thesis  preparation. Many graduate s t u d e n t s were i n v o l v e d i n some way d u r i n g the p r o j e c t , a i d i n g i n h a n d l i n g the animals or i n d i s c u s s i n g the experimental hypotheses.  My s p e c i a l t h a n k s go t o t h e  group o f graduate s t u d e n t s and f a c u l t y based a t t h e C e n t r a l A n i m a l Depot (now t h e Zoology V i v a r i u m ) , U n i v e r s i t y o f B r i t i s h Columbia.  The l o o s e l y o r g a n i z e d i n t e r d i s c i p l i n a r y  c o - o p e r a t i v e p r o v i d e d an i d e a l atmosphere f o r c o n d u c t i n g a project of t h i s nature. The B.C. F i s h and Game Branch gave p e r m i s s i o n each  year  f o r t h e c a p t u r e o f fawns f o r t h e purposes o f t h e e x p e r i m e n t s . At t i m e s when d i f f i c u l t y was e n c o u n t e r e d  i n capturing a  xi  s u f f i c i e n t number o f a n i m a l s , c o n s e r v a t i o n o f f i c e r s biologists  came t o my a i d t o h e l p f i l l  the  and  quota.  Crown Z e l l e r b a c k o f Canada L t d . were v e r y generous i n p r o v i d i n g u n l i m i t e d a c c e s s t o the s t u d y a r e a which s i s t e d o f t h e i r W o l f Lake l o g g i n g o p e r a t i o n a t British  con-  Courtenay,  Columbia.  I would e s p e c i a l l y l i k e t o thank Canadian I n d u s t r i e s Ltd.  and t h e H.R.  M a c M i l l a n F a m i l y f o r the  financial  support t h e y p r o v i d e d d u r i n g the course o f t h e work.  xii G l o s s a r y o f Terms The f o l l o w i n g l i s t  c o n t a i n s e x p r e s s i o n s o f energy i n t a k e  a r e used i n t h i s p r e s e n t a t i o n .  F o r the sake o f b r e v i t y ,  which units  have been o m i t t e d i n most areas o f d i s c u s s i o n but at a l l t i m e s t h e y are i m p l i e d as shown below.  apparent d i g e s t i b l e energy:  g r o s s energy i n a u n i t o f i n g e s t e d  f e e d l e s s the energy l o s t i n the c o r r e s p o n d i n g f e c e s . apparent t o t a l energy i n t a k e : d i g e s t i b l e energy;  not t o be confused w i t h apparent  i t i s the t o t a l energy i n t a k e  (see  below) e s t i m a t e d f o r an animal by one o r a l l parameters o f growth, as c o n t r a s t e d w i t h the measured, or t r u e ,  total  energy i n t a k e o f t h a t a n i m a l . c u m u l a t i v e energy i n t a k e  ( t o t a l energy i n t a k e ) :  suramated  energy i n t a k e s , determined as apparent d i g e s t i b l e in kilocalories  daily  energy  ( C a l o r i e s ) , over the e x p e r i m e n t a l p e r i o d  f o r each fawn; i . e . , from the day at which a body weight o f 10 pounds was in  reached, t o the end o f the time  interval  question.  t o t a l energy i n t a k e :  used synonymously  w i t h cumulative energy t r u e t o t a l energy i n t a k e : added t o emphasize intake.  and i n t e r c h a n g e a b l y  intake. as i n t o t a l energy i n t a k e , w i t h  the c o n t r a s t w i t h 'apparent* t o t a l  'true  1  energy  1  Introduction:  T h i s study was conducted t o determine i f t h e growth and development columbianus  o f B l a c k - t a i l e d deer fawns,  (Richardson),  O d o c o i l e u s hemionus  could q u a n t i t a t i v e l y r e f l e c t the  d i e t a r y energy l e v e l s t o which t h e animals had a c c e s s .  The  u l t i m a t e o b j e c t i v e was t o make p o s s i b l e t h e e v a l u a t i o n o f t h e energy regime o f w i l d fawns through an u n d e r s t a n d i n g o f t h e m o d i f i c a t i o n s induced i n t h e normal developmental p a t t e r n by time s p e c i f i c energy r e s t r i c t i o n s . and development  i s modified  Provided  t h a t t h e growth  i n a p r e d i c t a b l e manner, and t h a t  t h e m o d i f i c a t i o n s a r e a t l e a t semi-permanent i n n a t u r e , an a n a l y t i c a l procedure can be developed t h a t w i l l a l l o w t i f i c a t i o n o f the e f f e c t s o f i n s u f f i c i e n t  energy  iden-  supply.  F o r such a t e c h n i q u e t o be o f g r e a t e s t v a l u e , measurements t a k e n a t t h e end o f a d e f i n e d p e r i o d o f time must information  concerning  the extent,  convey  d u r a t i o n and time o f  o c c u r r e n c e o f any energy r e s t r i c t i o n s t h a t may have during that  occurred  interval.  Such an i d e a i s c e r t a i n l y not unique and has been suggested as an avenue o f i n v e s t i g a t i o n by Riney, 1952.  Until  now, however, t h e r e has not been a t r u l y q u a n t i t a t i v e method f o r e v a l u a t i n g t h e energy regime o f a f r e e f e e d i n g a n i m a l . Q u a l i t a t i v e approaches i n c l u d e t h e many c o n d i t i o n i n d i c e s t h a t have been proposed t o i d e n t i f y animals s u f f e r i n g from some  2 degree o f n u t r i t i o n a l inadequacy ( P a r k and Day,  1942; Cheatum,  1949; Bandy e t a l , 1956; S e v e r i n g h a u s and G o t t l i e b ,  1959).  None o f t h e s e i s s u f f i c i e n t l y comprehensive t o a l l o w a s o p h i s t i c a t e d a n a l y s i s o f s e a s o n a l t r e n d s i n adequacy d i e t a r y energy.  Two  of  s t u d i e s , Bandy (1955) and K l e i n ( 1 9 6 4 ) .  bear a c l o s e resemblance t o t h e p r e s e n t work.  However, i n t h e  f i r s t case o n l y a l i m i t e d number o f parameters o f growth were c o n s i d e r e d , and i n the second, n u t r i t i o n a l d i f f e r e n c e s were not measured but i n f e r r e d .  As i n the p r e s e n t s t u d y , the  h y p o t h e s e s o f Bandy and K l e i n d e v e l o p e d out o f t h e e x t e n s i v e n u t r i t i o n a l s t u d i e s of a g r i c u l t u r i s t s i n v o l v e d i n the p r o d u c t i o n o f improved meat c a r c a s s e s . The e x i s t e n c e o f a x i a l growth g r a d i e n t s has l o n g been known t o t h e d e v e l o p m e n t a l b i o l o g i s t .  The g r a d i e n t s seem t o  be i m p l i c i t i n t h e w r i t i n g s o f Thompson (1917, 1942) a l t h o u g h t h e y are not d e f i n e d as s u c h .  Huxley (1932) r e f e r s t o t h e  work o f Scammon and C a l k i n s (1929) when he d i s c u s s e s t h e of  'Law  A n t e r o - P o s t e r i o r Development and i t s E f f e c t Upon Growth'.  Most o f t h e e a r l y s t u d i e s assumed a ' n o r m a l l y g r o w i n g ' a n i m a l and d i d not c o n s i d e r n u t r i t i o n a l e f f e c t s upon growth, but i t has now been w e l l documented t h a t the form and development  of  a g r o w i n g a n i m a l i s i n f l u e n c e d g r e a t l y by an i n t e r a c t i o n o f a x i a l growth p a t t e r n s and t i m e s p e c i f i c energy M o u l t o n e t a l (1921) conducted one o f the f i r s t  restrictions. studies  concerned w i t h the i n t e r a c t i o n o f energy i n t a k e and body form  3 but i t was n o t u n t i l a f t e r 1935 t h a t much c o n c e r n was g i v e n t o t h i s a s p e c t o f growth.  P a l s s o n (1955) summarizes t h e f i n d i n g  o f many p r e v i o u s workers w i t h t h e s t a t e m e n t : " D u r i n g l a t e f o e t a l l i f e t o m a t u r i t y , any p a r t ,  organ  or t i s s u e o f an a n i m a l ' s body i s p r o p o r t i o n a t e l y 5  most r e t a r d e d i n development by r e s t r i c t e d at t h e stage when i t has h i g h e s t n a t u r a l intensity.  nutrition  growth  R e s t r i c t e d n u t r i t i o n , d u r i n g any age  i n t e r v a l from t h e l a t e f o e t a l stage u n t i l  growth  c e a s e s , has an i n c r e a s i n g r e t a r d i n g e f f e c t on t h e d i f f e r e n t t i s s u e s and r e g i o n s o f an a n i m a l ' s body i n t h e d i r e c t o r d e r o f t h e i r m a t u r i t y ; t h e e a r l i e s t maturing p a r t s or t i s s u e s being l e a s t and t h e l a t e s t m a t u r i n g ones most a f f e c t e d " . U s i n g t h e s e p r i n c i p l e s , i t i s now p o s s i b l e f o r t h e meat p r o d u c e r t o m a n i p u l a t e t h e energy i n t a k e o f an a n i m a l t o e f f e c t a d e s i r e d form o f c a r c a s s (Hammond, 1932a, 1932b; McMeekan, 1940;  P a l s s o n and V e r g e s , 1951; P a l s s o n , 1955; A r c e l a y , 1963;  Anderson e t a l , 1965; Wardrop, 1966).  Within genetic l i m i t s ,  t h e c o n f o r m a t i o n o f a c a r c a s s can be m o d i f i e d i n a p r e d i c t a b l e manner by a p p l i c a t i o n o f energy r e s t r i c t i o n s a t c r i t i c a l t i m e s . I f t h e converse i s t r u e , then an e x a m i n a t i o n o f t h e growth and form o f an a n i m a l s h o u l d g i v e a p i c t u r e o f t h e p a t t e r n o f energy a v a i l a b l e t o t h e a n i m a l d u r i n g t h e growing p e r i o d . Emphasis i n t h i s s t u d y has been p l a c e d on t h e assessment o f s k e l e t a l growth.  The s k e l e t o n i s d i s t r i b u t e d a l o n g t h e  a x i a l g r a d i e n t s and may  i s e a s i l y measured.  Whereas o t h e r  diminish i n s i z e during periods of d e p r i v a t i o n ,  s k e l e t a l d i m e n s i o n s , once a c h i e v e d , t h e r e f o r e , i s not  complicated  are s t a b l e and  by n e g a t i v e  growth.  tissues  the the a n a l y s i s , Another  very  i m p o r t a n t r e a s o n f o r the s e l e c t i o n o f the s k e l e t o n i s t h a t i t i s an e a r l y m a t u r i n g t i s s u e ( J a c k s o n  and  Lowrey, 1912)  and  a h i g h growth p r i o r i t y i n the e a r l y p o s t n a t a l p e r i o d . study (Addison,  1966)  has  A pilot  demonstrated t h a t s k e l e t a l growth i s  s e n s i t i v e t o energy r e s t r i c t i o n s d u r i n g t h e f i r s t s i x months o f life.  This study a l s o provided  e v i d e n c e t h a t t h e r e might be  s u f f i c i e n t growth i n some s k e l e t a l d i m e n s i o n s d u r i n g the six  months t o be o f use  second  i n e v a l u a t i n g energy i n t a k e .  I n o r d e r t o measure the e f f e c t o f a p p a r e n t d i g e s t i b l e energy i n t a k e on s k e l e t a l growth, w i l d caught fawns were i n t h e l a b o r a t o r y under c o n t r o l l e d f e e d i n g r e g i m e s . animals provided compared.  reared  These  the s t a n d a r d s t o w h i c h w i l d fawns c o u l d  be  Fawns were chosen because t h e i r r a p i d growth r a t e  makes them h i g h l y s u s c e p t i b l e t o v a r i a t i o n s i n a v a i l a b l e energy, whereas i n o l d e r a n i m a l s the growth r a t e slows as mature s i z e i s r e a c h e d .  One  s l i g h t l i m i t a t i o n t o the use  of  fawns i s t h a t t h e y are not d i r e c t l y range dependent d u r i n g f i r s t month o f l i f e and  do not become e n t i r e l y independent o f  doe's m i l k u n t i l f o u r o r f i v e months o f age. the l a c t a t i o n a l performance o f the dam i n t a k e , as was  the  However, s i n c e  i s i n f l u e n c e d by  shown by W a l l a c e (194$) f o r sheep, and  fawns are known t o r e s o r t t o f o r a g e  energy  since  t o supplement a d i m i n i s h i n g  5  s u p p l y o f m i l k e a r l i e r t h a n s i x weeks o f age  (Cowan, i n  T a y l o r e t a l , 1956), the growth o f the fawn s h o u l d s t i l l  be  s e n s i t i v e t o range c o n d i t i o n s . The  use o f an a n i m a l  i n d e x method o f range e v a l u a t i o n  p r o m i s e s t o have a p p l i c a t i o n i n the e x i s t i n g s i t u a t i o n s on Vancouver I s l a n d deer r a n g e s .  B l a c k t a i l deer t h r i v e i n the  e a r l y stages of e c o l o g i c a l succession f o l l o w i n g burning  or  l o g g i n g o f the f o r e s t c o v e r i n the c o n i f e r o u s f o r e s t s o f the west c o a s t  (Cowan, 1945;  R o b i n s o n , 1957;  G a t e s , 1968).  A c c o r d i n g t o S m i t h (1968) the peak v a l u e o f t h i s range f o r d e e r i s r e a c h e d f o u r t o e i g h t y e a r s . a f t e r l o g g i n g and/or burning.  T h i s i s f o l l o w e d by a g r a d u a l d e c r e a s e i n t h e  number o f deer u s i n g t h e a r e a .  Patch logging p r a c t i c e s  c r e a t e an i n t e r m i x t u r e o f s e r e s , each o f d i f f e r e n t v a l u e deer.  The  to  s p a t i a l i n t e r a c t i o n s together with differences  o f e l e v a t i o n and exposure must c r e a t e a v a s t a r r a y o f d i f f e r e n t e n e r g e t i c environments.  Because t h e  population  i s g e n e r a l l y d i s p e r s e d , not a l l i n d i v i d u a l s can have  access  t o t h e same p o r t i o n o f t h e r a n g e , and i t would be d e s i r a b l e t o s e l e c t a scheme t h a t would a l l o w each a n i m a l t o  provide  an independent e s t i m a t e  derived  from.that  o f the energy w h i c h i t had  p a r t o f the range t o w h i c h i t was  exposed.  ,6 Methods:  I.  R e a r i n g the E x p e r i m e n t a l  Animals.  A l l o f t h e fawns used i n t h i s experiment  were c a p t u r e d  on the s t u d y a r e a a t W o l f Lake near C o u r t e n a y , o f one t o f i f t e e n days.  B.C.  They were r e a r e d i n the  a t an  age  wildlife  u n i t on t h e U n i v e r s i t y o f B r i t i s h Columbia campus u s i n g management t e c h n i q u e s w h i c h have become s t a n d a r d  procedures  i n t h i s l a b o r a t o r y (Wood e t a l , 1961). Fawns were housed u n t i l t h e y were weaned i n 2' x 4* plywood pens w i t h sawdust b e d d i n g .  At t h i s time t h e y were  moved t o 4* x 10* pens equipped w i t h s l a t t e d wooden f l o o r s . The  s l a t s were 2" x 3" s t o c k spaced o n e - h a l f i n c h a p a r t w i t h  the edge g r a i n on the upper s u r f a c e .  U r i n e and f e c a l m a t e r i a l  passed between t h e s l a t s a l l o w i n g the a n i m a l s t o remain and  clean  dry. E a r l y f e e d i n g was  b o t t l e equipped  with a  done w i t h a 1000 1  c c . narrow-necked  lamb n i p p l e * i n which a l a r g e h o l e  (lmm.) had been burned.  The  q u a n t i t y of milk taken  was  measured as the d i f f e r e n c e i n weight o f the b o t t l e p l u s r a t i o n b e f o r e and a f t e r f e e d i n g . a p p r o x i m a t e l y 35 C was m i l k was  found t h a t warm m i l k a t  t a k e n most r e a d i l y by t h e fawns so a l l  heated b e f o r e f e e d i n g i n a t h e r m o s t a t i c a l l y  c o n t r o l l e d water The  I t was  bath.  fawns were f e d a c c o r d i n g t o body weight  t h a t had been d e r i v e d from p r e v i o u s e x p e r i e n c e .  to a standard The  standard  was  s l i g h t l y l o w e r t h a n t h e c a l c u l a t e d maximum f e e d i n t a k e f o r  t h e fawns, b u t a t t h i s l e v e l r e g u l a r c l e a n - u p s were common, the deer grew r a p i d l y , and t h e i n c i d e n c e o f severe s c o u r i n g was  low. The f e e d i n g s t a n d a r d i s l i s t e d i n T a b l e l a . At  each f e e d i n g , t h e fawns were o f f e r e d o n e - q u a r t e r o f t h e d a i l y f e e d i n t a k e p r e s c r i b e d by t h e s t a n d a r d f o r t h e body weight t h a t day.  Leche ( 1 9 6 4 ) emphasized  t h e need f o r r e g u l a t i n g  the amount o f f e e d o f f e r e d a t each f e e d i n g t o a c h i e v e maximum intakes.  A young a n i m a l l e f t t o i t s own d e v i c e s t e n d s t o  gorge a t one f e e d i n g and skimp on t h e next l e a d i n g t o a depression o f the t o t a l feed i n t a k e .  F u r t h e r m o r e , t h e uncon-  t r o l l e d f e e d i n g tends t o f a v o r s c o u r i n g .  H.C. Nordan  (viva  v o c e ) m a i n t a i n s t h a t much o f t h e problem o f s c o u r i n g i s caused by u n d i g e s t e d m a t e r i a l i n t h e l o w e r d i g e s t i v e t r a c t a s u b s t r a t e f o r i n t e s t i n a l microorganisms.  providing  The r e s u l t s o f  Walker and F a i c h n e y ( 1 9 6 4 ) s u p p o r t t h i s view.  Even s p a c i n g  o f t h e f e e d i n g s o v e r 2 4 hours was found t o be l e s s  effective  t h a n t h e p a t t e r n o f 6 a.m., 1 1 : 3 0 a.m., 4 p.m., and 9-"30 p.m. The uneven s p a c i n g a l l o w e d one l o n g p e r i o d o f u n i n t e r u p t e d r e s t f o r both t e c h n i c i a n s and fawns. Weaning commenced a t t e n pounds. .A p e l l e t e d c o n c e n t r a t e r a t i o n , U.B.C. 3 6 - S - 6 3 , was made a v a i l a b l e t o t h e fawns so t h a t t h e y might become f a m i l i a r w i t h t h e presence o f t h e f e e d . S m a l l amounts, l e s s t h a n t e n grams, o f l e a f y a l f a l f a hay were s p r i n k l e d over t h e r a t i o n each day as an a d d i t i o n a l to take the p e l l e t e d r a t i o n .  incentive  The hay was t a k e n by most  8  T a b l e 1.  The pre-weaning and weaning d a i l y a l l o t m e n t s o f m i l k  Table 1 a)  Daily Total  Weight 4  Lb.  The pre-weaning f e e d i n g s t a n d a r d f o r f u l l  850 C a l .  P e r F e e d i ng (x4) 1  213 C a l .  1  Per Feeding (x4)  Daily Total  Weight  Lb. 1175 C a.l  9  294 C a l .  1  880  220  5  920  230  10  5.5  950  238  10.5  6  980  245  11  6.5  1010  253  7  1040  7.5  1200  300  1235  309  1265  316  1300  325  11.5  1330  333  260  12  1360  340  1075  269  12.5  1400  350  8  1110  278  13  1430  358  8.5  1140  285  13.5  1450  363  T a b l e 1 b)  Day  9-5  feeding  •  The weaning s c h e d u l e s t a r t i n g weight  Energy A l l o t m e n t Per Day  a t 13 pounds body  D a i l y Regime ( C a l o r i e s / F e e d i n g x Feedings/Day  1- 5  800 C a l o r i e s  200 x 4  or  267 x 3  6- 7  656  I64  x 4  or  211 x 3  8-11  440  220 x 2  300  150 x 2  12+  1  C a l o r i e s o f Apparent D i g e s t i b l e Energy For weight o f condensed m i l k ( 1 : 1 d i l u t i o n ) , by  1.441-  For weight o f m i l k r e p l a c e r ( 1 : 5 - 2 5 m u l t i p l y by 1 . 4 7 9 -  multiply  dilution),  1  9 a n i m a l s from t h e f i r s t day t h a t i t was  presented.  At t h i r t e e n  pounds, t h e m i l k a l l o w a n c e was reduced t o encourage to  seek t h e d r y r a t i o n .  the fawns  I t was found b e s t t o make the  first  r e d u c t i o n o f m i l k d r a s t i c w i t h s m a l l e r r e d u c t i o n s over the r e s t o f the two week weaning p e r i o d .  The g e n e r a l weaning  regime, based on a f r a c t i o n o f f u l l f e e d a t t h i r t e e n pounds, i s shown i n T a b l e l b .  I t s h o u l d be made c l e a r t h a t t h e r e i s a  g r e a t i n d i v i d u a l v a r i a t i o n i n t h e w i l l i n g n e s s t o wean and c a r e must be t a k e n not t o weaken t h o s e a n i m a l s t h a t r e q u i r e extended p e r i o d o f weaning.  an  On o c c a s i o n s , i t has been  necessary to realimentate t e m p o r a r i l y i n order to restore a fawn t o a h e a l t h y c o n d i t i o n b e f o r e weaning c o u l d p r o c e e d . G e n e r a l l y , any d e b i l i t a t i o n can be a v o i d e d i f time i s t a k e n t o encourage  t h e fawns t o eat t h e d r y r a t i o n by o f f e r i n g by hand  a few p e l l e t s a t a t i m e .  The hungry fawn w i l l r e a d i l y t a k e  any form o f food t h a t i s p l a c e d i n i t s mouth a t t h i s t i m e . Adherence t o t h i s p r o c e d u r e markedly  s h o r t e n s t h e weaning  period. Body w e i g h t s were t a k e n each day d u r i n g t h e e a r l y f e e d i n g p e r i o d and l e s s f r e q u e n t l y d u r i n g l a t e r phases o f the e x p e r i ments.  The fawns were weighed t o the n e a r e s t o n e - q u a r t e r  pound on a p l a t f o r m s c a l e ( F a i r b a n k s Morse, Model 5264). proved c o n v e n i e n t t o weigh t h e a n i m a l s b e f o r e t h e f e e d i n g each day.  It  morning  T h i s a l l o w e d the r a t i o n a l l o t m e n t t o be  d e t e r m i n e d , and t h e a c t i v i t y d u r i n g the w e i g h i n g  procedure  a r o u s e d the fawns making them more eager at t h e f i r s t f e e d i n g .  For n e a r l y e v e r y fawn t h e r e were o c c a s i o n s o f some s o r t was r e q u i r e d .  Of most f r e q u e n t  i n t e s t i n a l i n f e c t i o n which caused s c o u r i n g .  when m e d i c a t i o n  o c c u r r e n c e was an T h i s was seldom o f  a s e r i o u s n a t u r e and c o u l d u s u a l l y be c o n t r o l l e d by a s i n g l e o r a l dose o f 20 mg. o f c h l o r t e t r a c y c l i n e (Aureomycin, Cyanamid). I f n e c e s s a r y , a second dose was g i v e n t h e f o l l o w i n g day. I n s t u b b o r n c a s e s where p h y s i c a l c o n d i t i o n was b e g i n n i n g t o d e t e r i o r a t e , an i n t r a m u s c u l a r  i n j e c t i o n of Procain  Penicillin  G, P e n i c i l l i n G P o t a s s i u m , and S t r e p t o m y c i n S u l p h a t e (PenS t r e p , S q u i b b ) was g i v e n i n a 2cc. dose on two days.  consecutive  T h i s d r u g has proven an e f f e c t i v e p r o p h y l a c t i c  pneumonia t h a t o f t e n f o l l o w s weakness from s e v e r e  against  scouring.  A l t h o u g h e x t e r n a l p a r a s i t e s were n o t a problem, t h e p r e c a u t i o n was t a k e n t o dust each fawn o c c a s i o n a l l y w i t h a p e s t i c i d e (Methoxychlor (10%), L a t e r Chemicals L t d . ) .  II.  Rations.  Milk: Deer 'fawns i n 1963 and 1964 were f e d a commercial c a l f r a t i o n , Peebles V - l e r . evaporated milk  From 1965 t h r o u g h 1967, P a c i f i c brand  (Fraser V a l l e y M i l k Producers A s s o c i a t i o n ,  Vancouver, B.C.) was used when t h e former brand o f m i l k r e p l a c e r became u n a v a i l a b l e .  E v a p o r a t e d m i l k had been t h e  fawn r a t i o n used at- t h i s l a b o r a t o r y u n t i l t h e summer o f 1963. The  c a l f r a t i o n was s u b s t i t u t e d a t t h i s p o i n t i n an attempt t o  11 improve t h e growth r a t e o f t h e fawns.  K i t t s , e t a l (1956) had  demonstrated t h a t deer m i l k c o n t a i n e d more o f i t s energy i n the form o f f a t t h a n d i d cow's m i l k .  No improvement was noted  w i t h t h e h i g h f a t c a l f r a t i o n and t h e more c o n v e n i e n t evapor a t e d m i l k was used from 1965 on. F o r t h e purposes o f f e e d i n g , t h e ' V - l e r ' was mixed  with  w a t e r i n a r a t i o o f 1 p a r t d r y r a t i o n t o 5 . 2 5 p a r t s o f water by weight (16$ d r y m a t t e r ) .  The e v a p o r a t e d m i l k was d i l u t e d  1:1 by volume w i t h water t o t h e form o f r e c o n s t i t u t e d cow's milk.  A . J . Wood ( v i v a voce) recommended t h a t a minimum o f 1.1  grams o f water p e r a p p a r e n t d i g e s t i b l e C a l o r i e o f energy be a l l o w e d when t h e o n l y a v a i l a b l e w a t e r was i n t h e m i l k .  Both  r a t i o n s as mixed o f f e r e d i n e x c e s s o f 1.2 grams o f w a t e r p e r Calorie.  The apparent d i g e s t i b l e e n e r g i e s o f t h e r a t i o n s as  f e d were:  P e e b l e ' s V - l e r (1:5.25, 16% d r y m a t t e r )  0.676 Cal./gm.  P a c i f i c evaporated milk  0.694 Cal./gm.  (1:1 d i l u t i o n )  The e v a p o r a t e d m i l k was supplemented w i t h m i n e r a l s t o meet t h e recommendations  of 0 Keefe T  (1957).  o f t h e f o l l o w i n g mix was g i v e n p e r day:  One  millilitre  Mineral Mix magnesium c h l o r i d e  50.0  gm.  ferric  citrate  13-0  gm.  cupric  sulphate  0.6  gm.  6 3 . 6 gm. d i s s o l v e d i n 1 0 0 0 ml. aqueous  solution.  This l e v e l o f supplementation  prevented  t h e development o f  anemia o f t e n a s s o c i a t e d w i t h f e e d i n g h i g h l e v e l s o f cow's m i l k (Underwood, 1 9 6 2 ) . A d e p r e s s i o n o f growth r a t e o f some fawns i n 1 9 6 5 (not animals  from t h i s e x p e r i m e n t ) showed a marked response t o  Vitamin A supplementation. halibut  T h e r e a f t e r , 1 ml. o f d e o d o r i z e d  o i l containing approximately  A was added t o t h e m i l k each day.  100,000  i . u . of Vitamin  There was no e v i d e n c e o f  anemia o r V i t a m i n A d e f i c i e n c y when u s i n g t h e m i l k r e p l a c e r i n 1 9 6 3 and 1 9 6 4 .  D.A. Leckenby ( v i v a voce) found adequate  l i v e r s t o r a g e o f V i t a m i n A i n fawns r a i s e d  Weaner r a t i o n  on t h i s r a t i o n .  (U.B.C. 3 6 - S - 6 3 ) :  T h i s r a t i o n was f o r m u l a t e d t o p r o v i d e h i g h l e v e l s o f a l l n u t r i e n t s i n an e a s i l y d i g e s t i b l e r a t i o n f o r a monogastric as r e a d i l y a v a i l a b l e  form.  I t i s essentially a  a n i m a l c o n t a i n i n g most o f i t s energy  carbohydrates.  I t c o n t a i n s a supplemen-  t a t i o n o f B v i t a m i n s which have been demonstrated t o be  unnecessary  i n a d u l t a n i m a l s w i t h an a c t i v e  but e s s e n t i a l t o the weaning ruminant weaner r a t i o n was months o f age.  rumen  ( M o r r i s o n , 1956).  f e d t o fawns u n t i l t h e y had reached  The  i n Table 4.  three  requirements  sheep (U.S. N a t i o n a l Academy o f S c i e n c e s , The  d i g e s t i b i l i t y o f t h i s r a t i o n was  average v a l u e o f 75.0% Calories  The  formulation of t h i s r a t i o n i s l i s t e d i n  Table 2 and i t s c o m p o s i t i o n i s compared w i t h the f o r a growing  microflora,  w i t h a d i g e s t i b l e energy  1963)  taken at i t s  of  3.104  per gram o f f e e d as f e d ( 8 % m o i s t u r e ) .  Adult r a t i o n  (U.B.C.  36-57):  From e a r l y September on, fawns were f e d the r a t i o n used f o r a l l a d u l t deer i n the u n i t . p r o t e i n was  In t h i s r a t i o n , l e s s of the  added as f i s h meal, and c o r n meal p r o v i d e d the  major energy  source.  B v i t a m i n s were not added.  The  formu-  l a t i o n o f t h i s r a t i o n i s l i s t e d i n T a b l e 3 and i t s n u t r i e n t c o m p o s i t i o n i s compared w i t h the r e q u i r e m e n t s sheep i n Table 4.  The  d i g e s t i b i l i t y of t h i s r a t i o n  t a k e n at i t s average v a l u e o f 66.0% o f 2.746 C a l o r i e s  for a  with a digestible  growing was energy  per gram o f f e e d as f e d {9% m o i s t u r e ) .  R e g a r d l e s s o f the r a t i o n b e i n g f e d , each a n i m a l showed a good growth response of c o n d i t i o n ,  was  and i t s h a i r c o a t , a good  excellent.  The  a n i m a l s b e i n g f e d U.B.C. 36-57 was adequacy o f t h a t r a t i o n  criterion  a n t l e r growth i n a d u l t normal, and a t t e s t s t o the  ( F r e n c h e_t a l , 1956).  T a b l e 2.  The f o r m u l a t i o n o f weaner r a t i o n , U.B.C. 36-S-63  Ingredient  Amount  Ground barley-  200 pounds  Oat g r o a t s  390  Wheat b r a n  130  H e r r i n g meal  200  Soya meal  100  Skim m i l k  200  Dried  150  grass  D i c a l c i u m phosphate  10  Iodized s a l t  10  Brewer's y e a s t  20  Chromic  oxide  Irradiated Vitamin A  yeast  1 2 2,000,000 u n i t s  2,000 pounds  Table 3 .  The f o r m u l a t i o n o f a d u l t r a t i o n , U.B.C.  Ingredient  Amount  Corn meal  600  Ground wheat  250  Bran  275  Molasses  150  Beet p u l p  200  V i t a gras  200  Soya bean meal  175  H e r r i n g meal  110  Bone meal  20  Iodized s a l t  20  Chromic o x i d e  1  2,000  pounds  36-57  16  Table 4 .  N u t r i e n t c o m p o s i t i o n o f r a t i o n s , U.B.C. 3 6 - S - 6 3 and U.B.C. 36-57 compared w i t h t h e r e q u i r e m e n t s f o r g r o w i n g sheep ( U n i t e d S t a t e s N a t i o n a l Academy o f S c i e n c e s , 1963 ).  N.A.S. Requirements'  U.B.C.  U.B.C.  36-S-63  36-57  Nutrient  Units  Digestible protein^  mgm/Cal.3  Ca  mgm/Cal.  0.97  1.37  1.01  P  mgm/Cal.  0.87  1.92  1.63  Vitamin A  1 U/Cal.  I.84  Vitamin D  1 U/Cal.  0.5  36.3  46.2  32.0  +4 +4  1  C a l c u l a t e d from t o t a l d a i l y r e q u i r e m e n t s f o r a 60 pound lamb.  2  Crude p r o t e i n x 60%.  3  C a l o r i e s o f .apparent d i g e s t i b l e energy.  4  T o t a l c o n t r i b u t i o n s from a l l r a t i o n i n g r e d i e n t s n o t known, but a p a r t i a l t o t a l exceeds t h e N.A.S. requirements.  17 I I I . Determination of Ration D i g e s t i b i l i t y , Apparent D i g e s t i b l e Apparent rations:  M o i s t u r e , and  Energy.  d i g e s t i b l e energy c o n t e n t s o f t h r e e o f t h e  V - l e r , U.B.C. 3 6 - S - 6 3 ,  mined by d i g e s t i b i l i t y t r i a l s .  and U.B.C. 36-57 were d e t e r For the fourth r a t i o n ,  e v a p o r a t e d m i l k , t h e d i g e s t i b i l i t y and apparent  Pacific  digestible  energy were t a k e n from p r e v i o u s d i g e s t i b i l i t y t r i a l s w i t h deer fawns (O'Keefe,  1957).  M o i s t u r e c o n t e n t was measured f o r t h e two p e l l e t e d r a t i o n s a c c o r d i n g t o t h e recommendations o f H o r w i t z ( 1 9 5 5 ) . They were d r i e d f o r t h r e e hours a t 125 C and t h e n f o r one hour periods t o constant weight.  I t was n e c e s s a r y t o use a m o d i f i e d  method f o r t h e V - l e r s i n c e t e m p e r a t u r e s o v e r 90 C caused scorching o f the r a t i o n .  I t was d r i e d a t 85 C f o r 16 hours  and t h e n f o r f o u r hour p e r i o d s t o c o n s t a n t w e i g h t .  The  e v a p o r a t e d m i l k c o u l d n o t be d r i e d a d e q u a t e l y because a vacuum d r y i n g oven was n o t a v a i l a b l e . Digestibility  was e s t i m a t e d f o r each r a t i o n by t h e  r e c o v e r y o f a n o n - d i g e s t i b l e chromagen ( S c h u r c h e t a l , 1950). Chromic  o x i d e ( C ^ O ^ ) was added t o t h e r a t i o n a t t h e f o l l o w i n g  levels:  The  V-ler  0.5% o f a i r d r y w e i g h t ,  U.B.C. 36-S-63  0.05/o o f a i r d r y w e i g h t ,  U.B.C. 36-57  0.05% o f a i r d r y w e i g h t .  chromic o x i d e c o n c e n t r a t i o n s i n f e e d s and f e c e s were  18 determined  by the method o f C z a r r o c k e t a l (1961).  Chromic o x i d e was  mixed i n t o enough dry V - l e r t o l a s t  f o r t h e d u r a t i o n o f a t e n day t r i a l u s i n g f o u r fawns r e c e i v i n g a r e s t r i c t e d amount o f f e e d .  Only t h e s e f o u r were a v a i l a b l e  f o r t h i s a s p e c t o f the s t u d y .  The  f u l l y f e d fawns were  a l r e a d y b e g i n n i n g the weaning p r o c e s s b e f o r e p r e p a r a t i o n s c o u l d be made t o conduct  the d i g e s t i b i l i t y  study.  p o s s i b l e t h a t the r e s u l t s so o b t a i n e d may  It is  have o v e r e s t i m a t e d  the d i g e s t i b i l i t y o f t h e m i l k when f e d ad l i b i t u m .  Increased  d i g e s t i b i l i t y o f roughages when f e d i n l i m i t e d q u a n t i t i e s been demonstrated (1961).  by B l a x t e r e t a l (1955, 1956)  and by R e i d  Most o f t h i s d i f f e r e n c e can be accounted  change i n t h e time o f passage.  has  f o r by a  I t seems r e a s o n a b l e t o s u s p e c t  t h a t f o r a h i g h l y d i g e s t i b l e r a t i o n such as m i l k , t h e t i m e o f passage would not be a l i m i t i n g f a c t o r i n d i g e s t i b i l i t y . Leche (1964) found no i n d i c a t i o n o f d i f f e r e n c e s i n the d i g e s t i b i l i t y of milk fed at d i f f e r e n t l e v e l s to c a l v e s . d i g e s t i b i l i t y f a c t o r determined  The  i n the p r e s e n t s t u d y has been  a p p l i e d t o b o t h r e s t r i c t e d and u n r e s t r i c t e d fawns on the assumption The  t h a t any d i f f e r e n c e s i^ould be  chromic  o x i d e marked r a t i o n was  as the normal r a t i o n . second  small. f e d i n t h e same manner  F e c a l samples were c o l l e c t e d f o r the  f i v e days o f a t e n day t r i a l and t h o s e from a l l f o u r  a n i m a l s were lumped t o g e t h e r f o r a n a l y s i s .  