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The effect of vitamin A deficiency on some postmortem parameters of avian muscle Sundeen, Garfield Byron 1978

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THE EFFECT OF VITAMIN A D E F I C I E N C Y ON SOME POSTMORTEM PARAMETERS OF AVIAN MUSCLE  BY  GARFIELD BYRON SUNDEEN  B.Sc.  (Agr.),  University  of British  Columbia,  1973  A T H E S I S SUBMITTED IN P A R T I A L FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in THE FACULTY OF GRADUATE D e p a r t m e n t o f Food  STUDIES  Science  We a c c e p t t h i s t h e s i s a s c o n f o r m i n g required  t othe  standard  THE UNIVERSITY OF B R I T I S H COLUMBIA November, 1977 fc)  G a r f i e l d Byron  S u n d e e n , 1977  In p r e s e n t i n g an  this thesis  in p a r t i a l  advanced degree at the U n i v e r s i t y  the  Library  shall  f u l f i l m e n t o f the requirements f o r of B r i t i s h  make i t f r e e l y a v a i l a b l e  I f u r t h e r agree t h a t p e r m i s s i o n  Columbia,  f o r reference  I agree  that  and study.  f o r e x t e n s i v e copying o f t h i s  thesis  f o r s c h o l a r l y purposes may be granted by the Head o f my Department or by  h i s representatives.  It i s understood  of  t h i s t h e s i s f o r f i n a n c i a l gain  w r i t ten pe rm i ss i on .  Department o f FOOD SCIENCE The  University  of B r i t i s h  2075 Wesbrook Place Vancouver, Canada V6T 1W5  Date  December 29,1977  Columbia  shall  that  copying o r p u b l i c a t i o n  not be allowed without my  i ABSTRACT The e f f e c t o f t h e d i e t a r y s t a t u s o f v i t a m i n A on c a r b o h y d r a t e metabo l i s m , postmortem i s o m e t r i c t e n s i o n development and shear r e s i s t a n c e o f P e c t o r a l i s ma.jor muscle was s t u d i e d . Depletion  s t u d i e s , conducted o v e r a f i v e week p e r i o d ,  indicated a  d e f i n i t e i n f l u e n c e of v i t a m i n A d e f i c i e n c y on muscle c a r b o h y d r a t e metabolism. Mild hypovitaminosis  A induced an i n c r e a s e  i n glycogen d e p o s i t i o n whereas a  s e v e r e d e f i c i e n c y l e d t o a r e d u c t i o n o f these e l e v a t e d Vitamin  A def i c i e n c y d i d not a f f e c t t h e abi I i t y o f F\_ ma.jor s t r i p s  t o develop i s o m e t r i c t e n s i o n postmortem. cockerels generally required stages,  stores.  P. ma.jor s t r i p s sampled from d e f i c i e n t  longer t o reach maximum t e n s i o n and, i n t h e l a t e r  developed s i g n i f i c a n t l y g r e a t e r maximum t e n s i o n than those of c o n t r o l s .  The extended t i m e s t o maximum t e n s i o n r e f l e c t e d an i n c r e a s e d content.  A s i g n i f i c a n t increase  increased  m y o f i b r i l l a r c o n t r a c t i o n noted i n t h e l a t e r d e f i c i e n c y Cockerels  i n shear v a l u e  muscle glycogen  which had p r e v i o u s l y r e c e i v e d a c o m p l e t e l y  d e f i c i e n t r a t i o n f o r f i v e weeks were u t i l i z e d Though t h e r e was a d i s t i n c t delay  stages.  vitamin A  f o r the two week r e p l e t i o n s t u d y .  i n response t o t h e f e e d i n g of a v i t a m i n A  adequate r a t i o n , t h e muscle glycogen c o n t e n t , and  s i m i l a r l y corresponded t o t h e  isometric tension  parameters  s h e a r v a l u e s o f r e p l e t e d b i r d s were s i m i l a r t o t h o s e of c o n t r o l s w i t h i n  the two week p e r i o d .  TABLE OF CONTENTS Page ABSTRACT  i  LIST OF TABLES  v  ACKNOWLEDGEMENTS  '.-vll  INTRODUCTI ON  1  LITERATURE REVIEW  2  The e f f e c t of v i t a m i n A on c a r b o h y d r a t e metabolism  2  a)  V i t a m i n A and l i v e r glycogen  2  b)  V i t a m i n A and t i s s u e glycogen  4  R i g o r m o r t i s , metabolism and i s o m e t r i c t e n s i o n  4  Tenderness  5  and c o n t r a c t i o n s t a t e  MATERIALS AND METHODS  7  P o u l t r y source  7  D i e t f o r m u l a t i o n and e x p e r i m e n t a l design  7  Slaughter  procedure  10  I s o m e t r i c t e n s i o n measurement  10  Tenderness  11  evaluations  Sample p r e p a r a t i o n 1)  Ti ssue  2)  Liver  11 1 1  .  Chemical t e s t s 1) Blood g l u c o s e and P. major pH 2) M e t a b o l i t e a n a l y s i s 3) L i v e r vitamin A content S t a t i s t i c a l analysis  12 ,  12 12 13 13 13  EXPERIMENT 1 Body weight I s o m e t r i c t e n s i o n and thaw r i g o r Blood g l u c o s e , P. major pH .and ATP Liver EXPERIMENT 2 MortaIity  rate  Body weight Feed e f f i c i e n c y L i v e r weight and m o i s t u r e c o n t e n t I s o m e t r i c t e n s i o n parameters a)  Time t o maximum t e n s i o n  b)  Maximum i s o m e t r i c t e n s i o n  P. major metabol i t e I'evels a)  ATP  b)  Glycogen  Shear v a l u e s L i v e r glycogen and v i t a m i n A EXPERIMENT 3 Mortality rate Body weight Feed e f f i c i e n c y L i v e r weight and m o i s t u r e c o n t e n t I s o m e t r i c t e n s i o n parameters a) Time t o maximum t e n s i o n b) Maximum t e n s i o n  iv  Page P. major m e t a b o l i t e l e v e l s  38  a)  ATP  38  b)  Glycogen  38  Shear v a l u e L i v e r glycogen  38 and v i t a m i n A  '  41  DISCUSSION  44  SUMMARY AND CONCLUSIONS  53  LITERATURE CITED  54  V  LIST OF TABLES Table  I II III IV  V VI  Page  Wheat-soybean o i l meal basal d i e t  8  S u c r o s e - c a s e i n basal d i e t  9  The e f f e c t of d i e t a r y v i t a m i n A on body w e i g h t .  (Experiment 1)  15  Means and s t a n d a r d e r r o r s of time and t e n s i o n development p o s t mortem and d u r i n g thaw r i g o r f o r s t r i p s of C o r n i s h White Rock P. major muscle run i n phosphate b u f f e r . (Experiment 1)  16  Means and s t a n d a r d e r r o r s of blood g l u c o s e and P. major pH and ATP l e v e l . (Experiment 1)  17  initial  Means and s t a n d a r d e r r o r s o f l i v e r w e i g h t , m o i s t u r e and glycogen content.  (Experiment 1)  18  VII  Means and s t a n d a r d e r r o r s o f body w e i g h t .  VIII  The e f f e c t o f v i t a m i n A on feed e f f i c i e n c y  21  Means and s t a n d a r d e r r o r s of l i v e r weight and m o i s t u r e c o n t e n t Means and s t a n d a r d errors o f time and t e n s i o n development p o s t mortem f o r s t r i p s o f White Leghorn c o c k e r e l P. major muscle run in phosphate b u f f e r  22  Means and s t a n d a r d e r r o r o f P. major, ATP and glyqogen  26  IX X  XI XII XIII XIV XV  (Experiment 2)  • Means and s t a n d a r d e r r o r s o f shear v a l u e s f o r White c o c k e r e l P. major  levels  XVII  24  Leghorn  C o r r e l a t i o n m a t r i x f o r parameters s t u d i e d control birds  i n Experiment 2,  C o r r e l a t i o n m a t r i x f o r parameters s t u d i e d treatment b i r d s  i n Experiment 2,  28 29 29  The e f f e c t of v i t a m i n A d e f i c i e n c y on l i v e r glycogen and v i t a m i n A content  XVI  20  Means and s t a n d a r d e r r o r s of body weight. The e f f e c t of v i t a m i n A on feed e f f i c i e n c y .  31 (Experiment 3) (Experiment 3)  33 34  vi Table  XVIII XIX  XX XXL XXII XXIII  Page  Means and s t a n d a r d e r r o r s o f l i v e r weight and m o i s t u r e c o n t e n t . (Experiment 3)  35  Means and s t a n d a r d e r r o r s o f time and t e n s i o n development p o s t mortem f o r . s t r i p s of White Leghorn c o c k e r e l P. major muscle run i n phosphate b u f f e r  37  Means and standard e r r o r s of P. major, ATP and glycogen  39  levels  Means and s t a n d a r d e r r o r s of shear v a l u e s f o r White Leghorn c o c k e r e l P. major  40  C o r r e l a t i o n m a t r i x f o r parameters treatment b i r d s  41  s t u d i e d i n Experiment  The e f f e c t o f v i t a m i n A on l i v e r glycogen on v i t a m i n A  3, 43  ACKNOWLEDGEMENTS  The  a u t h o r would  like t o express h i s sincere appreciation  a d v i s o r , D r . J . F. R i c h a r d s , P r o f e s s o r ,  Department  o f Food  g u i d a n c e and e n c o u r a g e m e n t d u r i n g t h e c o u r s e o f t h i s He i s a l s o g r a t e f u l members o f h i s g r a d u a t e  Science f o r h i s  study.  f o r t h e a d v i c e and c o n t i n u e d i n t e r e s t o f t h e  committee:  D r . D. B. B r a g g , D e p a r t m e n t  of Poultry Science  Dr. J . V a n d e r s t o e p , Department and e x t e n d s a s p e c i a l  to his  o f Food  note o f thanks t o Dr. Bragg  Science f o rh i s assistance  in the  f o r m u l a t i o n a n d management o f t h e d i e t s . The  a u t h o r w i s h e s t o t h a n k Mr. Mel Hudson a n d t h e s t a f f  farm f o r t h e i r a s s i s t a n c e  in this  and M a r g a r e t Morrow f o r t e c h n i c a l computer  study.  He i s a l s o g r a t e f u l  of the poultry  t o Ms. E l l e n  a s s i s t a n c e and t o M i s s L y n n e R o b i n s o n f o r  assistance. I would  also  like  p a t i e n c e and u n d e r s t a n d i n g .  t o e x p r e s s my g r a t i t u d e t o my f r i e n d s f o r t h e i r  Holbek  1 INTRODUCTION The u l t i m a t e consumer a c c e p t a b i l i t y o f muscle systems as food i s i n f l u e n c e d by t h e t e x t u r a l c h a r a c t e r i s t i c o f t e n d e r n e s s . q u a l i t y has been r e s o l v e d i n t o two s t r u c t u r a l components:  In t u r n , t h i s  sensory  a 'background  tough-  ness' a t t r i b u t a b l e t o c o n n e c t i v e t i s s u e and o t h e r stromal p r o t e i n s and an 'actomyosin  toughness'  from t h e c o n t r a c t i l e apparatus.  latter's contribution will and  loss of e x t e n s i b i l i t y  The e x t e n t o f t h e  depend on t h e e v e n t s o f r i g o r m o r t i s - " t h e s t i f f e n i n g i n postmortem muscle"  (BendalI  T h i s s t i f f e n i n g and l o s s o f e x t e n s i b i l i t y  1973).  i n postmortem s k e l e t a l  muscle i s an o b v i o u s change a s s o c i a t e d w i t h a c o m p l e x i t y o f chemical  events.  One o f t h e s e i s t h e d i s a p p e a r a n c e o f glycogen v i a a n a e r o b i c g l y c o l y s i s , and both t h e r a t e and e x t e n t o f t h e c o n c o m i t a n t pH d e c l i n e have been r e l a t e d t o muscle t e n d e r n e s s . A s h o r t e n i n g o r c o n t r a c t i o n i s a n o t h e r p h y s i c a l event t h a t has been demonstrated  i n p r e r i g o r , postmortem muscle and s i m i l a r l y r e f l e c t s t h e chemical  e v e n t s which o c c u r .  T h i s s h o r t e n i n g can be e a s i l y monitored  by i s o m e t r i c t e n s i o n measurement, a t e c h n i q u e which important a n a l y t i c a l t o o l f o r measuring  quantitatively  i s becoming an i n c r e a s i n g l y  t h e ante and postmortem f a c t o r s o f  tenderness. The d i e t a r y s t a t u s o f v i t a m i n A i s one n u t r i t i o n a l e x e r t s an e f f e c t on c a r b o h y d r a t e metabolism  factor  which  and t h i s s t u d y was conducted t o  examine t h i s e f f e c t i n r e l a t i o n t o postmortem i s o m e t r i c t e n s i o n development and u l t i m a t e muscle t e n d e r n e s s .  