Representative  samples o f r a t i o n were c o l l e c t e d from the f e e d each day t o g i v e an average f i g u r e f o r t h e c o n c e n t r a t i o n o f chromic  oxide  19 i n the feed. The  marker was added t o t h e commercial p e l l e t e d  during t h e i r preparation.  rations  E a r l i e r t r i a l s w i t h these  rations  had demonstrated t h a t t h e marker was p o o r l y mixed and t h e concentration varied  from bag t o bag and even i n two samples  from t h e same c o n t a i n e r .  I n order t o minimize  this error,  t e n grams o f f e e d were c o l l e c t e d from t h e f e e d pan o f each a n i m a l a f t e r t h e d a i l y a l l o c a t i o n s had been made.  These d a i l y  f e e d samples were accumulated t o g i v e an average chromagen l e v e l f o r the determination o f the d i g e s t i b i l i t y o f the r a t i o n by each a n i m a l . I n o r d e r t h a t apparent r a t i o n c o u l d be c a l c u l a t e d , a l l f e e d and f e c a l samples.  d i g e s t i b l e energy c o n t e n t o f each g r o s s e n e r g i e s were d e t e r m i n e d f o r These g r o s s energy v a l u e s were  combined w i t h d i g e s t i b i l i t y d a t a t o y i e l d apparent  digestible  energy: A.D.E. = G.E.  feed  '- G.E.  feces  (1 - c o e f f .  To f a c i l i t a t e t h e c o n v e r s i o n t o apparent i n t a k e from weight  of d i g e s t i b i l i t y ) ,  digestible  energy  o f f e e d consumed, t h e d i g e s t i b l e energy  c o n t e n t o f each r a t i o n was c o r r e c t e d t o t h e average  moisture  l e v e l o f t h e f e e d as i t was f e d .  These v a l u e s have a l r e a d y  been mentioned i n t h e d e s c r i p t i o n  of the r a t i o n s .  t  20  IV.  Experimental Treatments. The e x p e r i m e n t a l p e r i o d r a n from t h e time t h a t t h e fawns  r e a c h e d 10 pounds u n t i l 322 days l a t e r .  T h i s time p e r i o d  was  d i v i d e d i n t o 5 i n t e r v a l s o f 4 9 , 63, 63, £4 and 63 days d u r a t i o n respectively.  The  first  i n t e r v a l o f 4 9 days, s t a r t i n g a t 10  pounds body w e i g h t , can be c o n s i d e r e d t o be a 63 day from b i r t h , assuming o f 1/3  pound per day.  interval  a r a t e o f weight g a i n s l i g h t l y i n e x c e s s T h i s e s t i m a t e was  based on t h e  growth  r a t e s o f t h r e e s e t s o f t w i n s (Nos. 101, 102; 3 5 0 , 351;  352,  353) born t h c a p t i v e does i n t h e w i l d l i f e u n i t . F o r t y male fawns and s i x t e e n female fawns were r a i s e d f o r the purposes o f t h i s s t u d y .  These a n i m a l s were a r r a n g e d  groups so t h a t the n u t r i t i o n a l t r e a t m e n t i n each o f t h e  into first  t h r e e i n t e r v a l s c o u l d be one o f t h r e e d i f f e r e n t l e v e l s o f energy 1.  intake: h i g h p l a n e (H)  -- ad l i b i t u m i n t a k e o f a h i g h energy  2.  ration;  medium p l a n e (M) -- a r e s t r i c t i o n t o 8 5 % o f h i g h p l a n e a t any body w e i g h t ;  3.  low p l a n e (L)  -- a r e s t r i c t i o n t o 70% o f h i g h p l a n e a t any body w e i g h t .  A l l a n i m a l s c a r r i e d t o an age g r e a t e r t h a n 175 days were s u b j e c t e d t o a r e s t r i c t i o n t o low p l a n e d u r i n g the next 84  day  interval.  T h i s p e r i o d s e r v e d as a s t a n d a r d i z e d moderate  w i n t e r regime.  Only two n u t r i t i o n a l t r e a t m e n t s  during the l a s t ( f i f t h ) i n t e r v a l :  f  H  f  and ' L . 1  were  considered  The o b j e c t i v e s  d u r i n g t h i s p e r i o d were f i r s t l y , t o d e t e r m i n e i f growth d i f f e r e n c e s i n t h e s k e l e t o n c o u l d be i n d u c e d r e s t r i c t i o n s a t t h i s age, and s e c o n d l y , s k e l e t a l d i f f e r e n c e s induced  by energy  t o determine i f  by e a r l i e r r e s t r i c t i o n s c o u l d be  removed by compensatory growth. The males were a r r a n g e d  i n s i x t e e n regimes  comprising  e i g h t n u t r i t i o n a l sequences (see F i g u r e 1 ) . A minimum o f two fawns were k i l l e d a t t h e time o f each n u t r i t i o n a l change and a t t h e end o f each sequence. i n t o f i v e treatments two  The female fawns were d i v i d e d  and two n u t r i t i o n a l sequences.  These  sequences were i d e n t i c a l t o two o f t h e male sequences, and  were i n c l u d e d t o measure s e x d i f f e r e n c e s i n t h e r e s p o n s e o f the s k e l e t o n t o d i e t a r y r e s t r i c t i o n s . Although treatments  the animals  were f e d a c c o r d i n g t o t h e proposed  as c l o s e l y as p o s s i b l e , i n d i v i d u a l v a r i a t i o n s made  i t necessary  t o r e c l a s s i f y some a n i m a l s  a t t h e end o f t h e  experiment.  T h i s w i l l be d i s c u s s e d l a t e r i n t h e R e s u l t s  section.  V.  C o l l e c t i o n of Field  Specimens.  W i l d fawns were c o l l e c t e d from t h e V/olf Lake a r e a on Vancouver I s l a n d a t d a t e s c o r r e s p o n d i n g f o r t h e l a b o r a t o r y r e a r e d fawns.  to the slaughter dates  Fawns o f e i t h e r s e x were  22  Figure 1  The Proposed Energy Regimes F o r Male (Females D u p l i c a t e HHH' 1  Fawns  and «LLL»).  ;EE|E.:!:::; '".'.V~'.V.'.'.'\ t':'E" :~i'r.: -  11  : ::i:,:: f  f i l l I  l f t i l F  ;  -:. E':.;: E . . E E ; E E E : E  •  lEEiEE 'EEiEE  E . E E E .j.i-li-J li:.:  EEI.EE:  :.::!.:::  :.-:: 1  isiOil  DIM i n i ...  6  EE  3 Ibj  (  1  1  ^  :: E E E  ;  .—,  :E|::::  •rE-jrEr E-EE'-E-  TE  i::: -  l (  ::-JEEE  • i i i  :":::.[.•:•.:.:  <  -.:•:•:[:::;  . . . : _ L _ -7  TEEJ::E : EEJE;E  :.:E  : E  E  E  E. EE:  T T E I E E ; E . : : | E E  ::.:~|.:.v: :::::::. .... 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TE :  EEEEE EEEE  EN  EE  TT'Tir  TTE  TTU:1::_:::  EE  EE  1- : • •  ETIETT:  :  EEE^EE E H ' E r i EE :7:T"h.;:F ......)........  9 ^ Q  ~-|TT:—  T T T T I T : —  .TT:T .|T-;:  :  TTTT.JTTTr.  . . : | : E T  :::: r.:::  .::  TTTTJT::TT.  E  E Ei'EE' —  TTT^(::TT::  | ' - - EEITEE;  T;E  ;L  1 1 H i  Hi-EEEE  ll y.  :T:TT  ••--,)  EEirEE ----j-:-!— T : E I E . : T E E ! . : : r; i: :r: EE-1.EE  _  TTTTfTTrT ITT H :  •E T T T - JTTTT;  EE  i.n.-.T  hi  TTlTpTTT ::.:.:q:TT-.  ;E EE  :  .r-tj-_j.  E . i'E :Ei:  if  \  :~:T.  :  ••:'!;••::  ....  :T:  EElEE  . E E I E E  T'TE  :  -  E : J ! T E ' EEEpE- E T : t E E E E EE- \~-:~. EE : :': IETIETT E";; l" - irpr" • EET E:  E E - J E T T .  E r r j E E .  . -  ::.:.: h:.":-:  '"•"1" ::::i:-::  S  •EEEE  T" '"TT": E E i E E j  : : • : I:.: :  - k 1  :.-::T|T:TT  •.]:•••• •  EE  EEr jrir.:  »'.- . ' :  T E E T r E  ~:IET  : : E  EE  ::!»1 EETJEE EE-EE E E E T : : • • I T - . T : ;EE  b F  TTE  : : E ! T : E  E  ;  I:::::T  T.,"T:  EEJEE  EEjEE  ::.:: j : . ; : :  :  :.-:  -EEjET.:  EEIEE  ~.:rr:\::.:.:  :  1  :  Q i7  _  %I  ::  .....  rE^Ef  E  ET-  ; •.-/:-  .-  ... i  E;E E ; EEEE i E .•EE;;;-  E.::L:::J  E< • EpE  : :::TITT  l E E ^ E EEjEE- . E T : J T E ~  EE  :. •  :;::).:.:  •:.: T - T T !  : • \ •  EEILE:  ::.._i.Ei.: :E':i EE  !EEJ:E EE'E'EE  • E E E E !; E j E •'.  :  ! . ,T  EEjEE E E  EiEIEEi  TTT-.f -:-:  •: •: i.:. •  r: ::--'•:::( ::::•:::•:]  :  t a k e n when a v a i l a b l e . be c o l l e c t e d in  1965,  I t was hoped t h a t enough fawns c o u l d  each y e a r t o a l l o w comparison o f the growth  1966 and 1967-  rates  However, t h e u n a v a i l a b i l i t y o f fawns  at some c r i t i c a l p e r i o d s has made t h i s d i f f i c u l t .  A compari-  son o f f i e l d and l a b o r a t o r y r e a r e d fawns had t o be made w i t h a pool of three years data. i s g i v e n i n T a b l e 5« collected  A l i s t of the animals  collected  A l a r g e number o f f o r e cannons were  from h u n t e r k i l l e d deer i n November o f each y e a r t o  a l l o w a more e x t e n s i v e a n a l y s i s o f t h e v a r i a b i l i t y i n t h e d i m e n s i o n s o f a s k e l e t a l component i n t h e p o p u l a t i o n . Where p o s s i b l e , t h e s e were used t o augment t h e i n f o r m a t i o n c o l l e c t e d from t o t a l s k e l e t o n s . g i v e n i n Table  A l i s t o f t h e cannon bones c o l l e c t e d i s  6.  VI.  Measurement o f Growth.  1.  Body w e i g h t : Body weight change i s the most commonly e n c o u n t e r e d  measure o f growth and has been g i v e n f i r s t c o n s i d e r a t i o n i n t h i s study.  A l t h o u g h body weight g i v e s no i n d i c a t i o n  o f the  s p a t i a l d i s t r i b u t i o n o f the growth, i t i s n e c e s s a r i l y a s s o c i a t e d w i t h a change i n s i z e .  Because i t can be measured  f r e q u e n t l y from each a n i m a l , weight d a t a are more abundant t h a n s k e l e t a l data.  Each a n i m a l c o n t i n u e s t o c o n t r i b u t e w e i g h t d a t a  u n t i l the time o f i t s d e a t h , w h i l e measurements o f  skeletal  d i m e n s i o n s are t e r m i n a l o n l y . Weights o f l a b o r a t o r y a n i m a l s were t a k e n d a i l y a t younger  Table 5 .  E s t i m a t e d c u m u l a t i v e energy i n t a k e s o f w i l d fawns based on body weight Body Weight (eviscerated)  Body Weight (estimated l i v e )  2  T o t a l Energy Intake  Sex  Age  YF17  M  116 Days  YF19  M  118  YF16  F  116  38.9  l b . no l i v e r  56.2 l b .  203,200 C a l o r i e s  YF18  F  117  30.6  l b . no l i v e r  44.2 l b .  159,500  YF20  F  118  36.7  l b . no l i v e r  53.0 l b .  191,500  WF 1  M  180  41.5  lb. liver i n  54.5  lb.  289,400 C a l o r i e s  WF 2  M  180  41.5  lb. liver i n  56.6  lb.  298,700  YF38  M  177  36.0 l b . no l i v e r  49-3  lb.  259,900  YF37  F  176  No.  1  39.8  l b . no l i v e r  lb.  -  -  -  57.5  . 207,900 C a l o r i e s  -  -  -  1  Age i s counted from June 24 — i . e . , t h e mean day a t w h i c h 10 pounds i s r e a c h e d when t h e mean b i r t h date i s June 10.  2  See pp. 20;  3  E s t i m a t e d from body weight a c c o r d i n g t o F i g u r e s 5 and 6 .  48-53-  T a b l e 6. No. XF 6* XF 7* XF 8* XF 9 XF10 XF11 XF12 XF13 XF14* XF15* XF16* XF17* YF21 YF22 YF23 YF24 YF36 YF40 YF41 YF42  Fore cannons c o l l e c t e d from h u n t e r k i l l e d fawn s Sex  Age  M M M M M M M M M M M M M M M M M M M M  142 142 143 150 150 150 150 150 156 156 156 157 142 142 142 142 142 162 162 162  1  No. YF43 YF44 YF45 YF46 YF47 YF48 YF49 YF50 YF51 XF 5 YF25 YF26 YF67 YF68 YF69 YF70 YF71 YF72  Sex  Age  M M M M M M M M M F F F F F F F F F  162 162 162 162 162 162 162 162 162 140 142 142 162 162 162 162 162 162  1  *From a n i m a l s o f known body w e i g h t .  NoXF 6 XF 7 XF14 XF15 XF16 XF17 XF 8  Body Weight (eviscerated) 39-0 l b . l i v e r i n 29-0 l b . l i v e r i n 34-0 l b . l i v e r i n , no head 44-0 l b . l i v e r i n 42-5 l b . l i v e r i n 38.0 l b . l i v e r i n , no head 41.0 l b . l i v e r i n  Body Weight (estimated l i v e )  ?  52.7 l b .  38.6 l b .  48.7 l b . 59.7 l b . 57-6 l b . 54-3 l b .  55-5 l b .  1  Age i s counted from June 24 -- i . e . , t h e mean day a t which 10 pounds i s r e a c h e d when t h e mean b i r t h date i s June 10.  2  See pp. 20; 48-53-  26 ages, becoming more i n f r e q u e n t l a t e r on.  F o r fawns o v e r s i x  months o f age, w e i g h i n g a t seven t o t e n day i n t e r v a l s was adequate  t o e n a b l e changes i n body weight t o be f o l l o w e d .  In  a l l o f t h e d a t a r e p o r t e d h e r e , body weight on a g i v e n day has been e s t i m a t e d from a smooth l i n e drawn t h r o u g h a graph o f t h e a c t u a l w e i g h t s t o h e l p t o m i n i m i z e d a i l y f l u c t u a t i o n s due t o 'fill . 1  I d e a l l y , e v i s c e r a t e d w e i g h t would p r o b a b l y be a b e t t e r  measure t h a n l i v e body weight because o f t h e problem o f v a r i a t i o n s i n gut c o n t e n t .  However, t h i s i n v o l v e s t h e same  problem as t h e c o l l e c t i o n o f s k e l e t a l d a t a ; t h e measurements are  t e r m i n a l o n l y and t h e sample s i z e i s t h e r e f o r e r e d u c e d . Although l i v e  body w e i g h t i s a c o n v e n i e n t l a b o r a t o r y  measurement, i t i s e x t r e m e l y d i f f i c u l t t o o b t a i n i n t h e f i e l d . Deer weighed a t 'check s t a t i o n s ' a r e always e v i s c e r a t e d , and even a n i m a l s f r e s h l y k i l l e d  and b e f o r e t h e y a r e e v i s c e r a t e d  s u f f e r from a v a r i a b l e degree o f b l o o d l o s s .  L a b o r a t o r y fawns  were weighed p r i o r t o s l a u g h t e r and t h e n a f t e r e v i s c e r a t i o n t o p r o v i d e a method f o r c o n v e r t i n g t h e f i e l d d a t a back t o l i v e weight.  A l t h o u g h weight d a t a from h u n t e r k i l l e d  deer were not  o b t a i n e d d u r i n g t h i s s t u d y , t h e s i t u a t i o n was f o r e s e e n where t h i s might be done.  Weight d a t a were accumulated  so t h a t compensation  c o u l d be made f o r t h i s organ whether l e f t  i n o r removed.  for livers  S h r i n k a g e o f c a r c a s s e s t a k e n i n t h e f i e l d was  not c o n s i d e r e d a problem because w e i g h t s were t a k e n u s u a l l y w i t h i n one day o f k i l l  from c a r c a s s e s which had been l e f t w i t h  t h e h i d e on and t h e v i s c e r a l i n c i s i o n p u l l e d s h u t .  Laboratory  a n i m a l s t r e a t e d i n the same manner and reweighed 24 hours a f t e r s l a u g h t e r showed not more t h a n o n e - h a l f pound o f weight l o s s . When, on o c c a s i o n , i t was n e c e s s a r y t o t a k e w e i g h t s o f a n i m a l s i n the f i e l d ,  a h o u s e h o l d bathroom s c a l e was used.  The  fawns were weighed by the d i f f e r e n c e i n w e i g h t between a p e r s o n and t h a t p e r s o n h o l d i n g t h e fawn.  The o v e r a l l a c c u r a c y was  w i t h i n - 1 pound. L i v e body w e i g h t s o f l a b o r a t o r y fawns were r e g r e s s e d a g a i n s t c u m u l a t i v e energy i n t a k e i n C a l o r i e s a t 49, 112, and 322 days o f age.  175,  These ages a r e t h e p o i n t s a t which  n u t r i t i o n a l changes were made.  S u f f i c i e n t w e i g h t s were not  a v a i l a b l e a t 259 days t o e s t a b l i s h a weight t o energy i n t a k e relationship.  The r e g r e s s i o n l i n e s were used t o e s t i m a t e t h e  t o t a l energy i n t a k e o f w i l d fawns.  2.  S k e l e t a l measurements: The components o f growth t h a t a r e o f g r e a t e s t i m p o r t a n c e  i n t h i s s t u d y are t h e changes i n s k e l e t a l d i m e n s i o n s .  The  c h o i c e o f t h e p a r t i c u l a r bones and t h e s e v e r a l d i m e n s i o n s used r e q u i r e s some e l a b o r a t i o n .  The w o r k i n g h y p o t h e s i s i s t h a t  energy i n t a k e w i l l i n f l u e n c e bone growth a l o n g a x i a l gradients.  growth  I t i s g e n e r a l l y conceded t h a t growth i n l e n g t h  p r e c e d e s growth i n w i d t h i n t h e l o n g bones (Hammond, 1932; P a l s s o n , 1955; P r a t t e t a l , 1964a; 1964b). over-simplification.  Walnut  T h i s may be an  (1967) s u g g e s t s t h a t growth o f a  l o n g bone can b e s t be d e s c r i b e d as a number o f v e c t o r s  28 proceeding point.  a t d i f f e r e n t v e l o c i t i e s from a s i n g l e s t a t i o n a r y  I f a l l v e c t o r s were o f t h e same r e l a t i v e  velocity,  then growth would be by m a g n i f i c a t i o n w i t h no change i n shape. T h i s i s n o t t h e case and shape does change.  The a x i a l  g r a d i e n t t h a t i s o b s e r v e d w i l l depend upon t h e p a r t i c u l a r v e c t o r b e i n g measured and t h e e q u i v a l e n c e  of t h i s  measurement  from one bone t o t h e n e x t . Bones were s e l e c t e d t h a t would b e s t demonstrate t h e i n t e r a c t i o n s o f a x i a l growth and n u t r i t i o n a l l e v e l .  The l o n g  bones o f t h e f o r e and h i n d l i m b s were an immediate c h o i c e . These bones a l l have a major a x i s t h a t i s f u n c t i o n a l l y e q u i v a l e n t f r o m one bone t o t h e n e x t .  Length was measured on each  o f t h e f o r e cannon, r a d i u s , humerus, s c a p u l a , h i n d cannon, t i b i a , femur and p e l v i s .  The s c a p u l a and p e l v i s were i n c l u d e d  because o f t h e i r a n a t o m i c a l bones on t h e l i m b s .  p o s i t i o n s r e l a t i v e t o the long  There a r e i n d i c a t i o n s i n t h e d a t a t h a t t h e  l e n g t h o f t h e s c a p u l a and t h a t o f t h e p e l v i s i n c r e a s e a t r a t e s commensurate w i t h t h e i r p o s i t i o n s i n t h e l i m b g r a d i e n t s . had  It  o r i g i n a l l y been p l a n n e d t h a t w i d t h o f a l l l i m b bones  s h o u l d be measured, b u t s a t i s f a c t o r y e q u i v a l e n c e not be f o u n d .  points  could  An e x c e p t i o n was t h e w i d t h o f t h e f o r e cannon  w h i c h c o u l d be r e p r o d u c e d s a t i s f a c t o r i l y because shape remained o r i e n t e d toward a f l a t p o s t e r i o r s u r f a c e .  Minimum w i d t h o f t h e  d i a p h y s i s and maximum w i d t h s a t t h e bases o f t h e p r o x i m a l and d i s t a l e p i p h y s e s were measured a c r o s s a plane l y i n g p a r a l l e l t o the f l a t  surface.  These measurements were i n c l u d e d m a i n l y t o  a l l o w a more e x t e n s i v e a n a l y s i s o f t h e cannon bones c o l l e c t e d from t h e h u n t e r k i l l e d  samples.  The a n t e r i o r - p o s t e r i o r a x i s o f the body e x h i b i t s a marked g r a d i e n t d u r i n g e m b r y o l o g i c a l development and p o s t n a t a l growth a p p e a r s t o f o l l o w a s i m i l a r p a t t e r n (Scammon and C a l k i n s , 1929; H u x l e y , 1932).  The v e r t e b r a e l i e d i r e c t l y a l o n g t h i s g r a d i e n t  and were used t o demonstrate growth i n t h i s d i m e n s i o n . l e n g t h and w i d t h were measured a t w e l l d e f i n e d points.  Both  equivalence  C e r v i c a l s 3, 5, 7; t h o r a c i c s 1, 5, 10; and lumbars 1,  3, and 5 were s e l e c t e d as r e p r e s e n t a t i v e o f t h e v e r t e b r a e . L e n g t h was measured from t h e e x t r e m i t i e s o f t h e zygopophyses. W i d t h was measured a c r o s s t h e t r a n s v e r s e p r o c e s s e s  o f the  c e r v i c a l and t h o r a c i c v e r t e b r a e and a c r o s s t h e a n t e r i o r ( p r e ) zygopophyses o f t h e l u m b a r s .  These measurements a r e shown i n  t h e s k e t c h e s i n Appendix 1. A l l o f t h e measurements were t a k e n from c l e a n , d r y s k e l e tons.  S e v e r a l methods were used t o c l e a n the s k e l e t o n s , a l l  o f w h i c h began w i t h d e f l e s h i n g w i t h a k n i f e .  In e a r l i e r  s t a g e s o f t h e s t u d y , some s k e l e t o n s were d r i e d and t h e n by d e r m e s t i d b e e t l e s . bones were b l e a c h e d  Upon r e m o v a l from t h e b e e t l e c o l o n y , t h e  and d r i e d .  w a t e r f o r 20 - 24 h o u r s . were p r e p a r e d  cleaned  O t h e r s were simmered i n 88-93 C  By f a r t h e m a j o r i t y o f the s k e l e t o n s  by p r e s s u r e c o o k i n g i n an a u t o c l a v e .  The  d e f l e s h e d s k e l e t o n was d i s j o i n t e d and c o v e r e d w i t h water i n deep m e t a l pans.  These were p l a c e d i n t h e a u t o c l a v e a t 15  pounds p r e s s u r e f o r 30 - 60 m i n u t e s , the o l d e s t a n i m a l s t a k i n g the l o n g e s t t i m e .  I n b o t h o f t h e l a s t two methods, t h e cooked  30  f l e s h was removed e a s i l y by hand.  D r y i n g a t 7 0 C f o r 2 4 hours  followed. One problem which a r o s e f r e q u e n t l y was the l o o s e n i n g o f epiphyses during cooking.  S e v e r a l s k e l e t o n s w h i c h had  s u r v i v e d p r o c e s s i n g i n t a c t were measured  and t h e n r e - c o o k e d  u n t i l the e p i p h y s e s f e l l o f f . These were g l u e d back on and t h e bones were remeasured.  No d i f f e r e n c e s c o u l d be d e t e c t e d  between the two s e t s o f measurements.  A l l other epiphyses  were g l u e d on and t h e s e bones were i n c l u d e d i n t h e d a t a w i t h o u t f u r t h e r mention.  V I I . Treatment o f t h e D a t a . The u l t i m a t e o b j e c t i v e was t o o b t a i n an e s t i m a t e o f t h e t o t a l energy i n t a k e o f any w i l d fawn and an i n d i c a t i o n o f the p a t t e r n o f energy r e s t r i c t i o n w h i c h may have been imposed. These e s t i m a t e s were made by i n f e r e n c e from t h e growth behavior of laboratory c o n t r o l s . growth were used.  B o t h body weight and  skeletal  Body w e i g h t and s k e l e t a l d i m e n s i o n s were  graphed a g a i n s t c u m u l a t i v e energy i n t a k e , e x p r e s s e d as a p p a r e n t d i g e s t i b l e energy i n C a l o r i e s (A.D.E.) f o r each o f t h e l a b o r a t o r y fawns. unknowns.  These p r o v i d e t h e b a s i s f o r a n a l y s i s o f w i l d  The t e c h n i q u e s w h i c h were used t o i n t e r p r e t t h e s e  graphs w i l l be e x p l a i n e d as t h e y a r e e n c o u n t e r e d i n R e s u l t s and Discussion.  Results: I.  C u m u l a t i v e Energy I n t a k e . The d a i l y energy i n t a k e o f each fawn i n t h e l a b o r a t o r y  s t u d y was c a l c u l a t e d from t h e amount o f f e e d consumed.  The  measured f e e d i n t a k e , i n grams, was m u l t i p l i e d by t h e a p p r o p r i a t e c o n v e r s i o n f a c t o r (see pp. 10-13) f o r t h e r a t i o n b e i n g f e d a t t h a t t i m e i n o r d e r t o e x p r e s s t h e i n t a k e as k i l o c a l o r i e s ( C a l o r i e s ) o f a p p a r e n t d i g e s t i b l e energy.  The  d a i l y energy i n t a k e s were summed o v e r each o f t h e e x p e r i m e n t a l i n t e r v a l s ; i . e . , 0-49, 50-112, 113-175, 176-259 and 260-322 d a y s .  T h i s a l l o w e d t h e e v a l u a t i o n o f energy i n t a k e  by i n t e r v a l and i n t o t a l .  The t o t a l energy i n t a k e s d e t e r m i n e d  i n t h i s manner p r o v i d e t h e u n i t s o f t h e x t  o f body w e i g h t and s k e l e t a l d i m e n s i o n s .  l  axes i n a l l  graphs  The c u m u l a t i v e energy  i n t a k e s f o r each a n i m a l a r e g i v e n i n T a b l e s 7 and 8. Some o f t h e fawns i n t h i s s t u d y were i n c l u d e d i n a p r e s e n t a t i o n o f p r e l i m i n a r y work ( A d d i s o n , 1966).  The  c u m u l a t i v e energy i n t a k e s shown i n T a b l e s 7 and 8 a r e a t v a r i a n c e w i t h t h o s e p r e s e n t e d e a r l i e r , b e i n g l o w e r by 1 2 % a t f o u r months and 14% a t s i x months.  Subsequent  t o the e a r l i e r  work, a c o r r e c t i o n was made i n t h e v a l u e s f o r t h e apparent d i g e s t i b l e energy o f t h e p e l l e t e d r a t i o n s w i t h b o t h b e i n g l o w e r e d by 14%. The energy v a l u e s f o r t h e r a t i o n s had been based o r i g i n a l l y on f i g u r e s from a g r i c u l t u r a l f e e d t a b l e s w h i l e t h e c o r r e c t e d f i g u r e s were d e r i v e d from a c t u a l b i l i t y t r i a l s with the deer.  The apparent d i g e s t i b l e  digestienergy  T a b l e 7.  Cumulative energy i n t a k e m a l e s , by i n t e r v a l  No.  (Calories) of laboratory  I n t e r v a l (Days) 113-175 176-259  ~~0^49  50-112  U 7  51,917 51,917  166 , 8 4 8 218 ,765  226,647 445,412  u 8  50,485 50,485  149 , 5 4 8 200 ,033  210,777 410,810  U15  31,813 31,813  58 , 0 7 3 89 , 8 8 6  94,216 184,102  U30  37,471 37,471  75 ,183 112 654  120,251 232,905  U33  44,237 44,237  133 553 177 790  222,832 400,622  V 8  37,050 37,050  125 3 6 3 1 6 2 413  120,972 283,385  V10  38,263 38,263  6 7 710 105 973  89,315 195,288  Vll  57,723 57,723  171 939 229 6 6 2  V12  32,821 32,821  60 576 93 397  V19  31,038 31,038  63 94  V20  32,530 32,530  65, 1 7 5 97, 705  V21  38,155 38,155  127, 220 165, 375  V24  35,176 35,176  130, 040 165, 216  172,580 337,796  V25  33,212 33,212  6 6 , 584 9 9 , 796  135,846 235,642  92,392 185,789  322 360  260-322  32  I 33 T a b l e 7.  (Continued)  No.  Interval  (Days) 176-259  0-49  50-112  V26  54,015 54,015  131 909 185, 924  122 248 308 172  V27  32,308 32,308  62 7 3 0 95 038  1 2 1 699 216, 7 3 7  V28  32,671 32,671  69 102  942 613  130 233  801 414  113- -17?  260-322  X  4  40,770 40,770  142 183  898 668  209 392  148 816  231,448 624,264  195,674 819,938  X  5  69,017 69,017  138 207  690 707  188 753 396 460  247,115 643,575  221,876 865,451  X  6  38,359 38,359  158 196  201 560  138 335  203,819 539,093  279,011 818,104  X  7  33,762 33,762  115 517 1 4 9 ,279  163 852 313 ,131  209,405 522,536  194,811 717,347  X  8  38,106 38,106  140 883 1 7 8 ,989  136 846 315 ,835  200,071 515,906  161,235 677,141  X  9  39,612 39,612  135 ,982 175 594  146 ,658 322 ,252  216,422 538,674  171,665 710,339  X10  33,683 33,683  137 ,442 1 7 1 ,125  190 , 8 1 3 361 ,938  222,110 584,048  174,848 758,896  Xll  41,954 41,954  154 , 5 0 7 196 ,461  172 ,771 369 ,232  232,480 601,712  249,202 850,914  X14  42,595 42,595  151 ,891 194 ,486  208 ,305 402 ,791  227,345 630,136  177,382 807,518  X16  42,773 42,773  125 ,792 168 , 5 6 5  133 , 4 6 3 302 , 0 2 8  196,603 498,631  202,232 700,863  714 274.  T a b l e 7.  No.  (Continued)  0-49  50-112  113-175  53,322  120,079 173,401  145,935 319,336  39,223  39,223  111,596 150,819  Y 3  36,608 36,608  83,101 119,709  Y 4  34,921 34,921  129,432 164,353  Y 5  46,666 46,666  109,290 155,956  Y 6  37,190 37,190  102,316 139,506  Y 8  39,443 39,443  109,737 149,180  Y10  45,490 45,490  Yll  53,561 53,561  Y13  47,323 47,323  151,899 199,222  177,633 376,855  Y21  45,901 45,901  147,770 193,671  195,230 388,901  267,285 656,186  Y24  48,129 48,129  150,662 198,791  208,693 407,484  280,000 687,484  Y 1 Y 2  53,322  135,771 291,727  T a b l e 8.  C u m u l a t i v e energy i n t a k e f e m a l e s , by i n t e r v a l  (Calories) of laboratory  I n t e r v a l (Days) 50-112 113-175 176-259  -  0-49  U10  41 , 6 7 1 41 , 6 7 1  115 ,943 1 5 7 ,614  179,685 337,299  Ull  42 725 42 , 7 2 5  128 ,410 171 ,135  181,794 352,929  U14  50 ,222 50 222  135 913 186 , 1 3 5  194,289 380,424  U32  27  613 27 6 1 3  65  932  93 545  104,982 198,527  U38  31 119 31 119  67 1 4 6 98 265  104,486 202,751  V  7  48 147 48 147  1 4 7 813 195 960  V  9  35 4 6 1 35 461  67 0 6 7 1 0 2 528  91,017 193,545  V13  52, 445 52 4 4 5  133 627 186 072  171,915 357,987  V14  35 7 5 6 35 7 5 6  63 1 3 0 98 8 8 6  VI5  35 030 35 030  97  N o  62  900 930  V16  3 6 253 36 253  65 929 1 0 2 182  95,537 197,719  V17  23 4 0 6 23 4 0 6  58 919 82 3 2 5  84,418 166,743  Y 7  4 6 377 46 377  169 7 3 9 216 116  260-322  o f m i l k remained t h e same i n b o t h s t u d i e s . months a c o n s i d e r a b l e  Since at four  f r a c t i o n o f t h e t o t a l energy i n t a k e  had been c o n t r i b u t e d by m i l k , t h e c o r r e c t e d c u m u l a t i v e i n t a k e s a r e o n l y 12% l o w e r .  At s i x months, where t h e m i l k  had been a much s m a l l e r component o f t h e t o t a l f e e d the cumulative  II.  energy  intake,  energy i n t a k e s were \K% l o w e r .  Variations i n Nutritional  Plane.  As mentioned e a r l i e r , t h e d i f f e r e n c e s i n t h e l e v e l o f energy i n t a k e o v e r each i n t e r v a l a r e o f major importance i n t h i s study.  S i n c e t h e s i g n i f i c a n c e o f a c e r t a i n energy i n t a k e  depends upon t h e weight o f t h e a n i m a l a t t h a t t i m e , energy i n t a k e a l o n e i s o f l i t t l e v a l u e . t o f i n d a method o f e x p r e s s i n g  cumulative  I t i s necessary then  energy i n t a k e i n r e l a t i o n t o  body w e i g h t . The energy r e q u i r e m e n t s o f an a n i m a l can be e x p r e s s e d i n a v a r i e t y o f ways.  The most f r e q u e n t l y e n c o u n t e r e d  reference  i s t h a t o f ' b a s a l heat p r o d u c t i o n '  (B.H.P.).  Conditions f o r  m e a s u r i n g t h i s have been d i s c u s s e d  i n d e t a i l elsewhere  (Brody,  1945; K l e i b e r , 1961) and w i l l be mentioned o n l y b r i e f l y h e r e . The two c r u c i a l p o i n t s a r e t h a t t h e a n i m a l s must be i n t h e post-absorptive  s t a t e and must be p h y s i o l o g i c a l l y  Another reference production'  i n frequent  (M.H.P.).  quiescent.  use i s 'maintenance heat  T h i s i s t h e energy e x p e n d i t u r e  animal that i s being f e d at a l e v e l t o j u s t maintain weight.  The magnitude o f M.H.P. i s e s t i m a t e d  o f an body  t o be 2 x B.H.P.  37 ( G a r r e t t §_t a l , 1959).  A t h i r d reference  p o i n t t h a t may be  encountered i s the l e v e l o f feed i n t a k e o f a growing animal on u n r e s t r i c t e d f e e d s u p p l y . ( K l e i b e r , 1961).  This i s approximately  5 x B.H.P.  I t i s evident that i f the l e v e l o f feeding  o f a g r o w i n g a n i m a l i s e x p r e s s e d as a m u l t i p l e o f b a s a l heat p r o d u c t i o n , t h e r e s u l t i n g f i g u r e w i l l f a l l i n t h e range o f 2 - 5 x B.H.P. The  m u l t i p l e o f B.H.P. was used as t h e b a s i s f o r  d e s c r i b i n g energy i n t a k e s i n t h i s s t u d y .  The method was used  w i t h s a t i s f a c t o r y r e s u l t s by Leche (1964) t o document t h e e n e r g y i n t a k e s o f young g r o w i n g c a t t l e .  B l a x t e r and Graham  (1955) have p o i n t e d out t h a t t h e use o f m u l t i p l e s o f B.H.P. i s not a v a l i d e x p r e s s i o n  o f energy r e q u i r e m e n t s because t h e  amount o f energy g o i n g t o a p a r t i c u l a r need i s n o t a p r o p o r t i o n o f t h e t o t a l energy i n t a k e . t h i s i s used s t r i c t l y  constant  However, as l o n g as  as an i n d e x o f energy i n t a k e and not as  a measure o f any one energy compartment, i t p r o v i d e s a convenient  reference.  B.H.P. i s most commonly e x p r e s s e d i n terms o f body w e i g h t . Brody (1945) d e r i v e d t h e f o l l o w i n g e q u a t i o n consideration o f the metabolic  from a  r a t e s o f mature a n i m a l s  r e p r e s e n t i n g a d i v e r s i t y o f s p e c i e s and body  weights:  B.H.P. = 70.5 W°/ <- C a l o r i e s p e r 24 h o u r s ; 73Z  kg.  or r e w r i t i n g in.pounds, B.H.P. - 39.65 W°* 34 C a l o r i e s p e r 24 h o u r s . lb. 7  38  These e q u a t i o n s g e n e r a l l y a l l o w a r e l i a b l e e s t i m a t i o n o f B.H.P. f o r mature a n i m a l s  o f any body  weight.  T h i s c o n s t a n t r e l a t i o n s h i p o f b a s a l m e t a b o l i s m t o weight r a i s e d t o t h e 0.734 power i n mature a n i m a l s  cannot be expected  t o a p p l y , and does not a p p l y t o r a p i d l y growing  animals.  Brody (1945) d e s c r i b e d t h e weight r e l a t i o n s h i p o f B.H.P. i n young a n i m a l s  o f s e v e r a l s p e c i e s and i t i s e v i d e n t t h a t t h e r e  i s no s i m p l e e x p r e s s i o n o f t h i s r e l a t i o n s h i p i n t h e young as there i s i n the adult animal.  Each s p e c i e s e x h i b i t s a  d i f f e r e n t p a t t e r n w i t h t h e B.H.P. o f some b e i n g h i g h e r t h a t p r e d i c t e d by t h e a d u l t e q u a t i o n and o t h e r s b e i n g  than lower.  Much o f t h i s v a r i a b i l i t y can l i k e l y be accounted f o r by t h e d i f f e r e n t degrees o f m a t u r i t y a t b i r t h .  Although  B.H.P. f o r  young growing deer has not been d e s c r i b e d , Nordan e t a l (1970) have measured t h e r e s t i n g heat p r o d u c t i o n o f t h e s e  animals.  They argue t h a t s i n c e , by d e f i n i t i o n , a g r o w i n g a n i m a l conform t o t h e r e q u i r e m e n t s p r o d u c t i o n , another production  f o r measuring b a s a l heat  term must be used.  R e s t i n g heat  (R.H.P.) as d e f i n e d by Nordan e t a l i s measured on  a q u i e s c e n t a n i m a l e i g h t hours a f t e r i t s l a s t f e e d i n g . these and,  cannot  Under  c o n d i t i o n s , t h e heat p r o d u c t i o n w i l l be above B.H.P. because o f t h e r a p i d growth r a t e , s h o u l d a l s o be g r e a t e r  than M.H.P. S i n c e B.H.P. i s b e i n g used here as t h e b a s i s f o r t h e e x p r e s s i o n on energy l e v e l , an attempt has been made t o use the R.H.P. d a t a t o e s t i m a t e B.H.P. o f t h e younger a n i m a l s .  Nordan e t a l d e s c r i b e d t h e R.H.P. o f growing male deer by t h e f o l l o w i n g equations: 1.  3 - 1 0 kg. ( 6 . 6 - 2 2  R.H.P. =  l b .)  1 1 2 . 7 W°-76  = 61.9  kg.  2.  10-100  C a l o r i e s per lb. ^ 2 4 hours  W°-76  kg. ( 2 2 - 2 2 0 l b . )  R.H.P. =  164-0  W°- 3 7  C a l o r i e s per  = 9 2 . 3 W°-73  kg.  lb.  24  hours.  I f t h e assumption can be made t h a t R.H.P. i s a c o n s t a n t m u l t i p l e o f B.H.P. a s i s M.H.P. and maximum f e e d i n t a k e , t h e n t h e comparison o f R.H.P. and B.H.P. o v e r a weight range where b o t h can be measured w i l l produce t h e n e c e s s a r y f a c t o r t o i n t e r c o n v e r t t h e two. E q u a t i o n 2 e x p r e s s e s R.H.P. a s a f u n c t i o n o f body weight t o t h e 0 . 7 3 power.  That i s , o v e r t h e  w e i g h t range o f 2 2 t o 2 2 0 pounds R.H.P. i s a c o n s t a n t m u l t i p l e (2.2645)  o f B.H.P.  equation  1  i s also  Assuming t h a t t h e R.H.P. d e s c r i b e d by 2.2645  t i m e s B.H.P., an e s t i m a t e o f t h e  B.H.P. o f t h e fawns i n t h i s w e i g h t range can be made.  This  a l l o w s a comparison between B.H.P. o f t h e young growing fawn w i t h t h a t p r e d i c t e d by t h e a d u l t B.H.P. e q u a t i o n . I t i s now p o s s i b l e t o b e g i n t o d e r i v e an i n d e x o f p l a n e o f n u t r i t i o n t h a t w i l l a p p l y over a l l ages and body w e i g h t s encountered d u r i n g the experiment.  One e q u a t i o n f o r t h e  e x p r e s s i o n o f B.H.P. can be d e r i v e d so t h a t t h e i n t e r c e p t remains c o n s t a n t and t h e weight exponent v a r i e s t o account f o r  t h e change i n t h e p r o p o r t i o n o f m e t a b o l i c a l l y a c t i v e mass w i t h change i n body w e i g h t . i n d e x can be d e t e r m i n e d  From t h i s e q u a t i o n t h e energy  s i m p l y by d i v i d i n g  t h e energy i n t a k e  per m e t a b o l i c pound p e r day by t h e i n t e r c e p t 39*65. A problem  a r i s e s i n t h a t t h e two e q u a t i o n s o f Nordan e t  a l a r e d i s c o n t i n u o u s a t 22 pounds and t h e r e f o r e cannot be f i t t e d by a s i n g l e e x p r e s s i o n .  I n the o r i g i n a l data, the  r e g i o n between 15 and 30 pounds showed c o n s i d e r a b l e v a r i a b i l i t y and i t was o b v i o u s t h a t t h e .slopes o f t h e two l i n e s were b e i n g governed by t h e d a t a o v e r t h e remainder w e i g h t range.  Because o f t h e v a r i a b i l i t y ,  o f the  i t was d e c i d e d t o  use e q u a t i o n 2 above 30 pounds, and t o make an a p p r o x i m a t i o n t o t h e d a t a below t h i s w e i g h t .  B.H.P. (R.H.P./2.2645) f o r t h e  fawns was d e s c r i b e d by t h e e q u a t i o n B.H.P. = 39.65 W3f, C a l o r i e s p e r 24 h o u r s . lb. U s i n g t h i s e q u a t i o n t o approximate deer  the metabolic data o f the  (Nordan e_t a l ) gave a v a l u e o f  y = 0.734 - 0.00012 (30 - V / ) 2  2  from 6.6 t o 30 pounds,  and y.= 0.734 o v e r 30 pounds. The range o f ' y  1  i s from 0.60 t o 0.734 ( F i g u r e 2 ) .  The meta-  b o l i c r a t e d e s c r i b e d by t h i s e x p r e s s i o n i s compared w i t h t h a t d e s c r i b e d by Nordan*s e q u a t i o n s 1 and 2 i n F i g u r e 3-  The  R.H.P. was d i v i d e d by 2.2645 t o compare t o t h e B.H.P. e q u a t i o n .  Figure 2  V a r i a t i o n s i n t h e Exponent »y* Required t o F i t Heat P r o d u c t i o n Data o f Deer When t h e Slope, 3 9 . 6 5 ,  i s Held  Constant;  i . e . , B.H.P. » 3 9 . 6 5 W  y lb.  42  Figure 3  R e s t i n g Heat P r o d u c t i o n o f Deer D e s c r i b e d As a M u l t i p l e o f B a s a l Heat P r o d u c t i o n ; y i . e . , R.H.P. = 2.2645 B.H.P. = 2.2645 (39.65)W lb.  A l t h o u g h t h e new e q u a t i o n does not c o i n c i d e v e r y w e l l o v e r t h e range o f e q u a t i o n 1, i t does f a l l w i t h i n t h e v a r i a b i l i t y o f the  d a t a a t t h e s e body w e i g h t s . In  o r d e r t o d e l i n e a t e t h e t h r e e energy r e g i m e s i n t h i s  e x p e r i m e n t , t o t a l a p p a r e n t d i g e s t i b l e e n e r g i e s f o r each i n t e r v a l were e x p r e s s e d as C a l o r i e s p e r u n i t m e t a b o l i c a l l y a c t i v e t i s s u e p e r day; i.e.,  B.H.P. = 39.65 C a l o r i e s per pound^ p e r day.  The f i g u r e r e s u l t i n g from t h i s c a l c u l a t i o n s h o u l d range from 2 t o 5 t i m e s 39-65 o r from 79.30 t o 198.24 C a l o r i e s p e r pound^ per day.  The t o t a l energy i n t a k e o v e r an i n t e r v a l  can  be d i v i d e d by t h e number o f days i n t h e i n t e r v a l and, by t h e average m e t a b o l i c body w e i g h t o v e r the i n t e r v a l .  Dividing  t h i s q u o t i e n t by 39.65 l e a d s t o t h e 'energy i n d e x ' , a d i m e n s i o n l e s s number w h i c h s h o u l d f a l l between t h e l i m i t s o f 2 and 5.  I n t h e c a l c u l a t i o n o f t h e energy i n d i c e s f o r t h e  a n i m a l s i n t h i s s t u d y , growth o v e r each i n t e r v a l was assumed to  be l i n e a r and a mean o f t h e l o g a r i t h m s o f mean w e e k l y body  w e i g h t s o v e r t h a t i n t e r v a l was used i n t h e c a l c u l a t i o n o f m e t a b o l i c a l l y a c t i v e mass. In  t h e e x p e r i m e n t a l p l a n , i t had been proposed t h a t i n  each time i n t e r v a l t h e l e v e l o f f e e d i n g c o u l d be h i g h (H), medium (M), o r low (L) a c c o r d i n g t o t h e f l o w s h e e t i n F i g u r e For  s e v e r a l r e a s o n s , r e - e v a l u a t i o n o f t h e regimes  necessary.  Originally,  'M'  was  and 'L' were based on 85% and  70%  1.  r e s p e c t i v e l y o f t h e average ad l i b i t u m , o r 'H', f e e d i n t a k e a t a given weight.  The v a r i a t i o n w i t h i n t h e *H  1  t r e a t m e n t was  g r e a t and i n many c a s e s t h i s t r e a t m e n t approached  levels  had been o r i g i n a l l y d e f i n e d as 'M'.  'energy  The average  which  i n d e x ' f o r t h e 'H' t r e a t m e n t i n i n t e r v a l s 2 and 3 was 4 . 1 7 'M' p l a n e d e f i n e d on t h i s b a s i s would be would be 2 . 9 2 .  3•54  and 'L' p l a n e  F u l l f e e d i n g s h o u l d have r e s u l t e d i n a maximum  'energy i n d e x ' o f 5.00, and some o f t h e 'H' fawns d i d r e a c h this  level. S i n c e i t was not p o s s i b l e t o c o n s i d e r t h e plane o f  n u t r i t i o n as a continuum,  t h r e e d i s t i n c t ranges o f t h e energy  i n d e x had t o be d e f i n e d which would i n c l u d e t h e v a r i a b i l i t y i n the l e v e l s o f treatment f o r a l l animals.  The range o f t h e  energy i n d i c e s c a l c u l a t e d f o r a l l i n t e r v a l s and a l l a n i m a l s was  from  2.42  to  4.95•  T h i s range was a r b i t r a r i l y  divided  i n t o t h r e e i n t e r v a l s by t a k i n g 5 as a maximum and l e t t i n g any v a l u e s o f 8 0 % o f t h i s o r o v e r ( 4 o r above) be d e f i n e d as 'H', 60% o r over ( 3 o r above) as 'M' and l e s s t h a n 6 0 % as ' L'. The n u t r i t i o n a l t r e a t m e n t s o f t h e e x p e r i m e n t a l a n i m a l s , r e - e v a l u a t e d i n t h i s manner, a r e shown i n T a b l e 9 .  It will  be seen t h a t t h e o r i g i n a l p l a n has been d i s r u p t e d , and e v a l u a t i o n o f t h e t r e a t m e n t s made more d i f f i c u l t because o f m i s s i n g treatments or s i n g l e animals per treatment. The  f i r s t time i n t e r v a l has p r o v i d e d a c o n s i d e r a b l e  problem i n a n a l y s i s .  During the processes o f adaptation t o  c a p t i v i t y and o f weaning,  v a r y i n g degrees o f u n p r e v e n t a b l e  Table 9.  No. 0^49  R e - e v a l u a t i o n o f energy i n d i c e s and r e g i m e s o f l a b o r a t o r y - r e a r e d fawns  Energy Index by I n t e r v a l (Days) 50-112 113-175 176-259 260-122 Males  Regime  U 7  3.61  4.21  3.83  MHM  U 6*  3-69  4-31  4.03  MHH  U15  2.77  2.43  2.84  LLL  U30  2.87  2.42  2.81  U33  2.21  3-72  4-05  LMH  V 8  3.10  4.15  2.83  MHL  V10  3.08  2.53  2.68  MLL  Vll  4.29  4.90  V12  2.96  2.76  V19  2.74  2.65  LL  V20  2.87  2.92  LL  V21  3.26  4-32  MH  V24  2.97  4.19  3.59  LHM  V25  2.92  2.79  3.83  LLM  V26  4-05  3-98  2.82  HML  V27  2.87  2.62  3-54  LLM  V28  2.87  2.94  3.80  LLM  X 4  3.26  4.24  3.95  MHM  1  See page 43-  HH 2.99  LLL  T a b l e 9.  (Continued)  No.  Energy Index by I n t e r v a l (Days) 50-112 260-322 113-175 176-259 0-49 Males  Regime  X 5  4.76  3-60  3.53  2.98  3 .21  HMMLM  X 6  3.15  4.95  2.98  2.91  4-71  MHLLH  X 7  2.88  3.93  3.73  3.07  3-45  LMMMM  X 8  3.13  4-42  3.01  3.05  3 .10  MHMMM  X  9  3-34  4.28  3-04  3.01  2.99  MHMML  X10  2.85  4.43  3.97  3.06  3.11  LHMMM  Xll  3.54  4-72  3.30  2.94  3-85  MHMLM  XH  3.32  4.44  4-04  2.96  3.09  MHHLM  X16  3-46  3.95  2.99  3.09  3-84  MMLMM  Y 1  3.92  3.46  3-16  Y 2  3.15  3-55  MM  Y 3  3-13  3 .26  MM  Y 4  2.84  4.06  LH  Y 5  3-54  3-32  Y 6  3.09  3.61  MM  Y 8  3-19  3-54  MM  Y10  3-51  M  Yll  3.97  M  Y13  3-60  4-23  3.54  Y21  3.66  4.50  3.97  3-36  MHMM  Y24  3.60  4.21  4.01  3-49  MHHM  MMM  3.16  MMM  MHM  T a b l e 9-  (Continued)  No.  Energy Index by I n t e r v a l (Days) 50-112 113-175 176-259 260-322 0-49 Females  Regime  U 7  3.16  4-59  3-52  MHM  Ull  3-48  3.99  3.75  MMM  U14  3.69  3 .81  3.80  MMM  U32  2.82  3 .20  3.18  LMM  U38  2.91  3.08  3.14  LMM  V 7  3.70  4.38  V 9  3 .08  2.89  3.07 •  MLM  V13  3.95  3-94  3-65  MMM  V14  3.05  2.62  ML  VI5  2.99  2.67  LL  V16  3.06  2.66  2.98  MLL  V17  2.07  2.58  2.84  LLL  Y 7  3.55  4.96  MH  MH  r e s t r i c t i o n s o f f e e d i n t a k e have o c c u r r e d . common treatment  must be c l a s s i f i e d as ' L . f  In f a c t , the most It i s possible  i n most i n s t a n c e s t o regroup animals f o r a n a l y s i s o f . t h e e x p e r i m e n t a l t r e a t m e n t s , but d i f f e r e n c e s between f i e l d  and  l a b o r a t o r y fawns caused by t h i s e a r l y r e s t r i c t i o n remain unknown e n t i t y .  T h i s problem  w i l l be encountered  an  later in  attempts t o e v a l u a t e the n u t r i t i o n a l p l a n e s o f w i l d fawns.  I I I . V a r i a t i o n s o f Body Weight w i t h Cumulative The  v a r i a t i o n s i n body weight  Energy  w i t h cumulative energy i n t a k e  were c o n s i d e r e d a t the end o f each i n t e r v a l .  A l l d a t a were  i n c l u d e d r e g a r d l e s s o f the p a t t e r n o f t r e a t m e n t . animal c o n t r i b u t e d weight  Intake.  Since  each  d a t a at every i n t e r v a l up u n t i l  the  time o f s l a u g h t e r , more d a t a p o i n t s a r e a v a i l a b l e a t the younger ages. weight  F i g u r e s 4 t o 7 show the r e s u l t i n g graphs  v e r s u s energy  intake.  I t was  of  p o s s i b l e t o f i t the d a t a  a t each age w i t h a l i n e a r r e g r e s s i o n l i n e .  The d e r i v e d  e q u a t i o n s f o r the l i n e s o f b e s t f i t are as f o l l o w s : 49  days  Y = 6.723 + 37.012 X  112  days  Y = 4.145  175  days  Y = 4.627 + 17.286 X  322 days  s  + 25.082 X  Y = -26.463 + 13.62  where Y •= body weight  yx  lb.  3.05  lb.  3.53  lb.  6.22  lb.  s  yx V X  =  s  yx  i n pounds  X = cumulative energy i n t a k e i n C a l o r i e s x 10" (A.D.E.) 5  = 3.05  Figure  4  The R e l a t i o n s h i p Between Body Weight and T o t a l Energy Intake o f L a b o r a t o r y Reared Fawns a t 49 Days o f Age.  •ill  its  Figure  5  The R e l a t i o n s h i p Between Body Weight and T o t a l Energy Intake o f L a b o r a t o r y Reared Fawns a t 112 Days o f Age.  Figure  6  The R e l a t i o n s h i p Between Body Weight and T o t a l Energy Intake o f L a b o r a t o r y Reared Fawns a t 175 Days o f Age.  Figure  7  The R e l a t i o n s h i p Between Body Weight and T o t a l Energy Intake o f L a b o r a t o r y Reared Fawns a t 322 Days o f Age.  53 I n terms o f t h e a n i m a l p r o d u c e r , t h i s i m p l i e s a c o n s t a n t f e e d efficiency. Winchester  The r e s u l t s a r e i n agreement w i t h t h o s e o f and Howe (1955) who found t h a t c a t t l e ,  which  r e a l i m e n t e d a f t e r 6 months o f r e s t r i c t i o n , t o o k t h e same amount o f energy t o r e a c h 1000 pounds a s d i d u n r e s t r i c t e d controls.  A l t h o u g h i n t h e p r e s e n t work a f i x e d i n t e r v a l o f  t i m e was c o n s i d e r e d r a t h e r t h a n a f i x e d body w e i g h t , t h e constant feed e f f i c i e n c y p r e v a i l e d .  The s t r a i g h t l i n e  relation-  ship w i t h a high c o r r e l a t i o n c o e f f i c i e n t provides a reasonable e s t i m a t i o n o f c u m u l a t i v e energy i n t a k e i r r e s p e c t i v e o f p a t t e r n of r e s t r i c t i o n . A l l o f t h e d a t a c o n s i d e r e d so f a r have d e a l t w i t h l i v e body w e i g h t .  S i n c e much o f t h e f i e l d d a t a a r e a v a i l a b l e o n l y  as e v i s c e r a t e d w e i g h t s , a c o n v e r s i o n t a b l e has been  developed  u s i n g l i v e and e v i s c e r a t e d w e i g h t s o f l a b o r a t o r y a n i m a l s .  The  f o l l o w i n g p e r c e n t a g e s r e l a t e l i v e body w e i g h t t o 'hog d r e s s e d ' weight;  i . e . , a w e l l bled carcass with a l l viscera  including  l i v e r and k i d n e y s removed.  Age 49 days  No. o f A n i m a l s 3  E v i s c e r a t e d / L i v e Weight 69  percent percent-  112  13  69.2*1.59*  175  24  73.0^1.56*  322  10  76.5 1.33* +  *P = 0.95  The a p p l i c a t i o n o f t h e s e c o n v e r s i o n f a c t o r s i s extremelyu s e f u l , because c a l c u l a t e d l i v e body w e i g h t s a r e t h e n e x p r e s s e d i n terms o f l a b o r a t o r y w e i g h t s . to  T h i s a v o i d s c o m p l i c a t i o n s due  d i f f e r i n g c o n t r i b u t i o n s o f rumen weight when the a n i m a l s are  b e i n g f e d c o n c e n t r a t e r a t i o n s as opposed t o when t h e y are b e i n g fed  forages. The i n c r e a s e i n d r e s s i n g percentage w i t h age agrees w i t h  t h e f i n d i n g s r e p o r t e d f o r many a n i m a l s ( J a c k s o n and Lowrey, 1912; Hammond, 1932;  P a l s s o n , 1955).  This i s usually explained  on t h e b a s i s t h a t v i s c e r a have r e a c h e d a r e l a t i v e l y h i g h p r o p o r t i o n o f mature weight a t b i r t h and as growth  proceeds,  t h e r e s t o f t h e body assumes a l a r g e r f r a c t i o n o f t h e mass.  total  One would expect a s i m i l a r i n c r e a s e i n d r e s s i n g p e r -  centage i n l a r g e r a n i m a l s a t any one age, but sample s i z e s i n t h i s s t u d y were t o o s m a l l t o demonstrate  such an i n c r e a s e .  L i v e body w e i g h t s c a l c u l a t e d f o r f i e l d k i l l e d deer i n t h i s s t u d y are  shown i n T a b l e 5 and t h e i r c u m u l a t i v e energy i n t a k e s are  estimated. In a n t i c i p a t i o n o f t h e f u t u r e use o f i n f o r m a t i o n from h u n t e r k i l l e d d e e r , l i v e r w e i g h t s have been r e c o r d e d so t h a t c o r r e c t i o n s can be made upon t h e w e i g h t s o f a n i m a l s t h a t are brought t o t h e c h e c k i n g s t a t i o n w i t h t h e l i v e r s t i l l a t t a c h e d . Head w e i g h t s were a l s o r e c o r d e d , i n the event t h a t t h e s e would be removed a l s o (see F i g u r e 8 ) .  IV.  S k e l e t a l Dimensions  o f L a b o r a t o r y Reared  Deer.  F o r each o f the s k e l e t a l d i m e n s i o n s d e s c r i b e d on page 28,  Figure  F i g u r e 8a.  8  Changes With Age i n t h e  L i v e r Weights o f L a b o r a t o r y  F i g u r e 8b.  Fawns.  Changes With Age i n the  Head Weights o f L a b o r a t o r y W i l d Fawns.  and  a graph was c o n s t r u c t e d r e l a t i n g t h e magnitude o f t h a t d i m e n s i o n t o c u m u l a t i v e energy i n t a k e a t 112, 175 and 322 days. F i g u r e s 9 t o 50 show t h e s e g r a p h s .  A l l s k e l e t a l measurements  have been t a b u l a t e d and a r e i n c l u d e d i n T a b l e s 10-56 (pp. 100146). The t r e a t m e n t o f t h e d a t a a t 112 and 175 days w i l l be considered f i r s t .  Only a r e s t r i c t e d number o f t r e a t m e n t s are  i n c l u d e d a t 322 days and t h e s e a r e b e s t c o n s i d e r e d l a t e r .  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Wi. .... .EE EPLEE  .ipE-  ;  ;  :  :.-:-| -:;  J  • ]•  :"-P  3 2 2 r) A Y S  E n e r g y ; Intake, C alories x I 0 1A.D.E.) ::.•'•  .. E:..  EEpEE:  EEpr-:  3-ri~-|r-.-i--r  :': E E X E X X  o EEiEiE  ES  i E i i E E E-Ei-EE —r  E.E:|pE  ..L:J.-::_ i::,4^iiA:i^^  V  . : . :.j . . : i. P . .r x  ---E!-E;  TE|~  f  : i :i:.: •:.:::::::::•:] . x . E . : :  =  i ..::.:..: E L.EiEE:  :  EE:IEE< EEEEE :ELEE.E.  -  •_  " •;. ii • : PPE ii  O.:...  E E E  ' E:E'  :  : :!: ::.:.  :  -  iiiiii'  .:.: :: I::  1  .  : P  :  E:P:".".El  E  Leaf 99 omitted i n page numbering.  100  Table 1 0 A n i m a l No. Sex  VII  T o t a l E . I . 229,662 1  Age 1 1 2 Days  M  Calories Pattern E.I,  Deviation  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  14.65 15.3 15.3 12.5  1.76 1.78 1.82 1.85  length length length length  h i n d cannon tibia femur pelvis  22.0 17.9 17.9  17.65  1.78 1.86 1.87 2.27  -0.52  length length length length length length length length length  cervical  4.32  1.75  3-55  I.83  -0.55 -0.34 -0.47  3.19  1.78 2.28 2.30 2.28  thoracic lumbar  5 10 1 3 5  width width width  cervical thoracic lumbar  length width,  3 5 7 1  distal  fore fore fore fore  5 5 3  cannon cannon cannon cannon  2  4.06  2.87  3.06  3.95 4.18  3-96  Intercept"^"  HH  1.96  2.30 2.25  x  10  5  -0.54 -0.52  -O.48 -0.45  -0.44 -0.43 -0.30  -0.52 -0.02 0.00 -0.02 0.00 -0.05  3 .61 3.88  1.78 2.28  2.74  2.30  -0.52 -0.02 0.00  1.76 1.73 1.37 1.54  -  14.65 2.70 2.51  1.19  0 0 0 0  1  Apparent d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  . . . .  54 57 93 76  x  1  10  5  T a b l e 11 A n i m a l No. Sex  V19  M  Total E . I .  1  94,360  Age 112 Days  P a t t e r n E . I . LL Intercept  1 1 Deviation  12.35 12.9 12.6 10.1  1.00 x 105 0.91 0.91 0.92  +0.06 -0.03 -0.03 -0.02  15.25  0.90 0.91 0.92 0.93  -0.04 -0.03 -0.02 -0.01  1.15  +0.21 +0.01 +0.13 +0.13 +0.25 +0.26 -0.01 0.00 +0.09  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  length length length length  h i n d cannon tibia femur pelvis  18.9  length length length length length length length length length  cervical  3  Calories  2  15.0 14.6  5  3.68 3.29 2.96 2.56 2.36 2.54 3.01 3.28 3-29  5 5 3  3.20 3.04 2.07  0.92 0.91 1.15  -0.02 -0.03 +0.21  cannon cannon cannon cannon  12.35 2.33 2.36 1.12  1.00 0.96 0.92 .123  +0.06 +0.02 -0.02 +0.29  thoracic  5 7  1 5  lumbar  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  10 1 3  0.95 1.07 1.07 1.19 1.20 0.93 0.94 1.03  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  Table 1 2 A n i m a l No. Sex  V20  M  Total E.I.  1  9 7 , 7 0 5i  Age 1 1 2 Days  Calories P a t t e r n E . I . LL  Intercept  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  13.25 13.4 13.1 10.63  1.23 1.22  length length length length  h i n d cannon tibia femur pelvis  15.9  length length length length length length length length length  cervical thoracic lumbar  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  3 5 7 1 5 10 1 3 5 5 5 3  cannon cannon cannon cannon  2  x  1  105  Deviat  1.08  +0.25 +0.24 +0.10  1.12  +0.14 +0.15  15.35  1.13 1.07 1.03  15-3  1.12  +0.14  0.92  -0.06  19.4  3-43  3.29 2.87  2.69 2.06 2.27 3.05 3-32 3.22  0.95 0.94  1.08 0.92 0.92 0.98 0.98 0.95  +0.09 +0.05  -0.03 -0.04 +0.10 -0.05 -0.05 0.00 0.00 -0.03  3.22 3.17  0.97  2.19  1.26  -0.01 +0.08 +0.28  1.23 0.94 0.98 1.18  +0.25 -0.04 0.00 +0.20  13.25 2.32 2.36 1.11  1.06  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  x  10  13  Table A n i m a l No. V21 Sex  Total E.I.  165,375  P a t t e r n E . I . MH  Age 112 Days  M  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  14.3 15.0  length length length length  h i n d cannon tibia femur pelvis  length length length length length length length length length  cervical  3 5 7 thoracic 1 5 10 lumbar 1 3 5  3.87 3.50 3.18 2.77 2.88 3.69 3.86 3.87  width width width  cervical thoracic lumbar  3.46 3-53 2.43  length width,  fore fore fore fore  distal  5 5 3  cannon cannon cannon cannon  Calories  2  14.95  12.45  Intercept  1  Deviation  1.63 x 105 1.69 1.70  -0.02 x 1 0 -0.04 +0.05 +0.27  1.82  17.2 21.6 17.75 17.5  1.62  -0.03 +0.08  1.82 I.83  +0.17 +0.18  4.15  1.59  -0.06 +0.06 +0.12 +0.12  14.3  2.64 2.66 1.18  1  1.73  1.71 1.77 1.77 1.80  1.81 1.75 1.68  +0.15  +0.16 +0.10  1.80  +0.03 +0.15  1.46 1.57 1.70  -0.19 -0.13 +0.05  1.63 1.61 1.77 1.50  -0.02 -0.04 +0.12 -0.15  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  5  104  14  Table A n i m a l No. Y2 Sex  Total E.I.  1  1  150,819  Age 1 1 2 Days  M  Pattern E.I.  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  15.0 15.3 15.6  length length length length  h i n d cannon tibia femur pelvis  18.15 22.5 18.5  length length length length length length length length length  cervical  <  thoracic lumbar  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  3 5 7 1  Calories  2  12.25  1  Deviation  Intercept 1.88 1.78 1.93 1.75  1.96  x  10  MM  5  +0.37 +0.27 +0.42 +0.24  2.01 2.08  +0.45 +0.50 +0.57  1.57 1.77  +0.01 +0.26  15.0  1.88  +0.37  2.67 1.26  1.80 1.86  +0.29 +0.35  4.O8  3.91  5 10 1 3 5  5 5 3  cannon cannon cannon cannon  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  x  1  10  5  105  15  Table A n i m a l No. Y3 Sex  M"  Total E.I.  119,709  Age 1 1 2 Days  Calories Pattern E.I.  ?  1 1 Deviation  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  14.15  14.25  14.5 11.6  1.57 1.47 1.55 1.50  length length length length  h i n d cannon tibia femur pelvis  17.35 20.8 17-0  1.67 1.48 1.58  +0.47 +0.28 +0.38  length length length length length length length length length  cervical  3  3.92 3.59  1.37  +0.17  1.35  +0.15  5 7  thoracic  Intercept  MM  x 105  +0.37 +0.27 +0.35 +0.30  1  5 10  lumbar  1  3 5  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  5 5 3  cannon cannon cannon cannon  3.59  14.15 2.49 2.57 1.23  1.73  +0.53  1.57 1.30  +0.37 +0.10  1.53 1.72  +0.33 +0.52  1  Apparent d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  x  10-  106 Table A n i m a l No. Sex  Y4  M  T o t a l E.I .  1  16  164,353  Calories P a t t e r n '.E . I . LH  Age 112 Days  Intercept  1  14.7  1.77 x 1 0  5  15.4 12.2  1.85 1.72  +0.21 +0.08  17.3 21.8  1.66 1.80 2.03  +0.02 +0.16 +0.39  3.71  1.80 1.99 2.05  +0.16 +0.35 +0.41  5 5 3  3.95  2.49  +0.85  cannon cannon cannon cannon  14.7 2.79 2.91 1.29  1.77 1.92 2.43 1.99  +0.13 +0.28 +0.79 +0.35  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  length length length length  h i n d cannon tibia femur pelvis  length length length length length length length length length  3 5 7 thoracic 1 5 10 lumbar 1 3 5  4.37  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  cervical  2  18.4  4.08  Devia +0.13  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  Table 1 7 A n i m a l No. Y6  T o t a l E.I ^  139,506  Calories P a t t e r n E . I . MM  Sex M  Age 1 1 2 Days  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  13.7 14-35 14-55 11.55  1.40 1.57 1.47  +0.17 +0.07  length length length length  h i n d cannon tibia femur pelvis  16.55  1.38 1.52  -0.02 +0.12  1.71 1.43  +0.31 +0.03  length length length length length length length length length  cervical  4.10  1.56  +0.16  2.09  +0.69  1.22  3 5  3.41 2.35 2.70 3-42 3.84 3.81  1.37 1.43 1.61  -0.18 -0.03 +0.03 +0.21  1.69  +0.29  5 5 3  3.53 2.10  1.53 1.17  +0.13  1.40  0.00 +0.52 -0.13 -0.20  thoracic  3  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  20.9 17.4  16.4  Intercept  1.49  x  10  Deviat  1  5  0.00  +0.09  5 7  1  5 10  lumbar  2  1  cannon cannon cannon cannon  13.7  2.79  1.92  2.46 1.11  1.27  1.19  -0.23  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  108 Table A n i m a l No. Y8 Sex  M  18  T o t a l E . I . 149,180 Age 112 Days  Calories P a t t e r n E . I . MM  p  i Intercept  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  14.1 14-35 14.65 11.75"  1.55 x 105 1.49 1.60 1.55  +0.16 0.00 +0.11 +0.06  length length length length  h i n d cannon tibia femur pelvis  16.85 20.5 17.3  1.49 1.40 1.67  0.00 -0.09 +0.18  length length length length length length length length length  cervical  3 5 7 thoracic 1 5 10 lumbar 1 3 5  4.18 3.83 3.35  1.62 1.66 1.57  +0.13 +0.17 +0.08  width width width  cervical thoracic lumbar  3.71  1.98  +0.49  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  1.55  +0.16  1.40 1.23  -0.09 -0.26  5 5 3  cannon cannon cannon cannon  14.1 2.52 1.12  Deviation  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  i  109 Table A n i m a l No. U 7  Total E.I.  Sex M  Age 1 7 5 Days  Dimension  Bone  length length length length  f o r e cannon radius humerus scapula  length length length length  h i n d cannon tibia femur pelvis  length length length length length length length length length  cervical thoracic lumbar  445,412  Calories Pattern E.I.  Size  3 5 7 1  19  p  16.3  17.2  17.4 15.3  19.8  25.7 21.3  21.9 5.25 4.81  4.19  Intercept 4.45 4.40 4.15 4-52 4 • 48 4.50 4.24 4.8O 4.87 4.72 4.46  3.68  5 10 1  4.61  3.17 3.77 4.70.  4.73  3 5  4.92 4.78  4.90 4.73 4.52 4.59  width width width  cervical thoracic lumbar  5 5 3  4.62 4.49 2.89  4.30  length width, d i s t a l width, proximal w i d t h , minimum  fore.cannon f o r e cannon f o r e cannon f o r e cannon  16.3 3-04 2.77 1.45  4.45 4.62 3.68 4.47  4.73 3.25  x  10  1  5  MHM  Deviation 0.00 -0.05 -0.30 +0.07 +0.03 +0.05 -0.21 +0.35 +0.42 +0.27 +0.01 +0.16 +0.28 +0.45 +0.28 +0.07 +0.14 -0.15 +0.28 -1.20 0.00  +0.17 -0.77 +0.02  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  110 T a b l e 20 A n i m a l No. U8 Sex  M  T o t a l E.I -  1  410,810  P a t t e r n E .1.  Age 175 Days Size  Dimension  Bone  length length length length  f o r e cannon radius humerus scapula  15.9 16.9 17.1  length length length length  h i n d cannon tibia femur pelvis  19.4  length length length length length length length length length  cervical  3 5 7 thoracic 1 5 10 lumbar 1 3 5  4.82  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  5 5 3  cannon cannon cannon cannon  Calories  14.8  25-2  21.3 21.0 4.91  4.68 4.15 3.37  2  Intercept^ 3.52  3.80 3.60 4.05 3.75 4.10  x 10  5  MHH  Deviation  -0.59 x 1 0 -0.31 -0.51 -0.06 -0.36 -0.01  4.25  +0.14  4.00  -0.11 +0.19  4.21  4.30  +0.10  4.26 3-47 4.75  +0.15  4.74  3.98 4.08  4.00  -0.13 -0.03 -0.11  4.64 4.14 2.78  4.35 3.48 3.H  +0.24 -0.63 -1.00  3.52  -0.59 -1.13 -0.94  3-37 3.64  4.39  15.9  2.86 2.68 1.40  4-52  2.98 3.17 4.07  1  -0.64 +0.64 +0.41  -0.04  1  Apparent d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  5  Ill Table A n i m a l No. Sex  U15  M  Total E.I.  1  21  184,102  Calories Pattern E.I.  Age 175 Days  Intercept  LLL  Deviation  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  13-45 14.2 14.0 11.55  1.85 x 105 1.94 1.80 1.82  +0.01 x 1 0 +0.10 -0.04 -0.02  length length length length  h i n d cannon tibia femur pelvis  16.65 20.3 16.8 17.1  1.99 1.75 1.93 1.94  +0.15 -0.09 +0.09 +0.10  length length length length length length length length length  cervical  3 5 7 thoracic 1 5 10 lumbar 1 3 5  4.15 3.74 3-35 2.82 2.66 2.84 3.65 3.81 3.88  2.17 2.13 2.01 2.17 2.41 2.29 2.21 2.06 2.33  +0.33 +0.29 +0.17 +0.33 +0.57 +0.45 +0.37 +0.22 +0.49  width width width  cervical thoracic lumbar  5 5 3  3.51 3.54 2.43  2.27 2.19  +0.43 +0.35 +0.96  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  cannon cannon cannon cannon  13.45 2.44  1.85 2.12 2.22 1.85  +0.01  2  2.38 1.08  2.80  1  +0.28 +O.38  +0.01  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  1  5  112  Table A n i m a l No. U33 Sex  M  22  T o t a l E . I . 400,622  Calories P a t t e r n E . I . LMH  Age 175 Days p  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  15.5  length length length length  h i n d cannon tibia femur pelvis  length length length length length length length length length  cervical  3 5 7 thoracic 1 5 10 lumbar 1 3 5  4.96 4.57 4.00 3.63 3.15 3.37  width width width  cervical thoracic lumbar  length width, d i s t a l  fore fore fore fore  14.9  3.25 x 1 0 3.44 3.42 4.15  19.0  3.28  24.5  3.50  16.6 16.8  20.9 20.7  1  Intercept  3.90 4.01  Deviation 5  -0.76 x 10-0.57 -0.59  +0.14  -0.63 -0.51 -0.11 0.00 +0.10 -0.05  4-87 4.69  4.11 3.96 3.85 4-42 3.90 3.74 3.70 4.39 4.32  5 5 3  4.28  3.52  4.15 2.80  3.51 3.13  -0.49  cannon cannon cannon cannon  15.5 2.97 2.74 1.40  4-30 3.38 4.08  4.28  3.25  1  -0.16 +0.41 -0.11 -0.27 -0.31 +0.38 +0.31 -0.50  -0.88 -0.76 +0.29 -0.63 +0.07  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  113 Table A n i m a l No. Sex  V8  M  Total E.I.  23  283,385  Age 1 7 5 Days  Calories Pattern E.I.  MHL  Dimension  Bone  Size  Intercept  length length length length  f o r e cannon radius humerus scapula  14.8 15.9 16.1  2.76 2.99 3.01  13.95  3.26  +0.43  length length length length  h i n d cannon tibia femur pelvis  18:0 22.5 19.2  2.74 2.56  -0.17  length length length length length length length length length  cervical thoracic  3 5 .7  1  5  lumbar  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  10 1  19.4 4.55 4.16 3-75 3.36  10  5  -0.07  +0.16 +0.18  -0.09 +0.09  3.13  +0.30  3.05 2.77 3-05 3-44  +0.22 -0.06 +0.22  +0.61  2.90  3.16  +0.33  3.17  3.12 2.81  +0.29  3 5  3.91 4.09 3.96  5 5 3  3.55 4.08 2.22  cannon cannon cannon cannon  2.92  x  Deviation  14.8 2.64 2.48  1.26  2.55 2.45  -0.02 -0.28 -0.38  2.31 3.25 2.61  -0.52 +0.42 -0.18  2.76 2.78  -0.07 -0.05 -0.28 +0.11  2.55 2.94  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  x  10  5  114 Table A n i m a l No. V10 Sex  M  Total E.I.  1  24  195,288  P a t t e r n '.E . I .  