2  LITERATURE REVIEW The  e f f e c t of v i t a m i n A on c a r b o h y d r a t e metabolism  S i n c e the s i m u l t a n e o u s d i s c o v e r y of v i t a m i n A by McCollum Davis (1913) and Osborne and Mendel (1913),  numerous i n v e s t i g a t o r s have demon-  s t r a t e d t h a t a l l v e r t e b r a t e s r e q u i r e t h i s f a t - s o l u b l e compound. n a t u r a l source i s f i s h  l i v e r o i l s b u t i the  and the v i t a m i n A a c t i v i t y of the  and  initial  l a t t e r i s due  The  most  important  source i s p l a n t m a t e r i a l s  t o t h e i r c o n t e n t of p r o v i t a m i n  A  carotenoids. R e t i n p l , r e t i n a l , r e t i n o i c a c i d and  some of t h e i r  stereoisomers  possess v i t a m i n A a c t i v i t y f o r t h e c h i c k e n and the v a r i o u s p r o v i t a m i n s converted  i n t o t h e s e compounds i n the The  vision  intestinal  A are  mucosa.  e s s e n t i a l r o l e of v i t a m i n A s t e r e o i s o m e r s  in the p r o c e s s of  i s w e l l known, but s e v e r a l o t h e r b o d i l y f u n c t i o n s are a l s o a f f e c t e d  by the d i e t a r y s t a t u s of t h i s v i t a m i n .  These f u n c t i o n s i n c l u d e growth, the  p r e v e n t i o n of s e v e r e a t a x i a , maintenance of the normal  i n t e g r i t y of mucous  membrane, r e p r o d u c t i o n , the p r o p e r growth of c a r t i l a g e m a t r i x and t h e maintenance of normal c e r e b r o s p i n a l  f l u i d pressure  ( S c o t t e t a I . , 1969).  Several  workers  have a l s o i n v e s t i g a t e d the r o l e of v i t a m i n A i n c a r b o h y d r a t e metabolism, a)  V i t a m i n A and  l i v e r glycogen  In v i t a m i n A d e f i c i e n t r a t s , Johnson and i n c o r p o r a t i o n of  labelled acetate,  l a c t a t e and  and e s s e n t i a l l y no glycogen d e p o s i t i o n i n the d i d not a f f e c t l a b e l l e d g l u c o s e c o n t r a d i c t e d these r e s u l t s .  Wolf (1960) found a  lowered  g l y c e r o l i n t o l i v e r glycogen liver.  Vitamin A d e p r i v a t i o n  i n c o r p o r a t i o n but subsequent i n v i t r o stud i e s  These a u t h o r s suggested t h a t the e f f e c t of  a v i t a m i n o s i s A on glycogen metabolism from a c e t a t e r e f l e c t e d a adrenalectomy w i t h r e s p e c t t o g l u c o c o r t i c o i d b i o s y n t h e s i s .  chemical  3 Stoewsand and  S c o t t (1964) s t u d i e d the e f f e c t of a d i e t a r y s t r e s s  on v i t a m i n A metabolism i n c h i c k s .  They r e p o r t e d t h a t h i g h p r o t e i n d i e t s  produced a s t r e s s s i t u a t i o n , e v i d e n c e d by adrenal  hypertrophy and h y p e r a c t i v i t y .  Though d a i l y i n j e c t i o n s of c o r t i c o s t e r o n e a l l e v i a t e d t h e s e symptoms, they g e n e r a l l y decreased  l i v e r v i t a m i n A s t o r e s and c o n c o m i t a n t l y  L i v e r glycogen f o r m a t i o n  a f t e r a 1 day  f a s t d i d not appear t o be  v i t a m i n A d e f i c i e n c y upon f o u r hours of r e f e e d i n g . p e r i o d was  extended t o 24 h o u r s , the amount of  feed consumed was The by Perek and (1971b). m a t t e r was  The  significantly  Kendler  However, when t h i s  by  refeeding  l i v e r glycogen formed/gm t o t a l  decreased.  (1969),  was  not s u b s t a n t i a t e d by N o c k e l s and  Phillips  l a t t e r a u t h o r s found t h a t the amountiof l i v e r glycogen/gm dry  the c o n t r o l d i e t .  i n 4 week o l d v i t a m i n A d e f i c i e n t b i r d s than i n t h o s e r e c e i v i n g For t h o s e b i r d s maintained  units vitamin A palmitate/kg  on the basal d i e t supplemented  r a t i o n t h i s d i f f e r e n c e was  whereas f o r t h o s e which r e c e i v e d a lower l e v e l was  impaired  A.  d e c r e a s e i n l i v e r glycogen d u r i n g v i t a m i n A d e f i c i e n c y observed  greater  w i t h 600 USP  i n c r e a s e d plasma v i t a m i n  (300 USP  significant,  u n i t s / k g ) , the d i f f e r e n c e  n o n s i g n i f i c a n t when compared t o c o n t r o l s . Apparently  c o n t r a d i c t o r y r e s u l t s were l a t e r r e p o r t e d  a u t h o r s (Nockels e t a j _ . , 1973).  In t h i s study a v i t a m i n o s i s A i n i t i a l l y  plasma c o r t i c o s t e r o n e s i g n i f i c a n t l y . c o r t i c o s t e r o n e and  percent  T h i s was  Two  improvement was  progressed.  noted  in c o r t i c o s t e r o n e  l i v e r glycogen of 7 week o l d b i r d s .  r e c e n t s t u d i e s w i t h r a t s q u e s t i o n the r e s e a r c h emphasis on  adrenocortical action in vitamin A deficiency.  increased  f o l l o w e d by a decrease in plasma  l i v e r glycogen as the d e f i c i e n c y  D e s p i t e a s c o r b i c a c i d s u p p l e m e n t a t i o n , no l e v e l o r i n the p e r c e n t  by t h e s e same  Janson and H a r i l l  (1974)  r e p o r t e d t h a t a v i t a m i n A d e f i c i e n c y led t o a decrease in l i v e r g l y c o g e n  4 concentration  i r r e s p e c t i v e of the d i e t a r y N source.  without a s i g n i f i c a n t a l t e r a t i o n  The decrease was noted  i n serum c o r t i c o s t e r o n e l e v e l s .  Dileepan  e t aj_. (1974) found t h a t /.avitaminosis A d i d n o t i n f l u e n c e h e p a t i c l e v e l s o r adenyl activity.  c y c l a s e a c t i v i t y but d i m i n i s h e d  hepatic  phosphodiesterase  They suggested t h a t t h e a l t e r a t i o n s of glycogen  due t o changes i n h e p a t i c c y c l i c adenosine monophosphate b)  V i t a m i n A and t i s s u e Although  glycogen  metabolism a r e p a r t l y levels,  glycogen  t h e f o r m a t i o n o f glycogen  occurs  i n p r a c t i c a l l y every t i s s u e  of t h e body and p r i m a r i l y i n l i v e r and muscle, o n l y t h e r e p o r t o f one study which s p e c i f i c a l l y  i n v e s t i g a t e d t h e e f f e c t o f v i t a m i n A on t i s s u e  glycogen  c o u l d be found. N o c k e l s and P h i l l i p s  (1971a) examined t h e p r o g r e s s o f a v i t a m i n o s i s A  in c h i c k e n s f o r 4, 8, 16, and 24 weeks.  They r e p o r t e d t h a t s t a t i s t i c a l l y  t h e r e were no d i f f e r e n c e s i n muscle glycogen  content  between v i t a m i n A d e f i c i e n t  b i r d s and c o n t r o l s ; however, t h e r e was a d e f i n i t e t r e n d muscle glycogen  with  indicating  increased  i n c r e a s i n g v i t a m i n A d e f i c i e n c y i n 8 and 16 week o l d  c h i c k e n s . A s i m i l a r p a t t e r n was observed f o r muscle ATP l e v e l s . R i g o r m o r t i s , metabolism and i s o m e t r i c t e n s i o n Bendall of e x t e n s i b i l i t y  (1973) has d e f i n e d r i g o r m o r t i s as t h e s t i f f e n i n g and l o s s i n postmortem muscle.  a s s o c i a t e d w i t h a c o m p l e x i t y of chemical  I t i s an o b v i o u s postmortem change events t h a t r e a l i z e s the disappearance  of N p h o s p h o r y l c r e a t i n e , ATP and glycogen ammonia and l a c t i c a c i d (deFremery, 1966). from a n a e r o b i c  and t h e appearance o f i n o s i n i c a c i d , The a c c u m u l a t i o n  of l a c t i c acid  g l y c o l y s i s causes t h e pH of t h e muscle t o f a l l  central  f a c t o r u n d e r l y i n g t h e p h y s i c a l changes o f r i g o r m o r t i s  1948).  Both t h e r a t e and e x t e n t of postmortem pH f a l l  u l t i m a t e muscle t e n d e r n e s s (Newbold and H a r r i s , 1972).  and t h i s i s . a . ;••:.! (Bate-Smith,  have been r e l a t e d t o  5 In a d d i t i o n t o l o s i n g e x t e n s i b i l i t y , u n r e s t r a i n e d muscle s h o r t e n s d u r i n g r i g o r development  and t h e e x t e n t o f t h i s c o n t r a c t i o n  t h e presence o f ATP and on t e m p e r a t u r e .  i s dependent on  The mechanism o f t h i s  s h o r t e n i n g i s b e l i e v e d t o be t h e same as muscular  contraction  postmortem in vivo, the  presently-known d e t a i l s o f which can be found i n r e c e n t r e v i e w s (Murray and Weber, 1974; Cohen, 1975; Mannherz and Goody, 1976).  T h i s occurance has l e d  t o t h e development o f a new t e c h n i q u e t o f o l l o w t h e t i m e - c o u r s e o f r i g o r : isometric tension  development.  Busch e t aj_. (1967) and Jungk e t §_[_• (1967) f i r s t r e p o r t e d t h e p a t t e r n of postmortem muscle.  i s o m e t r i c t e n s i o n development and d e c l i n e i n bovine and r a b b i t  T h i s p a t t e r n has been f u r t h e r demonstrated t o be a widespread phenomenon  o c c u r i n g i n bovine (Busch e t a]_., 1967, 1972; Jungk e t aj_., 1974), c h i c k e n (Khan, 1974; Wood and R i c h a r d s , 1974; Khan and Kim, 1975), p o r c i n e (Schmidt etaj_.,  1970a, b; Busch e t aj_., 1972), r a b b i t (Busch e t aj_., 1972; Jungk e t aj_.,  1974), and t u r k e y muscle (Jungk and M a r i o n , 1970; M a r i o n , 1971; Vanderstoep and R i c h a r d s , 1974). The most f r e q u e n t l y r e p o r t e d parameters o f i s o m e t r i c t e n s i o n a r e t h e t i m e t o maximum t e n s i o n and t h e maximum t e n s i o n d e v e l o p e d , and most o f t h e i n d i c a t e d s t u d i e s i n v e s t i g a t e d t h e response t o v a r i o u s t r e a t m e n t s as w e l l as t h e r e l a t i o n s h i p t o m e t a b o l i t e l e v e l s . Tenderness and c o n t r a c t i o n  state  In t h e i r reviews on t h e b i o c h e m i c a l and b i o p h y s i c a l i n muscle f o o d s , both Cassens (1977) and Marsh of meat i s t h e r e s u l t o f two components:  basis of tenderness  (1977) emphasized t h a t t h e toughness  a 'background toughness' a t t r i b u t a b l e  t o c o n n e c t i v e t i s s u e and o t h e r stromal p r o t e i n s and an 'actomyosin toughness' from t h e c o n t r a c t i l e a p p a r a t u s .  The e a r l y s t u d i e s which e s t a b l i s h e d t h e a s s o -  c i a t i o n between c o n t r a c t i o n and t e n d e r n e s s used sarcomere  length as t h e index  6  of c o n t r a c t i o n s t a t e ( H e r r i n g et_ aj_., 1965; Marsh and L e e t , 1966; Howard and Judge, 1968). Several  r e c e n t s t u d i e s have attempted t o r e l a t e i s o m e t r i c t e n s i o n  parameters and shear r e s i s t a n c e .  Busch e t a]_. (1967) r e p o r t e d t h a t  isometric  t e n s i o n parameters and shear were somewhat r e l a t e d a t 2°C i n beef semi t e n d i n o s u s but not a t 16° o r 37°C.  They suggested t h a t a t h i g h e r t e m p e r a t u r e s o t h e r  f a c t o r s b e s i d e s s h o r t e n i n g may be more important.  In an e x p e r i m e n t conducted  by Wood (1973) t o i n v e s t i g a t e t h e e f f e c t o f e p i n e p h r i n e tenderness,  i n j e c t i o n s on meat  s i g n i f i c a n t p o s i t i v e c o r r e l a t i o n s were found f o r shear  w i t h both t i m e and t e n s i o n .  values  Khan (1974) r e p o r t e d t h a t t h e u l t i m a t e shear f o r c e  of p o u l t r y b r e a s t meat cooked a t 20 hours postmortem.appeared t o be d i r e c t l y p r o p o r t i o n a l t o t h e maximum t e n s i o n developed. Kim  A l a t e r study  by Khan and  (1975) found t h a t t r e a t m e n t s which a c c e l e r a t e d g l y c o l y s i s and t h e o n s e t  of r i g o r m o r t i s , measured i s o m e t r i c a l l y , induced toughness i n c h i c k e n The  l a c k o f i n f o r m a t i o n on t h e e f f e c t o o f v i t a m i n A d e f i c i e n c y on  muscle g l y c o g e n , and  muscle.  