Age 175 Days  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  14.5  length length length length  h i n d cannon tibia femur pelvis  21.5 16.9  length length length length length length length length length  cervical thoracic lumbar  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  3 5  7 1 5 10 1 3  Calories  2  13.9 14-55  12.1 17.1  17.2 4.05 3.55  3.37 2.64  2.38 2.70  3.42 3.67  Intercept 2.16 x 1 0 2.11 2.13 2.10 2.24 2.23 1.95  1.98 2.00 1.84 2.05 1.84 I.84  2.06  5  3.62  1.89 1.84 1.93  5 5  3.52  2.28  3.53  2.17  3  cannon cannon cannon cannon  2.19 13.9  2.56 2.40 1.19  2.59  2.16 2.41 2.31 2.42  1  5  MLL  Deviation  1  +0.21 x 105 +0.16 +0.18 +0.15  +0.29 +0.28 0.00 +0.03 +0.05 -0.11 +0.10 -0.11 -0.11 +0.11 -0.06 -0.11 -0.02 +0.33 +0.22 +0.64  +0.21 +0.46 +0.36  +0.47  1  Apparent d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  25  Table A n i m a l No. V12 Sex  M  Total E.I.  185,789  Calories P a t t e r n E . I . LLL  Age 175 Days  Intercept  1  2.09 x 1 0 1.86 1.91  5  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  13.8 14.1 14.2  11.95  2.03  length length length length  h i n d cannon tibia femur pelvis  16.7 20.5 16.6 16.9  1.84 1.85 I.84  length length length length length length length length length  cervical  3 5 7 thoracic 1 5 10 1 lumbar  width width width  cervical thoracic lumbar  length width, d i s t a l  fore fore fore fore  3  5 5 5  3  2  3.92 3.67 3.26 2.79 2.48  2.69 3.48  3.68 3.61 3.65 3.29  2.19  cannon . 13.8 cannon 2.44 cannon 2.23 cannon 1.10  2.02  Deviation +0.23  0.00 +0.05 +0.17 +0.16 -0.02 -0.01 -0.02  2.02 1.79 1.83 2.11 2.05 2.04 1.97 1.86 1.91  +0.19 +0.18 +0.10 0.00 +0.05  2.40 1.77 2.59  +0.54 -0.09 +0.73  2.09  +0.23  2.12 1.79 1.96  +0.16 -0.07 -0.03 +0.25  +0.26 -0.07 +0.10  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  x  10  116  26  Table A n i m a l No. V 2 4 Sex  M  Total E.I.  337,796  Calories  Age 1 7 5 Days  Pattern E.I. ?  LHM  1  Dimension  Bone  Size  Intercept  length length length length  f o r e cannon radius humerus scapula  15.9  16.8 16.8  3 3 3 3  .53 .60 .42 .65  length length length length  h i n d cannon tibia femur pelvis  3 3 3 3  • 34 .06 .38 .67  -0.04 -0.32 0.00 +0.29  length length length length length length length length length  cervical  3 3 3 3 3 3 3 3  .81 .39 .50 • 56 .62 .27 .29 .00  +0.43 +0.01 +0.12 +0.18 +0.22 -0.11  3 5  4.84 4.34 3.86 3.53 3.03 3.33 4.10 4.40 4.21  5 5 3  3.94 4.25 2.23  2 .75 3 .89 2 .62  -0.63 +0.51 -0.76  cannon cannon cannon cannon  15.9 2.79 2.67 1.27  3 3 3 3  +0.15 +0.10 -0.25  thoracic lumbar  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  3 5 7 1 5 10 1  14.4  19.1 23.9 20.3 20.2  • 53 .48 .13 .02  x  105  Deviation +0.15 +0.22 +0.04 +0.27  -0.09 -0.38  -O.36  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  27  Table A n i m a l No. V25 Sex  M  Total E.I.  1  235,642  Age 1 7 5 Days  Calories Pattern E. 1 .  Deviation  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  14.45 14.95 15.4 12.9  2.54 2.40  length length length length  h i n d cannon tibia femur pelvis  17.35 21.8  2.37 2.31 2.46 2.32  +0.01  length length length length length length length length length  cervical  2.12 2.25 2.66 2.66 2.55 2.50 2.14  -0.24 -0.11 +0.30 +0.30  thoracic  3 5 7 1 5 10 1  2  18.1 17.9 4.12 3.82 3.63 3 .08  3 5  2.73 2.94 3.60 3.87 3-89  5 5 3  cannon cannon cannon cannon  lumbar  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  Intercept  LLM  2.60 2.53  x  10  1  5  +0.18 +0.14 +0.24 +0.17  -0.05 +0.10 -0.04  +0.19  2.34  +0.14 -0.22 -0.20 -0.02  3.83 3.74 2.41  2.58 2.52 2.78  +0.22 +0.16 +0.42  14.45 2.59 2.45 1.21  2.54 2.56 2.46 2.53  +0.18 +0.20 +0.10  2.16  +0.17  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  x  1  10  Table A n i m a l No. V26 Sex  M  28  T o t a l E.I  Calories  Age 1 7 5 Days  P a t t e r n E . I . HML ?  Bone  Size  length length length length  f o r e cannon radius humerus scapula  15.25 15.90  14.05  3.08 x 1 0 3.00 3.12 3.32  length length length length  h i n d cannon tibia femur pelvis  18.25 23-5 19.4 19.1  2.88 2.92 3.00 2.92  -0.20 -0.16 -0.08 -0.16  length length length length length length length length length  cervical  3 5 7 thoracic 1 5 10 lumbar 1 3 5  4.42 4.07 3.79 3.22 2.87 2.98 3.88 4.16 3.97  2.73 2.64 3.18 2.92 3.02 2.61 2.74 2.72 2.50  -0.35 -0.44 +0.10 -0.16 -0.06 -0.47 -0.34 -0.36 -0.58  width width width  cervical thoracic lumbar  5 5 3  4.08 3.84 2.32  3.08 2.68 2.70  0.00 -0.40 -0.38  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  cannon cannon cannon cannon  15.25 2.60 2.63 1.23  3.08 2.61 3.01 2.70  0.00 -0.47 -0.07 -0.38  16.3  Intercept  1  Dimension  5  Deviation 0.00 -0.08 +0.04 +0.24  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  Table A n i m a l No. Sex  V27  M  Total E.I.  1  29  2 1 6 , 7 3 7'  Age 1 7 5 Days  Calories P a t t e r n E . 1 . LLM  Intercept  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  14.7  2.71  15.0 12.3  2.44 2.31 2.22  length length length length  h i n d cannon tibia femur pelvis  17.8  2.62  21.7  2.27  length length length length length length length length length  cervical  3  thoracic  1  lumbar  5 7 5  10 1 3  2  14.9  17.8 17.8 4.27 3.84 3.47 2.99 2.58 2.77 3.63  2.34 2.27 2.35 2.28 2.25 2.50 2.25 2.17 2.18  x  10  Deviation  1  5  +0.54  +0.27 +0.14  +0.05 +0.45 +0.10 +0.17 +0.10 +0.18 +0.11 +0.08 +0.33 +0.08 0.00 +0.01  3.90 3.90  2.36  +0.19  3  3.82 3.59 2.61  2.57 2.27 2.97  +0.40 +0.10 +0.80  cannon cannon cannon cannon  14.7 2.53 2.48 1.21  2.71 2.34 2.55 2.53  +0.54 +0.17  5  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  5 5  +O.38 +O.36  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  x  1  10  120 Table A n i m a l No. V28 Sex  M  Total E.I.  30  233,414  Age 1 7 5 Days  P a t t e r n E . I . LLM Intercept  1  Deviation  5  12.2  2.13 x 1 0 2.19 2.36 2.16  -0.20 x 10 -0.14 +0.03 -0.17  16.9 21.8 17.4 17.7  2.13 2.30 2.17 2.22  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  13.85  length length length length  h i n d cannon tibia femur pelvis  length length length length length length length length length  cervical  3 5 7 thoracic 1 5 10 lumbar 1 3 5  4.26 3.96  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  5 5 3  cannon cannon cannon cannon  Calories  2  14.6  15.0  2.33 2.46 2.27  -0.20 -0.03 -0.16 -0.11  0.00  2.92 2.60 2.60  +0.13 -0.06 +0.29 +0.59 +0.27 +0.27  4.20  2.96  +0.63  4.02 3.63 2.72  2.93  +0.60 +0.92 +0.72  2.13  -0.20 +0.41 +0.47 +0.28  3.48  3.06 2.84  2.98 3.83 4.13  13.85  2.63 2.56 1.22  2.62  3.25 3.05  3.25 2.80  2.61  1  Apparent d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  1  T a b l e 31 A n i m a l No. Sex  Yl  M  Total E.I.  1  319,336  Calories P a t t e r n E,. 1 .  Age 1 7 5 Days  Intercept  MMM  Deviation  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  16.0 16.75  3.60  17.2 13.6  3.65  length length length length  h i n d cannon tibia femur pelvis  19.3 2.46  3.57 3.56  +0.38 +0.37  20.5 20.05  3.55 3.55  +O.36 +O.36  length length length length length length length length length  cervical  4.66 4.47 3.72 3.23 3-04  3.35 3.65 2.93  -0.26  thoracic lumbar  3 5 7 1 5 10 1 3 5  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  5 5 3  cannon cannon cannon cannon  2  3.50  2.90  2.96 3.60  x  10  1  5  +0.41 +0.31 +0.46  -0.29  +0.16  +O.46 -0.23  3.32  3.59  4.10 4-37 4.30  3.19  +0.41 +0.40 0.00  3.24  +0.05  4.18  3.30  3.99 2.65  2.93 3.24  +0.21 -0.26 +0.05  16.0 2.75 2.70  1.29  3.60 3.28  +0.41  3.22 3.19  +0.03 0.00  +0.09  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  x  1  10  T a b l e 32 A n i m a l No. Y5  Total E.I.  291,727  Sex M  Age 175 Days  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  length length length length  h i n d cannon tibia femur pelvis  length length length length length length length length length  cervical  Calories P a t t e r n E . I . MMM Intercept  1 1 Deviation  16.2 16.28 13.2  3.32 x lO^ 3 .17 3 .12 2 .70  +0.40 x 10 +0.25 +0.20 -0.22  18.98 23.4 19.48 19.10  3 .28 2 .89 3 .04 2 .92  +0.36  3 5 7 thoracic 1 5 10 lumbar 1 3 5  4.45 3.69 3.22 2.89 3.24 4.09 4.21 4.09  2 .81 2 .74 2 .86 2 .92 3 .09 3 .36 3.24 2 .83 2 .65  -0.11 -0.18 -0.06 0.00 +0.17 +0.44  width width' width  cervical thoracic lumbar  5 5 3  3-93 3.91 2.84  2 .74 2 .80 3 .17  -0.18 -0.12 +0.25  length width,  fore fore fore fore  cannon cannon cannon cannon  15.6 2.61 2.77 1.21  3 .32 2 .66 3.58 2 .52  +0.40 -0.26 +0.66 -0.40  distal  7  15.6  4.14  -0.03  +0.12 0.00  +0.32  -0.09 -0.27  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  Table 3 3 A n i m a l No. Y13  Total E . I . 376,855 1  Calories  Sex M  Age 1 7 5 Days  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  15.5  3.25  16.45  3.34 3.53  length length length length  h i n d cannon tibia femur pelvis  length length length length length length length length length  cervical  fore fore fore fore  x  105  1  Deviation -0.52 -0.43  -0.24  14.15  3.40  -0.37  19.18 24.33  3.36  -0.41  -0.42  19.83  3.35 3.54 3.41  4.66  3.35  -0.42 -O.58  20.48  4.32 3.93 3.58  3-19  -0.23  -O.36  4.29 4 •44 4.26  3-48 3.14  5 5 3  4-32 4.20 2.67  3.62 3.68 3.02  -0.15  cannon cannon cannon cannon  15.5 2.85 2.83 1.41  3.25 3.75 4.28  -0.52 -0.02  lumbar  distal  17.0  Intercept  5 10 1 3 5  cervical thoracic lumbar  length width,  o  3.64 4.25 3.17 3-34 3.72  thoracic  width width ividth  3 5 7 1  P a t t e r n E . I . MHM  2.91 3.23  4.15  -0.13  +O.48 -0.60 -0.43 -0.05  -0.29 -0.63  -0.09 -0.75  +0.51  +O.38  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  x  1  10  Table 3 4 A n i m a l No. X 4  T o t a l E.I  1  819,938  P a t t e r n E . I . MHMLL  Sex M  Age  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  17.38  length length length length length length length length length length length length length  cervical  322  Calories  Day, 2  Intercept x  Deviation  19.6  9.15 8.58 9.00  16.15  8.17  -0.03  h i n d cannon tibia femur pelvis  21.48  9.70  +1.50  27-3  8.39  +0.19  23.3  8.62  +0.42  3  5-71 5.33  8.70  +0.50 +1.54 -0.13 +0.13 -0.29 +0.99 +0.33 +0.14 +0.01  5 7  thoracic  1  19.1  24.2  4.42 4.18  9.06  9.74 8.07 8.33  7.91 9.19  5 10  3.48 4.00 4.85  3 5  5.17 5-19  5 5 3  4-91  8.05  4.66 3 .01  10.00 7.86  cannon cannon cannon cannon  17.38 2.94 3 . 0 0  9.15  lumbar  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  1  I.64  8.53 8.34  8.21  8.07  9.32 10.25  10  1  5  +0.95  +O.38 +0.80  +0.86  -0.15 +1.80 -0.34 +0.95 -0.13 +1.12 +2.25  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  x  1  10  125 Table 3 5 A n i m a l No. X5 Sex  M  T o t a l E.I -  1  865,481  Age 3 22 Days  Calories P a t t e r n E . I . HMMLL  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  16.6  7.27 x 1 0  19.03 16.98  9.60 9.30  length length length length  h i n d cannon tibia femur pelvis  20.4 27.05  7.52  length length length length length length length length length  cervical  3 5 7 thoracic 1 5 10 lumbar 1 3 5  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  2  18.9  Intercept  8.32  8.12 8.49  23.6 23.75  9.02  5.79  9.08  4.76 4.35 3.75 3.91 4.72 5.14 5.30  9.33  3  5.11 4.62 3.10  8.61 9.49 8.50  cannon cannon cannon cannon  16.6 2.78 2.74 1.43  7.22 7.37 7.59  5 5  5.08  8.09  9.10 9.25  8.67 7.85 8.22 8.50  7.27  Deviation  1  5  1  -1.38 x 1 0 -0.33 +0.95 +0.65 -1.13 -0.53 -0.16 +0.37 +0.43 -0.56 +0.68 +0.45 +0.60 +0.02 -0.80  -0.43 -0.15 -0.04  +O.84  -0.15 -1.38 -1.43 -1.28  -1.06  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  5  126  Table 3 6 A n i m a l No. X 6  Total E.I.  818,104  Calories  Sex M  Age 3 2 2 Days  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  17.3 18.55 18.5 15.5  7.87 7.25 7.30  length length length length  h i n d cannon tibia femur pelvis  20.6  7.92  -0.26  8.04  22.63  7.45 7.71  -0.14 -0.73 -0.47  length length length length length length length length length  cervical thoracic lumbar  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  3 5 7  1  P a t t e r n E . I . LHLLH ?  26.98 22.3  Intercept 8.97  5.62 4.96  7.30  4.46 4.01  7.57  8.26 8.22  x  1  105  Deviation +0.79  -0.31  -0.93 -0.88  +0.08 -0.88 +0.04  -0.61  5 10  3.31  7.06  1  4.76.  3 5  5.26  8.06 8.70  5.12  8.03  -1.12 -0.78 -0.12 +0.52 -0.15  4.80 4.40  7.73 6.71 6.36  -0.45 -1.47 -1.82  8.97 8.29  +0.11  5 5 3  cannon cannon cannon cannon  3.69  2.80 17.3  2.98 2.82 1.46  7.40  7.97 7.97  +0.79 -0.21 -0.21  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  37  Table A n i m a l No. X 7 Sex  M  T o t a l E.I  1  717,347  Calories P a t t e r n E . I . LMMLM  Age 3 2 2 Days  Intercept  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  16.4  15.68  7.55  length length length length  h i n d cannon tibia femur pelvis  20.13 26.45 23.23 23.35  6.98 7.44 8.07 8.66  length length length length length length length length length  cervical  3  5.22  4.90  6.33  5 7  thoracic  1  5 10  1  2  17.65  18.6  4.07 3.97 3.37 3.79  6.79 6.69  7.60  6.91  x  10  Deviation  1  5  -0.38 -0.48 +0.43 +0.38  -0.19 +0.27  +0.90 +1.49 -0.84  -0.26  6.78 7.39 7.36  -0.39 +0.22  7.98  +0.81  +0.19  4.62  7.33  +0.16  3 5  4.94  7.41  +0.24 +0.70  5 5 3  4.57  7.08  -0.09  3.05  9.24 8.14  +2.07 +0.97  cannon cannon cannon cannon  16.4 2.77 2.68  6.79 7.17  6.92  -0.38 0.00 -0.25  1.33  6.33  -O.84  lumbar  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  5.06  4.60  7.87  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  x  1  10  128 38  Table A n i m a l No. X8 Sex  M  Total E.I.  1  677,141  Calories P a t t e r n E.,1. LHLL  Age 322 Days  Intercept  Deviation  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  16.35 17.93 18.05  6.67 x 105  15.08  6.73  -0.10 x 1 0 +0.29 +0.06 -0.04  length length length length  h i n d cannon tibia femur pelvis  19.93  6.57 6.75 6.50 6.26  -0.20 -0.02 +0.27 -0.51  length length length length length length length length length  cervical  3 5 7 thoracic 1 5 10 lumbar 1 3 5  7.68 8.09 7.26 6.49 7.11 6.36 6.28 6.28  +0.91 +1.32 +0.49 -0.28 +0.34 -0.41 -0.49 -0.49 -0.19  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  5  5 3  cannon cannon cannon cannon  2  25.83  21.7 20.68 5.50 5.08  4.20 3.77 3.32 3.51 4.42 4.66 4.57 4.24  4.58 2.79 16.35 2.64 2.67 1.33  7.06 6.83  6.58  1  6.15 8.99 6.29  -0.62 +2.22  6.67 6.47  -0.10 +0.30 +0.07 -0.44  6.84  6.33  -O.48  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  1  5  Table 3 9 A n i m a l No. X9 Sex  M  Total E . I . 710,339 1  Age 3 2 2 Days  Calories P a t t e r n E . I . MHLLL 1  Deviation  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  16.88 18.03 17.85 15.63  7.95 x 1 0 7.19 6.55 7.47  length length length length  h i n d cannon tibia femur pelvis  20.63  7.98  length length length length length length length length length  cervical  3 5 7 thoracic 1 5 10 lumbar 1 3 5  5.42 4.78 4.12 3.93 3.37 3.55 4.70 4.91 4.69  7.29 6.12 6.96 7.21 7.36 6.59 7.75 7.29 6.89  +0.19 -0.98 -0.14 +0.11 +0.26  width width width  cervical thoracic lumbar  4.82  7.79 4.05 7.00  +0.69 -3.05 -0.10  length  fore fore fore fore  7.95  +O.85  5 5 3  cannon cannon cannon cannon  2  25.9  22.08 21.8  4.19 2.89 16.88 2.71 2.71 1.44  Intercept  6.83  6.89 7.27  6.84  7.14  7.72  5  1  +0.85 x 10 +0.09 -0.55 +0.37 +0.88 -0.27 -0.21 +0.17  -0.51  +0.65 +0.19 -0.21  -0.26 +0.04 +0.62  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  Table A n i m a l No. Sex  X10  M  Total E . I .  1  40  758,896  Calories P a t t e r n E . I . LHMLL  Age 3 2 2 Days  Dimension  Bone  Size'  Intercept" "  length length length length  f o r e cannon radius humerus scapula  16.45 18.3  6.92 7.55 7.74  length length length length  h i n d cannon tibia femur pelvis  length length length length length length length length length  cervical  3  thoracic  5 7 1  lumbar  5 10 1 3 5  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  5 5  3  cannon cannon cannon cannon  I8.7 15.2  19.9 26.5 22.4 20.95 5.33 4.88 4.29 4.13  3.28 3.77 4.53 4.99 4.86  1  6.89  x  10  5  Deviation -0.67  -0.04 +0.15 -0.70  6.51 7.50 7.22  -1.08  6.50  -1.09  6.86  -0.73 -0.81 0.00 +0.52 -0.67  6.78 7.59 8.11 6.92 7.86 6.86  7.61 7.34  -0.09 -0.37  +0.27 -0.73 +0.02 -0.25  +0.40  4.89 4.27 2.88  7.99 5.06 6.93  -2.53 -0.66  16.45  6.92 7.59 7.07 7.34  -0.67 0.00 -0.52 .-0.25  2.85  2.70 1.41  1  Apparent d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  x  1  10  T a b l e 41 A n i m a l No. X l l Sex  M  T o t a l E . I ,. !  850,914  P a t t e r n E., 1 . MHMLM  Age 3 2 2 Days  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  17.65  length length length length  h i n d cannon tibia femur pelvis  21.4 27.88 24.08  length length length length length length length length length  cervical  3 5 7 1  Calories  2  19.4 19.68 16.48  23.53  Intercept 9.81 x 8.98 9.10 8.62 9.53 9.04 8.93 8.83  1 0  1  5  Deviation +1.30 +0.47 +0.59 +0.11 +1.02 +0.53 +0.42 +0.32  -0.06  8.45 9.54 8.30 8.56  +1.03 -0.21 +0.05  5 10 1 3 5  5.66 5.30 4.48 4.23 3.73 3.83 4.94 5.18 5.44  9.15  +O.64  5 5 3  5.13 4.57 3.14  8.67 8.86 8.79  +0.16 +0.35 +0.28  cannon cannon cannon cannon  17.65 3.08  9.81 8.82  2.95 1.46  8.95 7.97  +1.30 +0.31 +0.44  thoracic lumbar  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  8.21 9.01 8.38 8.87  -0.30 +0.50 -0.13 +0.36  +O.46  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  x  1  10  T a b l e 42 A n i m a l No. X14 Sex  M  Total E.I.  1  807,518  Calories P a t t e r n E .1. MHHLL  Age 322 Days Bone  Size  length length length length  f o r e cannon radius humerus scapula  16.4 18.63 18.7 16.18  6.79 x 1 0 7.97 7.74 8.22  length length length length  h i n d cannon tibia femur pelvis  20.38  7.48 8.15 7.66 7.32  -0.60 +0.07  length length length length length length length length length  cervical  3 5 7 thoracic 1 5 10 lumbar 1 3 5  5.02 4.30 3.98 3.52 3.80 4.88 5.11 5.10  7.63 7.70 7.63 7.43 8.11 8.03 8.69 8.10 7.97  -0.45 -0.45 -0.65 +0.03 -0.05 +0.61 +0.02 -0.11  width width width  cervical thoracic lumbar  4.99 4.49 3.35  8.27 7.85 10.29  +0.19 -0.23 +2.21  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  16.4 3.27 2.94 1.50  6.79  -1.29 +1.76 +0.79 +0.40  5 5 3  cannon cannon cannon cannon'  2  27.0'8 22.83  21.85 5.49  Intercept  Deviation  Dimension  9.84  8.87 8.48  1  5  1  -1.29 x 10 -0.11 -0.34 +0.14  -0.42  -0.76  -0.38  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  43  Table A n i m a l No. X16 Sex  M  Dimension  Total E.I. Age  322  700,863  Calories P a t t e r n E . I . MMLLM  Day Size  Bone  2  Intercept  1  Deviation  1  *  5  length length length length  f o r e cannon radius humerus scapula  16.5 18.28 18.33 15.05  7.03 x 10  +0.02 x 10  7.22 6.69  +0.21 -0.32  length length length length  h i n d cannon tibia femur pelvis  20.0 26.85 21.98 20.7  6.71 7.89 6.81 6.28  -0.30 +0.88 -0.20 -0.73  length length length length length length length length length  3 5 7 thoracic 1 5 10 lumbar 1  5.18 4.78 4.13 3.65  -0.87 -0.89 -0.01 -1.06  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  cervical  7.52  +0.51  5  4.61 4.56  6.14 6.12 7.00 5.95 6.17 6.99 6.23 6.07 6.55  5 5 3  4.42 4.35 2.78  6.66 6.08 6.21  -0.35 -0.93 -0.80  cannon cannon cannon cannon  16.5 2.77  7.03 7.17 6.92  +0.02 +0.16 -0.09 -0.43  3  3.13  3.62 4.41  2.68  1.35  6.58  -O.84  -0.02 -0.78 -0.94 -O.46  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  Table 4 4 A n i m a l No. V7  T o t a l E.I  1  195,960  Calories P a t t e r n E . I . MH  Sex F  Age 112 Days  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  14-45  15.3 15.18 12.65  1.68 x 1 0 1.78 1.78 1.90  length length length length  h i n d cannon tibia femur pelvis  17.13 21.78 18.48 18.2  1.59 1.80 2.00 2.53  length length length length length length length length length  cervical  4.27  1.70 1.82 1.66 1.85 1.54 2.13 2.10 1.87 2.42  thoracic  3 5 7  1 5  lumbar  10 1 3 5  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  5 5 3  cannon cannon cannon cannon  2  3.96 3.42 3.24 2.68 3.00 3.86 3.97 4.00 3.57 3.76 2.32  14-45  2.62 2.50 1.23  Intercept  1.70 2.03  1.48 1.68 1.57 1.35 1.72  1  5  Deviation  1  -0.28 x 10 -0.18 -0.18 -0.06 -0.37 -0.16 +0.04 +0.57 -0.26 -0.14  -0.30 -0.11 -0.42 +0.17 +0.14  -0.09 +0.46 -0.26 +0.07 -0.48 -0.28 -0.39 -0.61 -0.24  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  T a b l e 45 A n i m a l No. V14  T o t a l E.I -  1  98,886  P a t t e r n E . I . ML  Sex -F  Age 112 Days  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  13.0 13.43  length length length length  h i n d cannon tibia femur pelvis  15.65 19.53  length length length length length length length length length  cervical thoracic  3 5 7  1 5  lumbar  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  10 1 3  2  13.2 10.6  Intercept  Deviation  1  1.14 x 1 0 1.22 1.12 1.11  5  15.03  +0.06 +0.11 +0.18 +0.05  3-76  1.22  1.03  +0.23 +0.10 +0.12 -0.11 +0.05 +0.20 +0.16 +0.08 +0.04  2.06  0.50 1.08 1.13  +0.09 +0.14  13.O 2.29 2.17 1.04  0.88 ^.48 0.88  +0.15 -O.ll -0.51 -0.11  15.B  3.40 3.00 2.55  2.18 2.53  3-19  3.39  3.29  5 5 3  3.19  2.99  1.09  1.11 0.88 1.04 1.19 1.15 1.07  1  +0.15 x 10 +0.23 +0.13 +0.12  1.05 1.10 1.17 1.04  5  cannon cannon cannon cannon  Calories  -0.49  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  136 Table 4 6 A n i m a l No. V15 Sex  F  Total E . I .  1  97,930  Age 112 Days  Dimension  Bone  length length length length  f o r e cannon radius humerus scapula  length length length length  h i n d cannon tibia femur pelvis  length length length length length length length length length  cervical  P a t t e r n E . I . LL Size  thoracic  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  ?  12.7 12.83 12.95 10.35  15.4  18.75 14.9 14.28  1.03 x 1.04 1.03 1.01  0.96 0.86 0.88 O.83  1.00 0.82 1.02 0.78  3 5  5 5 3  3.16 3.05 2.08  O.83  1  10 1  cannon cannon cannon cannon  12.7  2.24 2.18 1.09  1 1 Deviation  Intercept  3.52 3.19 2.93 2.47 2.29 2.36 3.02 3.19 3.19  3 5 7 5  lumbar  Calories  1.15  1.02 0.95 O.83  0.91 0.92 1.15  1.03 0.77 0.49 1.10  10  5  +0.05 x 1 0 +0.06 +0.05 +0.03 -0.02 -0.12 -0.10 -0.15  +0.02 -0.16 +0.04 -0.20 +0.17 +0.04 -0.03 -0.15  -0.07 -0.15  -0.06 +0.17 +0.05 -0.21  -0.49  -0.12  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  5  Table 4 7 A n i m a l No. Y7  Total E.I.  Sex F  Age 112  Dimension  Bone  length length length length  f o r e cannon radius humerus scapula  length length length length  h i n d cannon tibia femur pelvis  length length length length length length length length length  cervical  216,116  Days  P a t t e r n E . I . MH Size  thoracic  3 5 7  1  Calories  o  1  Intercept x  -O.46  14.5 15.4  1.70 1.81  16.05  2.07 2.19  -0.09 + 0.03  1.64  -0.52 -0.13 +0.14  1.90  -0.26 +0.34  13.4 17.3. 22.55  19.3 4 • 48 4.49  3-92 3.06  2.03 2.30  2.50  2.32 1.60  10  Deviation  5  -0.35  +0.16 -0.56  5 10  lumbar  1 3 5  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  5 5 3  cannon cannon cannon cannon  3.55  1.65  -0.51  14.5 2.86 2.58 1.31  1.70 2.07 1.56 2.05  -O.46  -0.09 -0.60 -0.11  1  Apparent d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  1  Table 4 8 A n i m a l No.  U10  Total E.I.  1  337,299  P a t t e r n E . I . MHM  Sex F  Age 1 7 5 Days  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  15.85  -  Calories  2  16.68 17.0  Intercept 3.50 3.48 3-54  14.65  3.90  19.2-  3.40  x  Deviation  1  10  5  +0.13 +0.11 +0.17 +0.53  length length length length  h i n d cannon tibia femur pelvis  length length length length length length length length length  cervical  3  4.66 3.74  3.38 3.01  thoracic  5 7 1  3.40  3.58  3.06  3.67 4.22  +0.21 +0.30 +0.85  lumbar  5 10 1 3 5  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  24.8 20.98 20.8  4.40  3.54 4.51 4.74 4.53  4.40  3.74 3.96 4.07  3.36  +0.03 +0.37 +0.59 +0.70 -0.01 +0.01  -O.36  4.27  +0.90  4.10 3.88  +0.73 +0.51  3.80  +0.43 -0.47 -0.30  5 5 3  3.98  2.90  2.73  3.07  cannon cannon cannon cannon  15.85 2.65 2.66 1.36  3.50 2.83 3.10 3.75  +0.13 -0.54  -0.27 -O.38  1  Apparent d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  x  1  10  139 Table A n i m a l No. U l l Sex  F  Total E.I.  1  49  352,929  Calories P a t t e r n E .1.  Age 175 Days  m  Dimension  Bone  Size  2  Intercept  1  Devia  length length length length  f o r e cannon radius humerus scapula  15.68 16.7 17.05 14.6  3.39 x 1 0 3.49 3-57 3.85  5  -0.14 x 10' -0.04 +0.04 +0.32  length length length length  h i n d cannon tibia femur pelvis  18.95 24-23 20.93 20.88  3.25 3.25 3.92 4.12  -0.28 -0.28 +0.39 +0.59  length length length length length length length length length  cervical  3 5 7 thoracic 1 5 10 lumbar 1 3 5  5.06 4.52 3.80 3-42 3.13 3-35 4-14 4.48 4.12  4.38 3.77 3.22 3.65 3.91 3.68 3.36 3.47 2.73  +O.85 +0.24 -0.31 +0.12 +0.3-8 +0.15 -0.17 -0.06 -0.80  width width width  cervical thoracic lumbar  5 5 3  4.15 4.12 2.71  3.23 3.38 3.06  -0.30 -0.15 -0.47  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  cannon cannon cannon cannon  15.68 2.76 2.63 1.25  3.39 3.34 3.01 2.86  -0.14 -0.19 -0.52 -0.67  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  1  Table 5 0 A n i m a l No.  Total E.I.  U14  380,424  1 7 5 Days  Sex F  Age  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  15.48  length length length length  h i n d cannon tibia femur pelvis  18.68  length length length length length length length length length  cervical thoracic  3 5 7 1 5 10 1  Pattern E.I. 2  16.15 17.3  14.7  24.10 20.93 20.4 5.06 4.63 4.18 3.70 3.22  fore fore fore fore  3.80 4.38 4.12 4.44 4.68 4.21 4.22  10  5  -0.55 -0.65 -0.05 +0.15  -0.69 -0.66 +0.12 0.00 +0.58 +0.32  +O.64 +0.88  +0.41  3.93 3.83 2.94  +0.13 +0.03 -0.86  15.48 2.78 2.63 1.32  3.25 3.42 3.01  -0.55 -0.38  cannon cannon cannon cannon  length width, d i s t a l  3.H 3.14 3.92  x  4.46 4.24 2.58  5 5 3  cervical thoracic lumbar  3.25 3.15 3.75 3.95  Deviation  1  3.65 3.63  3 5  width width width  Intercept  MMM  +0.42 -0.19 -0.15 -0.17  3.54 4.24 4.55 4.44  lumbar  Calories  3.61  3.35  -0.79 -0.45  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  x  1  10  Table 5 1 A n i m a l No. U32  Total E.I.  198,$27  Sex F  Age 1 7 5 Days  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  13.98 14.78  15.03 12.25  length length length length  h i n d cannon tibia femur pelvis  21.75 17.9 17.33  length length length length length length length length length  cervical  3 $ 7 thoracic 1 $ 10 lumbar 1 3 $  width width width  cervical thoracic lumbar  length width, d i s t a l  fore fore fore fore  Calories P a t t e r n E . I . LMM  2  Intercept" '  Deviation  2.21 x 1 0 2.29  +0.30  1  2.38  2.18  5  +0.22 x 10  +0.39 +0.19  2.10 2.28  +0.11 +0.29 +0.39 +0.06  4.07 4.00 3.42 2.96 2.78 3.13 3.73 4.00 4.02  2.04 2.46 2.15 2.44 2.72 3.05 2.37 2.37 2.55  +0.05 +0.47 +0.16 +0.45 +0.73 +1.06  $ 5 3  3.59 3.48 2.45  2.35 2.08  +0.36 +0.09  cannon cannon cannon cannon  13.98 2.39 2.33 2.14  2.21 2.00 2.10 2.16  16.83  2.38 2.05  2.82  1  +O.38 +0.38 + O.56  +0.83  +0.22 +0.01 +0.11 +0.17  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  Table 5 2 A n i m a l No. U38  T o t a l E.I  202,751  Calories P a t t e r n E,. 1 .  Sex F  Age 1 7 5 Days  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  13.8 14.45 14.73 12.1  2.10 x 10  length length length length  h i n d cannon tibia femur pelvis  16.98 21.35 17.78 17.9  2.17  length length length length length length length length length  cervical  3 5 7 thoracic 1 5 10 lumbar 1 3 5  4.25 4.08  2.32 2.58  3.15  2.80  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  5 5 3  cannon cannon cannon cannon  2  3-38 2.73 2.79 3.83  Intercept  1  5  2.08  2.20 2.11 2.15  2.33  2.31  2.07  2.55 2.21 2.61  LMM  Deviation  +0.07 x 10 +0.05 +0.17 +0.08  +0.14 +0.12 +0.30 +0.28 +0.29 +0.55 +0.04 +0.77 +0.52  +0.18  +O.58  2.47  +0.29 +0.44  3.53 3-73 2.55  2.29 2.50 2.91  +0.26 +0.47 +0.88  13.8 2.34 2.43  2.10 1.87  +0.07 -0.16 +0.22 +0.67  3.97 3.97  1.23  2.32  2.25  2.70  1  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  Table 5 3 A n i m a l No. V9  Total E.I.  193,545  Calories P a t t e r n E . I . MLM  Sex F  Age 175 Days  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  13.68 14.13  2.00 x 1 0 1.90  11.53  length length length length  h i n d cannon tibia femur pelvis  16.8 20.8 16.73  length length length length length length length length length  cervical  3 5 7 thoracic 1 5 10 lumbar 1 3 5  width width width  cervical thoracic lumbar  length width,  fore fore fore fore  distal  2  14.1  15.55  3.76 3.54 3.03 2.87  2.47 3.26 3.52  3.58  Intercept  1  Deviation  1  I.85  1.80  +0.06 x 10 -0.04 -0.09 -0.14  2.08 1.93 1.90 1.20  +0.14 -0.01 -0.04 -0.74  1.52  -0.42  5  1.81 1.36 2.27 2.03  -0.13 -0.58 +0.33 +0.09  1.66 1.61 I.87  -0.28 -0.33 -0.07  5 5 3  3.07 3.32  1.78  1.83 1.82 2.21  -0.11 -0.12 +0.27  cannon cannon cannon cannon  13.68 2.20 2.20 1.06  2.00 1.10 1.70 1.75  +0.06 -O.84  -0.24 -0.19  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  Table A n i m a l No.  V13  Total E . I .  1  54 Calories  357,987  Sex F  Age 1 7 5 Days  Dimension  Bone  length length length length  f o r e cannon radius humerus scapula  14.0  3.26  length length length length  h i n d cannon tibia femur pelvis  I8.58 24.15 20.75 19.33  3 3 3 3  length length length length length length length length length  cervical  P a t t e r n E . I . MMM Size  thoracic  3 5 7 1 5 10 1  2  15.33  16.7 16.98  4.67 4.35 3.79 3.33  2.96  Intercept 3.14 3.49 3.35  x lO^  1  Deviation -  0 0 0 0  . . . .  4 0 2 3  4 9 3 2  .05 .18 .76 .09  -0.53 -0.40 +0.18  3.38 3.23 3.18 3.32 3.33  -0.20  -0.49  -0.35  -0.40 -0.26 -  0 0 0 0  .25 .42 .41 .23  3.16  3 5  3.17 4.06 4.43 4.37  5 5 3  3.99 3.94 2.64  2.87 2.84 2.99  -0.71 -0.74 -0.59  cannon cannon cannon cannon  15.33 2.64 2.43  3.14 2.74 2.40 3.10  -0.84 -1.18  lumbar  width width vri. d t h  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  1.28  3.17 3.35 3.44  -0.14  -O.44  -O.48  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  x  1  10  145  Table 5 5 A n i m a l No. V16 Sex  F  T o t a l E.I  •  197,719  Age 1 7 5 Days  Pattern E.1-  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  13.88  length length length length  h i n d cannon tibia femur pelvis  length length length length length length length length length  cervical thoracic lumbar  3 5 7 1 5 10 1  17.35 21.1  16.98 16.3 3.78  1.96 2.05  1.96  2.38 2.04 1.87 1.56 1.56  x  10  5  Deviation +0.17 -0.02 +0.07 -0.02 +0.40 +0.06 -0.11 -0.42  1.82  -0.16  3.30 3.46 2.15  2.07 2.05 2.54  +0.09 +0.07  13.88 2.30 2.33 1.09  2.15 1.78 2.10 1.90  +0.17 -0.20 +0.12 -0.08  2.79 2.31 2.66  cannon cannon cannon cannon  fore fore fore fore  2.15  1  1.96  3.18  5 5 3  length width, d i s t a l width, proximal w i d t h , minimum  Intercept  MLL  -0.42 -0.07 -0.31 +0.14 -0.28 +0.02 -0.17 -0.02  3.61  3.37 3.74 3.55  cervical thoracic lumbar  2  14.25 14.45 11.85  3 5  width width width  Calories  1.91 1.67 2.12 1.70 2.00 1.81  +O.56  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  x  1  10  5  T a b l e 56 A n i m a l No. V17  Total E.I.  166,743  Calories P a t t e r n E .1. L L L  Sex F  Age 175 Day;  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  13.75  length length length length  h i n d cannon tibia femur pelvis  16.75 20.8 16.5 16.33  length length length length length length length length length  cervical  3 5 7 thoracic 1 5 10 lumbar 1 3 5  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  2  14.33  14.05  11.63  Intercept 2.06 x 1 0 2.01 1.82 1.86  Deviation  1  5  +0.39 x 10 +0.34 +0.15 +0.19  2.05  +O.38  1.94 1.80 1.56  +0.27 +0.13 -0.11  3.79 3.42 3.10 2.60 2.33 2.56 3.26 3.57  1.57 1.63  3.50  1.67 1.69 1.74  -0.10 -0.04 -0.17 +0.10 +0.07 +0.16 0.00 +0.02 +0.07  5 5 3  3.49 3.36 2.18  2.25 1.88 2.57  +0.21 +0.90  cannon cannon cannon cannon  13.75  2.06 2.00 2.37 2.11  +0.39 +0.33 +0.70 +0.44  2.39 2.42 1.13  1.50  1.77 1.74 1.83  1  +O.58  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  147 ( t e x t c o n t i n u e d from p. For  56)  each s k e l e t a l d i m e n s i o n a t 112 and 175 d a y s , t h e d a t a have  been bounded by a t r i a n g l e . The r e s u l t i n g f i g u r e bounds the d i s t r i b u t i o n o f d a t a i n a unique manner and has a t t r i b u t e s t h a t have proven u s e f u l f o r describing that d i s t r i b u t i o n . according to a s t r i c t of  The t r i a n g l e s were drawn  p r o c e d u r e and t h e consequent shapes are  considerable importance.  F o r t h e purpose o f i d e n t i f i c a t i o n  i n t h e f o l l o w i n g d e s c r i p t i o n , the lower-most apex w i l l be A, t h e upper l e f t B, and t h e upper r i g h t C.  called  The s i d e s a r e  i d e n t i f i e d by t h e i r a d j a c e n t a p i c e s as AB, BC and AC.  Apex A  was chosen t o c o r r e s p o n d t o an energy l e v e l below any a c h i e v e d by t h e l a b o r a t o r y a n i m a l s and p r o b a b l y a p p r o a c h i n g t h e c r i t i c a l energy i n t a k e f o r s u r v i v a l o f a w i l d  fawn.  The h e i g h t , w i d t h , and shape o f each bounding t r i a n g l e i s a p r o d u c t o f t h e i n t e r a c t i o n o f energy i n t a k e and t h e p r i o r i t y o f a p a r t i c u l a r bone t o grow.  The energy v a l u e s  chosen f o r A were 75,000 C a l o r i e s (A.D.E.) a t 112. days and 150,000 C a l o r i e s (A.D.E.) a t 175 days. to  t h e r i g h t o f t h e d i s t r i b u t i o n o f d a t a i n a manner t h a t  just included a l l data points. of  A l i n e AC was drawn  From the p o i n t o f i n t e r s e c t i o n  t h i s l i n e , AC, w i t h the chosen minimum energy v a l u e , A, a  l i n e AB was drawn t o the l e f t o f , and j u s t i n c l u d i n g , t h e • d i s t r i b u t i o n of data.  The l i n e BC, w i t h z e r o o r p o s i t i v e  s l o p e , was drawn t o bound the d a t a from above and t o complete the  triangular For  shape.  t h e purpose o f l o c a t i n g s p e c i f i c d a t a p o i n t s w i t h i n  t h e boundary, a r e f e r e n c e l i n e has been i n c l u d e d . d e s c r i b e s the mean c u m u l a t i v e energy  This l i n e  i n t a k e a t each v a l u e f o r  a s k e l e t a l measurement a c c o r d i n g t o the boundary o f the d a t a and not t o the i n d i v i d u a l d a t a p o i n t s .  T h i s method  was  chosen because the d i s t r i b u t i o n o f t r e a t m e n t s used i n the experiment treatments.  was  not c o n s i d e r e d r e p r e s e n t a t i v e o f a l l p o s s i b l e  The  a d d i t i o n o f a new  t r e a t m e n t may  l i n e f i t t e d t o t h e d a t a p o i n t s but i t w i l l be  change a  demonstrated  l a t e r t h a t t h e a d d i t i o n o f t h i s t r e a t m e n t s h o u l d not a l t e r the boundaries.  I d e a l l y t h e r e f e r e n c e l i n e s h o u l d be  an  a s y m p t o t i c curve d e s c r i b i n g growth on i n v a r i e n t p a t t e r n s , but d i f f e r e n t magnitudes o f energy  intakes.  However, s u f f i c i e n t  d a t a were not a v a i l a b l e t o e s t a b l i s h such l i n e s .  A straight  l i n e i s no l e s s u s e f u l f o r e s t i m a t i n g p a t t e r n o f  energy  i n t a k e t h a n i s the i d e a l c u r v e , f o r i t i s t h e p a t t e r n o f d i s p l a c e m e n t which i s o f i n t e r e s t ; but i n t e r p r e t a t i o n o f t h e d a t a i s made more d i f f i c u l t . On each graph are shown b o t h male and female d a t a .  Only  the male d a t a were used i n f i t t i n g t h e bounding l i n e s , w i t h t h e female d a t a b e i n g added a f t e r w a r d s .  T h i s was  done i n  o r d e r t o a c c e n t u a t e any p o s s i b l e sex d i f f e r e n c e s i n the response  to a treatment.  Both male and female d a t a are  cussed a c c o r d i n g t o d i s p l a c e m e n t  dis-  from t h e same r e f e r e n c e l i n e .  At 322 days, a d i f f e r e n t approach  was n e c e s s a r y .  ments a t e a r l i e r d a t e s i n c l u d e d an a r r a y o f energy r a n g i n g from t h e l o w e s t t o the h i g h e s t extremes,  Treat-  intakes  along with  many i n t e r m e d i a t e p o s s i b i l i t i e s .  A l l of these c o n t r i b u t e d to  e s t a b l i s h i n g the t r i a n g u l a r boundaries.  W i t h the  limited  number o f t r e a t m e n t s c o n s i d e r e d at 3 2 2 days, the t r i a n g l e s c o u l d not be drawn.  F o r example, a c o n t i n u a t i o n o f c o n s t a n t  'L' t o 3 2 2 days would have r e s u l t e d i n a c u m u l a t i v e i n t a k e o f around  420,000  energy  C a l o r i e s , whereas t h e l o w e s t a c t u a l  t r e a t m e n t c o n s i d e r e d r e s u l t e d i n a c u m u l a t i v e energy i n t a k e of  675,000  Calories.  A l i n e was  f i t t e d t o the data p o i n t s  u s i n g a ' l e a s t squares r e g r e s s i o n ' . As mentioned  earlier,  t h i s t r e a t m e n t i s not v a l i d s t a t i s t i c a l l y because the v a r i a t i o n s are known not t o be random.  However, s i n c e the  same t r e a t m e n t s are i n v o l v e d i n each g r a p h , the l i n e p r o v i d e s a convenient r e f e r e n c e .  F o r each a n i m a l c a r r i e d t o 3 2 2 days,  t h e magnitude o f each s k e l e t a l d i m e n s i o n a t 1 7 5 days o f age c o u l d be e s t i m a t e d from t h e energy regime t h i s date.  e f f e c t i v e p r i o r to  T h i s a l l o w s t h e growth between 1 7 5 and 3 2 2 days t o  be equated t o t h e p r e v a i l i n g energy  treatment.  Once the graphs f o r the e x p e r i m e n t a l d a t a were drawn, i t was n e c e s s a r y t o d e v e l o p a method t h a t would a l l o w s k e l e t a l measurements from a n i m a l s o f unknown n u t r i t i o n a l h i s t o r y t o be used t o e v a l u a t e the energy regime o f t h e s e a n i m a l s .  For  each  o f t h e a n i m a l s r e a r e d at the l a b o r a t o r y , t o t a l energy  intake,  p a t t e r n o f i n t a k e , and a l l o f the s k e l e t a l measurements are known.  F o r t h e w i l d a n i m a l s , o n l y the s k e l e t a l measurements  and, i n some c a s e s , an e s t i m a t e o f t o t a l energy i n t a k e based on a body weight r e g r e s s i o n are known.  N o t h i n g i s known o f  150 t h e p a t t e r n o f energy i n t a k e . In t h e f o l l o w i n g s e c t i o n , u n l e s s o t h e r w i s e i n d i c a t e d , t h e procedure  w i l l r e f e r t o t h e a n a l y s i s o f a n i m a l s a t 175 days o f  age o r younger.. I t has a l r e a d y been suggested  t h a t a t i m e s p e c i f i c energy  r e s t r i c t i o n would a f f e c t each s k e l e t a l d i m e n s i o n  differently.  These d i f f e r e n c e s a r e m a n i f e s t e d i n d i s p l a c e m e n t  from t h e r e -  ference l i n e s .  The d i s p l a c e m e n t  s k e l e t a l dimension  i s the r e s u l t of p l o t t i n g a  a g a i n s t a known c u m u l a t i v e energy i n t a k e .  I f o n l y the s k e l e t a l dimension  i s known, t h e r e f e r e n c e  line  w i l l p r o v i d e an e s t i m a t e o f t h e c u m u l a t i v e energy i n t a k e . dimension  Each  would p r o v i d e an independent e s t i m a t e o f energy  i n t a k e , and t h e d i f f e r e n c e s among t h e s e e s t i m a t e s would depend upon t h e p a t t e r n o f energy r e s t r i c t i o n s w h i c h had o c c u r r e d . F o r t h e l a b o r a t o r y a n i m a l s , t h e d e v i a t i o n s between t h e t r u e and t h e e s t i m a t e d c u m u l a t i v e energy i n t a k e s have been measured f o r each d i m e n s i o n arranged  o f every animal.  The d e v i a t i o n s have been  i n s e r i e s as f o l l o w s :  1.  Length  o f f o r e l i m b elements,  d i s t a l to proximal;  2.  Length  o f h i n d l i m b elements,  d i s t a l to proximal;  3.  Length o f v e r t e b r a e , a n t e r i o r t o p o s t e r i o r ;  4.  Width o f v e r t e b r a e , a n t e r i o r t o p o s t e r i o r ;  5.  D i m e n s i o n s o f t h e f o r e cannon, a r r a n g e d  i n the  order of length, d i s t a l width, proximal width, minimum w i d t h . T h i s i n f o r m a t i o n i s shown i n T a b l e s 10 t o 56, a l o n g w i t h t h e  151 a p p r o p r i a t e energy regime, t o p r o v i d e t h e s t a n d a r d s f o r t h e e v a l u a t i o n o f t h e energy regimes o f t h e f r e e r a n g i n g w i l d fawns. Two s i t u a t i o n s may e x i s t i n t h e e v a l u a t i o n o f unknown data.  I n one c a s e , b o t h body weight and s k e l e t a l d i m e n s i o n s  are a v a i l a b l e , and i n t h e o t h e r , o n l y s k e l e t a l d i m e n s i o n s . When body w e i g h t i s a v a i l a b l e t o p r o v i d e an e s t i m a t e o f t o t a l energy i n t a k e , t h e n t h e t r e a t m e n t i s e s s e n t i a l l y t h e same as j u s t o u t l i n e d f o r the laboratory animals.  When o n l y  skeletal  d i m e n s i o n s a r e a v a i l a b l e , a l l t h a t can be o b t a i n e d i s a s e r i e s of i n t e r c e p t s with the reference l i n e s .  In the l a t t e r  s i t u a t i o n , a n a l y s i s becomes much more d i f f i c u l t because no base l i n e i s a v a i l a b l e .  The r e s u l t s o f t h e e v a l u a t i o n o f t h e  energy r e g i m e s o f t h e w i l d fawns a r e shown i n T a b l e s 57 t o 65.  The i n t e r c e p t s w i t h t h e r e f e r e n c e l i n e s a r e t a b u l a t e d  and, where a body w e i g h t e s t i m a t e o f t o t a l energy i n t a k e was a v a i l a b l e , t h e d e v i a t i o n s between t h e energy i n t e r c e p t s f o r each s k e l e t a l d i m e n s i o n and t h e energy i n t a k e e s t i m a t e d by body weight have been shown.  ( t e x t c o n t i n u e s on p. 161)  152 Table A n i m a l No. YF17  Total E.I.  57  197,000 C a l o r i e s  Estimated  Pattern E.I. Wild  Sex M  Age 116 Days  Dimension  Bone  Size'  Intercept  length length length length  f o r e cannon radius humerus scapula  14.63  15.9 12.95  1.74 x 105 1.78 2.02 2.00  -0.23 -0.19 +0.05 +0.03  length length length length  h i n d cannon tibia femur pelvis  17.7 22.7 19.0  1.80 2.08 2.18  -0.17 +0.11 +0.21  length length length length length length length length length  cervical  4.35 3.98  1.77 1.85  -0.20 -0.12  4.10  2.80  +O.83  1.74 2.00 1.45 2.03  -0.23 +0.03 -0.52 +0.06  3 5 7  thoracic  15.3  1  Deviation  1  5 10  lumbar  1  3 5  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  5 5 3  cannon cannon cannon cannon  14.63 2.83 2.54 1.30  1  Apparent d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  x  1  10'  153 Table 5 8 A n i m a l No. YF19 Sex  M  Total E.I.  205,000 C a l o r i e s  Estimated  Pattern E.I. Wild  Age 118 Days  Dimension  Bone  Size'  Intercept  length length length length  f o r e cannon radius humerus scapula  15-38  16.0 16.15 12.75  2.03 x 101.98 ,10 .93  -0.02 x 10-0.07 +0.05 -0.12  length length length length  h i n d cannon tibia femur pelvis  18.23 23.1 19.55  1.98 2.20 2.37  -0.07 +0.15 +0.32  length length length length length length length length length  cervical  4.32 4.08  1.74 1.98  -0.31 -0.07  5 5 3  4.13  2.02  +0.87  cannon cannon cannon cannon  15.38 3.07 2.84 1.33  2.03 2.50 2.25 2.16  -0.02 +0.45 +0.20 +0.11  3 5 7  thoracic  1  Deviation  1 5  lumbar  10 1 3 5  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  1  154  Table 5 9 A n i m a l No. YF16  T o t a l E . I . 193,000 1  Calories  Estimated  Pattern E.I. Wild  Sex F  Age 116 Days  Dimension  Bone  Size  Intercept  Deviation  length length length length  f o r e cannon radius humerus scapula  14- 95 15- 3 15.7 13.0  1.87 x 101.78 1.95 2.02  -0.11 x 10 5 -0.20 -0.03 +0.04  length length length length  h i n d cannon tibia femur pelvis  17.35 22.7 18.95  1.85 2.08 2.17  -0.13 +0.10 +0.19  length length length length length length length length length  cervical  4.32 4.12 3.60  1.74 1.90  -0.24 +0.05 -0.08  5 5 3  3.86  2.30  +0.32  cannon cannon cannon cannon  14.95 2.81 2.49 1.27  1.87  -  o  3 5 7  thoracic  1  1  2.03  5 10  lumbar  1  3 5  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  1.96 1.32 1.90  0 0 0 0  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  .11 .02 .66 .08  1  T a b l e 60 A n i m a l No. YF20  T o t a l E . I . 16*0,000 1  Sex F  Age 118 Days  Dimension  Bone  length length length length  f o r e cannon radius humerus scapula  length length length length  h i n d cannon tibia femur pelvis  length length length length length length length length length  cervical  Calories  Estimated  Pattern E.I. Wild Size'  Intercept  14.95 15-1  1.87  1.72  +0.07 -0.08  12.6  1.87  +0.07  18.03 22.8  1.92 2.11  18.55  2.05  +0.12 +0.31 +0.25  4.22 3.78  1.65 1.59  -0.15 -0.21  5 5 3  3.75  2.07  +0.27  cannon cannon cannon cannon  14.95 2.69 2.61 1.22  1.87 1.72 1.63 1.93  +0.07 -0.08  thoracic lumbar  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  3 5 7 1  x  10  5  1  Deviation  5 10 1 3 5  -0.17 +0.13  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  x  1  10  T a b l e 61 A n i m a l No.  YF18  Total E.I.  1  Calories  170,000  Estimated  Sex F  Age 1 1 7 Days  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  15.2  1.96 x  15.83 15.8  1.93 1.98  +0.26 +0.23 +0.28  12.5  I.84  +0.14  length length length length  h i n d cannon tibia femur pelvis  18.38 22.73 18.93 17.75  2.04 2.08 2.16 2.07  +0.34 +0.38 +0.46 +0.37  length length length length length length length length length  cervical  4.18  r.62  3-83 3.49 2.98  1.78  I.48  -0.08 +0.08 +0.03 -0.22  1.92  +0.22  thoracic lumbar  3 5 7 1 5 10 1  P a t t e r n ;E . I . W i l d 2  2.92 3.90  1.94  +0.24  2.51  1.86  +0.16  1.96  +0.26 +0.06 -0.30 -0.24  cannon cannon cannon cannon  15.2 2.71 2.52 1.17  fore fore fore fore  +O.48  3-69  5 5 3  length width, d i s t a l width, proximal w i d t h , minimum  2.18 1.78  5  Deviation'  +0.08 -0.39  3.93 3-51  cervical thoracic lumbar  1.75  10  1  1.31  3 5  width width width  Intercept  1.76 1.40 1.46  1  Apparent d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  x  10  T a b l e 62 A n i m a l No. WF1  T o t a l E . I . 310,000  Sex M  Age 180 Days  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  15.28  length length length length  h i n d cannon tibia femur pelvis  length length length length length length length length length  cervical  3 5 7 thoracic 1 5 10 lumbar 1 3 5  width width width  cervical thoracic lumbar  length width,  distal  fore fore fore fore  5 5 3  cannon cannon cannon cannon  Calories  Estimated  Pattern E.I. Wild 2  15.9 16.65 12.9  Intercept  1  3.10 x 1 0 3.00 3.33  5  Deviation  0.00 x 10 -0.10  2.28  +0.23 -0.82  3.06 2.88 3.29 2.92  -0.04 -0.22 +0.19 -0.18  4.51 4.17 3-74  2.97  -0.13 -0.39 -0.08 +0.07 +0.13 +0.06  4.46 4.23  3.43  +0.33  4.13 3.73  3.18 2.50  2.46  2.83  +0.08 -0.60 -0.27  15.28  3.10 3.29 3.04 3.59  0.00 +0.19 -0.06 -0.49  18.6  23.4 20.1 19.1  2.71  3.02 3.17 3.23 2.93 3.17 . 3.16 3.29  2.75 2.64 1.34  3-05  1  -0.05  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  Table 6 3 A n i m a l No. WF2  T o t a l E . I . 320,000 1  Calories  Estimated  Pattern E.I. Wild  Sex M  Age 180 Days  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  15.20 16.1 16.2 13.65  3.04  length length length length  h i n d cannon tibia femur pelvis  18.3  2.90 2.96 3.13  -0.30 -0.24 -0.07  length length length length length length length length length  cervical  3 5 7 thoracic 1 5 10 1 lumbar 3 5  4.54 4.43 3.76 3.39 2.78 3.22 3-84 4.36 4.28  3.04 3.49 3.08 3.54 2.71 3.30 2.73 3.18 3.18  -0.16 +0.29 -0.12 +0.34 -0.49 +0.10 -0.47 -0.02 -0.02  width width width  cervical thoracic lumbar  5 5 3  4.23 4.05 2.65  3.42 3.13 3.00  +0.22 -0.07 -0.20  cannon cannon cannon cannon  15.20 2.85 2.74 1.42  3.04 3.75 3.38 4.23  -0.16 +0.55 +0.18 +1.03  length width,  distal  fore fore fore fore  2  23.6  19.7  Intercept  3.12 3.07 2.93  x 10  Devial  1  5  -0.16 x 10 -0.08 -0.13 -0.27  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  64  Table A n i m a l No.  YF38  Total E.I.  320,000  Estimated  Pattern E. 1 .  Sex M  Age 1 7 7 Days  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  15.9 16.5 16.2  length length length length  h i n d cannon tibia femur pelvis  length length length length length length length length length  cervical  3 5 7 1  Calories  2  Intercept 3.53  x  10  1  5  Wild  Deviation +0.33 +0.17 -0.13  12.85  3.37 3.07 2.51  19.25  3.50  23.7  2.99  +0.30 -0.21  19.3  2.96  -0.69  -0.24  4.13 4.54  2.13 3.83  -1.07 +0.63  5 5 3  3.66  2.42  -0.78  cannon cannon cannon cannon  15.9 2.88  3.53 3.88 3.38 3.19  +0.33 +0.68 +0.18 -0.01  thoracic lumbar  5 10 1 3 5  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  2.74 1.29  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  x  1  10  :  160 Table 6 5 A n i m a l No. YF37  T o t a l E.I •  1  265,000  C a l o r i e s Estimated P a t t e r n E .1. W i l d  Sex F  Age 176 Days  Dimension  Bone  Size  length length length length  f o r e cannon radius humerus scapula  14.58 15.35 15.23 12.85  2.64 2.64  length length length length  h i n d cannon tibia femur pelvis  18.03  2.75  length length length length length length length length length  cervical thoracic lumbar  width width width  cervical thoracic lumbar  length width, d i s t a l width, proximal w i d t h , minimum  fore fore fore fore  2  23.08 18.9  I8.48  Intercept  2.51 2.51  2.77 2.79  2.60  3  4.12  5 7  3.99  1  3.00  5  2.64 2.95  2.37  3.79  2.03  10 1 3 5 5 5 3  cannon cannon cannon cannon  3.39  3.90  3.69 3.73  2.20 14.58  2.54  2.40 1.23  2.12 2.45 2.09 2.51 2.52  Deviation  1  x 10  5  -0.01 x 105 -0.01 -0.14 -0.14  +0.10 +0.12 +0.14  -0.05  -0.53  -0.20 -0.54  -0.14 -0.28  -0.13 -0.62  2.34  -0.21  2.45 2.50  -0.15  2.59  2.64 2.37  2.32 2.70  1  -0.20 -0.06 -0.01  -0.28  -0.33 +0.05  1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  2  A c t u a l bone d i m e n s i o n i n c e n t i m e t e r s .  (text continued  from p.  151)  Discussion: The e v a l u a t i o n o f t h e n u t r i t i o n a l regime t o which an a n i m a l has been exposed can be approached from two r e l a t e d , but d i s t i n c t a s p e c t s :  an e s t i m a t i o n o f t h e t o t a l amount o f  energy consumed, and t h e d e t e r m i n a t i o n energy i n t a k e .  o f the p a t t e r n o f  I n t h e f o l l o w i n g pages the a n a l y s i s o f  n u t r i t i o n a l regime w i l l p r o c e e d from the use o f s k e l e t a l growth and body w e i g h t t o e s t i m a t e t h e use o f t h e d i m e n s i o n a l  t o t a l energy i n t a k e , t o  differences i n selected  skeletal  e l e m e n t s t o demonstrate t h e p a t t e r n o f energy i n t a k e — i . e . t h e t i m e , d u r a t i o n and e x t e n t o f energy r e s t r i c t i o n s .  1.  E s t i m a t i o n o f C u m u l a t i v e Energy  Intake.  The e s t i m a t i o n o f t o t a l energy i n t a k e by  skeletal  measurements i s based on t h e premise t h a t t h e b o u n d a r i e s drawn on F i g u r e s 9 t o 50 i n c l u d e a l l p o s s i b l e v a r i a t i o n s i n s k e l e t a l s i z e a t each l e v e l o f energy i n t a k e .  This i s quite  u n l i k e l y since the boundaries simply define the d i s t r i b u t i o n o f a s m a l l sample o f a n i m a l s drawn from a p o p u l a t i o n o f unknown v a r i a b i l i t y and i n v o l v e a r e s t r i c t e d number o f t r e a t ments.  An e r r o r term w i l l be a s s o c i a t e d w i t h each measurer  ment, but t h i s w i l l not be c o n s i d e r e d  until  later.  An e x a m i n a t i o n o f F i g u r e s 9 t o 50 w i l l show t h a t f o r each measurement o f a s k e l e t a l d i m e n s i o n t h e r e i s a c e n t r a l v a l u e o f t o t a l energy i n t a k e d e f i n e d by t h e r e f e r e n c e  line a  162 a range d e f i n e d by t h e o u t e r boundary.  In this section i t i s  t h e range t h a t i s under c o n s i d e r a t i o n .  I f the dimensions o f  a s e r i e s o f s k e l e t a l components a r e used t o e s t i m a t e t o t a l energy i n t a k e f o r an a n i m a l , t h e ranges o f t h e independent e s t i m a t e s w i l l d i f f e r because o f t h e unique r e s p o n s e o f each d i m e n s i o n t o a p a r t i c u l a r n u t r i t i o n a l regime.  S i n c e each  r a n g e , by t h e o r i g i n a l p r e m i s e , must c o n t a i n t h e t r u e measure o f t o t a l energy i n t a k e , a l l must o v e r l a p f o r a t l e a s t a s m a l l amount.  Combining a l l o f t h e i n d i v i d u a l ranges y i e l d s a much  n a r r o w e r e s t i m a t e o f t o t a l energy i n t a k e by t h e e x c l u s i o n o f t h e end p o r t i o n s o f t h e energy regimes t h a t a r e n o t common t o all.  A d i a g r a m m a t i c example i s shown i n F i g u r e s 51 and 52  u t i l i z i n g l e n g t h measurements o f t h e f o r e l i m b and lumbar v e r t e b r a e f o r two 175 day o l d l a b o r a t o r y r e a r e d fawns.  The  c a l c u l a t e d t o t a l energy i n t a k e f o r each w i l d fawn, based on t h e a v a i l a b l e s k e l e t a l measurements f o r t h a t a n i m a l , . ( T a b l e s 57-65), a r e g i v e n i n T a b l e  66(p.l65).  When e x a m i n i n g t h e s k e l e t a l measurements o f t h e w i l d fawns, i n s t a n c e s o c c u r r e d where two d i f f e r e n t d i m e n s i o n s from t h e same a n i m a l i n d i c a t e d ranges o f t o t a l energy i n t a k e t h a t had no common p o i n t .  As mentioned e a r l i e r , t h e use o f t h e  b o u n d a r i e s d e r i v e d from t h e d i s t r i b u t i o n o f t h e l a b o r a t o r y d a t a i n v o l v e s an unknown e r r o r term.  Since a l l o f the  l a b o r a t o r y d a t a have been i n c l u d e d i n t h e e s t a b l i s h m e n t o f t h e s e b o u n d a r i e s , a r e g i o n o f o v e r l a p must n e c e s s a r i l y e x i s t f o r a l l measurements from t h e s e a n i m a l s - s p e c i f i c a l l y , t h e  Figure  51  The L i m i t s o f t h e Range o f P o s s i b l e T o t a l Energy I n t a k e s f o r L a b o r a t o r y Fawn V26 (HML Regime) as D e f i n e d by I t s S k e l e t a l Dimensions.  ........  .. 1  .....  •; ; ':::  :  ..'.iii;.  •  E  E  E  _:  x  ..:  ..::._:....  E  "  :..  XX:i.:  X  .  .  E  E  E  E  . .: :::..:.:; • • E  :;:-:  1  •EXE:  -rr-..  k J! \ D t U 5 :  ::  :  r.:  i-  r  ' x i i :  r.irr. .  .  :  .  .: •!..:• i .  ... E _..  :::;:;-•  ••  Xi-X.E  ::  :-Ei ...: 1  EX F X  :ii.  ... r r r  .Eii  EiiH  r . r  Eii.:i  ;  ii-  ]...  X  '. H X X X l X X  rr.  X E E 1 E  1 r ri 1:-: -:.: iiii:- •: i i - E i l E :-  •  Ei :i ii. 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X X .  :"• I:.:. : • ;•  OF .  >  .EiX...  1 NTE  : : • . 1  iiiii  ;  : •: •  XXi•  •;;: ,.i • •  ;:•_!:.:• ... ,. .  .;_;  : :::  ' .;  -;.  '  EF T S ;  --•'-  -  2 9 3 , DOO Co l o r ies i  :  .  •: r. •  :: i: •: : ;  X... - MEAN  X  : •:..  .:  ...i •  ..  .  Eii ::;..:::  ::"  ':..:::  E T  .....  ix-  : j *: ..... .  . r r r r r r . .....  1 •'  :.  *.x .  .  :x  i  i!EE  :1  .  ' i  :.:.::  •E.xx-  r  "  i " " E i  :  ...—... _  rr.. .  lEEX  :....:....:  F i g u r e 52  The L i m i t s o f t h e Range o f P o s s i b l e T o t a l Energy I n t a k e s f o r L a b o r a t o r y Fawn U33 (LMH Regime) as D e f i n e d by I t s S k e l e t a l Dimensions.  -T7TT  iiii iiii  1:::  .i iiii '|XV Hi iiiiiii' iiiiiiii Hi iiiiiii "I .".mil ix: . |H : : : ; n x 1 !••i::x :!:: i' :r::i;;: xxx: S : ::Xt: x:.i : :x. 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No.  The e s t i m a t e d t o t a l energy i n t a k e s o f w i l d fawns  Sex  Age  YF17  M  116 Days  197,000 C a l o r i e s  YF19  M  118  205,000  YF16  F  116  198,000  203,200  YF18  F  117  170,000  159,500  YF20  F  118  180,000  191,500  WF 1  M  180  310,000  289,400  WF 2  M  180  320,000  298,700  YF38  M  177  320,000  259,900  YF37  F  176  265,000  1  1  T o t a l Energy I n t a k e  2  T o t a l Energy  Intake^  207,000 C a l o r i e s  Age i s counted from June 24 -- i . e . , t h e mean day a t w h i c h 10 pounds i s r e a c h e d when t h e mean b i r t h date i s June 10.  2  Estimated  from . s k e l e t a l d i m e n s i o n s a c c o r d i n g t o pp. 161-  166. 3  Estimated  from body w e i g h t a c c o r d i n g t o F i g u r e s 5 and 6.  t r u e t o t a l energy i n t a k e .  T h i s i s not t r u e , however, f o r t h e  w i l d fawns and some measurements must be e x p e c t e d t h a t  would  f a l l o u t s i d e the b o u n d a r i e s i f t h e y were p l o t t e d a g a i n s t t h e t r u e t o t a l energy i n t a k e .  These d i f f e r e n c e s u s u a l l y  appear  t o be s i n g l e a b e r r a n t measurements w h i c h can be i d e n t i f i e d as f a i l i n g t o conform t o t h e p a t t e r n o f t h e o t h e r measurements. I n t h e e s t i m a t i o n o f t o t a l energy i n t a k e s i n T a b l e 6 6 , t h e s e measurements were e x c l u d e d .  T h i s problem w i l l be  e n c o u n t e r e d a g a i n when t h e r e l i a b i l i t y o f t h e e s t i m a t e s i s discussed. As shown i n an e a r l i e r s e c t i o n (pp. 48 t o 54 i n c l u d i n g F i g u r e s 4 t o 7 ) , body w e i g h t and t o t a l energy i n t a k e a r e r e l a t e d and f o r t h e l a b o r a t o r y a n i m a l s , r e g r e s s i o n l i n e s were d e r i v e d a t a number o f ages.  Two o f t h e s e r e g r e s s i o n  lines  were used t o e s t i m a t e t h e apparent t o t a l energy i n t a k e o f the w i l d fawns a t 112 and 175 days.  I n a comparison between  t h e a p p a r e n t t o t a l energy i n t a k e e s t i m a t e d by body w e i g h t and t h a t by t h e s k e l e t a l d i m e n s i o n s , a major d i s c r e p a n c y became apparent a t 175 days.  Both methods gave s i m i l a r r e s u l t s a t  112 days as shown i n T a b l e 66 ( p . 1 6 5 ) , but a t 175 days body w e i g h t i n d i c a t e d a l o w e r apparent t o t a l energy i n t a k e t h a n did  the skeleton.  A l i k e l y explanation  f o r this difference  i s t h a t between 112 and 175 days o f age t h e p l a n e o f n u t r i t i o n o f t h e f i e l d a n i m a l s was s u f f i c i e n t l y d i f f e r e n t from t h o s e o f t h e l a b o r a t o r y a n i m a l s t o u p s e t t h e body weight energy i n t a k e r e l a t i o n s h i p e s t a b l i s h e d by t h e 175 day regression l i n e .  The r e l a t i o n s h i p demonstrated by t h e  l a b o r a t o r y animals i s only v a l i d f o r those n u t r i t i o n a l planes w h i c h were used t o e s t a b l i s h i t ; but more p a r t i c u l a r l y , i t i s o n l y concerned w i t h zero or p o s i t i v e weight g a i n s . i n t a k e s from t h i s r e g r e s s i o n i n c l u d e a maintenance  The  energy  component  t h a t i s based on a f i n a l body w e i g h t w h i c h i s the h i g h e s t weight a c h i e v e d by t h e a n i m a l .  I f weight l o s s o c c u r s , the  f i n a l w e i g h t w i l l u n d e r e s t i m a t e the maintenance  requirement  and w i l l t h e r e f o r e u n d e r e s t i m a t e t h e t o t a l energy i n t a k e . There i s an i n d i c a t i o n t h a t weight l o s s might be o c c u r r i n g i n the  field  a n i m a l s as shown i n F i g u r e 53 •  However, t h e  s i z e i s s m a l l and the c r i t i c a l p e r i o d between 112 and days i s not w e l l r e p r e s e n t e d . the  magnitude  begins.  sample 175  I t i s not p o s s i b l e t o e s t i m a t e  o f t h i s l o s s o r t h e date a t w h i c h the l o s s  The b e s t i n d i c a t i o n o f a w e i g h t l o s s remains t h e  d i s c r e p a n c y between t h e a p p a r e n t t o t a l energy  intakes.  A l t h o u g h body w e i g h t i s shown here t o be an u n r e l i a b l e i n d i c a t o r o f t o t a l energy i n t a k e a t 175 days, i t p r o v i d e s u s e f u l i n f o r m a t i o n that w i l l a i d i n the i n t e r p r e t a t i o n o f the p a t t e r n o f energy i n t a k e .  I t appears t h a t t h e l a t e  fall  n u t r i t i o n a l regime w i l l not c o r r e s p o n d t o any o f the l a b o r a t o r y treatments, w i t h the l i k e l i h o o d t h a t i t w i l l even l o w e r t h a n ' L . f  be  When t h e s k e l e t a l measurements o f t h e  175 day o l d fawns a r e examined  f o r the t r e n d s i n d i c a t i n g  p a t t e r n o f n u t r i t i o n , t h e p o s s i b i l i t y o f a v e r y low l e v e l o f energy i n t a k e i n the t h i r d i n t e r v a l w i l l be  included.  168  Figure 53  The L i v e Body W e i g h t s o f W i l d  Fawns  C o l l e c t e d Between 49 and 270 Days o f Age.  i:.:r irii::ti:j-:it-(: iiii-Iiii '-Hi- >'-\i iH-' "it i Hii ::!it i!:h iiiii i iiii: itit i i i i ip iiiii ii::. i i .-:-:.[HiP ii: iti: t}.tt iii; ' ' 1 •IP i ]-;i iiii: l i iiii |pi l i p i i p i p i l t t M I I I I I l;:;f iittippi , iiii iii; i !.i; j ;•; ;ji i iii i t P i i i i l iiiiiiiii i P P i i i iP i f i i p l i l l f t l l iiii:IP Hir " l i : .-::::1:T:J I P HH iPiiiP iii T P i il i ... ip, ii ' • ' i:ii:|-ttpt ii.it irit ;:r: Mil- iiiii Pi iir: i HHiiiii Hi-iHn; iiiiiiiii iiii Iiii !;! i"rrii iii] iii! iiii 11 it ii si i.lj.r :itj.q-lj. 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H H  P,i',r -Hi-i  m m  -]  %  :  I f  H  i.|  ; izi  l-i-.-r- i r - i P i i  P'-  Hi-IE-J: i H i -i-i- -i U - - h H j - - H  i'-fl-|ii-'  i t  '••; • { ; i ' P l'..'.'.',']L'~~..  ;  iii:  ! i  M-rt  I.:T.-  ;  ii  ii;  f-  _tii:S,  :;;  iiii-  :.; !:i  l.i I-i.  m  l-(-H-  m  i  1  iij!-  -iiii  i;  l-iili-ii M i - 1-i-r -H--1-1-tltr --I --!  k  T  T  iiij  -j.i , :  f-H-t-  r-H-l- -f-  j.  : : ; r  ifij  •i -  x:  1.  i.i!  ;  i-i-i; vii  i.i h  ;  i iii  i:  :;  :  :::1  1 ••• 1 • ; • :  1-  !- - I -  J  ;  r  ;.; 1 . ii-l-i -itji  i' if  1  T P ;  :  ifri  !:  1 - • ;-  • ':: il "  liiifi  •H-rl  ; 11  111!  \\\\j.l.I j.  r,xv I.T::I:  .;:JT|-  :  t  :  i  ::::!:: I  "IT'S, i l:|\i-.l:  rr  _  |i  TrH"  vJTi-  ! I-i • !• -1-1 j- - | T  i ] ; i  •  :  • •  nm m  11  TTi-f-  |iH  IT  ;  r  1 t :.l  ii  iiii  I;.1 !  1  :  !•( i-j'  ;  :i;:l;ji: • ; i • • • •  f::ni'::  1  -i  ;i'  p i  : :  )  !  . . H  p •  I::.TT  •pi i  i J P : 1 : :  ••'  : : : :  rP'-  —Cr  r  ~sii  ii.p-  rxU..  m  i:Hj  11 i ;  i;i  : : T | b i :  ;•;;!  "i.li.:.I.U...I.l.... I.;  ii +!-  M  !  i  "i  !!'! i  .  i;;:::!: I  j..  II.  Determination The  raw  o f P a t t e r n o f Energy I n t a k e  (Energy Regime)  s k e l e t a l measurements are not i n a c o n v e n i e n t  f o r m t o compare growth p a t t e r n s .  I t i s necessary to  them t o 'apparent t o t a l energy i n t a k e s ' from the graphs ( F i g u r e s 9 t o 50). these f i g u r e s provides and  The  reference  a standard  t o t a l energy i n t a k e .  convert  appropriate  l i n e on each o f  r e l a t i o n s h i p between s i z e  I f the s k e l e t a l d i m e n s i o n s o f  a n i m a l w i t h a known t o t a l energy i n t a k e were p l o t t e d on g r a p h s at the a p p r o p r i a t e  the  l i n e s , a v a r i e t y o f apparent t o t a l energy i n t a k e s  would r e s u l t . estimates  lines.  i f the t o t a l energy i n t a k e f o r the a n i m a l were not  known and the s k e l e t a l d i m e n s i o n s were r e l a t e d t o reference  the  energy i n t a k e , t h e d i m e n s i o n s would  be l o c a t e d at v a r i o u s d i s t a n c e s from the r e f e r e n c e Conversely,  an  By e x a m i n i n g the d i s c r e p a n c i e s among t h e s e  o f t o t a l energy i n t a k e s a l o n g the  gradients  p r e v i o u s l y e s t a b l i s h e d (p. 150), t r e n d s can be n o t e d t h a t l e a d u l t i m a t e l y t o the d e t e r m i n a t i o n intake —  o f a p a t t e r n o f energy  i . e . , the n u t r i t i o n a l regime w i l l be  However, b e f o r e  proceeding  will  identified.  w i t h the e v a l u a t i o n o f  nutritional  r e g i m e s , t h e growth mechanism r e s p o n s i b l e f o r the v a r i a t i o n s i n the apparent t o t a l energy i n t a k e s w i l l be r e v i e w e d w i t h p a r t i c u l a r reference scapula.  