t h e recognized  i n t e r r e l a t i o n s h i p between glycogen  availability  muscle c o n t r a c t i o n , and t h e s e n s i t i v i t y o f t h e i s o m e t r i c t e n s i o n measurement  technique  t o changes i n postmortem s h o r t e n i n g p r o v i d e t h e b a s i s f o r t h e r e s e a r c h  presented  in this thesis.  7 MATERIALS AND METHODS P o u l t r y source Day-old b i r d s were o b t a i n e d from a commercial h a t c h e r y and t r a n s p o r t e d t o t h e U.B.C. p o u l t r y farm.  In t h e p r e l i m i n a r y s t u d y , d e s i g n a t e d  Experiment 1, C o r n i s h White Rock b i r d s o f both sexes were used.  Subsequent  s t u d i e s u t i l i z e d White Leghorn c o c k e r e l s . D i e t f o r m u l a t i o n and e x p e r i m e n t a l d e s i g n The basal wheat-soybean  oil  meal d i e t o f Experiment 1 i s shown i n  T a b l e 1 and i s s i m i l a r t o t h a t o f N o c k e l s and K i e n h o l z (1967).  The f o u r  l e v e l s o f v i t a m i n A s u p p l e m e n t a t i o n u t i l i z e d were 375, 750, 1125, and 5000 lU/kg r a t i o n , w i t h t h e h i g h e s t l e v e l s e r v i n g as t h e c o n t r o l d i e t .  Fifteen  d a y - o l d b i r d s were randomly a l l o t t e d t o each d i e t and were reared i n e l e c t r i c a l l y heated b a t t e r y brooders u n t i I 5 weeks o l d , when they were p l a c e d i n growing batteries.  They r e c e i v e d feed and water ad I i b i t u r n and were  weighed a t weekly  individually  intervals.  The p u r i f i e d c a s e i n - s u c r o s e d i e t shown i n T a b l e 2 was t h e basal d i e t f o r both Experiments 2 and 3, and was f o r m u l a t e d on a weekly b a s i s . In Experiment 2, one hundred f i f t y d a y - o l d c h i c k s were p l a c e d i n e i g h t e l e c t r i c a l l y heated b a t t e r y b r o o d e r s ; i n Experiment 3 one hundred b i r d s were used. In both s t u d i e s , t h e c h i c k s were m a i n t a i n e d on t h e basal d i e t supplemented 5000IU v i t a m i n A/kg r a t i o n u n t i l  with  5 weeks o l d , a t which t i m e they were randomly  a l l o t t e d t o e i t h e r a v i t a m i n A - d e f i c i e n t o r v i t a m i n A-adequate r a t i o n and p l a c e d i n growing b a t t e r i e s .  (5000t1U/kg)  In Experiment 3 t h o s e b i r d s s u c c e s s -  f u l l y c o m p l e t i n g 5 weeks o f v i t a m i n A d e f i c i e n c y were r e a s s i g n e d t o t h e c o n t r o l diet.  In both s t u d i e s feed and water were a v a i l a b l e ad I i b i t u r n and t h e b i r d s  were i n d i v i d u a l l y weighed a t weekly i n t e r v a l s .  B i r d s i n Experiments 2 and 3  r e c e i v e d an i n t r a o c u l a r i n n o c u l a t i o n a g a i n s t Newcastle d i s e a s e a t 7 and 5 weeks o f age r e s p e c t i v e l y .  8  T a b l e I: Wheat-soybean o i l meal basal  diet  wheat < 1 356P) soybean o i l meal, s o l v e n t process^C48;5#P) d i c a l c i u m phosphate Iimestone s a l t , iodized DL methionine animal f a t V i t a m i n s added/kg r a t i o n vitamin ^tocopherol acetate choline riboflavin vitamin K niacin (Ca) p a n t o t h e n i c a c i d vitamin B ]2  M i n e r a l s added/kg r a t i o n MnSO "H„0 ZnCO^ CuS0,-'5H '0 4 2 o  Percent 65.8 24.2 2.7 0.8 0.5 0.22 5.0 990 IU 12 IU 1980 mg 2.54 mg 1.16 mg 27 mg 10 mg 11.10 ug 80 31 4  mg mg mg  9  Table I I :  Sucrose-casein  basal d i e t  sucrose case? n gelatin refined s a f f l o w e r o i l ("SaffIo") a Ipha ceI I 2 L arginine hydrochloride g I yc i ne DL m e t h i o n i n e V i t a m i n premix nicotinic acid thiamine hydrochloride riboflavin f o l i c acid pyridoxine b i ot i n choline chloride santoqu i n d Ca p a n t o t h e n a t e menadione Na b i s u l f i t e vitamin vitamin vitamin E  / 1 0 0 H b I ration.: 57.54 lb 20.00 5.00 4.00 3.00 1 .50 1 .00 0.40  2.2680 gm 0.6804 0.6804 0.2722 0.2722 0.0272 90.7185 4.5359 0.9072 0.0680 6.8038 1.0206 4.5318  Mi neraI Ingred i e n t s dical Iimestone KH P0 2  4  NaHCO, MgSO FeSO^ 7H 0 MnSO^ H 0 ZnCO, Cuso; 5H 0 KI0, Na JioO.- 2H 0 Na^SeO^ CaCI 2  2  2  1 2  /100 l b ration 1.80 l b 1.90 1 .40 399.16 gm 136.08 22.68 15.876 6.8039 1 .3608 0.4536 0.3765 0.0907 0.0771  adapted from S c o t t e± aj_. (1969) L .argirvirie h y d r o c h l o r i d e s u p p l e m e n t a t i o n ceasecl a f t e r 10 weeks  10  S l a u g h t e r procedure The s a m p l i n g of t h e b i r d s i n Experiment 1 began when t h e y were e i g h t weeks o l d and c o n t i n u e d u n t i l  t h e s u p p l y was e x h a u s t e d . Four b i r d s , one from  each t r e a t m e n t , were k i l l e d each day.  B i r d s were p l a c e d  i n a metal funnel and  were e x s a n g u i n a t e d by an o u t s i d e neck c u t and a l l o w e d t o b l e e d f r e e l y .  In  a d d i t i o n t o t h a t p r o v i d e d by t h e f u n n e l , r e s t r i c t i o n of wing and l e g movement d u r i n g s l a u g h t e r was manually a i d e d . In Experiment 2, e i g h t b i r d s from each t r e a t m e n t were s e l e c t e d a t 6, 7, 8, 9, and 10 weeks o f age, c o r r e s p o n d i n g t o one through f i v e weeks o f t r e a t m e n t , and were k i l l e d by c e r v i c a l d i s l o c a t i o n .  Four b i r d s , two c o c k e r e l s  from each t r e a t m e n t , were k i l l e d each day f o r f o u r days. were manually r e s t r a i n e d d u r i n g s l a u g h t e r . procedure was u t i l i z e d until  An i d e n t i c a l  The wings and legs s e l e c t i o n and s l a u g h t e r  i n Experiment 3, b e g i n n i n g a t 9 weeks and c o n t i n u i n g  t h e b i r d s were 12 weeks o f age.  T h i s p e r i o d corresponded t o t h e f o u r t h  and f i f t h week o f d e f i c i e n c y and t h e f i r s t and second week o f r e p l e t i o n f o r t h i s study. I s o m e t r i c t e n s i o n measurement The development and d e c l i n e o f i s o m e t r i c t e n s i o n was monitored w i t h an E + M 6-channel p h y s i o g r a p h * f i t t e d w i t h i s o m e t r i c t r a n s d u c e r s .  P. major  muscle s t r i p s f o r t e n s i o n measurement were prepared a c c o r d i n g t o t h e method of  Wood and R i c h a r d s (1974) w i t h one m o d i f i c a t i o n :  t o 4 cm i n l e n g t h .  t h e muscle s t r i p s were c u t  Three s t r i p s f o r each b i r d were e x c i s e d and a l l o w e d t o  develop i s o m e t r i c t e n s i o n i n phosphate b u f f e r pH 7.2, i o n i c s t r e n g t h (Gomori of  1955).  0.15  The maximum t i m e lapse from e x s a n g u i n a t i o n t o t h e attachment  s t r i p s was 20 m i n u t e s .  *Narco-Bio-Systems I n c . , Houston, Texas  11  Thaw r i g o r development was measured  o n l y f o r t h o s e b i r d s i n Experiment  1. Three P. major muscle s t r i p s from each b i r d , e x c i s e d and prepared i n t h e manner p r e v i o u s l y d e s c r i b e d , were i n d i v i d u a l l y wrapped i n Saran Wrap and p l a c e d in a -37°C b l a s t f r e e z e r .  Twenty-four hours postmortem t h e s e s t r i p s were removed,  a t t a c h e d t o t h e t e n s i o n measurement system and a l l o w e d t o thaw.  To p r e v e n t  s e v e r e s u r f a c e d e h y d r a t i o n about 4 cm of phosphate b u f f e r was p l a c e d  in the  bottom of t h e chamber and the top o f t h e chamber was c o v e r e d w i t h aluminum  foil  a l l o w i n g a small space f o r the attachment o f t h e s t r i p t o t h e t r a n s d u c e r . Tenderness e v a l u a t i o n s The muscle samples used f o r i s o m e t r i c t e n s i o n measurements and m e t a b o l i t e a n a l y s e s were e x c i s e d from t h e same P. major of each b i r d . r e m a i n i n g , i n t a c t P. major was covered w i t h Saran Wrap, packed ice  and aged f o r 24 hours a t 2°C.  i n drained crushed  Tenderness measurements were performed i n  a manner s i m i l a r t o t h a t r e p o r t e d by deFremery and Pool (1960). muscle was e x c i s e d , p l a c e d between two aluminum t h i c k n e s s , immersed  The  The  breast  p l a t e s held apart a t a constant  In b o i l i n g water f o r 10 m i n u t e s , t h e n c o o l e d i n c o l d t a p  water f o r 5 minutes.  S t r i p s of p a r a l l e l  fibers,  1.0 cm wide, were prepared  and each s t r i p ' s r e c o r d e d t h i c k n e s s r e p r e s e n t s t h e average of measurements along i t s length.  Shear was measured  on an I n s t r o v U n i v e r s a l T e s t i n g Instrument  (Model 1122) f i t t e d w i t h an A l l o - K r a m e r s i n g l e blade s h e a r a t t a c h m e n t . A l l shears were performed w i t h a c r o s s h e a d speed of 100 mm/min and a f u l l d e f l e c t i o n of 10 kg.  A minimum of 6 s h e a r s per b i r d was  scale  obtained.  Sample p r e p a r a t i o n 1) T i s s u e Muscle samples f o r chemical t e s t s were t a k e n a f t e r t h e P. major s t r i p s had been e x c i s e d f o r i s o m e t r i c t e n s i o n measurement.  The samples were  12  f r o z e n i n l i q u i d n i t r o g e n (LN^) and s t o r e d  i n aluminum  foil  envelopes a t  -37°C and s u b s e q u e n t l y powdered a c c o r d i n g t o t h e method o f B o r c h e r t and B r i s k e y (1965) as m o d i f i e d by Vanderstoep and R i c h a r d s (1974). samples were removed and p u l v e r i z e d 1.5 min. a t 11000 rpm. at a l l t i m e s .  The f r o z e n  i n a macro model V i r t i s homogenizer f o r  Care was t a k e n t o ensure t h e sample remained  The powdered sample was r e p l a c e d  and s t o r e d a t -37°C u n t i l  i n aluminum  foil  frozen  envelopes  extracted.  2) L i v e r L i v e r s were removed, b l o t t e d on paper t o w e l s and weighed. immediately f r o z e n -37°C u n t i l  i n Lls^, p l a c e d  freeze-dried.  w i t h mortar and p e s t l e .  i n aluminum  foil  They were  e n v e l o p e s and s t o r e d a t  Dry l i v e r s were weighed and ground t o a powder The l i v e r powder was s t o r e d  in f o i l  envelopes i n a  dess i c a t o r . Chemical  tests  1) B l o o d q l u c o s e _ a r i d P. major pH Blood g l u c o s e d e t e r m i n a t i o n s were done f o r Experiment 1 o n l y . B l o o d was c o l l e c t e d a t t h e t i m e o f e x s a n g u i n a t i o n i n a t e s t tube c o n t a i n i n g 20 mg p o t a s s i u m o x a l a t e and 25 mg sodium f l o r i d e .  G l u c o s e was measured i n  whole, d e p r o t e i n a t e d b l o o d u s i n g t h e g l u c o s t a t * method (Washko 1969). Absorbance was measured  w i t h a Beckman DB s p e c t r o p h o t o m e t e r a t  420 nm. The  initial  pH o f t h e muscle sample  In Experiment 1 was determined  by m o d i f y i n g t h e method o f Cassens and Newbold (1967). was homogenized  1 - 2 g o f muscle t i s s u e  w i t h 20 ml n e u t r a l i z e d 0.005M sodium i o d o a c e t a t e i n a Waring  L-1.;•;•[, r b l e n d e r f o r 5 min. pH was determined w i t h a F i s h e r Accumet, Model 230 pH Meter.  *Worthington B i o c h e m i c a l Corp., F r e e h o l d , N. J .  