t o t h e l e n g t h s o f the f o r e cannon  Because t h e s e two  and  d i m e n s i o n s are s i m i l a r i n o v e r a l l  s i z e , t h e y can be compared w i t h o u t  c o r r e c t i o n f o r growing  mass or d i f f e r e n c e s i n mature s i z e . The  l e n g t h o f the cannon i s an e a r l y m a t u r i n g d i m e n s i o n  whereas the growth i n l e n g t h o f the s c a p u l a o c c u r s at a l a t e r  170 time ( c e n t r i p e t a l  gradient).  The r e s p o n s e o f t h e s e  dimensions  t o t h e v a r i o u s n u t r i t i o n a l regimes demonstrate two v e r y different The  p a t t e r n s o f growth as seen i n F i g u r e s 54 and 55'  d i s t r i b u t i o n o f t h e d a t a f o r t h e cannon growth y i e l d s a  much w i d e r and s l i g h t l y s h o r t e r p a t t e r n t h a n f o r t h e s c a p u l a . The  l i n e r e p r e s e n t i n g t h e maximum growth r a t e f o r t h e cannon  s u g g e s t s an a s y m p t o t i c  approach t o a maximum, w h i l e  t r e n d i s n o t apparent f o r t h e s c a p u l a .  These  this  observations  can be combined t o e x p l a i n t h e growth r e s p o n s e s o f t h e s e to n u t r i t i o n a l The  bones  level.  amount o f growth o f t h e f o r e cannon l e n g t h e l i c i t e d  by t h e d i f f e r e n t  nutritional levels  seems t o c h a r a c t e r i z e  t h e development o f a d i m e n s i o n t h a t has passed i t s g r e a t e s t growth p r i o r i t y a t b i r t h .  The b l o c k y d i s t r i b u t i o n  i n Figure  54 i s caused by a t o l e r a n c e t o moderate r e s t r i c t i o n and an i n t o l e r a n c e t o severe  r e d u c t i o n o f energy i n t a k e .  A moderate  energy r e d u c t i o n d u r i n g t h e i n t e r v a l between 112 and 175 days produces l i t t l e  o r no r e d u c t i o n i n growth r e l a t i v e t o t h a t o f  a f u l l - f e d animal.  At t h e l e v e l o f most severe  r e s t r i c t i o n very l i t t l e  energy  growth i s p e r m i t t e d a t any t i m e .  These o b s e r v a t i o n s a r e c o m p a t i b l e  w i t h a growth demand t h a t i s  t o o l o w t o f u l l y u t i l i z e a h i g h l e v e l o f energy i n t a k e , and i s a l s o t o o low t o compete under h e a v i e r r e s t r i c t i o n w i t h  regions  o f h i g h e r growth demands. According  t o d e s c r i p t i o n s o f a x i a l g r a d i e n t s a c t i n g from  d i s t a l to proximal r e g i o n s , the scapula should e x h i b i t i t s peak growth response a t a l a t e r date t h a n does t h e f o r e cannon.  Figure  54  A Diagram o f t h e Growth o f t h e F o r e Cannon R e l a t e d t o T o t a l Energy I n t a k e  .).... :.:.!:::: ::::!:::. !!iiii; ! ::::i:.::: :i:i i-iiiiii 1 i i i ji iii ::;:!::•: :.•::;::;. i i i i i . i l i i i i i i . . i t - i i i i i i . i - I i i i i i .iiiiiiii i i i i i iiiii:::-: i i "iiii •i: |..:. ••Iliiliii i i i j i i - i l : : •::•:.iiiiiii'TTiil :... : : : .i :|. .: : ;. . 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U" l i l :;;• ! i: | i: i: :::: i:; •: ;::: j:::; i m i l l • iiii ....!.!.. 1 1 •iiiii — . >.. , i! i l . i i l l ' [ :• M:i 7l : :i:'. . 7 i l iii l l i l . iiiiiii:; : i i i ; : : i i i i . | i i ii i i •pii-ii iiiiiii I-:" i l i i i l i ^ ... -ii---!il-r--iii.....xj:.:;.: —! ~~ i i i i i i i i i i i i i i !• ::|---.: :: i j::." i:: • ZZ~."•.••!-.1 :•!:•= i :. ...,. , iiiii .; i; j ; : ; : 7 iT7 i r. .. I i.. i :. i i i . • : • i . • 16i! i i i r iiii iiiii Iiiiiii iiiiii;! i.-'klillk i i i i i i i|ii!: i i i i i i i i i 1:111.1. iiiiii.:.:: iiiiiii; iiiiiii:.: TiT.irriT i7l]i|"! 1:7! 1~ .: :: ; ; irmiiii' i i i i i i i T i l i ' l i i i i i : : ! '•^"^ . i l i i i i i i : ill : i io 1 i :. -! 1 • 1 :. :i;ij;i.: i i i j i ! iiifjii: • 1 • • »^iil! . i: i... j:: i• i — i •— k- t *J • i 1 i : > ; i .... '-i • • 1! i!! :i!. [ \: 11 • ME AN t ^T 32 2 DA Y S ..iii..!::.. iiii ;i i:!:;:: l-i-i|ii iii j i j . 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J j : | :.: ' i 1 • i i j i : / ::::!;::: ....[. ........ :: " i i:.:i! ii ;• ;.;.;: :ilii- i : ••:• j!!.: • i i ! / // yy I• I i i j 1iii; 11: ; i i i i l l 1 -i-iiiii. ; ."ji : i: i i •:: • j:: i !: i . :;: i i: :.i: ' : i i i / y > lillii: i i i i i i i ii iii iii; :! •: ,-. i: I: i i: 1' 111111: iiiiiii.::r . / / A ::::!:!.•" j:° L i :: 1' i.i i: • ::.:]:.:!' :i::i:::: :::;!::•: ) U N D A R Y AT 1 Ii2" AY S iiiii.';; "f::?t'-:;|T HI 1"; 11 i: : .•• : xi !_i^ — £ '2 I L'.l.!.l.iii. .:x.:.,„i.:j. i :z..L-i-.- i •;: j • •;; i; rTlilTTl:" "';::] : I-i. . t.. . iil|iii i i ! ! i I ' i i ' T - i i i . i'; i i i iiiiiii: i i i i j i :ii |: i i iiiiiii!: I i i i i i i i i iii |ii i iliiliii iiii! jiil .... i . . . . :: -j.' ..;i:::; ::!: i:; •. ::.; i;;; i : -:; 1:.;: :::: i •:.. .i-iii-il.i ..! . • i i i i i l i i i ; : : i i: i i i i i l i i i Ill: i i i.i.i.i i i i . i l i i i : ...... .11:iiii :.. H i . L i . ' I n L i i i L .:::!:;:: iiii-: - i i T i t r r r r iii-i-^ii i i i r : v : ! rr;t iilili-ii ;;;;!:;;; . . . . . . . i7~! ~ ' i i i i i i i i i i i ii'"! i i " Iiiiiii: 1 I i j i il:; •, i: ,: • • i i — in .: i! :.: ,;i. .1: .I'i-v • ijilliii . i: I !!,••' M • • ' • " : . : ! : : : ; .: : : 1 > • : : : - - -1 ::::ii;: iiiiiii iii; i i i l i i l l Iiii' iiii::, : II 1. ...:ii.:_:..i_-:...:i:i.:..!.;.:...:. : i , i | . . . : .... i : . : : .iiiii !.;-4 i h i l j i :|;i iiii i i .- i .- i i i•i.i i ! :._ij-ii:.i. iilii:!:. : I'lj": -'I i i i i i i i i . i :]Ti 11 i l l 'I ' I I I i - |ii:;: •: i.::: 1 i : i . i • ' ! i i i : i . :::.!::•: Iii illliiii iiiiiii; i i i i i i; :;ii|•i:i I'lil — r — ... i .. ::::|;::. • .. j 7 " : - i i,: i ; ; i i .Vljii: i:::!;; i • i i i i i i i i i i i i i i i i i : . . : ] . : • ; I: 1 i 1 : i :• i i i i i i i iiiiiilH iilijiiii iiiii Iiii . : . 1. i i - i i i i i ;.i i ] i : :X-i.iii-i. — 1 .•;.. i-i-ii-•••i .ii; •.. .'. | • ;;; ! 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THP " i I: iiiiiii ' l i l i l iiilliili i i i I i i iiiiiii ." : | ; ' I "1 : ; • : i : • : . :;, • 1.,: •iiiiii.!; . 1 iiiiii ii iiiiii iiiiiii: C > l - i i . . ! i i . .iii 1 .Li Iiii ' i i l i s .iliiiiii: i i p j l i i j i i l : i i i jiff m i|i|i!-c c) l i :::. .1i • ..• - r r r i ~ W:^ iiiiiiii ::•!:-.: ^ i i :. • j.': • • i i i •. -F:;.! ::: ' i i : : i:; j.:;.:; : i::';:: •nii i i i i i i i i i i i ' ' I ;::.!!:;• ::;i|.:-. rri'i; — I -: .••:•: • : : ) - : : : : • : : Ener< ii|:i n t d l ^ e , C a l o r i e s > I 0 ( A . biiE-i: •iiiii.-:: iiiiiii;-. iiiii; 2 . --\ ....... . . . J l i i j i i i i-lrh!-:: . ir:: • l.i.I± I :: i•!::i. i i i i i i i l i i i - l : :: *ij;Hiin-!r iiiiiii:! :::!:-.. ... i. i i i j i i j l i i iiiiiii, . . . . i — i7ll|;lil iiiiiii! i i i i i i i * - -1 - • • • 1 !::•:: iiii i :i-ii.|:.iiiiiii — i — :.. i.: •. -. ;:,: ,, -i. .: 'iliiiiii | : . .iiii:'!! . : j:-- i i i i " •::!;::• | --j: -.I.:::!.:.;i i i i i : . i l i i l i i i i | h i i i i i ' l iiiiiii • H i ' i i i i i i i i i i i i i : : liiiiiijiiiii' H i i : |:ii iiii :i; l l i l ! ::.:!..: i i i i i ! | i i 11!:! i I i i i i i iiiiiiiiiiii iiiiiii :i! l l i l iiiii ... :;::!:"• : :;jl i i-. •!:": • IfTH"! ~ | : ~ •;" i i i . 'iiiiii: iii: i: iii liilililli'. iiiii !; i i j i! i I HI j iiiji '.:: i j:! i j:|i iiiiii , : ' l : •• .... 1 liiiii'ii .l-LLuL-L;., • • i i i-lj-i |i :-r:-|||"ll i i i i 1 ' l i l l i i i i i l i ii iiii'iiii'!' ::J!-i.:|--L-i-;.L j::|- - ;; ; ::: 1:.. - Tin:-; iiiii.:!:! i-iiilill : I i: •: I': lliijiii Iiiiiii i i i i i H i . : ... j . . . . . . 1 ! iir;-' ••i-i .itujiil: i . l i i . • • I I iiiii i l i ' L i I i i i i i i i  I  ;:  i i 1.1;,:.  :  i i _ i . : . i _ ! : : ; . [ . : -  1  :  !;  !!  :  ;  1  :  1  1|  T  :  r7  r  r  :  :  !  /  ::  m  ,  :;  ;;  :  :  :  :  :  7  :  ;;  :  :  :  : ;  :  :  :  ;  :  :  !  1  :  1  ;  1  :  7  ;  ;i  :  ;  :  :  : :  :  1  iii  1  ;;  :  :  :  :  :  1  :  :  ;  :  5  k  ;:!  Tr  :  1  :  :  !  ;  ;  !  -  :  :  :1  :  !  ;  ;  172  Figure  55  A Diagram o f t h e Growth o f t h e  Scapula  R e l a t e d t o T o t a l Energy I n t a k e .  T h i s i s borne o u t by t h e s l e n d e r d i s t r i b u t i o n p a t t e r n i n F i g u r e 55. Between 49 and 112 days, l e s s o f a r e s t r i c t i o n o f energy can be t o l e r a t e d growth i s d e c r e a s e d .  t h a n i n t h e case o f t h e cannon b e f o r e  Between 112 and 175 days any  r e s t r i c t i o n o f energy r e s u l t s i n a d e c r e a s e i n growth. the  T  L  f  t r e a t m e n t , growth i n any i n t e r v a l i s l e s s  t h a n t h a t o f t h e f o r e cannon.  On  restricted  The suggested mechanism i s  t h a t t h e v e r y h i g h growth demands can o n l y be s a t i s f i e d by a h i g h l e v e l o f f e e d i n g , and t h a t t h e h i g h demand r e s u l t s i n a r e l a t i v e l y h i g h growth r a t e under s e v e r e energy r e s t r i c t i o n . These a r e o n l y two examples o f t h e wide range o f growth r e s p o n s e s t h a t c o u l d be caused by d i f f e r e n c e s i n t h e time o f peak growth p r i o r i t y .  Some d i m e n s i o n s o f o t h e r  skeletal  components have p r i o r i t i e s t h a t c o i n c i d e w i t h t h o s e o f f o r e cannon o r s c a p u l a r l e n g t h , some a r e i n t e r m e d i a t e t o them, and some appear t o be even l a t e r m a t u r i n g t h a n s c a p u l a r l e n g t h . The w i d t h s o f t h e t h o r a c i c v e r t e b r a e f o r example, show a l a g phase i n t h e i r growth r e s p o n s e s i n d i c a t i n g t h a t some time e l a p s e s a f t e r b i r t h b e f o r e t h e demand t o grow b e g i n s t o r e a c h a significant level. By a r r a n g i n g t h e s k e l e t a l d i m e n s i o n s i n a f i x e d o r d e r , p a t t e r n s o f t o t a l energy e s t i m a t e s r e s u l t t h a t the n u t r i t i o n a l treatments.  characterize  I n T a b l e s 57 t o 65, t h e  measurements t a k e n from t h e fawns i n t h i s s t u d y a r e d i v i d e d i n t o f i v e groups, each a r r a n g e d i n o r d e r o f growth  gradients.  When examining t h e n u t r i t i o n a l l y i n d u c e d changes i n s k e l e t a l conformation of the l a b o r a t o r y animals, i t i s  c o n v e n i e n t t o use the d e v i a t i o n s between t h e t r u e and t h e apparent t o t a l energy i n t a k e s . produced  I n t h i s way  the p a t t e r n  i s not o n l y r e l a t i v e t o t h e d i s c r e p a n c i e s w i t h i n i t s  components, but g i v e s an a b s o l u t e measurement o f s h i f t s o f t h e e n t i r e spectrum o f measurements.  I n the next s e c t i o n o f t h i s  d i s c u s s i o n t h e e f f e c t o f t h e n u t r i t i o n a l regimes on the s k e l e t o n s o f t h e l a b o r a t o r y a n i m a l s w i l l be d e s c r i b e d and, f o r t h o s e regimes not i n c l u d e d i n t h e l a b o r a t o r y s t u d y , an e s t i m a t i o n w i l l be made o f the t y p e o f p a t t e r n t o be e x p e c t e d . Up t o t h i s p o i n t a l l t h a t has been a c h i e v e d i s the d e t e r m i n a t i o n o f an apparent t o t a l energy i n t a k e f o r t h e w i l d fawns a t 112 and 175 days.  In the f o l l o w i n g  sections  t h e s k e l e t a l measurements w i l l a g a i n be examined, t h i s time i n o r d e r t o determine p a t t e r n o f energy i n t a k e .  The  effects  o f t h e n u t r i t i o n a l t r e a t m e n t s upon t h e l a b o r a t o r y faivns up t o 112 days w i l l be d i s c u s s e d f i r s t ,  f o l l o w e d by an attempt t o  c l a s s i f y the w i l d fawns t o one o f t h e s e t r e a t m e n t s . a n i m a l s a t 175 days w i l l t h e n be t r e a t e d  The  similarly.  I n each o f t h e t h r e e i n t e r v a l s t h a t are b e i n g c o n s i d e r e d d u r i n g t h e f i r s t 6 months -- i . e . , 0-49  days, 50-112 days and  113-175 days - t h e n u t r i t i o n a l p l a n e might have been one T  H , T  t  M*,  o r L ' i f a l l v a r i a t i o n s are c o n s i d e r e d . 1  of  With  t h r e e p o s s i b l e n u t r i t i o n a l p l a n e s i n each i n t e r v a l , the comb i n a t i o n s ( r e g i m e s ) are 3 at 49 days, 9 a t 112 days, 27 a t 175 days.  Out o f t h e s e c o m b i n a t i o n s some w i l l be more l i k e l y  t h a n o t h e r s as p o s s i b l e a p p r o x i m a t i o n s t o t h e regimes o f the w i l d fawns.  By e x c l u d i n g L t  f  nutritional  i n the  first  i n t e r v a l and to  ,  H  I  i n the t h i r d , the c o m b i n a t i o n s are  2, 6, and 12 r e s p e c t i v e l y .  reduced  There seems t o be ample  j u s t i f i c a t i o n f o r these o m i s s i o n s .  The s i m p l e s t e v i d e n c e i s  i n the d i r e c t o b s e r v a t i o n o f the f i e l d  animals.  In the sample  k i l l e d i n August  (49 days) t h e s m a l l e s t weight was 24 pounds  (av.  31.0, 4 a n i m a l s ) which i s i n e x c e s s o f the  23.2$, max.  15.0 t o 17.8  pound weight range o f ' L  laboratory.  F u r t h e r m o r e , the 24 pound fawn was  f  t r e a t e d fawns i n the  because i t was t h e s m a l l e s t one seen and was  selected  conspicuous  because i t had not l o s t t h e w h i t e s p o t s o f i t s c o a t .  The  s i z e s o f the remainder o f the fawns were e q u a l t o those o f the f  M  f  o r H*  animals i n the l a b o r a t o r y .  ,  The  second type o f  e v i d e n c e i s t h a t p r o v i d e d by the s k e l e t a l measurements o f the field  c o l l e c t e d fawns.  The t o t a l energy i n t a k e s i n d i c a t e d by  t h e measurements from the 112 day o l d fawns p r e c l u d e an ' L  T  t r e a t m e n t i n the f i r s t i n t e r v a l (compare i n t a k e s i n T a b l e 66 v/ith energy regimes i n F i g u r e 56 (p. 17$).  Although milk  p r o d u c t i o n i n sheep d e c r e a s e s under an energy r e s t r i c t i o n , i t i s m a i n t a i n e d a t moderate l e v e l s even under c o n d i t i o n s s e v e r e enough t o cause weight l o s s i n the dam  ( W a l l a c e , 1948).  A milk  l e v e l s h o u l d be e x p e c t e d from t h e doe t h a t w i l l p r o v i d e more than an L T  T  plane o f n u t r i t i o n .  Added t o t h i s i s the f a c t  t h a t the fawns wean onto browse t h a t has not begun t o undergo t h e l o w e r i n g o f n u t r i t i v e q u a l i t y a s s o c i a t e d w i t h w i n t e r dormancy ( L l o y d e t a l , 1961; B l a x t e r et a l , 1955, C h r i s t i a n and W i l l i a m s , 1957; G a t e s ,  1968).  1956;  176 The  argument f o r o m i t t i n g *H*  not q u i t e so o b v i o u s . had * HH  i n the l a s t i n t e r v a l i s  I f the browse i n the f i r s t  intervals  p r o v i d e d o n l y a r e s t r i c t e d amount o f energy ( l e s s t h a n 1  o r ^H*)  t h e n the q u a l i t y o f v e g e t a t i o n c o u l d not  e x p e c t e d t o improve as w i n t e r approached. d i g e s t i b i l i t y and  The  be  decreased  i n c r e a s e d t i m e o f passage o f t h e f o o d  can  o n l y l e a d t o a d e c r e a s e i n energy i n t a k e by the a n i m a l . a n i m a l s t h a t had r e c e i v e d a h i g h energy i n t a k e d u r i n g f i r s t two  For  the  i n t e r v a l s , the q u a n t i t y o r q u a l i t y o f f e e d  a v a i l a b l e i n the t h i r d i n t e r v a l i s i m m a t e r i a l .  The  fawns  r e a c h a s t a t e o f p h y s i o l o g i c a l m a t u r i t y where an o b l i g a t o r y d e p r e s s i o n o f growth r a t e o c c u r s effective  *L  f  plane r e s u l t s .  e x c l u s i o n o f an *H  T  The  treatment  t h e l o w e r end o f the H* T  (Bandy, 1955) field  i n the t h i r d i n t e r v a l .  treatment  52.8  o n l y 2.2 ,  H*  Even a t  That means a g a i n  and 175  days o f  age.  c o l l e c t e d fawns i n O c t o b e r weighed on t h e average  pounds ( 4 9 . 5 t o 57.5)  pounds  the  r a n g e , a weight g a i n o f a t  i n w e i g h t o f o v e r 20 pounds between 112 field  an  evidence supports  l e a s t l / 3 pound per day would be produced.  The  and  (49.3  to  and  i n December averaged  55.0  T h i s l e a v e s an average d i f f e r e n c e o f  58.9).  pounds; a l o n g way  from the growth e x p e c t e d on  the  l e v e l o f energy i n t a k e . J u s t e x c l u d i n g the  and the  ' H' t r e a t m e n t  * L* t r e a t m e n t  i n the f i r s t  interval  i n the t h i r d has c o n s i d e r a b l y reduced  t h e number o f regimes t h a t must be c o n s i d e r e d .  F o r any w i l d  fawn b e i n g examined, t h i s number can be r e s t r i c t e d  even  f u r t h e r because t h e t r u e energy i n t a k e must be w i t h i n the ranges estimated  contained  by i t s s k e l e t a l d i m e n s i o n s (pp.  161 t o 162 and F i g u r e s 51 and 5 2 ) . F o r each o f t h e n u t r i t i o n a l regimes t h e r e i s a l i m i t t o t h e energy i n t a k e w i t h w h i c h i t can be a s s o c i a t e d . f o r 175 days on a c o n s t a n t  f  That i s , an a n i m a l r a i s e d  L * regime cannot e q u a l t h e  energy i n t a k e o f one r a i s e d on c o n s t a n t  f  H , and s i m i l a r l y f  f o r each t r e a t m e n t l y i n g between, t h e r e a r e l i m i t s .  These  l i m i t s a r e shown i n F i g u r e s 56 and 57 (112 and 175 d a y s ) . A l l o w a n c e s were made f o r t h e range o f energy i n t a k e p e r metabolic  pound p e r m i s s i b l e on each n u t r i t i o n a l p l a n e and  f o r t h e v a r i a t i o n s i n growth r a t e s produced by t h e s e differences.  A l l o f t h e n u t r i t i o n a l regimes s t u d i e s i n t h e  l a b o r a t o r y p l u s a l l l i k e l y w i l d r e g i m e s have been i n c l u d e d i n t h e s e t a b l e s and a r e l i s t e d i n rank o r d e r o f energy i n t a k e . I f no p a r t o f t h e energy range c a l c u l a t e d f o r a fawn w i t h t h e energy range a s s o c i a t e d w i t h a p a r t i c u l a r  coincides  nutritional  r e g i m e , t h e n t h a t regime can be e l i m i n a t e d as a p o s s i b i l i t y f o r t h a t fawn.  F o r example, an a n i m a l a t 112 days w i t h a  t o t a l energy i n t a k e between 130,000 and 235,000 C a l o r i e s c o u l d not have been f e d on t h e regimes HL* o r 'ML*. T  four choices:  »HH», «MH», «HM» o r •MM .  energy i n t a k e a t 175 days between excludes  This  Similarly,  1  leaves a total  300,000 and 400,000 C a l o r i e s  »MML», »HLL» and »MLL», l e a v i n g HHM', »MHM», »HMM», 1  «MHL», *MMM», »HLM», »MLM«, »HHL», and »HML». I n o r d e r t o f u r t h e r d e f i n e t h e n u t r i t i o n a l regime i t i s n e c e s s a r y t o make use o f t h e d i f f e r e n t i a l r e s p o n s e s o f t h e  178  Figure  56  The L i m i t s o f T o t a l Energy I n t a k e s Within  Possible  Each Energy Regime a t 112 Days.  : . .  i i i i  ii!!;!  L  —iii-ill-  •:. 1:: • Liiiii  ..-  .. : j : : . .  :: I i i i i  ':.!::'•  Hi. i l l "  i : ! i i  11. i l l  ;  II1 i•"•' • ._l..i_l_l. i n . : : : • •: i H i l l  i^-jlll  il'!!: Li :.  •::. |-  1  . • •: i : : : : : . . ! . •: i l i l i . 1': i i i i i i i i i'l ' i i i i i : •:: i. ;: . I ' l l : :••[:!.: .::: i : : . : : . i | ' ' ; i i i i i i . :.:::: : : l i : .i-1. : . . ! : : : . •. • i • : : : ::: i i : :  • . !.. :  —'  i|i!  | i  ;  iii - - i i i  ;:  i "i-i-.:  ::.:.. i_.i. _  :::: t: "1 ..•.):::• ••!•!••  iiiiii:  i  i  n i l :  :.|I:  1  i l l  •iij-!--- • i - i l i i - i i i  -11-  ::: h::" i l l  i  f  |  i  I l i - L  iiiijiii;  i l l  Iliiliii! ~ M H ; — . j.... i  liijiii.;  — i . —. : i:::i.i::: :::::j — II > H H M . 1: ::::  l  l  1  i !1 r.~.i  ..... 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' : . . i r  :  ...  . . • • : . :.!:.:::  .;  •: . : . .•:  i i i i i i • i i i -  i ;  - i ; ~ | H : j . - r  -iiii"  i ~ i i  : : i i .. i..: i : i . i i i i i . _:...:::._.|..u:.ii: |li  i  :••:.-]:.: :  i i i i i .  i  T  . : . . i •• • • i : i ! i : . j l i i  j i i l i  1  i i i i  i i i li  I i i i i i i i 'ii :.:::.j::i: i i i j i i ' i i :::.;•]:::.-  :  -\.\ '"i  :zz:  1  !liii;i : :ix:i : -in i.:.:i..;i:: ::::::!:::.— . • :! :: -  ii-iliii -nil:::  r  i i i j i . 11  i i i i i i i - " I i i i i i i i i i i i i i i!' . : : : (.—;•: ::::i:i:i — i -H l | i i iiiiiii i i 1 illj.il" i i i liijiii -i  i i i i i i  :  •i  :  :  oI*, i i ' • - • ' i " i i - : x ;  i • v.D  E  :  o  ' '-.i-i  i i i i i  ii'Ll :.:: i-;::  i.i  -  11-1!! -' :  i' i'-i | i - i i i - i i - i  1  :!il !  :!:.:-.  i  :::.: j : : •  1  •11:  i ii  :!  !ll  - "  - i  ;  111 i 1'  ------  iii::: ii"  :  -  liii-ii : . i i - i  i i i i i .  i i  • " 1 i' 1:: iiLi.i! illi:: I  lil-l-l  i i i i  .j 1 '•-  ......  - i i i i i  .  -'-:i..rj:i::. ::. :i ' i : i i i i i i i  ::•:]::.: i l i i - i — i i i i • i i i i i •::::i::::  i-l-l i i i i i  i i i -  iiiiiii  . : - . . : J I : : I  1 i - l - -.-i-i i i l i - i  i;-i--i - . iiii  :  : : : : J : : . - : :  i  ll-  Iii  i i  i i j i i i  :  i i i i i i  -iiiii  T  i i  1  1  L i ;ii : |iI  i n i -ii-  I i i i i  i i i !  1--  iiiiiii  ii!:jill  lllliill ..  i i i i i 11  iiiliii i i  l l i i l i i i i i i : : : i - ! ;:-_••:. • i ' i i i i i i l i i j i i i i i i i1i i i ---i--- — i i r h i i i i i i ' .-iiiifc-.i. 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I i i i i i . • • ii— •  liijiii:  i i - i i  Iii  Iiiiiii  -UT  • •  i i i i :  iiiiiiii  iiiiiiii i S l i E .-:::: j.ii!i : . r : : i : :::  .:::: ii.:;:: l i i l  l i i i i j i i i III  •;":-  i i  "iiiiii i i j i i ! :...!:: •::.:::ii.-:z i l . i :-i:j: i i l i i l i i i Iiiiiii i — :::::l::i:: i i i i i i i -..::-;!:::-.: iiiiiiii i i j i i  rr  l i i 2  -  i i i i i i  :!• 1 ^ t ne r.c y In t a k e , C a o r i e s x l Cr  i - i i - i :.: i  -lilil  0 -i  ..i._.f  i i i i  ii- i l i l i !  i i i i  HiJiii iiiiii'  1 :-i:i.::-i • • i • • • • t -: . i : | i : : : i l i i i i i i : ::.;:]:::: i l i i l l iiiiiiii n:npi- l l 1 " '1 ::r:i.:::: :::;: i::::::: :.::i:j-:::i. i i i i i i . iiiiiii ii iiiiiliii I i i i i i ! . . . . !1• — 1 — i • • -—\ ii---iill I l i i i i i i ::.: :i:.:.:: - -t • • _ :::i: j i n :iii-ii:::i 11 i i i i i i i i i i i i i i : i i : iljilli ...... 1 ::::: :!.::::: ..I.i.... ::::::.!::i liijiii .il|ii! i l i i l i i i iiiiiii! L iiiiiliii.  i l l ji' i i i i • • i ' I i i i i i ' • • i •' L M ; .-- r .I-i... ii..l:.i.: 1 i ' i i i i ! i i i i i i — - i — , . ii:_-j:ir:: i i i i i i i i o l i i - i i '• . i n : • ;-- •:;:!;.:::: Hill.:::::: ..... . iiiiiii i i i i i ! i• l_l 1 H L -ll . 1. l i i i i ' i ililjill: :j::::i. ::; ii;;-.:: i i i i i . i i k l i il.;ijii i ; i i -—iiii i i i i i i i H i l l IIII •. iiii:i H i ' ! : : 11 • i i i i i i i " 'l/i I ... i M L : i l l ] H i ;.::.::i.:.:-.: 'iLiiii I i i i i l i i i i - - • f ••: : : : | : : : :::: :.r:..i i i i i i Tiiifiii: • • | : i : i i i . i * .iiiii::;:i l i i l i '•'•">-'• '• • I i i i i i i i i i iii : : : : . ! : : : : j :" .... 1. .. "1 — . . L. L ... :.::.:ix.::: i i j i i .!;:::... ::-: :•:. L i i j i i i iliiiiii ililjill "lilil: iiiiii! ::::.!:::.: ..:::!:::: in: :i :.:::].::: .: ::..:\ :. - - i i i i i i i i - i ^ n i I ~ • 11i i'j i i i : ill::..:  •  iiiiii  Ti  lil|lll  l i  I:.:! j : i l i.iiii iiiiii i i i i i i ' i i - i"""-T-  -liil-iii i i i i i i i  ;:::!::: .: 1 i:: : :  •iiii!!  ;  _:::.j.:.i:i  i L  iiiiili;i|- i i i i i i  • •i i i i i i i -  - i i  1' iiiiiii iiiiiii . 1. l l i i l : . . : i i i i i Iiiiiliii.  .....,  :::'j::::  •••:>  :::. i:::: iiiij Hi!  i i i i i i i i-iif|ii i . 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Ill  i l l . , i:  i i i i -  Figure  57  The L i m i t s o f T o t a l Energy I n t a k e s P o s s i b l e Within  Each Energy Regime a t 175  Days.  PpF Hiiiiiliriiini in!:;:; iii'iiii'i H ' - H . - i' ::IHH Hi-! .. •:::::::. 1 •!•••• rrr rrii r: :::: i:::: .:::!::: .:'..•: p i i i P j i P :::' j: •::  -  iiii-!: - I F FpiP  •.IP IP  r\i n n  P P P iiiPiill : . . . ; H - F ; I . : •.: j •:: . ; : : I :::: PUP! PiiPi IPiiPi ilp-p- Iiiiiiiii. FFl ; i:  Hp. - n;:i:i::. l..:i|..._ . in..:.. :  ;  :• .1- . .  -  .:.;.!.:....:  :: j: • . :  . • H  :  -PI-"-'  iii-P'F...  F  ..i•...  ii Hi .. j....  :  Pi-F-i- FH'-P  i'i:: •  H:'  i  H : :  I.;::  :  —— j " — : : " •: •: | " : : :  .:::!::::  i_Ui  ^ipiii : :.:.[::•:.  :: j:.;-. 1-iPPI ... ~T:.:.~ I f l .::p"":_. ip]P7 •;;;!';:•'; .:.:!;-:: :::!::: ;p;ph H M iiii | Hi: ::.:.: i:::: ipilp. -iii. - i - P P •pi ~:P ':~~\ ~~ F F ' P P P i i-ip-nl -rpi-Hp P F i i F . • • Flil i - r PPP! :•• .i 1HM-: .... ::::):::: Iv . . . . ! .. H I I I F : .::' ! : : : . ::::(:::: -i -i; i i P — i i-ptPP -iiijii-i•iPriP- ::: ~i::.: "•:-Ti'~ . i.i: illF : | : : : : : F : : HHH ::.: i...: HipH PIIIP • • • • \ ::-:|::.r: r MM n p:n ;• .*:: p:t:: PlPii Pri- -HF P iPiiiii FFi lliii^F ::::: 1::.: : : : : i":::::"i. FtiilF. rr.;: I: Hi ;F . •: i; ; n ".:-: j i ~ . ipjiiii I P | P i iiiiiiiii ••• • 1 ... i . . : Lr M H : :.H ;•:.. FF Iiiiiiiii ; -'".PH ; : :•.-.: i ::: : :.::i'..:: .;.:.: pin : : : : .ii; !:i iii:; P i P i P p i P i •:.:.: :.i:FFH p HF :" :7,i-: :.r -:r:-:|:::: :;: ;i;::; ...... i i ... . MM -1 :.-_:•_• l : . : : : i — :.:r.:\: i . ; Hi .::::.! ..:i: r:n; ii.vj::::iHi:;;:; HliiF!PP • I FiriFir !i; P i -H; !•!: Hp P i p p :-:.: :]•:-.:: IFFiilH ri,:j.::: P i p P ::.:: |.: ~: FFillpi i! il! iiiiiii ......... np-rp .:'-z'::iii:::i" _ HHiJiii: ~l .. iiiiiiiii Piii-Pi Pii Piii.;.:.! F I . ; ; Pi F i i -iii-' rPiilP Pi 'H-iiiiiii i f P P iiiiiiPP :::P:iP iill il 1:.'. IFIillii P i Hii ::r::|-:::: » — H L M F H I F H Hii •iiiiljp i t: rrr-ii; ;:r _ PiipiiiPii iiii Pi Pi i •• FlilFF' IFHFF .... \ i.:.. :!::.:-' •:-~|--.:-: 1 5>—M L M r• ..! . .1 . . . .:.:::::"ji~T"' :\::sJ:::.:.:. Hii FiiiniiPiiii -•:-ii.-.:--iiiipii. III; Hi! iiiii iipiiiiii P I I iHitH:: 1 1 • • • • i cr -iii- ±i • "-ri-jiiH. Hiii-rr; FliiiFI i4 i ; > i n i_ '-•'—-•-T~ •- .--i_i::-r >> Pii'lii r i - : i ; i ; i i i i i i F H i ~ i~\ :r.;-;: r: :-:.!•:.-.: - : . - : | T ~ - ::-..-.j • : : - : FiHFFi 1 FFlFH iiiiiiiii i-piijiPr H P H F FililFi; • - P - c : -.- : . - " & - : : r : - : j : : i - ; Hii A t_l 1 : ::-p:-.-_ .... 1; .. ... iu... m i ii_ Fiij-n.;: Flij " Fii !:i iiH.: iFiiF.;; iiiiiiiii . i i l IFi i..:. :.i-r]--::-: LLi= FII HI IF:'::!:. r:: r. : i | . : : - iiFjiii; P i i P i i : F n : FIllFli fiiiliTF r :•'•'' • L •i™ IT! . "TTt : 1 IVI t_ iii; i i i F i l Hi: i . i H L ;n ;j iP-PpPi j...:-.. Fii rF :-j-: H I iFF:-P — •:-rr:r.::: .... .:-;:t-v.:. ---"-' !" - '• il. ii!i:i !i!l: HiiF-'H ~:::r:::: l i D/!LTA NJOT v'A 1 A RI F 1 ;.. .U IV M T — I . ::::.: | : " : . ' IVI . t i ii::-1:::.;— i .:: :.::: r..'--'..'..\..:.~" r.-:::!:.: . \:t:pr—ll::|:::::: p£piipP iHi •iii'iii iiiiiiiii "I:.)::•::• i P i l i F r FF i iiiii: !P' 1 : ::~r' r u:| ' M F O FL L IMPT <? ON /ER r— :.: i:::; .: : L HL IVI H i i i - i . : : ' ! : : : : i HliilF :jii'n r;i; i i i i i- — P P P P P p F F iiripri:- .::::."!'.: r. !PpF 1 ( C A L O 5 C 0 , 0 ( )0 RIES ::.: t_ t_ FTTTTT^T" F i l l _: ilplH: i i i p : : iiiii ; nrirrri: :. |": Fiji::; i Fliili :~ii:.— PPPP F F H;:.: L7. " _: iiiiili PPiil :::;ii:i • i i i ' i ; ; -' IV»,1 I TTr7rrTJ™~r l_ L F-Fiin: P i p P -.::::}-.— LL IL:SA iiiijiiii -HlliP :::::.:;:::::' ""'I iipiiiiii PlHiP i-ipiiiP —i — ir;ii;;Fi :.":;r:r:~ iHllilli I —1 :::']::: iii;-;:; — L L-L ii" F: . . : : : i : . : : , i • • -iiiii;; ipiiiiii i .P_.Ppp _„4.:_i.: 11-: P P i l F i PiP. -.: I •:: : : •: iiiiiiiii :-_- i iFi.Hi IL' ""...[  P i P P  -ippi-i „:.;.i.nn i-l-P :...]-..:.  :: • : i.:::  ' ''  IFIHFH F-i-FiF  :  r  : : : : ! : : : :  •  ;:  ."."•:  : : : • ] : : : :  :  H H J ; ; . :  :•::  :  • • ; ' • ! " : : -  (  -4  IV1  ; H  :  ;  H : H  :  —  : :  — • : ] - : : : .  -  H  '  H  ;  _•::-•!.:..:.:-  ::.::L  -  H H ; ;  :.:::::|_:"-.  •'.:—.JT:.:::.  -•_..::::L":-  :  .:::;;  1  ":.-.::!  —  : . .  .I::-LJ.:-.":-;  I  • • • • ) - - — •  "•]-.7\"~.7:.:.  —TT.—  ; ; ;  .:::::!:--:::  !:::-"•-  :  :  :  :  I I H :  r  (  :  :  :  :  ;  :  : : : : ! : . : . :  : : . . ] : . : :  :'_  :::  :.:.:.:.I::.::J • • •• I  :  : ; ; ; - ) ; ; • ;  .•:)•:•:•::  :  1  •  •iPi Hp  • : i.::- 'ii|  :  -A , V  -• :::-:: : . . . . ; -  -F+F-  •  :::  \'  -  :  •  -: i t  •  2 •  i  E n e rgy  •  ~ •'. i '• - i n P P F-PPr ':Fi:'"" :  :  • •  1  :  ::•-  -. |.1  i  pp e ' i : ' L i i i : : ; ; : ii:;: ;.:Ji;wi:J:j::: i Intake, Calories x 10° (A.D.E.) . I F . P i ! , p. : : . | - : : : : ! : : : . | ; , i ; . | : | . : | !• • :  .  . .  :  I."  .  .,  •' i '  -pi..p  i_iii:.:.:: :: •  i . ' : :  iPli"::  ~PTV  : i. :.. . ' •• •t-• .::.'::::  : | i . .::•••  .'.  i  :'  :"':™: • r :  i:ii :.p.:..  :.!:!;.' •  r; i;• | • :• S H H i " r  ...iipi:: IF'i'l!' . . • 1 i::: • .iiii:'.-'  .  I.i:.:..:"i !" " :  : P P -Hiii. •i;pr  iipl" •  i . . .  "  | . . .:  :  !' •• r.. :  : :  ..  .  •:• p i .  .1 : • .  . . : ; : • : • . _  ' i "'  s k e l e t a l d i m e n s i o n s t h a t u n i q u e l y i d e n t i f y t h e v a r i o u s energy regimes.  I f l a r g e groups o f a n i m a l s r e p r e s e n t i n g a l l  p o s s i b l e n u t r i t i o n a l r e g i m e s were a v a i l a b l e as s t a n d a r d s , t h e energy regime o f any w i l d fawn c o u l d be d e t e r m i n e d Its  directly.  s k e l e t a l measurements would s i m p l y be compared w i t h t h e  s t a n d a r d s u n t i l t h e most s i m i l a r one was s e l e c t e d .  However,  t h e number o f s t a n d a r d s r e q u i r e d would be p r o h i b i t i v e .  Just  enough a n i m a l s were r e a r e d i n t h e l a b o r a t o r y t o t e s t t h e t h e o r i e s p e r t a i n i n g t o s k e l e t a l growth.  The a n i m a l s can a c t  as s t a n d a r d s f o r d i r e c t comparison, b u t o n l y i f t h e w i l d t r e a t m e n t happens t o c o r r e s p o n d . be e v a l u a t e d  Those f a l l i n g between must  by an i n t e r p r e t a t i o n o f t h e p a t t e r n o f growth  elicited.  I I I . Regimes o f N u t r i t i o n t o 112 Days. Of t h e s i x n u t r i t i o n a l regimes c o n s i d e r e d  a t 112 days as  p o s s i b l e representations o f w i l d d i e t s , l a b o r a t o r y animals were r a i s e d on o n l y f o u r .  One male fawn was r e a r e d on H H , t  l  one male and two f e m a l e s on 'MH', f o u r males on MM , and one T  female on 'ML*.  T  E a r l i e r , a mention was made o f an e x p e r i -  m e n t a l p l a n i n w h i c h e v e r y regime i n c l u d e d a t l e a s t two male fawns.  At t h a t time i t was a l s o i n d i c a t e d t h a t  difficulties  had  a r i s e n i n making a n i m a l s conform e x a c t l y t o t h e p l a n .  The  r e s u l t o f t h i s i s o b v i o u s here w i t h no a n i m a l s a t a l l  r e p r e s e n t i n g HM r  r  o r HL'. T  I n a d d i t i o n t o t h e s e s i x r e g i m e s , two o t h e r s a r e i n c l u d e d t o add f u r t h e r s u p p o r t t o t h e t h e s i s t h a t growth g r a d i e n t s a r e  131 b e i n g m a n i f e s t e d i n response t o n u t r i t i o n a l s t r e s s e s .  An  L L * regime was imposed upon two male and one female  fawns  f  and ' LH* upon one male fawn. The e f f e c t o f each p a t t e r n o f a l i m e n t a t i o n w i l l be desc r i b e d i n terms o f t h e s t a n d a r d t o t a l energy i n t a k e s t h a t a r e a s s o c i a t e d w i t h t h e s k e l e t a l d i m e n s i o n s a t t a i n e d by an a n i m a l on t h a t r e g i m e .  That i s , t h e s k e l e t a l d i m e n s i o n s w i l l be  used t o e s t i m a t e t o t a l energy i n t a k e e m p l o y i n g t h e r e f e r e n c e lines  ( p . 14S) t o r e l a t e t h e measurements t o t h e energy l e v e l .  