13 2) M e t a b o l i t e a n a l y s i s T i s s u e ATP d e t e r m i n a t i o n s were conducted  f o l l o w i n g t h e method o f  Lamprecht and T r a u t s c h o l d (1963) as m o d i f i e d by Wood  (1973).  L i v e r and t i s s u e glycogen was e x t r a c t e d from powdered samples by an enzymatic  method u t i l i z i n g amylo<*1, A-<, 6 g l u c o s i d a s e (Dalrymple and Hamm  1973) and determined All  as D-glucose by t h e method o f P f l e i d e r e r  (1963).  absorbance measurements were made w i t h an Unicam SP800 r e c o r d i n g  s pect rop hotomete r . 3) L i v e r v i t a m i n A c o n t e n t The a n a l y s i s f o r v i t a m i n A was performed on pooled each b i r d o f t h e same t r e a t m e n t c o n t r i b u t i n g 2 g o f powder.  liver  samples,  Triplicate  d e t e r m i n a t i o n s were o b t a i n e d a c c o r d i n g t o t h e method o f Dugan e t aj_. (1964), a c o l o r i m e t r i c determination with t r i f l u o r o a c e t i c  acid.  Absorbance was measured a t 616 nm w i t h a Beckman DB  spectrophotometer.  Statistical analysis D i f f e r e n c e s between t r e a t m e n t means were a n a l y z e d f o r s t a t i s t i c a l s i g n i f i c a n c e by t h e . t - t e s t a c c o r d i n g t o t h e method o f S t e e l and Torn ie (1960).  (  14 RESULTS Experiment 1 Body weight The e f f e c t o f d i e t a r y v i t a m i n A on body weight g a i n s i s p r e s e n t e d i n T a b l e I I I . Those c h i c k s r e c e i v i n g t h e lowest l e v e l o f v i t a m i n A supplementation  (375 lU/kg) had s i g n i f i c a n t l y h i g h e r (p<0.05) body w e i g h t s a t 1 week  of age than t h o s e on t h e c o n t r o l d i e t .  T h e i r mean v a l u e remained n u m e r i c a l l y  g r e a t e r f o r a n o t h e r two weeks, a f t e r which t h e t r e n d was r e v e r s e d . The body w e i g h t s o f t h e c h i c k s r e c e i v i n g t h e t h r e e lowest t r e a t m e n t l e v e l s were lower than c o n t r o l s a t 6, 7, and 8 weeks o f age; o n l y t h o s e c h i c k s on t h e lowest .level  (375 lU/kg) had s i g n i f i c a n t l y  l i g h t e r (p<0.05) body w e i g h t s  a t 8 weeks o f age. I s o m e t r i c t e n s i o n and thaw r i g o r The data f o r i s o m e t r i c t e n s i o n and thaw r i g o r development a r e shown in T a b l e IV. No s i g n i f i c a n t d i f f e r e n c e s o r d e f i n i t e t r e n d s i n e i t h e r t h e time t o maximum t e n s i o n o r t h e maximum t e n s i o n developed were e v i d e n t . Blood g l u c o s e , P. major pH and. ATP The data f o r t h e s e t h r e e parameters a r e p r e s e n t e d i n T a b l e V. Blood g l u c o s e l e v e l s were s l i g h t l y  lower and w h i t e muscle  initial  pH and ATP  v a l u e s were s l i g h t l y h i g h e r f o r t h o s e c h i c k s r e c e i v i n g d e f i c i e n t d i e t s no s t a t i s t i c a l  d i f f e r e n c e s were noted.=  though  Table I I I :  The e f f e c t of dietary vitamin A on body weight (Experiment 1)  Body Weight (g) Age in Weeks Dietary Vitamin A Day-old (lU/kg diet) 375 750 1125 5000(C)-  1 2 3  42±0.7a 40±0.9a 41±0.8a 39±1.1a  5 106±2.3b 100±2.8a 92±3.0a 96±2.8a  232±5.4a 213±4.6a 200±6.0b 221±6.1a  428± 8.1a 617± 8.2a 407±11.0a 642±19.8a 387±10.0a 597±13.2a 412± 7.8a 626±14.9a  7;  i  821±12.4a 907±28.0a 838±19.4a 855±20.6a  1127±21.2a 1147±33'.4a 1157±34.0a 1184±35.0a  mean and standard e r r o r means in same column followed by s i m i l a r l e t t e r s do not d i f f e r s i g n i f i c a n t l y (p<0.05) C = control  1383±28.2a 1375±47.5a 1435±44.9a 1466±46.5a  1627±38.4b 1738±55.0a 1789±64.3a 1801±68.8a  Table IV: Means and s t a n d a r d e r r o r s o f t i m e and t e n s i o n development postmortem and d u r i n g thaw r i g o r f o r s t r i p s o f C o r n i s h White Rock P. major muscle run i n phosphate b u f f e r . (Experiment 1)  I s o m e t r i c Tension Parameters Dietary Vitamin A (lU/kg d i e t ) 375 750 1 125 5000 (C) 1  Time t o Maximum T e n s i o n (hours) 4.93 5.02 5.20 4.79  Thaw R i g o r  Maximum Tension Developed  +  0.49a + 0.48a + 0.60a + 0.72a  means i n same column f o l l o w e d by s i m i l a r  63.1 54.8 59.5 62.6  +  Time t o Maximum  2  (gm/cm )  4.9a + 4.0a + 3.4a + 5.0a  ; i M a x ! m u m - T e n s i on 2 Tension (minutes) (gm/cm ) 8.8 7.7 9.6 8.4  +  0.6a + 0.8a + 0.9a + 0.5a  lower case l e t t e r s do n o t d i f f e r s i g n i f i c a n t l y  196.0 194.6 198.7 214.0  + + + +  (p<0.10)  12.9a 5.2a 7.4a 18.0a  17 T a b l e V:  Means and s t a n d a r d e r r o r s o f blood g l u c o s e and P. major pH and ATP l e v e l . (Experiment 1)  Dietary Vitamin A (Ill/kg d i e t )  375 750 1125 5000 (C)  1  Blood G l u c o s e Level (mg#)  170 174 170 181  ± ± ± ±  6.9a 6.6a 7.1a 7.6a  1  Initial pH  6.24 6.20 6.20 6.12  means i n same column f o l l o w e d by s i m i l a r s i g n i f i c a n t l y (p<0.10)  Li v e r  ± ± ± ±  0.05a 0.06a 0.06a 0.07a  lower case  initial  ATP (/imoles/gm)  6.17 6.28 6.74 5.87  ± ± ± ±  0.51a 0.48a 0.49a 0.62a  l e t t e r s do not d i f f e r  L i :r  On a weight b a s i s t h e c h i c k s r e c e i v i n g 375 and 750 IU v i t a m i n A/kg had s i g n i f i c a n t l y lower l i v e r w e i g h t s (p<0.10) than t h e c o n t r o l s  (TabIe V I ) .  Those r e c e i v i n g 750 lU/kg m a i n t a i n e d t h i s s i g n i f i c a n t d e f i c i t when l i v e r were e x p r e s s e d as a body weight  weights  percentage.  There d i d not appear t o be a s i g n i f i c a n t e f f e c t of v i t a m i n A d e p r i v a t i o n on  l i v e r m o i s t u r e c o n t e n t f o r t h o s e c h i c k s m a i n t a i n e d on t h e 375 and 750 lU/kg  rations.  However, a s i g n i f i c a n t l y h i g h e r (p<0.10) l i v e r m o i s t u r e c o n t e n t was  e x h i b i t e d by t h o s e r e c e i v i n g a moderate l e v e l of v i t a m i n A (1125 l U / k g ) . These b i r d s a l s o had a s l i g h t l y lower g l y c o g e n c o n t e n t but no s i g n i f i c a n t of v i t a m i n A d e f i c i e n c y on l i v e r g l y c o g e n  i s otherwise  demonstrated.  effect  Table VI:  Means and s t a n d a r d e r r o r s o f l i v e r weight, m o i s t u r e and glycogen c o n t e n t (Experiment 1)  L i v e r Weight Dietary Vitamin A (lU/kg d i e t ) 375 750 1125 5000 (C)  1  gm 44..2 44,.7 48..8 51,.4  +  1 .7b + 1 .8b + 2. 4a + 2. 4a  L i v e r moisture  L i v e r glycogen  {%)  (jumoles glucose/gm d r y wt.)  % body weight  1  2,.00 1,.87 2,.02 2,.05  +  0.04a 0.04b 0.11a + 0.05a + +  68.,7 69.,3 69..7 68.,4  +  0. 7a 0. 6a + 0. 6b + 0. 4a +  310.2 317.1 253.0 305.0  + + + +  28.3a 27.1a 35.2a 21.5a  means i n same column f o l l o w e d by s i m i l a r lower case l e t t e r s do not d i f f e r s i g n i f i c a n t l y (p<0.10)  19 Experiment Mortality  2 rate An u n u s u a l l y h i g h m o r t a l i t y r a t e of 27.4$ (42 o u t of 153 b i r d s )  recorded f o r t h e f i r s t week of t h e e x p e r i m e n t a l p e r i o d and t h i s r a t e s l i g h t l y t o 28.1$  i n t h e subsequent  was  increased  f o u r weeks p r i o r t o t r e a t m e n t a l l o c a t i o n .  By t h e end of t h e f i v e weeks o f t r e a t m e n t , c o r r e s p o n d i n g t o 6 through 10 weeks of age, c o n t r o l b i r d s (5000 lU/kg) e x h i b i t e d a 7.3$  m o r t a l i t y r a t e whereas t h o s e  r e c e i v i n g t h e a v i t a m i n o t i c A r a t i o n had 13$ m o r t a l i t y . As some c o n t r o l and a v i t a m i n o t i c A b i r d s showed p r e l i m i n a r y of Newcastle d i s e a s e , each group r e c e i v e d of age.  intraocular vaccinations  symptoms  a t 7 weeks  No deaths were r e c o r d e d i n e i t h e r group f o r t h e f o l l o w i n g two weeks.  Body weight T a b l e VII p r e s e n t s t h e average body w e i g h t s f o r t h e t o t a l  number  of b i r d s used from a t r e a t m e n t group, as w e l l as t h e weekly and t e r m i n a l w e i g h t s o f t h o s e b i r d s sampled  a t each s t a g e of t h e e x p e r i m e n t .  week t r e a t m e n t p e r i o d no s i g n i f i c a n t d i f f e r e n c e s  body  For t h e f i v e  i n body weight were o b s e r v e d .  Feed e f f i c i e n c y Feed e f f i c i e n c y v a l u e s f o r t h o s e b i r d s sampled in T a b l e V I I I .  In t h i s and subsequent  under t h e v i t a m i n A d e f i c i e n t heading. v a l u e s of t h e parameter  t a b l e s , two data columns a r e shown The  f o r a l l the sampled  f i r s t column r e p r e s e n t s t h e  Henceforth, these s h a l l  mildly deficient birds respectively.  average  b i r d s , w h i l e t h e v a l u e s which  in t h e second column are t h e means o n l y f o r t h o s e sampled p o s i t i v e weight g a i n s .  a t each s t a g e a r e prese  appear  b i r d s which showed  be r e f e r r e d t o as d e f i c i e n t and  T a b l e V l l : Means and s t a n d a r d e r r o r s o f body w e i g h t (Experiment 2)  Duration of Treatment (in weeks)  \  1 2  0 1 2 3 4 5  1  Weekly Weight of B i r d s used  V i t a m i n A D e f i c i e n t UContro 1  Control 337±;777a 413±9'.8a 546±13.3a 684±21.8a 793±28.6a 916±37.0a  Weekly Weight of Sampled B i r d s  2  351 ± :6.4a 435± :8.4a 565±13.4a 736118.8a 817±25.4a 880±50.4a  Week 0 c o r r e s p o n d s t o 5 weeks o f age means i n same row f o l l o w e d by s i m i l a r  394±24.4a 563±23.1a 668±51.3a 798±46.6a 916±37.0a  Terminal Weight of Sampled B i r d s  Vitamin A D e f i c i e n t  ControI  Vitamin A D e f i c i e n t  439±14.8a 546±32.1a 680±30.5a 812±29.9a 880±50.4a  427±27.9a 594±17.2a 680±50.8a 845±50.4a 966±48.1a  475±22.4a 580±29.2a 688±32.8a 833±32.7a 885±45.la  lower case l e t t e r s do not d i f f e r s i g n i f i c a n t l y  (p<0.05)  21 Vitamin A deficiency  d i d not s i g n i f i c a n t l y a f f e c t feed e f f i c i e n c y •  though t r e a t m e n t b i r d s had n u m e r i c a l l y stage.  lower v a l u e s than t h e c o n t r o l s  a t each  C o n t r o l and t r e a t m e n t b i r d s showed d e c r e a s i n g feed e f f i c i e n c y w i t h  increasing  age and when t h e data were s u b j e c t e d t o r e g r e s s i o n a n a l y s i s  signif-  i c a n t r e g r e s s i o n c o e f f i c i e n t s o f r = -.895 (p<0.02), r = -.981 (p<0.001), and r = -.988 (p<0.001) were o b t a i n e d f o r t h e c o n t r o l , d e f i c i e n t and m i l d l y birds  deficient  respectively.  Table V I M :  The e f f e c t o f v i t a m i n A on feed e f f i c i e n c y  Feed e f f i c i e n c y (gm/gm feed consumed) Vitamin A d e f i c i e n t Duration of Treatment ( i n weeks) 1 2 3 4 5 1 2 3  Control 0.394 0.410 0.369 0.357 0.345  ± ± ± ± ±  Deficient  0.02aA 0.02aA 0.03aA 0.02aA 0.02aA  0.418 0.404 0.356 0.341 0.320  ± ± ± ± ±  0.02aA^ 0.02aA 0.02aB O.OlaB O.OlaB  Mildly  deficient  0.366 ± 0.02aB 0.354 ± 0.01aB 0.326 ± 0.02aB  mean and s t a n d a r d e r r o r means i n same row f o l l o w e d by s i m i l a r lower case l e t t e r s do n o t d i f f e r s i g n i f i c a n t l y (p<0.10) means i n same column f o l l o w e d by s i m i l a r c a p i t a l l e t t e r s do n o t d i f f e r s i g n i f i c a n t l y from mean of Week 1 (p-'O.IO)  L i v e r weight and m o i s t u r e c o n t e n t No s i g n i f i c a n t d i f f e r e n c e s were r e a l i z e d u n t i l t h e f i f t h  i n l i v e r weight on a gm weight b a s i s  t r e a t m e n t week when t h e c h i c k s which were d e f i c i e n t  in v i t a m i n A showed s i g n i f i c a n t l y lower l i v e r w e i g h t s (p<0.10) than t h e controls.  (Table IX)  T a b l e IX: Means and s t a n d a r d e r r o r s o f l i v e r weight and m o i s t u r e c o n t e n t  Vitamin A d e f i c i e n t Duration of "Treatment ( i n weeks)  ' Control  Deficient  Mildly  deficient  16.6 17.6 19.7  + 1.7aA + 0.4aB + 1.4bB  L i v e r Weight (gm) 1 2 3 4 5  13.2 ± 0.9aA 16.1 ± 0.8aB 18.5 ± 1.4aB 19.9 i ±-1.8aB 26.5 ± 2.6aB  14.0 15.3 17.1 18.8 18.3  + + + + +  0.7aAi, 0.8aA .1.6aB 0.6aB 1.3bB  L i v e r Weiqht (as % body w e i q h t ) 1 2 3 4 5  3. 18 2.73 2.72 2.37 2.69  ± ± ± ± ±  0.3aA 0.2aA 0.1aA 0.2aB 0.2aA  2.96 2.65 2.50 2.31 2.11  + + + + +  0.1aA 0.1aA 0.2aA 0.2aB 0. IbB  2.33 2.04 2.21  + 0.2bB + 0.1aB + 0. IbB  L i v e r moisture content (decimal) 1 2 3 4 5  1 2  0.709 0.720 0.714 0.721 0.695  means i n same significantly means i n same significantly  ± ± ± ± ±  0.008aA 0.004aA 0.006aA 0.004aA 0.012aA  0.730 0.713 0.725 0.699 0.711  ± ± ± ± ±  0.004bA 0.005aB 0.004aA 0.011aB 0.007aB  row f o l l o w e d by s i m i l a r lower case (p<0.10) column f o l l o w e d by s i m i l a r c a p i t a l from mean o f Week 1 (p<0.10)  0.725 ± 0.004aA 0.698 ± 0.004bB 0.707 ± 0.018aB  l e t t e r s do not d i f f e r l e t t e r s do not d i f f e r  23  A d e f i n i t e trend of increasing  l i v e r weight w i t h  increasing  was noted f o r c o n t r o l and t r e a t m e n t b i r d s and s i g n i f i c a n t r e g r e s s i o n  age coefficients  of r = .965 (p<0.002), r = .962 (p<0.005) and r = .995 (p<0.001) were c a l c u l a t e d for  t h e c o n t r o l , d e f i c i e n t and m-ildly d e f i c i e n t groups  respectively.  When l i v e r weight i s e x p r e s s e d as p e r c e n t body w e i g h t , d e f i c i e n t and m i l d l y d e f i c i e n t b i r d s had s i g n i f i c a n t l y l i g h t e r l i v e r s than c o n t r o l s  (p<0.10)  a t t h e f i f t h t r e a t m e n t week; a d d i t i o n a l l y , m i l d l y d e f i c i e n t b i r d s had s i g n i f icantly  l i g h t e r l i v e r s than c o n t r o l s A d e f i n i t e trend  mildly deficient birds i n c r e a s i n g age.  (p<0.10) a t t h e t h i r d t r e a t m e n t week.  can be seen w i t h  r e s p e c t t o age; i n d e f i c i e n t and  l i v e r weight as p e r c e n t body weight decreased  S i g n i f i c a n t regression  c o e f f i c i e n t s o f r = -.993 (p<0.001)  and r = -.948 (p<0.005) were o b t a i n e d f o r t h e d e f i c i e n t and m i l d l y groups r e s p e c t i v e l y .  No r e c o g n i z a b l e t r e n d  Neither control with  with  i s indicated  deficient  f o r the control  nor t r e a t m e n t groups d i s p l a y a c o n s i s t e n t  group.  pattern  r e s p e c t t o l i v e r m o i s t u r e ; d e f i c i e n t b i r d s had a s i g n i f i c a n t l y h i g h e r  l i v e r (p<0.10) m o i s t u r e c o n t e n t a t t h e f i r s t week whereas t h e m i l d l y b i r d s had a s i g n i f i c a n t l y l o w e r ( p < 0 . l O ) m o i s t u r e c o n t e n t than c o n t r o l s  deficient after  f o u r weeks o f t r e a t m e n t . Isometric tension a)  parameters  Time t o maximum  tension  Samples from t r e a t e d isometric tension  birds generally  required  than samples from t h e c o n t r o l s  l o n g e r t o develop maximum  but t h i s d i f f e r e n c e  was s i g n i f -  i c a n t (p<0.10) o n l y f o r d e f i c i e n t b i r d s a f t e r 2 weeks on t h e a v i t a m i n o t i c  A  r a t i o n and f o r those b i r d s which were m i l d l y d e f i c i e n t a f t e r f o u r weeks ( T a b l e X ) . Samples from c o n t r o l significantly  birds  i n t h e f o u r t h t r e a t m e n t week  l e s s time t o reach maximum t e n s i o n  required  than t h o s e c o n t r o l s o f t h e  T a b l e X:  Means and s t a n d a r d e r r o r s o f t i m e and t e n s i o n development postmortem f o r s t r i p s of White Leghorn c o c k e r e l P. major muscle run i n phosphate b u f f e r .  Time t o Maximum T e n s i o n (minutes) Duration of Treatment ( i n weeks) 1 2 3 4 5  1 2  Control  272.2 244.4 266.5 175.2 212.4  + +  40.9aA 50.5aA + 40.4aA + 36.0aB + 38.0aA  Deficient  295.2 391 .1 261 .4 300.8 296.2  Mildly  deficient  + +  29.4aA* 35.IbB + 45.9aA + 78.1aA + 58.6aA  290.7 + 40.7aA 375.0 + 83.5bA 320.8 + 76.7aA*.  Maximum Control  96.66 82.10 83.15 80.95 73.21  + +  9.2aA 7.8aA + 5.0aA + 7.1aA + 3.9aB  I s o m e t r i c T e n s i o n (qm/cm ) Deficient  84.64 88.44 90.09 77.16 100.22  + + +  7.2aA 8.8aA 7.1aA + 6.0aA + 3.9bB  Mildly  deficient  94.44 + 6.4aA 80.93 + 7.4aA 99.50 + 6.4bA  means i n same row f o l l o w e d by s i m i l a r lower c a s e l e t t e r s do not d i f f e r s i g n i f i c a n t l y (p<0.10) means i n same column f o l l o w e d by s i m i l a r c a p i t a l l e t t e r s do not d i f f e r s i g n i f i c a n t l y from mean a t Week 1 ( (p<0.10)  25  f i r s t week (p<0.10).  The average time t o maximum t e n s i o n  b i r d s sampled i n t h e second week was s i g n i f i c a n t l y  longer  f o r the d e f i c i e n t (p<0.10) than f o r  t h o s e sampled i n t h e f i r s t week, b)  Maximum i s o m e t r i c Vitamin  tension  A d e p r i v a t i o n d i d not appear t o i n f l u e n c e t h e maximum  t e n s i o n developed i n t h e e a r l y s t a g e s o f d e f i c i e n c y . ( T a b l e  X). After  isometric  five  weeks of t r e a t m e n t however, d e f i c i e n t and m i l d l y d e f i c i e n t b i r d s developed s i g n i f i c a n t l y g r e a t e r maximum t e n s i o n than c o n t r o l s (p<0.10).  The f i f t h week  v a l u e f o r d e f i c i e n t b i r d s was a l s o s i g n i f i c a n t l y h i g h e r than t h a t developed by d e f i c i e n t b i r d s a t week 1 (p<0.10). When t h e data was s u b j e c t e d  to regression analysis a s i g n i f i c a n t  n e g a t i v e c o r r e l a t i o n c o e f f i c i e n t of r = -.896 (p<0.02) was o b t a i n e d c o n t r o l group. established  No r e l a t i o n s h i p between maximum t e n s i o n and age c o u l d be  f o r t h e t r e a t m e n t group.  P. major m e t a b o l i t e a)  f o r the  levels  ATP Elevated  P. major ATP l e v e l s appear i n t h e f i r s t f o u r weeks o f v i t a m i n  A d e f i c i e n c y and f o r t h e d e f i c i e n t b i r d s o f t h e second week t h i s  difference  i s s i g n i f i c a n t (p<0.10) (Table X I ) . Nbcother t r e a t m e n t o r t i m e e f f e c t i s i n d i c a t e d . b)  Glycoqen D e f i c i e n t b i r d s of t h e f i r s t week had s i g n i f i c a n t l y g r e a t e r  tissue  glycogen l e v e l s than c o n t r o l s (p<0.10) but no o t h e r t r e a t m e n t e f f e c t can be observed.  The c o n t r o l s o f t h e second week had s i g n i f i c a n t l y g r e a t e r  v a l u e s and t h e d e f i c i e n t b i r d s o f t h e f i n a l  week had s i g n i f i c a n t l y  v a l u e s than t h e i r r e s p e c t i v e f i r s t week v a l u e s (p<0.10).  glycogen  lower glycogen  T a b l e X I : Means and s t a n d a r d e r r o r o f P. major,.ATP and glycogen l e v e l s ATP(jumoles/gm)  Glycogen (jumoles glucose/gm);  Vitamin A d e f i c i e n t Duration of Treatment ( i n weeks) 1 2 3 4 5  1 2  Control 6.08 5.78 5.78 5.26 7.56  +  0.6aA 0.7aA + 0.9a A + 0.9aA + 0.9a A +  Deficient 7.57 7.92 6.46 5.32 5.83  + + + +  LlaA 0.4bA 0.8aA 0.9aA + 0.6aA  Mildly deficient  1  6.98 ± 0.7aA 6.14 ± :1.0aA 6.20 ±.0.8aA  Vitamin A d e f i c i e n t Control 7.92 11.42 9.22 8.78 9.01  + +  0.4aA 1.5aB + 0.8aA + l.laA + 0.9aA  Deficient 11.58 11.61 8.15 9.07 7.82  Mildly  deficient  + +  1.6bA l.laA + 1.3aA/ + 1.8aA + l.laB  8.83 + 1.2aA 9.60 + 2.4aA 6.88 + 1.4aB  means i n same row f o l l o w e d by s i m i l a r lower case l e t t e r s do not d i f f e r s i g n i f i c a n t l y (p<0.10) means i n same column f o l l o w e d by s i m i l a r c a p i t a l l e t t e r s do not d i f f e r s i g n i f i c a n t l y from mean of Week 1 (p<0.10)  27  Shear v a l u e s The Table X I I .  e f f e c t of vitamin  No s t a t i s t i c a l  A d e f i c i e n c y on shear v a l u e i s p r e s e n t e d i n  differences  i n shear v a l u e were e v i d e n t u n t i l t h e  f i f t h t r e a t m e n t week when d e f i c i e n t and m i l d l y d e f i c i e n t b i r d s had s i g n i f i c a n t l y g r e a t e r shear v a l u e s than c o n t r o l s Control increasing  (p<0.10).  and t r e a t m e n t groups a l l i n d i c a t e d a g e n e r a l t r e n d o f  shear w i t h  i n c r e a s i n g age.  S i g n i f i c a n t regression  c o e f f i c i e n t s of  r = .839 (p<0.05), r = .963 (p<0.005)and r = .911 (p<0.02) were c a l c u l a t e d f o r t h e c o n t r o l , d e f i c i e n t and m i l d l y d e f i c i e n t b i r d s r e s p e c t i v e l y . icant regression  Signif-  c o e f f i c i e n t s o f r = .955 (p<0.005) and r = .920 (p<0.01)  were o b t a i n e d f o r d e f i c i e n t and m i l d l y d e f i c i e n t b i r d s when shear was e x p r e s s e d as kg/cm^. A s i m p l e c o r r e l a t i o n a n a l y s i s was performed on t h e d a t a o b t a i n e d f o r t h e P. major muscle and t h e c o r r e l a t i o n m a t r i x f o r t h e c o n t r o l  and f o r t h e  d e f i c i e n t group i s p r e s e n t e d i n T a b l e X I I I and T a b l e XIV r e s p e c t i v e l y . In t h e c o n t r o l  group, t h e r e was a s i g n i f i c a n t c o r r e l a t i o n between  the time t o maximum t e n s i o n  and ATP, as w e l l as between t h e maximum  tension  developed and shear v a l u e . In t h e d e f i c i e n t group t h e time t o maximum t e n s i o n  was s i m i l a r l y  s i g n i f i c a n t l y c o r r e l a t e d w i t h ATP; a d d i t i o n a l l y i t e x h i b i t e d  a significant  association  with  glycogen.  The d e f i c i e n t group showed a s i g n i f i c a n t c o r r e l a t i o n  between ATP and glycogen and between ATP and s h e a r v a l u e . i c a n t l y d i f f e r e n t than those a s s o c i a t i o n s control  group (p<0.05).  Both a r e s i g n i f -  between t h e same parameters o f t h e  T a b l e XII:.  Means and s t a n d a r d e r r o r s o f shear v a l u e s f o r White Leghorn c o c k e r e l P. major  kg  kg/cm'  Vitamin A d e f i c i e n t Duration of Treatment ( i n weeks) 1  2 3 4 5  Control  1.82 1.97 2.06 2.40 2.19  + 0.2aA + 0 . laA + 0 . laA + 0.2aB + 0.2aA  Deficient  1.89 1.91 2.35 2.40 2.69  + + + + +  Mildly deficient  0.2aAi,  O.laA 0.4aA 0.2aA  0.2bB  1.99 2.30 2.78  + 0.2aA + 0.2aA + O.IbB  Vitamin A d e f i c i e n t  Control  3.04 3.29 3.43 3.99 3.51  + + + + +  0.3aA 0.2aA 0.3aA 0.3aB 0.4aA  Deficient  3.17 3.19 3.92 4.00  4.33  + + + + +  0.3aA 0 . laA 0.7aA 0.3aB  0.2bB  Mildly  3.32 3.84 4.48  deficient  + 0.3aA + 0.4aA + 0.3bB  1 means i n same row f o l l o w e d by s i m i l a r lower case l e t t e r s do not d i f f e r s i g n i f i c a n t l y ( p < 0 . 1 0 ) 2 means i n same column f o l l o w e d by s i m i l a r c a p i t a l l e t t e r s do not d i f f e r s i g n i f i c a n t l y from mean a t Week 1 (p<0.10)  CO  29  Table X I I I :  C o r r e l a t i o n m a t r i x f o r parameters s t u d i e d control birds  TMT MT ATP Gl ycogen Shear (kg) ^ Shear (kg/cm ) * **  TMT  MT  ATP  1.000 -0.097 0.359* 0.179 -0.185 -0.168  1 .000 -0.131 -0.119 -0.317* -0.294  1 .000 -0.274 0.019 0.003  Glycogen  1 .000 0.044 0.038  Shear (kg) Shear (kg/cm )  1 .000 0.992**  1 .000  p<0.05 p<0.01  T a b l e XIV:  C o r r e l a t i o n m a t r i x f o r parameters s t u d i e d treatment birds TMT  TMT MT ATP Gl ycogen Shear (kg) ^ Shear (kg/cm ) * **  i n Experiment 2,  p<0.05 p<0.01  1 .000 0.236 0.542** 0.343* -0.244 -0.246  MT 1.000 0.249 0.236 -0.191 -0.221  in Experiments,  ATP  Glycogen  Shear (kg)  1 .000 0.533** -0.383* -0.381*  1.000 -0.295 -0.285  1.000 0.996**  Shear (kg/cm )  1.000  30  L i v e r glycogen and v i t a m i n  A  When c a l c u l a t e d on p e r gram dry m a t t e r b a s i s , s i g n i f i c a n t l y greater  (p<0.10) f o r d e f i c i e n t b i r d s o f t h e f o u r t h t r e a t m e n t week  when compared t o c o n t r o l s  ( T a b l e XV).  h i g h e r l i v e r glycogen than c o n t r o l s avitaminoic  l i v e r glycogen was  M i l d l y d e f i c i e n t b i r d s had s i g n i f i c a n t l y  (p<0.10) o n l y a f t e r f i v e weeks on t h e  A r a t i o n though an e l e v a t e d  l e v e l was noted  in the fourth  treatment  week. D e f i c i e n t b i r d s of t h e f o u r t h and m i l d l y d e f i c i e n t b i r d s o f t h e f i f t h t r e a t m e n t week both m a i n t a i n e d s i g n i f i c a n t l y d i f f e r e n t v a l u e s when glycogen  l e v e l s were e x p r e s s e d as p e r c e n t t o t a l dry l i v e r ; t h e m i l d l y d e f i c i e n t  b i r d s of t h e f o u r t h week had s i g n i f i c a n t l y h i g h e r p e r c e n t l i v e r glycogen than c o n t r o l s  (p<0.10).  The e f f e c t o f d i e t a r y v i t a m i n presented i n T a b l e XV. had  levels  lower h e p a t i c  Those c h i c k s  s t o r e s than c o n t r o l  was i n d i c a t e d f o r t h e t r e a t e d  A on h e p a t i c  vitamin  r e c e i v i n g the vitamin  A stores  A deficient ration  b i r d s a t each t r e a t m e n t s t a g e .  b i r d s , but a gradual i n c r e a s e  i s also  No  trend  in l i v e r vitamin  was noted f o r t h e c o n t r o l b i r d s from 6 through 8 weeks o f age, a f t e r which a plateau  was reached.  A  T a b l e XV:  The e f f e c t o f v i t a m i n A d e f i c i e n c y on l i v e r glycogen and v i t a m i n A c o n t e n t Glycogen (jumoles glucose/gm d r y weight)  Glycogen ( P e r c e n t t o t a l dry l i v e r )  Vitamin A d e f i c i e n t Duration of Treatment ( i n weeks) 1 2 3 4 5  1 2 3  Control 253. 232. 257. 188. 272.  8±25 • 7aA 4±41 .4aA 5±51 • OaA 3±34 .3aA 1 ±28.3aA  Deficient 243.8±25.9aA^ 263.0+55.3aA 239.0±59.2aA 379.1±69.9bB 318.4±65.0aA  Vitamin A d e f i c i e n t  Mi Idly d e f i c i e n t  Control  236.8±59.2aA 333.0±89.5aA 403.0±75.7bB  4.57±0.5aA 4.18±0.6aA 4.62±0.9aA 3.39±0.5aA 4.90±0.5aA  Vitamin A (lU/gm d r y wt)  Deficient Mildly^defieientvControl 4 .39±0. 5aA 4 •73±1. OaA 4 .30±0. 9aA 6 . 82±.1.1 bB 5 .7311. 2aA  4.30±1 . laA 6.12±1 • 3bA 7.25±1 • 4bB  46.7±7.2 117.616.4 136.2±6.2 147.5±6.2 136.012.4  Deficient 6.910.8 6.110.8 4.912.7 1.610.4 3.210.7  mean and s t a n d a r d e r r o r means i n same row f o l l o w e d by s i m i l a r lower case l e t t e r s do not d i f f e r s i g n i f i c a n t l y (p<0.10) means i n same comumn f o l l o w e d by s i m i l a r c a p i t a l l e t t e r s do not d i f f e r s i g n i f i c a n t l y from mean o f Week 1 (p<0.10)  32 Experiment 3 Mortality  rate In Experiment 3 t h e m o r t a l i t y r a t e i n t h e f i r s t f i v e weeks p r i o r t o  d i e t a l l o c a t i o n was r e l a t i v e l y  low a t 1.9$.  By t h e end of t h e seven week  e x p e r i m e n t a l p e r i o d , 3.8$ of t h e c o n t r o l b i r d s d i e d whereas t h e group the t r e a t m e n t r a t i o n e x p e r i e n c e d a 34.6$ r e d u c t i o n .  receiving  N o t a b l y , t h e l a r g e r number  of deaths o c c u r r e d d u r i n g t h e f i f t h and s i x t h t r e a t m e n t weeks f o r which m o r t a l i t y r a t e s o f 23.1$ and 5.8$ were r e c o r d e d . Body weight No s i g n i f i c a n t d i f f e r e n c e s  i n weekly body weight were observed between  c o n t r o l and v i t a m i n A d e f i c i e n t b i r d s from 5 through 9 weeks o f age; however, t r e a t e d b i r d s were s i g n i f i c a n t l y  l i g h t e r (p<0.10) than c o n t r o l s a t 10, 11 and  12 weeks o f age ( T a b l e X V I ) . S i m i l a r t r e n d s s w e r e noted body w e i g h t s of t h e sampled  b i r d s , but t h e t e r m i n a l  i n t h e weekly and t e r m i n a l  weights of those  A b i r d s s a c r i f i c e d a t 9 weeks o f age were a l s o s i g n i f i c a n t l y  lighter  avitaminotic (p<0.10)  than c o n t r o l s . Feed e f f i c i e n c y Feed e f f i c i e n c y v a l u e s f o r t h o s e b i r d s sampled are p r e s e n t e d i n T a b l e X V I I .  a t each t r e a t m e n t s t a g e  The two columns a p p e a r i n g under t h e e x p e r i m e n t a l  heading a r e d e s i g n a t e d as d e f i c i e n t and m i l d l y d e f i c i e n t .  These two groups  were e s t a b l i s h e d a c c o r d i n g t o t h e manner d e s c r i b e d i n Experiment 2. b i r d s sampled  Additionally,  d u r i n g t h e f i r s t and second weeks o f r e p l e t i o n were d e s i g n a t e d  m i l d l y d e f i c i e n t i f they had a n e t p o s i t i v e weight g a i n , though some b i r d s had p r e v i o u s l y e x p e r i e n c e d weight l o s s . Feed e f f i c i e n c y v a l u e s were markedly  i n f l u e n c e d by t r e a t m e n t ; d e f i c i e n t  and m i l d l y d e f i c i e n t b i r d s had s i g n i f i c a n t l y ! l o w e r (p<0.10) feed e f f i c i e n c y  T a b l e XVI:  Means and s t a n d a r d e r r o r s o f body weight  (Experiment  Weekly Weight o f B i r d s Used  Age ( i n weeks) 5 6 7 8 9 10 11 12  1 2  Duration of Treatment^ ( i n weeks) 0 AD 2D 3D 4D 5D IR 2R  Control (gm)  320 467 566 710 821 988 1 106 1 158  + + + + + + + +  8.0a 11.4a 14.4a 15.4a 21.2a 26.0a 35.5a 64.8a  Weekly Weight of Sampled B i r d s  Experimental (gm)  309 456 571 700 778 844 870 951  3)  + + + + + + + +  L'6.4a 911a 10.4a 13.0a 18.6a 27.0b 40.9b 24.2b  Control (gm)  Terminal Weight of Sampled B i r d s  Expermintal (gm)  Control (gm)  Experimental (gm)  r  882 1045 1159 1158  + + +  37.6a 37.1a 35.4a + 64.7a  786 842 913 951  + + + +  35.1a 50.6b 75.6b 24.2b  D, R i n d i c a t e d e p l e t i o n and r e p l e t i o n as d e s c r i b e d i n t e x t means i n same row f o l l o w e d by s i m i l a r lower case l e t t e r s do n o t d i f f e r s i g n i f i c a n t l y  952 1097 12.1-3 1182  ± ± ± ±  39.'7a 44.3a 35.8a 77.6a  (p<0.05)  807 861 934 988  +  4,1.5b 57.2b 85.3b + 16Hb + +  34  v a l u e s a t each t r e a t m e n t s t a g e . feed e f f i c i e n c y w i t h i n c r e a s i n g analysis,  C o n t r o l and t r e a t m e n t b i r d s showed d e c r e a s i n g age and when t h e data were s u b j e c t e d t o r e g r e s s i o n  s i g n i f i c a n t r e g r e s s i o n c o e f f i c i e n t s o f r = -.985 (p<0.005), r = -.959  (p<0.02) and r = -.986 (p<0.005) were c a l c u l a t e d and  ml I d l y d e f i c i e n t b i r d s  Table X V I I :  f o r the c o n t r o l ,  respectively.  The e f f e c t o f v i t a m i n A on feed e f f i c i e n c y Feed E f f i c i e n c y  9 10 11 12 1 2 3  D u r a t i o n of Treatment. ( i n weeks) 4D 5D 1R 2R  Experimental  Control 0.403 0.386 0.340 0.300  ± ± ± ±  (Experiment 3)  (gm/gm feed consumed)  0 ! trr-j Age (in weeks)  deficient  Deficient  0.021aA 0.015aA O.OIlaB 0.020aB  0.345 0.288 0.271 0.249  ± ± ± ±  0.015bA^ 0.018bB 0.025bB 0.007bBB  Mildly  deficient  0.359 ± 0.007bA 0.313 ± 0.017bB 0.296 ± 0.024bB  as i n T a b l e XVI means i n same row f o l l o w e d by s i m i l a r lower case l e t t e r s do n o t d i f f e r s i g n i f i c a n t l y (p<0.10) means i n same column f o l l o w e d by s i m i l a r c a p i t a l l e t t e r s do n o t d i f f e r s i g n i f i c a n t l y from mean o f Week 4 (p<0.10)  L i v e r weight and m o i s t u r e c o n t e n t On a gram weight b a s i s d e f i c i e n t b i r d s had s i g n i f i c a n t l y l i g h t e r l i v e r w e i g h t s (p<0.10) than c o n t r o l s  a t 9, 10 and 11 weeks o f age ( T a b l e  XVIII);  m i l d l y d e f i c i e n t b i r d s had s i g n i f i c a n t l y l i g h t e r (p<0.10) l i v e r w e i g h t s than controls  a t 9 and 10 weeks of age.  Compared t o t h e average l i v e r weight a t  9 weeks o f age, s i g n i f i c a n t i n c r e a s e s i n l i v e r weight were observed f o r 11 and  12 week-old t r e a t e d  between c o n t r o l  group  birds.(p<0.10). birds.  No s i g n i f i c a n t d i f f e r e n c e s  were noted  35  Table X V I I I :  Means and s t a n d a r d e r r o r s of l i v e r weight and m o i s t u r e c o n t e n t (Experiment 3) Experimental  Age (in weeks)  D u r a t i o n of Treatment. ( i n weeks)  Control  Deficient  Mildly  deficient  L i v e r Weight (gm) 9 10 11 12  4D 5D 1R 2R  26.7 29.2 28.4 27.0  ± ± ± ±  1.5aA 0.9aA I.OaA 1.5aB  19.9 20.9 24.1 24.9  ± ± ± ±  1.2bA* 1.4bA .1 -5bBl.laB  20.8 22.8 25.3  + + +  0.8bA 1 .6bA 1.7aB  +  0.1aA 0.2aA 0.4aA  L i v e r Weight (as ?5 body weight) 9 10 11 12  4D 4D 1R 2R  2.79 2.61 2.35 2.23  ± ± ± ±  0.2aA O.laA O.laB 0.1aB  2.46 2.40 2.71 2.52  ± ± ± ±  O.laA O.laA 0.3aA 0:ibA  2.48 2.38 2.60  ± + ± ±  0.008aA 0.015aA 0.014aA 0.005aA  0.704 ± 0.008aA 0.693 ± 0.023aA 0.682 ± 0.008aB  + _±  L i v e r moisture content 9 10 11 12 1 2 3  4D 5D IR 2R  0.698 0.666 0.701 0.702  ± ± ± ±  0.011aA 0.019aA 0.006aA 0.007aA  0.706 0.702 0.701 0.707  as i n T a b l e XVI means i n same row f o l l o w e d by s i m i l a r lower case l e t t e r s do n o t d i f f e r s i g n i f i c a n t ! (p<0.10) means i n same column f o l l o w e d by s i m i l a r c a p i t a l l e t t e r s do not d i f f e r s i g n i f i c a n t ! from mean of Week 4 (p<0.10)  36 Expressed as p e r c e n t body w e i g h t , cantly different until greater  12  l i v e r weights were not s i g n i f i -  weeks of age when d e f i c i e n t b i r d s had s i g n i f i c a n t l y  l i v e r weight v a l u e s - t h a n c o n t r o l s In c o n t r o l b i r d s ,  (p<0.10).  l i v e r weight as p e r c e n t body weight  decreased  w i t h i n c r e a s i n g age and a s i g n i f i c a n t r e g r e s s i o n c o e f f i c i e n t o r r = (p<0.002) was o b t a i n e d f o r t h i s group.  -.991  No d e f i n i t e t r e n d was i n d i c a t e d f o r  the t r e a t e d groups. No s i g n i f i c a n t d i f f e r e n c e s were noted w i t h r e s p e c t t o l i v e r m o i s t u r e content. Isometric tension a)  parameters  Time t o maximum t e n s i o n P. major samples from b i r d s of the t r e a t e d group r e q u i r e d  t o develop  longer  i s o m e t r i c t e n s i o n than samples from c o n t r o l s a t 10 and 11 weeks of  age (Table X I X ) .  