By t a b u l a t i n g t h e d i f f e r e n c e s between t h e s e apparent v a l u e s and t h e t r u e t o t a l energy i n t a k e , a c h a r a c t e r i s t i c p a t t e r n o f d e v i a t i o n s can be produced f o r each r e g i m e . I t must be p o i n t e d o u t t h a t t h e p a t t e r n s o f d e v i a t i o n s d e s c r i b e d can be, and have been, p r e d i c t e d a c c o r d i n g t o a t h e o r y o f growth t h a t encompasses a l l r e g i m e s . the  *HH  T  F o r example,  regime i s r e p r e s e n t e d by one a n i m a l o n l y , b u t t h e  t y p e o f response t o be e x p e c t e d i s d e t e r m i n e d a c c o r d i n g t o a t h e o r y t h a t i s s u p p o r t e d by t h e t w e l v e a n i m a l s t h a t were r a i s e d on t h e o t h e r r e g i m e s .  The a n i m a l o r a n i m a l s on a  p a r t i c u l a r energy regime a i d i n e s t a b l i s h i n g t h e magnitude o f the  r e s p o n s e and a l s o p r o v i d e a check on t h e methods o f  i n t e r p r e t a t i o n b e i n g employed. On page 150 t h e s k e l e t a l measurements were a r r a n g e d i n t o six  groups and an i n d i c a t i o n was made o f t h e d i r e c t i o n o f  progress o f the a x i a l gradients.  I n t h e f o l l o w i n g diagram,  t h e s e same g r a d i e n t s a r e d e p i c t e d i n an attempt t o demonstrate  the r e l a t i v e t i m e s o f o c c u r r e n c e  o f t h e peak growth r a t e s  (see F i g u r e 5$)« T h i s diagram i s i n t e n d e d o n l y as a summary o f t h e o b s e r v a t i o n s on s k e l e t a l growth.  N e i t h e r the l e n g t h s nor  t h e a b s o l u t e p o s i t i o n s o f t h e arrows r e p r e s e n t i n g t h e g r a d i e n t s a r e known t o be e x a c t , but t h e y s e r v e t o show t h e d i v i s i o n o f t h e g r a d i e n t s i n t o t h e e a r l y , moderate and l a t e regions.  Throughout t h e f o l l o w i n g d i s c u s s i o n t h e f o r e l i m b  w i l l o f t e n be r e f e r r e d t o as e a r l y m a t u r i n g ,  but as can be  seen i n t h i s d i a g r a m , t h e d i s t a l end ( f o r e cannon) i s much e a r l i e r t h a n t h e p r o x i m a l end ( s c a p u l a ) .  Similarly,  there  i s a span i n each g r a d i e n t t h a t s h o u l d e n a b l e t h e d e t e c t i o n o f growth r e s p o n s e s ,  i n a d d i t i o n t o t h e use o f comparisons  between t h e groups o f measurements. The width.  s i x t h gradient i s that of the increase i n v e r t e b r a l Although  t h i s aspect  i s l a t e maturing,  a  fairly  r a p i d growth r a t e i s e x h i b i t e d d u r i n g t h e peak r e s p o n s e i n the moderate r e g i o n s .  D u r i n g t h i s time t h e l e n g t h t o w i d t h  r a t i o s of the vertebrae  remain n e a r l y constant.  I t i s not  u n t i l much l a t e r t h a t t h e r e i s an i n c r e a s e i n t h e growth p r i o r i t y f o r w i d t h and a change i n p r o p o r t i o n s .  Because o f  t h i s t y p e o f r e s p o n s e t h e r e i s n o t h i n g t o be g a i n e d by d i s c u s s i n g t h i s g r a d i e n t a t 1 1 2 days and i t w i l l be i n c l u d e d only at the l a t e r  dates.  As t h e d i f f e r e n t t r e a t m e n t s  a r e d i s c u s s e d , i t w i l l be  n o t e d t h a t t h e v a r i o u s r e g i o n s o f t h e s k e l e t o n change i n t h e  Figure  5#  A Diagram D e p i c t i n g t h e P o s s i b l e  Relative  Peaks o f Growth P r i o r i t y Through t h e Growth G r a d i e n t s That Have Been Examined.  ore Camnon, Lli^li4ilLLJji.L_ Di m e n s i o n s  w idth  e n g th dist  prox.  mm...  limp  Ve r tebra G o lu m n , Width  post  UJ 21 - V e r t e b r a < Column, Length 0 0  ° -Hind  r  dist  L i mb,  a: - - - , — ^ Length  i  -  a n t.  prox  -'  ore i Li mb,  dist  _ength  prox .:P;:rPiPL  AGE RELATIVE  i. • • • i  T ME- O F  P E A K ..GROWTH  IL  . ...  r  RESPONSE  184 p r i o r i t y l e v e l t o which t h e y a r e a l l o c a t e d .  T h i s i s because  o f t h e . u s e o f c h r o n o l o g i c a l time t o d e s c r i b e a p h y s i o l o g i c a l function.  By 4 months o f age an a n i m a l f e d t o a 'HFP  s t a n d a r d has a l r e a d y s a t i s f i e d t h e major growth o f i t s e a r l i e s t p r i o r i t y components and advanced t o t h e n e x t , on t h e 'LL' t r e a t m e n t  the e a r l y maturing  s t r i v i n g t o grow a t t h e most r a p i d r a t e .  while  regions are s t i l l On t h i s  treatment  (•LL*), t h e l e n g t h s o f t h e t h o r a c i c and lumbar v e r t e b r a e would be i n c l u d e d a l o n g w i t h t h e w i d t h s o f t h e f o r e cannon as b e i n g i n a p r e - p r i o r i t y phase o f growth.  Since the  r e l a t i o n s h i p between n u t r i t i o n a l l e v e l and p h y s i o l o g i c a l a g i n g has n o t been d e s c r i b e d f o r t h e growth o f t h e s k e l e t a l system, i t i s n e c e s s a r y have been reached  Regime *HH*  t o i n f e r which l e v e l s o f p r i o r i t y  by e x a m i n i n g t h e l a b o r a t o r y d a t a .  Animal V l l  Energy I n t a k e 230,000 C a l o r i e s  Because t h e boundary around t h e d a t a p o i n t s o f an e a r l y maturing  d i m e n s i o n has a h o r i z o n t a l upper (BC) s i d e (pp. 147-  148 and F i g u r e 9 ) , t h e r e f e r e n c e l i n e f a l l s t o t h e l e f t o f t h e *HH  T  treatment  data.  T h i s means t h a t t h e s e d i m e n s i o n s ,  as t h e l e n g t h s o f t h e l i m b bones, would be e x p e c t e d e s t i m a t e t o t a l energy i n t a k e .  such  t o under-  T a k i n g V l l a s an example, t h e  l e n g t h s o f t h e bones o f t h e f o r e l i m b and t h e l e n g t h s o f t h e c e r v i c a l vertebrae underestimate 34,000 t o 55,000 C a l o r i e s .  t o t a l energy i n t a k e by  I n t h e i n t e r m e d i a t e range o f  p r i o r i t i e s i n c l u d i n g t h e l e n g t h s o f t h e t h o r a c i c and lumbar  v e r t e b r a e , the upper b o u n d a r i e s  around the d a t a p o i n t s have  p o s i t i v e s l o p e s (e.g. F i g u r e s 14 t o 17) t e n d i n g t o move the r e f e r e n c e l i n e s t o w a r d the HH ,  t  data.  Except f o r T-j_, the  t h o r a c i c and lumbar l e n g t h s o f V l l g i v e a v e r y c l o s e o f t o t a l energy i n t a k e .  estimate  T]_ d e m o n s t r a t e s the same h o r i z o n t a l  BC s i d e t o i t s d i s t r i b u t i o n o f d a t a as do the l i m b bones and cervicals.  T h i s i s p r o b a b l y a r e f l e c t i o n o f t h e shape o f  T^  w i t h the p o s i t i o n o f t h e pre-zygopophyses b e i n g s i m i l a r t o t h a t of the c e r v i c a l s .  Very l a t e m a t u r i n g  d i m e n s i o n s such as  the p r o x i m a l and minimum w i d t h s o f the f o r e cannon a g a i n underestimate  t o t a l energy i n t a k e because the l a g i n growth  r e s p o n s e c r e a t e s a h o r i z o n t a l s l o p e t o the BC boundary, s h i f t i n g the r e f e r e n c e l i n e t o the l e f t . i n V l l by a 9 3 , 0 0 0 and 7 6 , 0 0 0 C a l o r i e  Regime MM t  A n i m a l Y2  l  T h i s i s demonstrated  underestimate.  Energy I n t a k e 151,000 C a l o r i e s  »  Y3  "  "  120,000  "  »  Y6  »  "  140,000  "  "  Y8  »  "  149,000  "  There i s an o v e r e s t i m a t i o n o f t o t a l energy i n t a k e by e a r l y maturing  d i m e n s i o n s on t h i s t r e a t m e n t  i n c a u s a t i o n t o the u n d e r e s t i m a t i o n The  on the  the  that i s s i m i l a r •HH'  treatments.  r e f e r e n c e l i n e b i s e c t s the h o r i z o n t a l upper boundary  l e a v i n g the  •I'M  1  d a t a t o the l e f t  o f the l i n e .  T h i s means  t h a t the r e f e r e n c e l i n e w i l l a s s o c i a t e t h a t s i z e o f measurement w i t h a h i g h e r energy i n t a k e .  Y2 and Y3 both show the  186 l i m b d i m e n s i o n s o v e r e s t i m a t i n g t o t a l energy i n t a k e by 20,000 to  50,000 C a l o r i e s w h i l e Y6 and Y8 show much s m a l l e r o v e r -  estimates.  These two p a i r s o f a n i m a l s are s e p a r a t e d here  because b o t h Y2 and Y3 adhered t o the 'MM' t r e a t m e n t a s i n t e n d e d , w h i l e Y6 and Y8 d i d n o t .  They a v e r a g e d * MM f  f  f  because o f p e r i o d i c l a p s e s o f growth r a t e due t o i n t e s t i n a l i n f e c t i o n s and c o n c u r r e n t l o s s o f a p p e t i t e , i n t e r s p e r s e d w i t h periods o f f u l l feeding.  The c o m p l e x i t y o f t h e v a r i a t i o n s i n  energy l e v e l f o r t h e s e a n i m a l s i s beyond the scope o f t h i s paper and the d i s c u s s i o n o f the MM ?  T  t r e a t m e n t w i l l be con-  f i n e d t o Y2 and Y3. The t h o r a c i c and lumbar v e r t e b r a e were not a v a i l a b l e f o r t h e s e a n i m a l s and, t h e r e f o r e , t h e e f f e c t upon t h e s e d i m e n s i o n s cannot be shown.  It i s likely,  however, t h a t  d u r i n g t h e second p o r t i o n o f t h e *MM' regime t h e s e v e r t e b r a e would have r e a c h e d a h i g h e r growth p r i o r i t y and would have become more s u s c e p t i b l e t o t h e r e d u c t i o n . will  Although there  s t i l l be a tendency t o o v e r e s t i m a t e t o t a l energy  intake  t h e d e v i a t i o n s w i l l be s m a l l e r t h a n t h o s e shown by t h e l i m b bone l e n g t h s .  The d i s t a l w i d t h o f t h e f o r e cannon, w h i c h  c o r r e s p o n d s i n p r i o r i t y t o the i n t e r m e d i a t e d i m e n s i o n , shows a s m a l l o v e r e s t i m a t e o f 10,000 C a l o r i e s f o r Y3 w i t h t h i s measurement f o r Y2 m i s s i n g . In t h e l a t e m a t u r i n g d i m e n s i o n s -- i . e . , the f o r e cannon w i d t h s , r a p i d growth has not begun a t t h i s time and, o n l y a s m a l l r e d u c t i o n i n growth can be e x p e c t e d .  therefore,  The r e s u l t  187 i s a h o r i z o n t a l BC s i d e t o the d a t a boundary and a r e f e r e n c e l i n e t h a t f a l l s t o the r i g h t o f the 'I'M' d a t a . e s t i m a t e i s demonstrated  The  over-  by Y2 and Y3 t o be l a r g e , i n t h e s e  a n i m a l s r a n g i n g between 29,000 and 52,000 C a l o r i e s .  Regime 'ML' The  A n i m a l V14  Energy I n t a k e 99,000 C a l o r i e s  r a p i d growth p e r m i t t e d i n the e a r l y m a t u r i n g  d u r i n g t h e 'M'  p o r t i o n of t h i s treatment  o v e r e s t i m a t i o n o f t o t a l energy  intake.  regions  s h o u l d l e a d t o an In the intermediate  r e g i o n , t h e growth i s more s e v e r e l y r e s t r i c t e d by t h e  'L'  part of the treatment which f a l l s at a time o f r e l a t i v e l y high p r i o r i t y .  However, t h e b o u n d a r i e s around t h e measure-  ments f o r t h e s e d i m e n s i o n s  have s l o p e d BC s i d e s and a  r e s u l t a n t l e s s e r s l o p e t o the AB s i d e .  The r e f e r e n c e l i n e s  move t o t h e r i g h t s t i l l a l l o w i n g an o v e r e s t i m a t e o f t o t a l energy  i n t a k e by t h e s e measurements.  The  l a t e maturing p a r t s  do not r e a c h a s t a t e o f h i g h p r i o r i t y by 112 days a t t h i s l e v e l o f energy i n t a k e and, t h e r e f o r e , cannot u t i l i z e p o r t i o n o f the t r e a t m e n t  f o r r a p i d growth.  The  the  'M'  increase i n  t o t a l energy i n t a k e w i t h o u t a c o r r e s p o n d i n g i n c r e a s e i n growth moves t h e d a t a p o i n t s t o t h e r i g h t o f the r e f e r e n c e l i n e s c a u s i n g an u n d e r e s t i m a t i o n o f t o t a l energy The measurements o f V14  intake.  s u p p o r t t h e above i n t e r p r e t a t i o n .  S i n c e i t w i l l be shown l a t e r t h a t sex d i f f e r e n c e s i n growth responses  s h o u l d o n l y be e x p e c t e d on t r e a t m e n t s l e a d i n g t o a  r e l a t i v e l y h i g h energy  i n t a k e , the p a t t e r n e x h i b i t e d by  V14  188 can be used as a r e f e r e n c e f o r male fawns a l s o .  Regime  'MH  1  A n i m a l V21  Energy I n t a k e 165,000 C a l o r i e s  tt  V7  tt  tt  196,000  tt  tt  Y7  it  tt  216,000  tt  There i s a t e n d e n c y f o r an i n c r e a s i n g e s t i m a t e o f t o t a l energy i n t a k e p r o x i m a l l y t h r o u g h t h e l i m b g r a d i e n t s caused by the  a b i l i t y o f the r e l a t i v e l y  l a t e r m a t u r i n g p a r t s t o respond  t o t h e 'H' t r e a t m e n t i n t h e second i n t e r v a l .  V21 under-  e s t i m a t e s t o t a l energy i n t a k e s l i g h t l y by t h e cannon bones, r i s i n g t o an o v e r e s t i m a t e by t h e s c a p u l a and femur.  V7 and  Y7 show t h e same p a t t e r n b u t a much l o w e r range o f e s t i m a t e s . The d i f f e r e n c e s between t h e e f f e c t e x h i b i t e d by V21 and by V7 are  caused by t h e d i f f e r e n c e s i n t o t a l energy i n t a k e .  When  one r e c a l l s t h e p o s i t i o n i n g o f t h e r e f e r e n c e l i n e s f o r t h e e a r l y m a t u r i n g d i m e n s i o n s , i t i s seen t h a t a t h i g h e r l e v e l s o f energy i n t a k e 'MH' d a t a would t e n d t o f a l l f u r t h e r t o t h e right  a c c o u n t i n g f o r t h e e f f e c t seen h e r e . The r e g i o n s r e a c h i n g a h i g h growth p r i o r i t y a t an i n t e r -  mediate t i m e u t i l i z e t h i s p a t t e r n o f energy i n t a k e v e r y w e l l because t h e 'H' p o r t i o n o f t h e t r e a t m e n t c o i n c i d e s w i t h t h e i n c r e a s e d a b i l i t y t o grow. intake For not  An o v e r e s t i m a t e o f t o t a l energy  results. t h e l a t e m a t u r i n g r e g i o n s , a h i g h growth p r i o r i t y has  y e t been r e a c h e d .  The r e l a t i v e l y h i g h l e v e l o f energy  i n t a k e cannot be u t i l i z e d f u l l y r e s u l t i n g i n an u n d e r e s t i m a t e  189 o f t o t a l energy i n t a k e by the f o r e cannon There i s no a p p a r e n t e v i d e n c e t h a t any  widths. sex  differences  are o c c u r r i n g i n t h e growth r e s p o n s e s o f the v a r i o u s  skeletal  e l e m e n t s b u t , because the energy i n t a k e s o f the male  and  f e m a l e s do not  c o i n c i d e , a d e f i n i t e statement t o t h i s  cannot be made.  Since  any  d i f f e r e n c e s t h a t might o c c u r would  be e x p e c t e d t o be enhanced by the onset o f s e x u a l t h e s e s h o u l d be g r e a t e s t The t  MH  d i f f e r e n c e s are not t  maturity,  i n the p h y s i o l o g i c a l l y o l d e s t  regime w h i c h i s the t r e a t m e n t h a v i n g the h i g h e s t  available.  ,  I  i n the r e s t the  s t i l l r e m a i n s t h a t t h e y might  Regime 'LH'  of  A n i m a l Y4  differences possibility  occur.  Energy I n t a k e  164,000 C a l o r i e s  T h i s t r e a t m e n t i s s i m i l a r i n e f f e c t t o the  'MH  t h a t m a t u r a t i o n w i l l have been d e l a y e d s l i g h t l y . Y4 i s not a s t r o n g example o f the  'LH'  o f the regime.  However, t r e n d s  treatment.  The  except  1  The  animal  t r e a t m e n t because i t  i s j u s t s l i g h t l y o v e r the boundary from 'M'  i n the  are e x h i b i t e d t h a t  'L'  part  identify  b e s t i n d i c a t i o n i s shown i n the  l e n g t h s o f the l i m b . b o n e s . estimate  level  female s t a n d a r d s are  i n t o t a l energy i n t a k e , sex  not be e x p e c t e d , w h i l e  'LH'  the  I t i s p r o b a b l y s a f e t o say t h a t i n a l l t r e a t m e n t s  f a l l i n g below MH should  animals.  i n d i c a t e d , a t l e a s t n o t i c e a b l y , on  energy i n t a k e f o r w h i c h b o t h male and  the  effect  There i s an i n c r e a s i n g o v e r -  p r o x i m a l l y from the f o r e cannon t h r o u g h the humerus,  f o l l o w e d by a d e c r e a s e i n the s c a p u l a . i n t h i s animal,  but the i n c r e a s e  The  radius i s missing  i n the o v e r e s t i m a t e  along  the  190 hind limb g r a d i e n t supports the t r e n d .  I f t h e p e l v i s was  a v a i l a b l e , i t s l e n g t h l i k e l y would a l s o show a overestimate.  decreased  Although the only vertebrae t h a t are  a v a i l a b l e f o r t h i s a n i m a l a r e t h e c e r v i c a l s , t h e s e showed a marked o v e r e s t i m a t e t h a t i n c r e a s e s p o s t e r i o r l y .  This i s i n  accordance w i t h the p r i o r i t i e s f o r these dimensions,  being  s i m i l a r t o t h o s e o f t h e l i m b bone l e n g t h s . The w i d t h s o f t h e f o r e cannon o v e r e s t i m a t e t o t a l energy i n t a k e , m a i n l y because t h e y a r e a f f e c t e d v e r y l i t t l e by t h e t  L  t  treatment  d u r i n g t h e f i r s t i n t e r v a l and grow t o s i z e s  s i m i l a r t o t h a t reached  on  'MH'.  The i n t e r m e d i a t e r e g i o n s would p r o b a b l y g i v e a r e a s o n a b l e e s t i m a t e o f t o t a l energy i n t a k e , d e c r e a s i n g s l i g h t l y along the gradient.  The t h o r a c i c and lumbar  v e r t e b r a e a r e n o t a v a i l a b l e f o r Y4 so t h i s cannot be v e r i f i e d .  Regime LL»  Animal V19  T  Energy I n t a k e 94,000 C a l o r i e s  "  V20  "  "  98,000  "  "  VI$  "  "  98,000  "  On t h e 'LL' t r e a t m e n t , a l l measurements must c o i n c i d e f a i r l y c l o s e l y w i t h t h e r e f e r e n c e l i n e s i n c e t h e range becomes more and more narrow a t t h e l o w e s t energy i n t a k e s . r a i s e d on L L ,  t  Animals  t r e a t m e n t s a r e c h a r a c t e r i z e d by a v a r i a b i l i t y  i n e s t i m a t e s o f t o t a l energy i n t a k e t h a t a r e g r e a t e r o r s m a l l e r t h a n t h e t r u e v a l u e , but s t r o n g t r e n d s a r e not apparent.  191 T h i s completes t h e coverage o f t r e a t m e n t s a t 112 days f o r which t h e r e a r e l a b o r a t o r y c o u n t e r p a r t s .  There a r e two more  t r e a t m e n t s t h a t a r e o f i n t e r e s t because o f t h e l i k e l i h o o d o f t h e i r o c c u r r e n c e under f i e l d  conditions.  An attempt w i l l now  be made t o a n t i c i p a t e t h e e f f e c t s o f t h e s e n u t r i t i o n a l regimes on s k e l e t a l  growth.  Regime HM*  No l a b o r a t o r y a n i m a l s  f  The growth o f each o f t h e e a r l y m a t u r i n g  dimensions  s h o u l d n o t be e x p e c t e d t o d i f f e r much from t h a t on t h e H H t  t r e a t m e n t and, t h e r e f o r e , s h o u l d n o t u n d e r e s t i m a t e energy i n t a k e t o t h e same e x t e n t . for  t  total  A reasonably close value  t o t a l energy i n t a k e s h o u l d r e s u l t d i s t a l l y w i t h a s l i g h t  i n c r e a s e p r o x i m a l l y because o f t h e s h i f t o f t h e r e f e r e n c e l i n e to  the r i g h t .  The t h o r a c i c and lumbar v e r t e b r a e , because o f  t h e i r h i g h e r p r i o r i t i e s f o r growth d u r i n g t h e second  interval,  w i l l be more a f f e c t e d and w i l l t e n d t o u n d e r e s t i m a t e  total  energy i n t a k e .  The f o r e l i m b w i d t h s w i l l be l i t t l e  affected  by t h i s t r e a t m e n t and w i l l g i v e a s m a l l o v e r e s t i m a t e o f t o t a l energy i n t a k e -- i . e . , t h e s i z e villi at  a l o w e r t o t a l energy  Regime HL* f  be s i m i l a r t o H H but T  T  intake.  No l a b o r a t o r y a n i m a l s  The s k e l e t a l d i m e n s i o n s o f a n i m a l s r a i s e d on t h i s  treat-  ment s h o u l d a l l t e n d t o u n d e r e s t i m a t e t o t a l energy i n t a k e .  In  the l i m b g r a d i e n t s , t h e d i s t a l r e g i o n s w i l l c l o s e l y approximate the t r u e energy i n t a k e , but d e c r e a s e i n v a l u e p r o x i m a l l y . most pronounced u n d e r e s t i m a t e s lumbar r e g i o n s because o f the time o f h i g h growth a c t i v i t y .  The  w i l l e x i s t i n t h e t h o r a c i c and l  L  t  r e s t r i c t i o n o c c u r r i n g at a  I n the v e r y l a t e r e g i o n s , no  pronounced e f f e c t s h o u l d be noted by t h i s t r e a t m e n t  and  the  e s t i m a t e o f t o t a l energy i n t a k e s h o u l d approach the t r u e v a l u e .  IV.  Energy I n t a k e s o f W i l d Fawns t o 112  A n i m a l YF17  Age  YF19  "  YF16  i n Days 116  11  YF18  "  YF20  11  » " " "  Days.  Energy I n t a k e  1  197,000 C a l o r i e s  118  «  »  205,000  "  116  »  "  198,000  "  117  "  118  "  "  170,000  "  "  180,000  "  Each o f the above a n i m a l s i s r e p r e s e n t e d by the major l i m b bones except t h e p e l v i s , and by c e r v i c a l s t h r e e and A n i m a l YF18 Although  five.  i s complete w i t h p e l v i s and v e r t e b r a l column.  o n l y a p a r t i a l s e r i e s o f measurements i s a v a i l a b l e ,  f o u r out o f f i v e can be p l a c e d i n the HM ,  confidence. evidenced  t  o r MH* regime w i t h t  Assuming t h a t sex d i f f e r e n c e s i n growth are  a t t h i s age  (see pp. 61-62), each o f the  not  field  animals  can be compared t o the l a b o r a t o r y s t a n d a r d s .  For  animals  YF17,  regimes  1  Estimated to  166.  YF19,. YF16  and YF20, t h e r e are o n l y two  from s k e l e t a l d i m e n s i o n s a c c o r d i n g t o pp.  161  t h a t would be c o m p a t i b l e w i t h t h e i r s k e l e t a l d i m e n s i o n s -- *HM  t  and  'MH . 1  The t o t a l energy i n t a k e s p r e v i o u s l y e s t i m a t e d  for  the a n i m a l s (see T a b l e 5) f a l l w i t h i n the energy i n t a k e s  cal-  c u l a t e d f o r these regimes. I n the l a b o r a t o r y ,  ' HM'  was not r e p r e s e n t e d  and t h e  p a t t e r n o f i n t e r c e p t s f o r t h i s regime c o u l d o n l y be d e s c r i b e d in  generalities.  As t h e y were d e s c r i b e d  there  i s very  d i f f e r e n c e i n the p a t t e r n s o f i n t e r c e p t s between t h e the  'MH r e g i m e s . T  that suggests that  little  * HM* and  There i s , however, one p i e c e o f e v i d e n c e '1'  i s the regime t h a t was f u n c t i o n a l i n  p r o d u c i n g the growth o f t h e s e fawns.  The w e i g h t s o f the  fawns  c o l l e c t e d i n August ( 4 9 days) i n d i c a t e a h i g h p l a n e o f n u t r i t i o n i n the f i r s t i n t e r v a l (see p 17$).  I f i t can be  assumed t h a t t h o s e fawns c o l l e c t e d a t 4 9 days were t a t i v e o f the p o p u l a t i o n  represen-  from w h i c h the 112 day o l d fawns were  c o l l e c t e d , t h e n HM* i s the most p r o b a b l e regime a c t i n g on t h e T  w i l d fawns. The  p a t t e r n o f i n t e r c e p t s produced by the s k e l e t a l  measurements o f a n i m a l Y F l B d i f f e r s from t h o s e o f the f o u r fawns.  other  The s k e l e t a l measurements p r e s e n t a p a t t e r n o f  i n t e r c e p t s t h a t i s c h a r a c t e r i s t i c o f the MM' regime b u t a t T  a t o t a l energy i n t a k e t h a t i s s l i g h t l y above the maximum c a l c u l a t e d f o r t h i s regime.  I t must be remembered, however,  t h a t the t o t a l energy i n t a k e f o r the w i l d fawns was e s t i m a t e d from a body w e i g h t r e g r e s s i o n and from the  s k e l e t a l measure-  ments, n e i t h e r o f w h i c h g i v e a p r e c i s e answer.  I f instead of  the estimated  170,000 C a l o r i e s the t o t a l energy i n t a k e had  been  160,000 C a l o r i e s , the a n i m a l would have f a l l e n w i t h i n the c l a s s i f i c a t i o n and  the p a t t e r n o f i n t e r c e p t s r e s u l t i n g from  t h i s change would s t i l l be c o m p a t i b l e w i t h the T h i s a n i m a l has, t i v e o f the  'MM'  'MM*  t h e r e f o r e , been c l a s s i f i e d as a  regime. representa-  •MM* r e g i m e .  I t w i l l be n o t i c e d t h a t the w i d t h s o f the f o r e cannon have not been used i n t h i s e v a l u a t i o n . t h i s age  These measurements a t  appear t o be v e r y s u b j e c t t o i n d i v i d u a l v a r i a t i o n s  which r e s u l t i n large d i f f e r e n c e s i n estimates intake.  T h i s w i l l be d i s c u s s e d  o f t o t a l energy  at g r e a t e r l e n g t h s when the  measurements o f the cannons c o l l e c t e d from h u n t e r k i l l e d are  V.  examined.  Regimes o f N u t r i t i o n t o 1 7 5 From b i r t h t o 112  Days.  days o f age  regime o f a fawn can be e v a l u a t e d by the use o f body weight and the t i m e t o 175  days o f age  days i n t r o d u c e s  a c c u r a t e l y and  Extending  f a c t o r s which tend to  diminish 112  For animals reared to  delineate  i n t e r v a l s of time w i t h  p o s s i b l e energy regimes b e i n g At 175  energy  conveniently  19 s k e l e t a l measurements were used t o  the n u t r i t i o n a l p l a n e s over two  or L ' ) .  i t appears t h a t t h e  s k e l e t a l measurements.  the s e n s i t i v i t y o f the a n a l y s i s .  six  deer  considered  (H T  o r M;  only H,  M  days t h e r e are t w e l v e energy regimes r e s u l t i n g  from the c o m b i n a t i o n s o f the p o s s i b l e t r e a t m e n t s i n each o f the t h r e e i n t e r v a l s ('H  o r M;  H, M o r L; M o r L»):  the  19  s k e l e t a l measurements must now s o l v e t h e n u t r i t i o n a l  regime  over t h r e e i n t e r v a l s , r e s u l t i n g i n a l o s s o f p r e c i s i o n .  The  a n a l y s i s i s f u r t h e r hampered by t h e f a c t t h a t t h e r e i s a f i n i t e l i m i t t o t h e growth o f each s k e l e t a l d i m e n s i o n , and i n the e a r l i e r maturing p o r t i o n s o f the s k e l e t o n t h i s l i m i t i s b e i n g approached by 175 days.  These d i m e n s i o n s a r e e f f e c t i v e l y  removed as i n d i c a t o r s o f n u t r i t i o n a l regime, e s p e c i a l l y a t h i g h e r l e v e l s o f energy i n t a k e . Because o f t h e c o m p l e x i t y o f t h e i n t e r a c t i o n s o f energy l e v e l , growth p r i o r i t y , and compensation, i t has n o t been p o s s i b l e t o t a k e each n u t r i t i o n a l regime and d e s c r i b e t h e r e s u l t i n g o r expected s k e l e t a l p a t t e r n s .  Only 6 o f t h e 12  regimes under c o n s i d e r a t i o n a r e r e p r e s e n t e d by l a b o r a t o r y a n i m a l s , and o n l y 3 o f t h e s e by more t h a n one a n i m a l ( T a b l e 9, pp. 4 5 - 4 7 ) .  The r e m a i n i n g 11 a n i m a l s a r e d i s t r i b u t e d among  6 a d d i t i o n a l t r e a t m e n t s w h i c h , a l t h o u g h u n l i k e l y as n a t u r a l f e e d i n g r e g i m e s , were i n c l u d e d t o h e l p t o d e t e r m i n e growth priorities.  U s i n g an e m p i r i c a l a p p r o a c h , i t must be assumed  t h a t t h e s k e l e t a l p a t t e r n g e n e r a t e d by each o f t h e s e a n i m a l s somehow r e p r e s e n t s t h e n u t r i t i o n a l regime on w h i c h t h e a n i m a l was r e a r e d .  However, when t h e s k e l e t a l p a t t e r n s were examined  i t was found t h a t no s t r o n g g e n e r a l i z a t i o n s c o u l d be made. P a r t o f t h e problem i s i n a l l p r o b a b i l i t y r e l a t e d t o t h e wide range o f energy i n t a k e s p e r m i s s i b l e w i t h i n each i n t e r v a l o f the regime t h a t , by t h e end o f t h r e e i n t e r v a l s , c r e a t e s many complex i n t e r a c t i o n s .  F o r example, t h e * HML  f  regime can have  196 a t o t a l energy i n t a k e r a n g i n g  from 210,000 t o 315,000 C a l o r i e s  and t h i s d i f f e r e n c e can be p a r t i t i o n e d i n many ways.  The  p a t t e r n o f i n t e r c e p t s produced by t h e s k e l e t o n from an a n i m a l w i t h an energy i n t a k e a t one end o f t h e spectrum need n o t n e c e s s a r i l y be r e p r e s e n t a t i v e  therefore  o f the pattern a t the  o t h e r end. The p a t t e r n s o f energy i n t e r c e p t s g e n e r a t e d by t h e l a b o r a t o r y fawns a r e shown i n Table 6 7 . Not o n l y a r e t h e r e no d i s t i n c t  d i f f e r e n c e s among t h e t r e a t m e n t s ,  any n o t i c e a b l e treatments.  there are not  s i m i l a r i t i e s among t h e a n i m a l s on t h e same  I t has n o t been p o s s i b l e t o p r o v i d e  any f u r t h e r  a n a l y s i s o f t h e growth p a t t e r n s o f t h e l a b o r a t o r y fawns, n o r , as i t w i l l be seen, has i t been p o s s i b l e t o r e l a t e any o f t h e w i l d fawns t o t h e l a b o r a t o r y  VI.  treatments.  P a t t e r n s o f Energy I n t a k e s  o f W i l d Fawns t o 175 Days.  There has a l r e a d y been enough e v i d e n c e accumulated i n earlier  s e c t i o n s t o suggest t h a t t h e sample o f 175 day o l d fawns  from t h e f i e l d w i l l f a l l i n t o one o f t h r e e r e g i m e s : o r MML . t  l  1.  B r i e f l y , t h i s evidence i s : t h e t o t a l energy i n t a k e e s t i m a t e d  for wild  fawns a t 49 and 112 days (see pp. 192-193) 2.  t h e p a t t e r n s o f s k e l e t a l i n t e r c e p t s a t 112 days (see pp. 193-194)  3.  t h e disagreement a t 175 days between t o t a l energy i n t a k e e s t i m a t e d  b y body weight and  t  HML , MHL t  t  t  197  T a b l e 6 7 . The d e v i a t i o n s o f energy i n t e r c e p t s o f t h e s k e l e t a l d i m e n s i o n s from t h e e s t i m a t e s  of actual  energy i n t a k e f o r l a b o r a t o r y and w i l d fawns  A n i m a l No.  U8  M  Sex Total E . I .  1  Pattern E.I.  4.11  MHH  U7  M 4-45  MHM  Bone Length  M 3.77  MHM  UIO  U14  V13  F  F  F  3.37  MHM  3.80  MMM  3.58  MMM  Deviation  F o r e cannon Radius Humerus Scapula  -0.59 -0.31 -0.51  Hind cannon Tibia Femur Pelvis  -0.36 -0.01  Cervical 3 Cervical 5 Cervical 7 Thoracic 1 Thoracic 5 Thoracic 1 0 Lumbar 1 Lumbar 3 Lumbar 5  Y13  0.00 -0.05 -0.30 +0.07  -0.52  -0.41  +0.14  +0.03 +0.05 -0.21  +0.10  +0.35  -0.11  +0.42 +0.27 +0.01  -0.42 -0.58  +0.16  +0.48 -0.60  -0.06  +0.19 +0.15 -0.64 +0.64  +0.41 -0.13  -0.03  +0.28 +0.45 +0.28  -0.43 -0.24 -0.37  -0.42 -0.23 -0.36  +0.13 +0.11  +0.17 +0.53 +0.03 +0.37 +0.59  +0.70  -0.55 -0.65 -0.05 +0.15  -0.69 -0.66 +0.12 0.00  -0.44  -0.09 -0.23 -0.32 -0.53 -0.40 +0.18 -0.49  +0.01 -0.36 +0.21 +0.30 +0.85  +0.32  +0.90  -0.19  -0.41  +0.73 +0.51  -0.15  -0.23  -0.17  -0.14  +0.13 +0.03 -0.86  -0.71  -0.75  +0.43 -0.47 -0.30  -0.13 -0.43 -0.05  -0.11  +0.07 +0.14  -0.29  +0.24 -0.63 -1.00  -0.15 +0.28 -1.20  -0.15 -0.09  -0.63  +O.64 +0.88  +0.41 +0.42  -0.35 -0.40 -0.26 -0.25 -0.42  Bone W i d t h Cervical Thoracic Lumbar  5 5 3  Fore cannon, length) Fore cannon, distal Fore cannon, proximal Fore cannon, minimum 1  -0.74 -0.59  -0.59  0.00  -0.52  +0.13  -0.55  -0.44  -1.13  +0.17  -0.02  -0.54  -0.38  -O.84  -0.94  -0.77  +0.51  -0.27  -0.79  -1.18  -0.04  +0.02  +O.38  +0.38  -0.45  -O.48  Apparent d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  I  I  I  o oo  • • • •p- H-"^>  +  1  +  *  •  1 1  WHH U i M W  +  1 1  J  I I  O OO •  v^J-p-O OOOO +  O O O » » • \_n O O ON<3VO + + +  OOO  +  I  +  +  • • • ONMV*j -p-rOUJ  +  I  +  I  o ooo  +  o  o o o o o o o  *  •  I  +  +  +  I  I  I  +  I  I  I  I  +  +  +  +  I  +  1  l  l  l  l  l  +  l  l  o• o o o o o o o o • • • • • • • •  u i W U f - O H H P ^ CO ON-p- <3 O N O N O -P" vn I  I  I  +  I  +  I  I  1  1  1  +  1  1  +  1  •  •  +  •  1  +  OHO^ OM^O  1  1  +  +  I  W  O  H  +  +  o o o o o o o o o  • • • • • • • • • O H O H H H H H O M H O ^ H H H O H U i  I  OOOO • • • • O  ovo-^ivo  I  I  I  I  H O H M O N CO O N O +  +  O O O O • • • • -P" M 0-P~ M +  o  H  O +  I  I  O O M M-p-.p-.p-  I  +  +  +  •  O +  o o  • • • O M JO V J J COMD  »  •  •  K>.p-VO-FvOOH H I  +  +  +  o• o• •o •o  W M M -f" WOmO  1  1  1  +  OOOO * • • » t— O O O -P~ vO-P~ ON 1  +  +  +  I  o o o o  • • • • -P-1- H - o 1  1  V*J C O O N - J  +  o •o• o• o•  I  +  o ooo  o o o o  o ooo • • • •  I  O O O O O O O O » » • » • • • • H O H O M H W O ONro-j ro co-p- H - - ^ 1  +  OOOO • « • • - J O O H -P--P- M-p-  OOOOOOOOO • • • « • • • • • w r o o f o ^ o r o o M co.0Q.rovow t— ro O N ro  l  •  1  OOOO • • • • O^uiH SO^> COVAJ  O ^ N •VJVJJOOI  +  +  VJOV^JVKJ'OJ ONQN-J CO-  I  J\>OV>J-p-|—'OOr-'H -<3 MD M - P - - O O O CO-M  I  +  oooo  o o o o o o o o o • • • • • • • • *  I  l + l  +  +  OOO  OOO • M-p-vi co.ro M  +  +  \o oa co  I  O OO •  OOO • WHH -O f\) H  l  O O -P" -P~ fO ts) -p- \-> V-n O O M O N W ON ON  OMW u i O H +  I  +  ooo o  I  o o o o o o o o  + I  o o o o M O O O -P~ -P~ OO O 1  +  1  +  o o o o • • • • O O O I—' ro-oro-o +  +  +  +  o o o o H H P W \_n CO ON t—'  199 T a b l e 67-  (Continued)  WFl  WF2  YF38  YF37  U33  V24  M  M  M  F  M  M  T o t a l E.I 1  3.10  3.20  3 .20  2.65  4.01  3.38  Pattern E.1.  