D e f i c i e n t and m i l d l y d e f i c i e n t 11 week-old b i r d s showed s i g -  n i f i c a n t l y extended t i m e s t o maximum t e n s i o n when compared t o c o n t r o l s  (p<0.10);  they were a l s o s i g n i f i c a n t l y s l o w e r in r e a c h i n g maximum t e n s i o n than t r e a t e d b i r d s a t 9 weeks o f age  (p<0.10).  No s t a t i s t i c a l d i f f e r e n c e s were noted between t h e d e f i c i e n t and the m i l d l y d e f i c i e n t b i r d s of a p a r t i c u l a r t r e a t m e n t s t a g e ; however, m i l d l y d e f i c i e n t b i r d s had n o m i n a l l y e l e v a t e d t i m e s t o maximum t e n s i o n when compared to t h e i r respective d e f i c i e n t b i r d s . b)  Maximum t e n s i o n D e f i c i e n t b i r d s developed s i g n i f i c a n t l y g r e a t e r maximum t e n s i o n  than c o n t r o l s a t 10 and 11 weeks of age (p<0.10), whereas m i l d l y d e f i c i e n t b i r d s developed s i gn i fji c a n t I y g r e a t e r t e n s i o n than c o n t r o l s (p<0.10) o n l y a t 11 weeks of age. cantly higher 9 weeks of  The v a l u e s f o r d e f i c i e n t 11 week-old b i r d s were a l s o s i g n i f i -  (p<0.10) than those develeped by t h e i r r e s p e c t i v e groups a t  age.  T a b l e XIX: Means and s t a n d a r d e r r o r s o f t i m e and t e n s i o n development postmortem f o r s t r i p s o f White Leghorn c o c k e r e l P. major muscle run i n phosphate b u f f e r . Time t o Maximum Tension (minutes)  Maximum I s o m e t r i c T e n s i o n (gm/cm )  Experimental  Age ( i n weeks) 9 10 11 12  1 2 3  Duration of Treatment^ ( i n weeks) 4D 5D IR 2R  Contro1 232.8±42.9aA 199.8±45.8aA 201.1±41.6aA 258.7±22.7aA  Def i c i e n t 238.5134.2aA!l 299.6159.4aA 376.1155.6bB 259.4164.OaA-  M i l d l y def i c i e n t 243.4±39.1aA 320.6±55.6aA 426.5±59.7bB  Experimental Control 81.05110.7aA 76.871 4-3aA 81.291 4.7aA 79.381 7.3aA  Def i c i e n t  89.2H5.2aA 89.64i6.4bA 113.61l8.4bB 84.50±5.8aA  Mildly  deficient  87.67i5.8aA 85.02l8.2aA 106.3219.4bA  D, R as i n T a b l e XVI means i n same row f o l l o w e d by s i m i l a r lower case l e t t e r s do not d i f f e r s i g n i f i c a n t l y (p<0.10) means i n same column f o l l o w e d by s i m i l a r c a p i t a l l e t t e r s do not d i f f e r s i g n i f i c a n t l y from mean o f Week 4 (p<0.10)  38  P. major m e t a b o l i t e l e v e l s a)  ATP No s i g n i f i c a n t d i f f e r e n c e s o r r e c o g n i z a b l e t r e n d s were i n d i c a t e d  f o r P. major ATP v a l u e s (Table XX). b)  Glycoqen D e f i c i e n t and m i l d l y d e f i c i e n t 9 week-old b i r d s had s i g n i f i c a n t l y  h i g h e r t i s s u e glycogen v a l u e s than c o n t r o l s (p<0.10); h i g h e r glycogen  levels  were a l s o noted f o r the t r e a t e d b i r d s a t 10 and 11 weeks of age ( T a b l e XX). S l i g h t l y e l e v a t e d l e v e l s were observed when m i l d l y d e f i c i e n t b i r d s a t 9, 10, and  11 weeks of age were compared t o d e f i c i e n t b i r d s a t the same stage o f  t r e a t m e n t but no s t a t i s t i c a l  d i f f e r e n c e s were e v i d e n t .  A d e f i n i t e t r e n d of i n c r e a s i n g t i s s u e glycogen w i t h age was observed  increasing  f o r c o n t r o l b i r d s and a s i g n i f i c a n t r e g r e s s i o n c o e f f i c i e n t  of r = .916 (p<0.05) was c a l c u l a t e d . Shear v a l u e D e f i c i e n t b i r d s a t 10 weeks o f age and m i l d l y d e f i c i e n t b i r d s a t 11 weeks of age demonstrated  s i g n i f i c a n t l y g r e a t e r shear v a l u e s (kg) than t h e i r  r e s p e c t i v e c o n t r o l s (p<0.10) (Table X X I ) . T h i s p a t t e r n was m a i n t a i n e d when 2  shear v a l u e was expressed as kg/cm , though d e f i c i e n t b i r d s were a d d i t i o n a l l y s i g n i f i c a n t l y tougher than c o n t r o l s a t 11 weeks o f age (p<0.10). A s i m p l e c o r r e l a t i o n a n a l y s i s was performed f o r t h e P. major muscle. parameters  on t h e d a t a o b t a i n e d  As no s i g n i f i c a n t a s s o c i a t i o n s between any o f t h e  were o b t a i n e d f o r t h e c o n t r o l group, t h e c o r r e l a t i o n m a t r i x f o r  t h i s group i s not p r e s e n t e d . is presented  i n Table X X I I .  The c o r r e l a t i o n m a t r i x f o r t h e d e f i c i e n t group  Table XX:  Means and s t a n d a r d e r r o r s o f P. major ATP and glycogen  levels  ATP (jumoles/gm)  Age ( i n weeks)  Duration of Treatment^ ( i n weeks)  9 10 11 12  4D 5D 1R 2R  1 2 3  Control 4.99±0.7aA 5.88±0.7aA 6.75±0.6aB 6.07±0.2aA  Mildly  Def i c i e n t 6.02±0.6aA 5.80±0.8aA 6.34±0.8aA 4.78±0.7aA  Glycogen  2  def i c i e n t  5.81±0.6aA 6.,T9±1.4aA 6.78±0.9aA  (jumoles glucose/gm)  Control  Def i c i e n t  5.36±0.6aA 8.72±1.2aB 8.51±0.8aB 10.42±1.0aB  9.51±1.4bA 11.32±2.5aA 9.98±1.7aA 8.83±0.6aA  Mi I d l y def i c i e n t 9.64±1.6bA 14.12±4.0aA 11.32±2.0aA  D, R as i n T a b l e XVI means i n same row f o l l o w e d by s i m i l a r lower case l e t t e r s do not d i f f e r s i g n i f i c a n t l y (p<0.10) means i n same column f o l l o w e d by s i m i l a r c a p i t a l l e t t e r s do not d i f f e r s i g n i f i c a n t l y from mean o f Week 4 (p<0.10)  UJ  VO  Table XXI:  Means and s t a n d a r d e r r o r s o f s h e a r v a l u e s f o r White Leghorn c o c k e r e l P. major Shear (kg)  Shear (kg/cm )  Exper imenta I Age ( i n weeks)  Duration of Treatment^ ( i n weeks)  9 10 11 12  4D 5D IR 2R  1 2 3  Control  2.12±0.2aA 1.92±0.2aA 1.98±0.1aA 2.23±0.2aA  Deficient  2.45±0.8aA^ 2.44±0.2bA 2.28±0.2aA 2.37±0.1aA  Mi I d l y  <:2. _ r i Exper imenta I deficient  2.31±0.2aA 2.42±0.2bA  Control  3.28±0.6aA 3.08±0.2aA 3.05±0.1aA 3.37±0.3aA  Def i c i e n t  3.68±1.1aA 3.95±0.3bA 3.87±0.3bA 3.75±0.2aA  Mildly  def i c i e n t  3.70±0.4aA 3.79±0.4bA  as i n T a b l e XVI means i n same row f o l l o w e d by s i m i l a r lower case l e t t e r s do not d i f f e r s i g n i f i c a n t l y (p<.0.10) means i n same column f o l l o w e d by s i m i l a r c a p i t a l l e t t e r s do not d i f f e r s i g n i f i c a n t l y from mean of Week 4 (p<0.10)  o  41  In t h e d e f i c i e n t group, t h e r e was a s i g n i f i c a n t c o r r e l a t i o n between the time t o maximum t e n s i o n and g l y c o g e n , as w e l l as between ATP and shear valuev  No o t h e r s i g n i f i c a n t a s s o c i a t i o n s  Table XXII:  C o r r e l a t i o n m a t r i x f o r parameters s t u d i e d • treatment birds  TMT MT ATP GIycogen Shear (kg) ^ Shear (kg/cm )  ATP  MT  TMT  * **  are evident.  1.000 0.115 0. 184 0.470** -0.049 -0.107  1 .000 0.191 -0.061 -0.135 -0.109  Glycogen  1.000 1 .000 0.168 -0.398* .,-0.060 -0.527** -0.132  i n Experiment 3,  Shear (kg) Shear (kg/cm )  1 .000 0.904**  1.000  p<0.05 p<0.01  L i v e r glycogen and v i t a m i n A When c a l c u l a t e d on per gram d r y m a t t e r b a s i s , s l i g h t l y  elevated  liver  glycogen v a l u e s were noted f o r d e f i c i e n t and m i l d l y d e f i c i e n t b i r d s a t 9 and 10 weeks o f age, and t h e d i f f e r e n c e s  between t h e glycogen v a l u e s o f m i l d l y  d e f i c i e n t b i r d s and c o n t r o l s were g r e a t e r deficient birds.  than those between c o n t r o l s and  A t 11 weeks o f age lowered l i v e r glycogen v a l u e s were observed  f o r d e f i c i e n t b i r d s , whereas m i l d l y d e f i c i e n t b i r d s and c o n t r o l s had s i m i l a r glycogen l e v e l s .  A s i m i l a r pattern  expressed as p e r c e n t t o t a l influenced  by v i t a m i n  was d i s p l a y e d  dry l i v e r .  when l i v e r glycogen i s  L i v e r glycogen was n o t s i g n i f i c a n t l y  A deficiency or vitamin  A r e p l e t i o n (Table X X I I I ) .  An age-dependent decrease i n l i v e r glycogen s t o r a g e was observed f o r c o n t r o l and m i l d l y d e f i c i e n t b i r d s .  S i g n i f i c a n t regression  coefficients  42  of r = -.966 (p<0.01) and r = -.947 (p<0.02) were c a l c u l a t e d f o r c o n t r o l and m i l d l y d e f i c i e n t b i r d s r e s p e c t i v e l y when l i v e r glycogen was e x p r e s s e d as jumoles/gm dry m a t t e r . H e p a t i c v i t a m i n A s t o r e s were d r a m a t i c a l l y a f f e c t e d by d i e t a r y vitamin  A . D e f i c i e n t b i r d s a t 9 and 10 weeks o f age had markedly reduced  l e v e l s when compared t o c o n t r o l s .  When p r e v i o u s l y d e f i c i e n t b i r d s  received  the v i t a m i n A - a d e q u a t e r a t i o n f o r a 2 week p e r i o d , t h e r e was a gradual in  increase  l i v e r v i t a m i n A s t o r e s , though t h e l e v e l s i n t h e r e p l e t e d b i r d s remained  lower than t h e c o n t r o l a v a l u e s .  43  Table X X I I I :  The e f f e c t o f v i t a m i n A on l i v e r g l y c o g e n and v i t a m i n A ExperimentaI  Age ( i n weeks)  D u r a t i o n of Treatment, ( i n weeks)  Control  Deficient  Mildly  deficient  L i v e r glycogen (umoles q l u c o s e / q m dry weiqht 9 10 11 12  4D 5D IR 2R  310.8 283.6 248.0 243.2  ± + ± ±  57.2aA 15.7aA 22.6aA 41.2aA  L i v e r glycogen ( P e r c e n t 9 10 1 1 12  4D 5D IR 2R  5.58 5.08 4.46 4.38  355. 1 300.5 206.2 250.6  t o t a l dry  ± • 0V7aA ± 0.2aA ± 0.4aA ± 0.7aA  ±151.5aA? 53.1aA + 56.9aB + 15.0aB  +  389.8 353.9 254.0  + +  44.0aA 60.9aA + 64.7aA  1 iver) 6.41 5.41 3.71 4.51  + + + +  0.9aA I.OaA 1 .OaB 0.3aA  7.04 + 6.37 + 4.59 +  Vitamin A (1U/qm d r y wt.) 9 .10 11 12  1 2 3  4D 5D 1R 2R  160.5 143.0± 186.0 151.6  ± 11.3 ±1.1.4 ± 11.7 ± 2.4  4.4 3.7 31.4 75.7  + + + +  1.5 0.5 1.0 1 .5  as i n T a b l e XVI means i n same row f o l l o w e d by s i m i l a r lower case l e t t e r s do not d i f f e r s i g n i f i c a n t l y (p<0.10) means i n same column f o l l o w e d by s i m i l a r c a p i t a l l e t t e r s do n o t d i f f e r s i g n i f i c a n t l y from mean of Week 4 (p<0.10)  0.8aA 1. 1aA 1.2aA  44  DISCUSSION Experiment 1 was c o n c e i v e d as a p r e l i m i n a r y d i e t formulation  study t o i d e n t i f y t h e  b e s t s u i t e d f o r subsequent s t u d i e s and t o e x e c u t e and e v a l u a t e  the a n a l y t i c a l t e c h n i q u e s . The data i n d i c a t e d s i g n i f i c a n t l y week-old b i r d s which r e c e i v e d  l i g h t e r body weight f o r t h e 8  375 IU v i t a m i n  A/kg and s i g n i f i c a n t l y l i g h t e r  l i v e r weights f o r t h e b i r d s maintanined on t h e two lowest l e v e l s o f v i t a m i n A s u p p l e m e n t a t i o n (373 and 750 l U / k g ) .  However, as no s i g n i f i c a n t d i f f e r e n c e s  or recognizable  t r e n d s were i n d i c a t e d f o r any o f t h e o t h e r parameters examined,  t h i s study w i l l  not be d i c u s s e d  further.  The e x t r e m e l y high m o r t a l i t y r a t e encountered i n t h e f i r s t week o f Experiment 2 i s d i f f i c u l t in t h e f o u r week p e r i o d  t o e x p l a i n but as no f u r t h e r deaths were recorded p r i o r t o treatment a l l o c a t i o n , the s u r v i v i n g  were c o n s i d e r e d s u i t a b l e f o r subsequent s t u d y . assumption as some c o n t r o l and v i t a m i n  t h e second week of t r e a t m e n t .  