Wild  Wild  Wild  Wild  LMH  LHM  Fore cannon Radius Humerus Scapula  0.00 -0.10 +0.23 -0.82  -0.16 -0.08 -0.13 -0.27  +0.33 +0.17 -0.13 -0.69  -0.01 -0.01 -0.14 -0.14  -0.76 -0.57 -0.59 +0.14  +0.15 +0.22 +0.04 +0.27  Hind cannon Tibia Femur Pelvis  -0.04 -0.22 +0.19 -0.18  -0.30 -0.24 -0.07  +0.30 -0.21 -0.24  +0.10 +0.12 +0.14 -0.05  -0.63 -0.51 -0.11 0.00  -0.04 -0.32 0.00 +0.29  -0.39 -0.08 +0.07 +0.13 +0.06  -0.20 -0.54 -0.14 -0.28 -0.13  +0.33 -0.05  +0.29 +0.63 -0.12 +0.34 -0.49 +0.10 -0.47 • -0.02 -0.02  -0.62 -0.21  +0.10 -0.05 -0.16 +0.41 -0.11 -0.27 -0.31 +0.38 +0.31  +0.43 +0.01 +0.12 +0.18 +0.22 -0.11 -0.09 -0.38  +0.08 -0.60 -0.27  +0.22 -0.07 -0.20  -0.78  -0.20 -0.15 -0.06  -0.49 -0.50 -0.88  -0.63 +0.51 -0.76  0.00  . -0.16  +0.33  -0.01  -0.76  +0.15  +0.19  +0.55  +0.68  -0.28  +0.29  +0.10  -0.06  +0.18  +0.18  -0.33  -0.63  -0.25  -0.49  +1.03  -0.01  +0.05  +0.07  -0.36  A n i m a l No • Sex  Bone Length  Cervical 3 Cervical 5 Cervical 7 Thoracic 1 Thoracic 5 T h o r a c i c 10 Lumbar 1 Lumbar 3 Lumbar 5 Bone W i d t h Cervical Thoracic Lumbar  5 5 3  (Fore cannon, • length) F o r e cannon, distal F o r e cannon, proximal F o r e cannon, minimum 1  A p p a r e n t d i g e s t i b l e energy i n t a k e i n C a l o r i e s .  Table 6 7 -  V25  V27  V28  U32  U38  U15  V12  V17  M  M  M  F  F  M  M  F  2.17  2.33  1.99  2.03  1.84  1.86  1.67  LLM  LLM  LLM  LMM  LMM  LLL  LLL  LLL  +0.18 +0.14 +0.24 +0.17  +0.54 +0.27 +0.14 +0.05  -0.20 -0. 14 +0.03 -0. 17  +0.22 +0.30 +0.39 +0.19  +0.07 +0.05 +0.17 +0.08  +0.01 +0.10 '- -0.04 -0.02  +0.23  +0.05 +0.17  +0.39 +0.34 +0.15 +0.19  +0.01 -0.05 +0.10  +0.45 +0.10 +0.17 +0.10  -0. 20 -0. 03 -0. 16 -0. 11  +0.11 +0.29 +0.39 +0.06  +0.14  +0.15 -0.09 +0.09 +0.10  +0.16 -0.02 -0.01 -0.02  +0.38 +0.27 +0.13 -0.11  -0.24  +0.18 +0.11 +0.08 +0.33 +0.08 0.00 +0.01  0. 00 +0. 13 -0. 06 +0. 29 +0. 59 +0. 27 +0. 27  +0.47 +0.16 +0.45 +0.73 +1.06  +0.33  +0.19  +0. 63  +0.17 +0.33 +0.57 +0.45 +0.37 +0.22 +0.49  +0.16 -0.07 -0.03 +0.25 +0.19 +0.18 +0.10 0.00 +0.05  -0.04 -0.17 +0.10 +0.07 +0.16 0.00 +0.02 +0.07  +0. 60 +0. 92 +0. 72  +0.36 +0.09 +O.83  +0.26 +0.47 +0.88  +0.43 +0.35 +0.96  +0.54 -0.09 +0.73  +O.58  +0.42  +0.40 +0.10 +0.80  +0.18  +0.54  -0. 20  +0.22  +0.07  +0.01  +0.23  +0.39  +0.20  +0.17  +0. 41  +0.01  -0.16  +0.28  +0.26  +0.33  +0.10  +0.38  +0. 47  +0.11  +0.22  +O.38  -0.07  +0.70  +0.17  +0.36  +0.28  +0.17  +0.67  +0.01  +0.10  +0.44  2.35  -0.04 -0.11  +0.30 +0.30 +0.19 +0.14 -0.22 -0.20 -0.02  +0.22  +0.16  +0.38 +O.38  +0.56  +0.12 +0.30 +0.28 +0.55 +0.04 +0.77 +0.52 +0.18 +0.58 +0.29 +0.44  +0.29  0.00  +0.21 +0-.90  201  by s k e l e t a l d i m e n s i o n s (see pp. 166-168) The s k e l e t a l measurements from t h e f o u r w i l d fawns c o l l e c t e d i n December w i l l be examined t o d e t e r m i n e i f p a t t e r n s e x i s t  that  w i l l a l l o w f u r t h e r r e f i n e m e n t o f t h e e v a l u a t i o n o f the energy regime. Three o f t h e fawns, WFl, WF2 and Y F 3 8 ,  a l l males, were  e s t i m a t e d t o have v e r y s i m i l a r energy i n t a k e s o f between 3 1 0 , 0 0 0 and 3 2 0 , 0 0 0 C a l o r i e s .  I t i s apparent from f i r s t  glance that t h e patterns o f i n t e r c e p t s f o r these three are not at  a l l similar.  More p a r t i c u l a r l y , none i s s i m i l a r t o t h e one  l a b o r a t o r y example o f HML t  (V26) w h i c h , a t 3 0 8 , 0 0 0 C a l o r i e s ,  t  i s v e r y near i n energy i n t a k e , n o r t o t h e example o f MHL' T  (V8).  The l a t t e r , however,  1  o n l y r e a c h e d an energy i n t a k e o f  2 8 3 , 0 0 0 C a l o r i e s and so t h e comparison may n o t be v a l i d .  No  l a b o r a t o r y r e p r e s e n t a t i v e o f 'MML'* i s a v a i l a b l e f o r c o m p a r i son,  b u t t h i s regime i s e x c l u d e d by t h e h i g h energy i n t a k e s  of t h e s e fawns —  i.e.,  t h e upper l i m i t f o r t h e MML t  t  regime  i s 290,000 C a l o r i e s . The f o u r t h fawn, YF37, a f e m a l e , was much s m a l l e r t h a n the  o t h e r s and had an energy i n t a k e e s t i m a t e d t o be 26$,000  Calories.  A t t h i s energy i n t a k e , t h e regime o f t h e fawn  c o u l d have been any o f t h e t h r e e b e i n g c o n s i d e r e d . s k e l e t a l development or  ,  HML  t  The  d i d n o t c o r r e s p o n d t o e i t h e r t h e MHL f  T  l a b o r a t o r y examples, and 'MML' was n o t r e p r e s e n t e d  as p r e v i o u s l y mentioned.  Because t h e o n l y female i s so  d i f f e r e n t from t h e t h r e e males i t i s t e m p t i n g t o suggest t h a t t h i s may be a s e x - r e l a t e d d i f f e r e n c e .  Hoiirever,  i t soon  becomes apparent t h a t t h i s i s n o t s o .  I t has a l r e a d y been  demonstrated t h a t a t 112 days t h e r e i s n o t an apparent d i f f e r e n c e between males and f e m a l e s , so any d i f f e r e n c e have t o o c c u r i n t h e t h i r d i n t e r v a l .  would  The male fawns, d u r i n g  t h e t h i r d i n t e r v a l , r e f l e c t energy i n t a k e s t h a t a r e below t h e i r maintenance  requirements.  I n o r d e r f o r t h e female t o  be s m a l l e r t h a n t h e male, i t s energy i n t a k e would have t o be d e p r e s s e d w e l l below maintenance.  A • v o l u n t a r y * decrease i n  energy i n t a k e , i . e . , one t h a t i s c o n t r o l l e d from w i t h i n i t h e a n i m a l r a t h e r t h a n by an i n a d e q u a t e s u p p l y o f f o o d , t o such a l o w l e v e l i s n o t borne out by f e m a l e s r e a r e d i n t h e l a b o r a t o r y , n o r i s i t r e a s o n a b l e t o expect such i n an a n i m a l t h a t must s u r v i v e t h e r i g o r s o f w i n t e r .  I t must be t h e  a v a i l a b i l i t y o f f e e d o n l y t h a t i s c o n t r o l l i n g energy  intake  i n t h e t h i r d i n t e r v a l and t h e l o w e s t i m a t e d energy i n t a k e o f t h i s female must be a c c e p t e d as e v i d e n c e o f r e s t r i c t e d  feeding.  T h i s l e a v e s t h e energy regime o f each o f t h e f o u r fawns s t i l l without s o l u t i o n . or  t  MHL  t  Three have been narrowed down t o * H M L  on t h e b a s i s o f energy i n t a k e , not s k e l e t a l d i m e n s i o n s ,  w h i l e t h e f o u r t h c o u l d have been any one o f t h e t h r e e regimes * HML',  T  'MHL*  or M M L . T  t  No m a t t e r how t h e w i l d fawns were com-  pared among t h e m s e l v e s and w i t h t h e l a b o r a t o r y fawns no c o n s i s t e n t s i m i l a r i t i e s c o u l d be found.  Even when t h e e s t i -  mated t o t a l energy i n t a k e s o f t h e w i l d fawns were a l l o w e d t o  203  v a r y , t h e a d j u s t e d i n t e r c e p t s d i d not f a l l i n t o l i n e i n any combination.  V I I . The  Skeletal  Growth o f Fawns t o 3 2 2 Days.  Nine male fawns were r e a r e d t o 3 2 2 days o f age on d e s i g n e d t o t e s t the r e c o v e r y from n u t r i t i o n a l i n f l i c t e d d u r i n g the f i r s t  1 7 5 days o f l i f e .  regimes  inadequacies S i n c e i t was  p o s s i b l e t o demonstrate a dependence between s k e l e t a l  con-  f o r m a t i o n and energy regime a t 1 7 5 days, i t would not  be  p o s s i b l e t o show d i r e c t l y a r e c o v e r y at a l a t e r d a t e . same end was  a c h i e v e d , however, i n a manner not  i n the design of the t e s t . energy  The  anticipated  Regardless of the p a t t e r n of  i n t a k e w i t h i n t h e l i m i t e d number c o v e r e d by  group o f a n i m a l s , t h e s k e l e t a l dimensions s i z e and r e m a r k a b l y  not  this  were s i m i l a r i n  s i m i l a r i n p r o p o r t i o n or  conformation.  Thus, a l t h o u g h i t was not p o s s i b l e t o measure t h e amount o f r e c o v e r y between 1 7 5 and 3 2 2 days, t h e s i m i l a r i t y a t 3 2 2 days showed t h a t r e c o v e r y had  occurred.  W i t h i n the c o n f i n e s o f the t r e a t m e n t s i n c l u d e d i n t h i s p a r t o f t h e s t u d y , i t appears t h a t many r o u t e s may one u l t i m a t e s i z e and c o n f o r m a t i o n o f s k e l e t o n . n e c e s s a r y t o i n c l u d e regimes f  H  t  such as c o n s t a n t  ,  L  lead to  I t would be t  or constant  f o r l o n g e r p e r i o d s o f t i m e t o t e s t the u n i v e r s a l i t y  of'  t h i s s t a t e m e n t , but from the l i m i t e d number o f regimes s t u d i e d h e r e , i t appears t h a t the u l t i m a t e growth and development o f a deer w i l l not be l i m i t e d by i t s s k e l e t o n .  The  variations  i n c o n f o r m a t i o n a t e a r l i e r ages w h i c h r e s u l t from changes i n r e l a t i v e r a t e s o f growth must be e r a s e d by a r e l a t i v e p r o l o n g i n g o f t h e g r o w i n g phase o f t h o s e d i m e n s i o n s p r e v i o u s l y most r e s t r i c t e d .  The e x i s t e n c e o f such a mechanism i s  s u p p o r t e d by t h e work o f McCay e t a l (1935; 1939) on r a t s and by many o t h e r w o r k e r s on d o m e s t i c s t o c k . paper on compensatory  In t h e i r review  growth, W i l s o n and Osbourn  (i960) do  not s i n g l e out t h e s k e l e t o n , b u t i n d i c a t e t h a t t h e r e i s a t e n d e n c y f o r compensation t o l e a d toward t h e c o n f o r m a t i o n and s i z e o f an u n r e s t r i c t e d a n i m a l i n a wide v a r i e t y o f species.  I f i t can be assumed t h a t s i z e and form have s u r -  v i v a l v a l u e , and t h a t s e l e c t i v e p r e s s u r e s f a v o r a s i n g l e c o n f o r m a t i o n i n d e e r , t h e n t h e s t r i v i n g toward a normal form w i l l be o f advantage t o t h e deer i n t h a t u n u s u a l i r r e g u l a r f l u c t u a t i o n s i n d i e t a r y energy w i l l n o t l e a v e t h e a n i m a l s t u n t e d o r deformed t o a p o i n t t h a t would l e a v e i t a t a d i s a d vantage .  V I I I . The Dimensions o f Fore Cannons C o l l e c t e d from HunterK i l l e d Fawns i n 1966 and 1967A l i s t o f t h e f o r e cannons t h a t were c o l l e c t e d was g i v e n i n T a b l e 6. the  The purpose o f t h i s c o l l e c t i o n was t o demonstrate  amount o f v a r i a b i l i t y t o be found i n t h e 4 d i m e n s i o n s  measured on t h i s bone; i . e . , l e n g t h , maximum b r e a d t h toward the  d i s t a l end, maximum b r e a d t h toward t h e p r o x i m a l end, and  minimum b r e a d t h o f t h e d i a p h y s i s .  I n a d d i t i o n , a comparison  205 was  made t o determine  i f a d i f f e r e n c e could be d e t e c t e d  between the y e a r s i n any o f the measurements. ments from these bones are shown i n Table  The  measure-  68.  Because the samples were c o l l e c t e d over a p e r i o d o f 21 days, the dependence o f s i z e on time was dimension  i n each o f the t h r e e c a t e g o r i e s (1966 males,  males and 1967  females).  regression line  1967  In each case the s l o p e o f the  ( S t e e l and T o r r i e , I960, Chp.  significantly different all  t e s t e d f o r each  from zero (p = 0.90)  data i n each c a t e g o r y were grouped  9) was  not  and t h e r e f o r e  for analysis.  The  l a c k o f growth over these t h r e e weeks i s reasonable when one c o n s i d e r s t h a t a very low plane o f n u t r i t i o n was f o r t h i s p e r i o d by the measurements taken from c a r c a s s e s a t 175 days. t h a t growth had  The  indicated  entire  above c a l c u l a t i o n s d i d not  ceased completely i n these dimensions,  prove but  simply t h a t changes caused by growth were s m a l l e r than c o u l d be d e t e c t e d w i t h i n the v a r i a b i l i t y i n the p o p u l a t i o n . U s i n g the grouped using student s t test f  data, no d i f f e r e n c e s were d e t e c t e d ( S t e e l and T o r r i e , I960, Chp.  any o f the 4 dimensions between males i n 1966 between males i n 1967  and females i n 1967  5) f o r  and 1967  (p = 0.90).  or The  l a c k o f d i f f e r e n c e between the males and females l e n d s support t o the view taken e a r l i e r t h a t t h e r e are not  any  important s k e l e t a l d i f f e r e n c e s between the sexes, at moderate v  n u t r i t i o n a l l e v e l s , by the age o f 6 months and t h a t male and female  l a b o r a t o r y standards may  be  grouped.  Ex N 1966  n = 12  n = 18  1967  n =  8  X  N  S  2  S  Sx  tSx (p = 0.95)  2800.0031  233.3333  0.517  0.719  0.208  0.457  2  7.8165  2.796  94.0739  7.8395  0.023  0.152  0.044  0.097  3  7. H 6 7  2.673  86.0637  7.1720  0.025  0.159  0.046  0.101  1.6555  1.287  19.9630  1.6636  0.008  0.090  0.026  0.057  fcl  219.8631  14.828  3964.5019  220.2501  0.387  0.622  0.147  0.309  fed  7.3682  2.714  133.0037  7.3891  0.021  0.145  0.034  0.072  fcp  6.6078  2.571  119.3630  6.6313  0.024  0.153  0.036  0.076  fcra  1.5653  1.251  28.2939  1.5719  0.007  0.081  0.019  0.040  fcl  216.8256  14.725  1738.6180  217.3272  0.502  0.708  0.250  0.592  fed  6.9696  2.640  55.9873  6.9984  0.029  0.170  0.060  0.142  fcp  6.1939  2.439  49.6884  6.2111  0.017  0.131  0.046  0.110  fem  I.4884  1.220  12.0058  1.5007  0.012  0.111  0.039  0.093  fed fcp  4  =  2  -  2 0 1  t p = 0.975, v = 17 = 2.110 P  2  15.258  1  = 0.975, v = 11 t  x  2  232.8157  fcl  fem  1967  Ex  2  = 0.975, v =  7  =  2  '  3 6 5  1 2 3 4  Fore.cannon Fore cannon Fore cannon Fore cannon  length b r e a d t h near d i s t a l end b r e a d t h near p r o x i m a l end minimum b r e a d t h a t d i a p h y s i s  A c l o s e r examination from males i n 1966  and  o f the measurements o f t h e  cannons  i n d i c a t e s t h a t t h e r e may  be some  1967  r e a l d i f f e r e n c e s between the two an e x a m i n a t i o n frequency indication  y e a r s t h a t were masked by  o f the means o f t h e measurements.  distributions  The  o f measurements i n F i g u r e 59  give  t h a t t h e r e i s not a normal d i s t r i b u t i o n and  t h e r e i s a change i n the shape o f the d i s t r i b u t i o n measurements between t h e two  years.  an  that  of  There i s o n l y a s l i g h t  tendency f o r a s h i f t o f t h e modes, b e i n g one  i n t e r v a l or  l e s s i n each c a s e , but the d i s t r i b u t i o n o f p o i n t s about t h e mode shows a downward s h i f t i n 1967.  This s h i f t increases  i n magnitude a l o n g the g r a d i e n t from l e n g t h t o d i s t a l w i d t h , t o p r o x i m a l w i d t h , t o minimum w i d t h .  In s p i t e o f the  l i m i t e d amount o f d a t a used t o e s t a b l i s h  very  the d i s t r i b u t i o n  measurements, and t h e r e f o r e the l a c k o f c o n f i d e n c e  of  i n the  a c t u a l shape, the t e n d e n c y f o r t h e s h i f t t o f o l l o w a p a t t e r n t h a t would be p r e d i c t e d by a c o n s i d e r a t i o n o f the growth g r a d i e n t l e n d s support  t o t h e e x i s t e n c e o f the s h i f t .  Furthermore, t h a t the g r e a t e s t e f f e c t occurs i n the  latest  m a t u r i n g d i m e n s i o n l a b e l s the c a u s a t i v e agent as a l a t e o c c u r r i n g energy r e s t r i c t i o n . d u r i n g the autumn o f 1967,  The  s u g g e s t i o n would be  that  the n u t r i t i v e q u a l i t y o f t h e  v e g e t a t i o n began t o d e c r e a s e e a r l i e r o r w i t h g r e a t e r s e v e r i t y than i n  1966.  I t i s important  t o emphasize here t h a t the modes d i d not  s h i f t markedly between the two  years.  That i s , the  skeletal  208  F i g u r e 59  The Frequency D i s t r i b u t i o n by I n t e r v a l o f t h e Fore Cannon Measurements From Hunter-Killed  Male Fawns i n 1966 . and 1967.  210 d i m e n s i o n s i n d i c a t e t h a t many o f t h e fawns i n 1966 and i n 1967 r e c e i v e d e q u a l n u t r i t i o n .  E v i d e n t l y , the f a c t o r which  caused t h e d i f f e r e n c e between t h e y e a r s d i d n o t a f f e c t a l l p a r t s o f t h e range e q u a l l y , and t h e d e e r , because o f t h e i r wide d i s t r i b u t i o n o v e r t h e r a n g e , would n o t have been affected  equally.  The s k e l e t a l d i m e n s i o n s cannot be used t o g i v e a d i r e c t e s t i m a t e o f t o t a l energy i n t a k e because t h e l a b o r a t o r y s t a n d a r d s a r e d e v e l o p e d f o r 112 and 175 days o n l y . s i n c e no measureable  However,  growth was i n d i c a t e d between 142 and  175 d a y s , t h e p r o j e c t e d energy i n t a k e s can be c a l c u l a t e d f o r t h e s e a n i m a l s a t 175 days.  I f t h e a s s u m p t i o n i s made t h a t  t h e energy i n t a k e a t t h i s t i m e i s v e r y near  maintenance  l e v e l s , a t t h e average body weight o f 55 pounds, t h e t o t a l energy i n t a k e e s t i m a t e d f o r 175 days can be c o r r e c t e d by r e d u c i n g i t be about 19,760 C a l o r i e s f o r 162 days and by about 50,160 C a l o r i e s f o r 142 days ( a l l o w i n g 2.0 x B.H.P. for  maintenance).  Rather than t r y t o estimate these  c o r r e c t i o n s f o r t h e v a r i o u s body w e i g h t s , t h e s u c c e e d i n g f i g u r e s w i l l assume t h a t a l l o f t h e fawns were 175 days o l d when k i l l e d .  By d o i n g t h i s , an a p p r o x i m a t i o n can be made  c o n c e r n i n g t h e d i v e r s i t y o f energy l e v e l s e x p e r i e n c e d i n t h e d i e t s o f t h e w i l d fawns. The energy i n t a k e f i g u r e s i n T a b l e 69 a r e based on a v e r a g e s o f t h e i n t e r c e p t s o f t h e f o u r measurements p e r bone for  each a n i m a l .  These e s t i m a t e s a r e based s t r i c t l y on t h e  211 T a b l e 69.  T o t a l energy i n t a k e s o f fawns based on measurements o f t h e f o r e cannon.  T a b l e 69 a )  The energy i n t a k e s f o r w i l d fawns i n 1966 and 1967.  1966 M a l e s Minimum 228,000  x - ts. 282,000  323,000  x + t s_  ^maximum  364,000  443,000 C a l o r i e s  X  1967 M a l e s minimum  x - ts-  207,000  262,000  x  x + ts290,000  X  318,000  maximum 373,000 C a l o r i e s  Where x = mean E . I . based on 4 i n t e r c e p t s from t h e f o r e cannon,  T a b l e 69 b)  No.  Regime  A c o m p a r i s o n between t h e energy i n t a k e e s t i m a t e d from t h e f o r e cannon w i t h t r u e energy i n t a k e f o r l a b o r a t o r y fawns. True Energy I n t a k e  E s t i m a t e d Energy I n t a k e  8  MHL  283,385 C a l o r i e s  276,000 C a l o r i e s  V26  HML  208,172  283,000  U 7  MHM  445,412  431,000  V24 "  LHM  337,796  329,000  Y13  MHM  376,855  386,000  V  212 T a b l e 69 c )  A comparison between t h e energy i n t a k e e s t i m a t e d from t h e f o r e cannon w i t h t h e b e s t energy i n t a k e e s t i m a t e d from a l l a v a i l a b l e measurements f o r w i l d fawns.  No.  Energy I n t a k e A l l Measurements  Energy I n t a k e , Fore Cannons  WF 1  310,000 Calories  326,000 Calories  WF 2  320,000  360,000  YF38  320,000  350,000  YF37  265,000  251,000  a v e r a g e s o f t h e energy i n t e r c e p t s t a k e n from F i g u r e s 35 and 36 and a r e n o t n e c e s s a r i l y t h e t r u e energy i n t a k e s .  I t must  be remembered t h a t f o r each magnitude o f a p a r t i c u l a r dimension there  i s a range o f p e r m i s s i b l e energy i n t a k e s and  t h a t t h e v a r i a t i o n s a r e n o t randomly d i s t r i b u t e d about t h e t r u e energy i n t a k e .  R a t h e r , t h e y a r e an e x p r e s s i o n o f t h e  p a t t e r n o f energy i n t a k e , and on c e r t a i n regimes may l e a d t o an e s t i m a t e  o f t o t a l energy i n t a k e t h a t i s as much as 50,000  C a l o r i e s above o r below t h e t r u e l e v e l .  From o n l y t h e f o u r  measurements o f t h e cannon i t i s n o t p o s s i b l e t o make a good estimate  o f t h e p a t t e r n o f energy i n t a k e , —  entire skeleton i t i s d i f f i c u l t , —  even w i t h an  b u t as mentioned  earlier,  t h e r e appears t o have been a l a t e o c c u r r i n g r e s t r i c t i o n o f d i e t a r y energy. decreasing estimated  I n t h e l a b o r a t o r y s t a n d a r d s showing a  n u t r i t i o n a l l e v e l w i t h t i m e , t h e energy i n t a k e s from t h e cannon measurements were c l o s e t o t h e  t r u e energy i n t a k e s (Table 69 b ) .  W i t h w i l d fawns f o r w h i c h  e n t i r e s k e l e t o n s were a v a i l a b l e , t h e energy i n t a k e s  estimated  by t h e cannons a l o n e d i d n o t d i f f e r much from t h o s e a c c e p t e d as t r u e energy i n t a k e based on a l l measurements (Table 69 c ) . On energy r e g i m e s t h a t a r e most p r o b a b l y  i n the w i l d , the  cannon d i m e n s i o n s s h o u l d g i v e e q u a l l y good e s t i m a t e s o f energy i n t a k e , and t h e e s t i m a t e s  s h o u l d be w i t h i n t 30,000  Calories. A l t h o u g h t h e e x a m i n a t i o n o f t h e s k e l e t a l measurements has a l r e a d y demonstrated t h a t t h e r e i s a d i v e r s i t y i n t h e  214 g r o w t h o f w i l d fawns, i t i s not u n t i l t h e s e measurements a r e e x p r e s s e d as energy i n t a k e s t h a t t h e magnitude o f t h e d i f f e r e n c e s becomes a p p a r e n t . a t l e a s t 150%  Some fawns must be r e c e i v i n g  o f the energy l e v e l e x p e r i e n c e d by t h e most  r e s t r i c t e d fawns.  T h i s a g a i n emphasizes t h e i m p o r t a n c e o f  e x a m i n i n g the a n i m a l and i t s r e l a t i o n s h i p t o i t s range on an i n d i v i d u a l l e v e l because i t i s t h e d i f f e r e n c e s among t h e members o f the p o p u l a t i o n t h a t d e f i n e t h e b i o l o g i c a l c h a r a c t e r i s t i c s o f the  population.  Summary and C o n c l u s i o n s . The f i n d i n g s o f t h i s work, summarized  i n the f o l l o w i n g  pages, have demonstrated t h a t t h e l e v e l o f energy i n t a k e  and  t h e p a t t e r n i n w h i c h i t i s a v a i l a b l e can a l t e r s k e l e t a l development  i n young fawns, and t h a t s k e l e t a l development i n  t u r n can r e f l e c t t h e energy r e g i m e .  Body w e i g h t a l s o  p r o v i d e s an e s t i m a t e o f energy i n t a k e , and when used i n conjunction  w i t h s k e l e t a l d i m e n s i o n s can l e a d t o an improved  i s o l a t i o n o f t h e p a t t e r n o f energy i n t a k e .  1.  L i v e body w e i g h t c o r r e l a t e d w e l l w i t h t o t a l energy i n t a k e f o r l a b o r a t o r y fawns a t 49, 112, 175 and 322 days, but the p r e d i c t i v e v a l u e d e c r e a s e d w i t h age.  The  relation-  s h i p was shown t o be d i s r u p t e d i n a n i m a l s w h i c h had  215  undergone w e i g h t l o s s . 2.  L i n e a r measurements o f t h e s k e l e t o n each p r o v i d e d estimate  an  o f energy i n t a k e , but were demonstrated t o be  b i a s e d away from t h e t r u e energy i n t a k e a c c o r d i n g t o the i n f l u e n c e o f t h e energy r e g i m e . 3.  S k e l e t a l e l e m e n t s i n v a r i o u s p a r t s o f t h e body responded d i f f e r e n t l y t o any p a r t i c u l a r energy r e g i m e , and t h i s d i f f e r e n c e was shown t o be r e l a t e d t o a x i a l  growth  gradients. 4.  I f the estimate  o f t o t a l energy i n t a k e from each  s k e l e t a l d i m e n s i o n i s g i v e n as a range w h i c h encompasses a l l p o s s i b l e energy r e g i m e s , t h e n an e s t i m a t e  can be  made based on an a r r a y o f measurements which i s l e s s b i a s e d from t h e t r u e energy i n t a k e t h a n would be the average o f t h e i n d i v i d u a l e s t i m a t e s measurements.  from t h e s e same  By t h e e x c l u s i o n o f t h o s e p o r t i o n s o f the  energy range not common t o a l l e s t i m a t e s , approximation  a better  t o t h e t r u e energy i n t a k e was shown f o r  t h e l a b o r a t o r y fawns, and t h e l i m i t s about t h i s mean were r e d u c e d . 5.  The d i f f e r e n t i a l growth o f t h e s k e l e t o n p r o v i d e d of determining  a means  t h e energy regime t o 112 days by two  intervals; 0 - 4 9  days and 50 - 112 days, w i t h an  e x a m i n a t i o n o f the s k e l e t o n o f the fawn at 112  days.  By  the n a t u r e o f the method employed, t o t a l energy i n t a k e i s e x p r e s s e d as a p r e c i s e v a l u e , but energy regimes are e x p r e s s e d i n terms o f w i d e r c a t e g o r i e s o f adequacy d e f i n e d as h i g h , medium o r low p l a n e s o f These c a t e g o r i e s 6.  i n t u r n have p r e c i s e  nutrition.  limits.  A l t h o u g h the t o t a l energy i n t a k e s o f fawns a t 175 c o u l d be e s t i m a t e d , not be  solved.  the p a t t e r n o f a v a i l a b i l i t y  could  However, t h e number o f p o s s i b l e regimes  c o u l d be markedly r e d u c e d . provide  days  The  cause o f t h e f a i l u r e  a s o l u t i o n appeared t o l i e i n the use  c a t e g o r i e s o f energy l e v e l i n each i n t e r v a l ,  to  of broad and not  in  the f a i l u r e o f t h e growth o f t h e s k e l e t o n t o respond t o t h e energy regime. 7.  By 322  days, d i f f e r e n c e s induced i n s k e l e t a l  conformation  a t e a r l i e r ages were reduced w i t h a l l s k e l e t o n s toward a s i m i l a r conformation. had  Compensatory p r o c e s s e s  so r e d u c e d the d i f f e r e n c e s t h a t an e v a l u a t i o n  energy regime was 8.  tending  considered  of  unfeasible.  An e x a m i n a t i o n o f f o r e cannons from w i l d fawns at a p p r o x i m a t e l y 150  days i n d i c a t e d a wide range i n t h e  l e v e l s o f energy i n t a k e s .  There was  a suggestion  d i f f e r e n c e i n the energy i n t a k e s between the y e a r s and  1967,  although the  sample s i z e was  not  of a 1966  sufficiently  217 large to support t h i s  observation.  The s t u d y o f the s k e l e t a l growth o f deer fawns has a c c o m p l i s h e d i t s o b j e c t i v e o f e s t a b l i s h i n g the e x i s t e n c e an a x i a l g r a d i e n t  - energy i n t a k e i n t e r a c t i o n .  The  of  infor-  m a t i o n o b t a i n e d from t h e l a b o r a t o r y r e a r e d fawns p r o v i d e s the b a s i s f o r i n t e r p r e t i n g the type o f n u t r i t i o n a l  regime  w h i c h produced an o b s e r v e d s k e l e t a l c o n f o r m a t i o n .  Before  a p p l y i n g t h e parameters f o r t h e e s t i m a t i o n o f energy  intake  t h a t were e s t a b l i s h e d w i t h t h e l a b o r a t o r y fawns t o the e s t i m a t i o n o f energy i n t a k e s o f w i l d fawns, c o r r e c t i o n s must be made f o r d i f f e r e n c e s i n e a r l y p o s t - n a t a l  nutrition.  There i s a l s o t h e problem o f t h e use o f i n t e r v a l s o f n u t r i t i o n a l regimes i n the l a b o r a t o r y .  A l t h o u g h t h i s method  was n e c e s s a r y t o a l l o w adequate c o n t r o l o v e r e x p e r i m e n t a l v a r i a b l e s , i t must be u n d e r s t o o d t h a t adherence t o t h e s e i n t e r v a l s cannot be e x p e c t e d from w i l d fawns. necessary to c o l l e c t a s e r i e s of skeletons  It will  be  from w i l d fawns  l i v i n g i n a r e a s o f d i f f e r i n g n u t r i t i o n a l adequacy,  o r from  one a r e a d u r i n g y e a r s o f d i f f e r i n g c o n d i t i o n s , i n o r d e r t o e s t a b l i s h the magnitude  o f s k e l e t a l d i m e n s i o n s t o be e x p e c t e d .  Once t h i s i s done, i n t e r p r e t a t i o n can be by comparison as i n the case o f t h e f o r e cannons from t h e w i l d fawns examined i n t h i s study. F o r a system f o r t h e e v a l u a t i o n o f t h e energy regimes o f  218  w i l d fawns t o be o f g r e a t e s t v a l u e , i t s h o u l d be conducted a t a t i m e when d a t a i s most r e a d i l y a t t a i n e d .  The h u n t i n g  season a l l o w s t h e c o l l e c t i o n o f s k e l e t a l p a r t s such as f o r e and h i n d cannons from fawns d u r i n g mid-November, o r a t about 150 days o f age.  T h i s date a l s o appears t o be a good t i m e  f o r i n t e r p r e t a t i o n o f s k e l e t a l growth because t h e d i f f e r e n c e s caused by t h e n u t r i t i o n a l r e g i m e s a r e s t i l l r e a d i l y a p p a r e n t . On t h e b a s i s o f t h e s k e l e t a l growth o b s e r v e d a t t h e o l d e r ages, i t does n o t appear t h a t n u t r i t i o n a l regimes can be r e c o g n i z e d much beyond 175 days.  Even when l i m i t e d t o  younger ages, however, t h e e x a m i n a t i o n o f f e e d i n g regimes s h o u l d be v e r y v a l u a b l e , f o r i t i n d i c a t e s d i f f e r e n c e s o c c u r r i n g d u r i n g t h e most c r i t i c a l g r o w i n g p e r i o d o f t h e y e a r .  219 Literature  Cited:  A d d i s o n , R.B. 1965- S k e l e t a l development i n t h e B l a c k t a i l deer. M.Sc. T h e s i s , U n i v e r s i t y o f B r i t i s h Columbia. Anderson, T.A., H.D. Fausch and J . G e s l e r . 1965. The e f f e c t o f r e s t r i c t e d a c c e s s t o f e e d on growth r a t e and body c o m p o s i t i o n o f swine. Growth 29: 213-218 A r c e l a y , C.L. 1963. The e f f e c t o f v a r i a t i o n s i n t h e energy and p r o t e i n l e v e l s o f t h e r a t i o n upon performance i n the p i g . Ph.D. T h e s i s , P e n n s y l v a n i a S t a t e U n i v e r s i t y . Bandy, P . J . 1955. 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Manage. 25: 295-302.  APPENDIX  Diagrams o f t h e S k e l e t o n o f the Deer, w i t h the Measurements  Used i n t h i s Experiment  FORE-CANNON:  R i g h t Leg  post.  ULNA-RADIUS:  R i g h t Leg  HUMERUS:  R i g h t Leg  HIND-CANNON:  R i g h t Leg  TIBIA:  R i g h t Leg  SCAPULA:  R i g h t Leg  A n t e r i o r View  FEMUR:  R i g h t Leg  PELVIS  D o r s a l View  ATLAS  Right Side  V e n t r a l View  D o r s a l View  Right  Side  Dorsal <  .  View .  THORACIC VERTEBRAE T 10  D o r s a l View  LUMBAR VERTEBRAE L 5  L 3  Right  L I  Side anterior  V e n t r a l View  D o r s a l View  

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