vaccination  T h i s may n o t have been a v a l i d  A d e f i c i e n t b i r d s e x h i b i t e d symptoms  of Newcastle d i s e a s e ( c o u g h i n g , b r e a t h i n g during  birds  w i t h a r a t t l i n g sound, weak necks)  T h i s o c c u r r a n c e and t h e implemented  procedure c o m p l i c a t e t h e i n t e r p r e t a t i o n o f t h e data o b t a i n e d -  data r e f l e c t i n g a n u t r i t i o n a l s t r e s s as w e l l as one due t o i n f e c t i o n .  No  e v i d e n c e of Newcastle d i s e a s e was noted f o r those b i r d s o f Experiment 3 but they a l s o r e c e i v e d  vaccinations.  In both e x p e r i m e n t s t h e m o r t a l i t y r a t e s of t h e c o n t r o l l e s s than those of t h e c o r r e s p o n d i n g t r e a t e d  b i r d s were  b i r d s and, as e x p e c t e d , t h e h i g h e s t  m o r t a l i t y r a t e s f o r t h e l a t t e r were noted i n t h e l a t e r s t a g e s o f d e f i c i e n c y . The e f f e c t of f e e d i n g was s t u d i e d  a v i t a m i n A adequate r a t i o n t o p r e v i o u s l y  deficient birds  i n Experiment 3 and i t would appear t h a t a s i n g l e week o f r e p l e t i o n  45 was not s u f f i c i e n t t o overcome t h e e f f e c t s o f a f i v e week p e r i o d o f d e f i c i e n c y : the second h i g h e s t m o r t a l i t y r a t e o f t h i s study was recorded  at this  time.  Both e x p e r i m e n t s i n d i c a t e t h a t a v i t a m i n o s i s A does not a f f e c t body weight g a i n d u r i n g t h e e a r l y s t a g e s d e f i c i e n c y progresses.  but s i g n i f i c a n t l y depresses growth as t h e  Harms e t aj_. (1955) r e p o r t e d t h a t t h e r e was no d i f f e r -  ence i n t h e average weight of 4 week-old b i r d s f e d 500 IU compared t o t h a t o f control  b i r d s which r e c e i v e d 4000 IU, w h i l e N o c k e l s e t a]_. (1973) r e p o r t e d  significantly  l i g h t e r body w e i g h t s f o r v i t a m i n A d e f i c i e n t b i r d s a t 5, 6 and  7 weeks o f age.  Dorr and B a l l o u n  (1976) confirmed  t h i s growth d e p r e s s i o n i n  a v i t a m i n o t i c A t u r k e y s o f both sexes and f u r t h e r suggested t h a t males may be more s e n s i t i v e t o v i t a m i n A t r e a t m e n t s .  Any d i s c r e p a n c i e s among t h e s e f i n d i n g s  may be a t t r i b u t e d t o v a r i a t i o n s i n t h e s e v e r i t y o f v i t a m i n A r e s t r i c t i o n and in t h e s t a g e o f development when t r e a t m e n t was i n i t i a t e d .  The l a g observed  in response t o v i t a m i n A r e p l e t i o n was not unexpected. Johnson and Wolf (1960) suggested t h a t o n l y t h e changes in a i mi Id d e f i c i e n c y should  observable  be thought i n d i c a t i v e o f t h e r e a l m e t a b o l i c  of v i t a m i n A and t h i s view has r e c e i v e d s u p p o r t from s e v e r a l o t h e r  function  workers.  (Nockels and P h i l l i p s 1971 a + b., Bcuckenta I e t aj_. 1974 and Krause e t aj_. 1975) The  a c t u a l c l a s s i f i c a t i o n o f which d e f i c i e n c y s t a t e t h e data presented  herein  r e p r e s e n t s , u t i I i z e d t h e c a t e g o r i e s proposed by Krause e_f aj_. (1975) who suggested t h a t r a t s were s e v e r e l y d e f i c i e n t when they no longer gained o r began t o lose weight but m i l d l y d e f i c i e n t when weight g a i n decreased w i t h respect t o control values.  F o r t h e purpose o f t h i s s t u d y , t h e ' d e f i c i e n t '  group (as i n d i c a t e d p r e v i o u s l y ) i n c l u d e s both s e v e r e l y and m i l d l y d e f i c i e n t bi r d s . Feeding a v i t a m i n A d e f i c i e n t r a t i o n lowered feed e f f i c i e n c y and  46  t h i s decrease was s i g n i f i c a n t f o r t h e t r e a t e d  b i r d s o f Experiment 3.  In both  e x p e r i m e n t s , t h e feed e f f i c i e n c y v a l u e s f o r m i l d l y d e f i c i e n t b i r d s f e l l those f o r c o n t r o l  and d e f i c i e n t b i r d s .  of d e c r e a s i n g feed e f f i c i e n c y w i t h Singh and Donovan  Both s t u d i e s  between  indicated a d e f i n i t e trend  i n c r e a s i n g age f o r a l l t h r e e groups o f b i r d s .  (1973) suggested t h a t a r e d u c t i o n  i n feed u t i l i z a t i o n was  due t o a lowered a b s o r p t i o n o f n u t r i e n t s , a normal a b s o r p t i o n f o l l o w e d by i n e f f i c i e n t m e t a b o l i c use o r a c o m b i n a t i o n o f both. The e f f e c t o f v i t a m i n  A d e f i c i e n c y on l i v e r weight appears s i m i l a r  t o i t s e f f e c t on body w e i g h t , w i t h and  l i t t l e o r no e f f e c t i n t h e e a r l i e r s t a g e s  a s i g n i f i c a n t d e p r e s s i o n f o r both d e f i c i e n t and m i l d l y d e f i c i e n t b i r d s  a f t e r f i v e weeks o f d e p l e t i o n .  The response t o r e f e e d i n g  of a vitamin A  adequate r a t i o n however was q u i c k e r than t h a t o f body weight r e s u l t i n g i n s i m i l a r l i v e r weight v a l u e s f o r c o n t r o l repletion.  and m i l d l y d e f i c i e n t b i r d s . a f t e r one week o f  When l i v e r weight was e x p r e s s e d as p e r c e n t body weight t h i s e f f e c t  of r e p l e t i o n was a c c e n t u a t e d . T r e a t e d and c o n t r o l l i v e r weight w i t h as c l e a r l y d e f i n e d  b i r d s showed t h e expected p a t t e r n  of increasing  i n c r e a s i n g age i n Experiment 2, but t h i s p a t t e r n  was n o t  i n Experiment 3 as t h e sampled b i r d s were o l d e r .  The f i n d i n g s o f t h e p r e s e n t study s u p p o r t t h e o b s e r v a t i o n by "  Krause e t aj_. (V975) t h a t r e t i n o I d e f i c i e n t r a t s had decreased l i v e r w e i g h t s . When expressed as p e r c e n t body weight however, s e v e r e l y significantly  d e f i c i e n t r a t s had  l a r g e r l i v e r s w h i l e m i l d l y d e f i c i e n t ones d i d not d i f f e r  i c a n t l y from c o n t r o l s .  This nonsignificant  l i v e r weight response has a l s o  been observed i n m i l d l y d e f i c i e n t c h i c k e n s ( N o c k e l s and P h i l l i p s L i v e r m o i s t u r e c o n t e n t i s not a f f e c t e d No e f f e c t o f v i t a m i n  A d e f i c i e n c y on t o t a l  signif-  by v i t a m i n  1971a).  A deficiency.  body water, e x p r e s s e d as p e r c e n t  47  body w e i g h t , was found e i t h e r i n r a t s (Mahant and Eaton 1976) o r i n c h i c k e n s (Lopez e t aj_. 1973), a l t h o u g h n e i t h e r of t h e s e s t u d i e s examined t h e m o i s t u r e content of s p e c i f i c  Organs.  There was a d e f i n i t e i n f l u e n c e o f v i t a m i n A d e f i c i e n c y on h e p a t i c c a r b o h y d r a t e metabolism and t h e response appears r e l a t e d t o t h e p r o g r e s s o f the d e f i c i e n c y .  M i l d h y p o v i t a m i n o s i s A :i,nduces an i n c r e a s e i n glycogen  d e p o s i t i o n whereas a severe d e f i c i e n c y stores.  leads t o a r e d u c t i o n o f t h e s e e l e v a t e d  The e f f e c t i s more apparent i n Experiment 3 where t h e r e s u l t s a r e  not masked by t h e s t r e s s o f i n f e c t i o n .  The lowered  l i v e r g l y c o g e n v a l u e observed  f o r t h e d e f i c i e n t b i r d s d u r i n g t h e f i r s t week o f r e p l e t i o n r e f l e c t s t h e cont r i b u t i o n o f t h o s e b i r d s which  lagged i n t h e i r response t o a v i t a m i n A  r a t i o n and m a i n t a i n e d a s e v e r e l y d e f i c i e n t s t a t u s . d e f i c i e n t b i r d s responded  adequate  On t h e o t h e r hand, m i l d l y  very r a p i d l y and e x h i b i t e d near normal  values f o r  the e n t i r e r e p l e t i o n p e r i o d , though they r e t a i n e d s l i g h t l y e l e v a t e d glycogen l e v e l s when e x p r e s s e d as p e r c e n t t o t a l  dry l i v e r .  These r e s u l t s g e n e r a l l y agree w i t h those p r e v i o u s l y r e p o r t e d , especially  i f c o n s i d e r a t i o n i s g i v e n t o t h e s e v e r i t y of v i t a m i n A r e s t r i c t i o n  and t h e age o f t h e b i r d s .  The r e p o r t by Janson and H a r r i I I (1974) t h a t a  decreased glycogen d e p o s i t i o n may be a s s o c i a t e d w i t h an a c u t e d e f i c i e n c y i n r a t s s u p p o r t s Johnson and Wolf  (1960) who found e s s e n t i a l l y no glycogen  d e p o s i t i o n i n a v i t a m i n o s i s A.  N o c k e l s and P h i l l i p s (1971 a + b) found h i g h e r  l i v e r glycogen v a l u e s f o r m i l d l y d e f i c i e n t c h i c k e n s but l a t e r r e p o r t e d a decrease  i f l i v e r glycogen was e x p r e s s e d on a p e r c e n t b a s i s ( N o c k e l s e t a I.  Perek and K e n d l e r (1969) a l s o r e p o r t e d t h a t l i v e r glycogen decreased  in v  vitamin A deficiency. In Experiment 3, both c o n t r o l and m i l d l y d e f i c i e n t b i r d s  indicated  1973).  48  an age-dependent decrease i n p e r c e n t weeks o l d . Kendler  l i v e r glycogen  f o r b i r d s 9 through  T h i s i s not i n accord w i t h the i n c r e a s e observed by Perek  (1969),  12  and  but t h e i r f i n d i n g s were o b t a i n e d w i t h 19 and 33 day o l d  ch i c k s . V i t a m i n A s t o r e s i n l i v e r are d e f i n i t e l y a v a i l a b l e when u r g e n t l y needed and  have been shown t o be a r e l i a b l e c r i t e r i o n f o r d e t e r m i n i n g  the  s t a b i l i t y , a v a i l a b i l i t y and u t i l i z a t i o n of, d i e t a r y v i t a m i n A (Harms e t aj_. 1955). V i t a m i n A d e f i c i e n t c o c k e r e l s showed a marked r e d u c t i o n in t h e i r though the f a c t t h a t the v i t a m i n A d e t e r m i n a t i o n s p r e c l u d e s the use of s t a t i s t i c a l was  anticipated,-the extremely  analysis.  low l e v e l s o f  were based on pooled  Although  samples  a f a i r l y r a p i d response  l i v e r v i t a m i n A encountered in  the f i r s t two weeks of d e f i c i e n c y i n Experiment 2 are p u z z l i n g . s t r e s s of i n f e c t i o n was  l i v e r stores  s u f f i c i e n t t o reduce these  Perhaps the  levels p r i o r to  treatment  a I location. The e f f e c t of r e p l e t i o n was in l i v e r v i t a m i n A s t o r e s was  a d r a m a t i c one  and the gradual  increase  expected.  V i t a m i n A d e f i c i e n c y does not s i g n i f i c a n t l y o f P. major muscle but does e x e r t an e f f e c t on  i n f l u e n c e the ATP  i t s glycogen  l e v e l , an  content  effect  t h a t i s perhaps b e s t demonstrated i n Experiment 3 where the r e s u l t s are l e s s confounded by e i t h e r an  i n f e c t i o n or a recent v a c c i n a t i o n .  i t s e f f e c t on h e p a t i c c a r b o h y d r a t e  Consistent  metabolism, m i l d h y p o v i t a m i n o s l s  glycogen  d e p o s i t i o n w h i l e a severe d e f i c i e n c y tends t o lower t h e s e  levels.  Although  i n c r e a s e d glycogen  A  with induces  elevated  l e v e l s were noted f o r the m i l d l y d e f i c i e n t  b i r d s the d i f f e r e n c e s from the c o n t r o l v a l u e s were n o n s i g n i f i c a n t , p o s s i b l y due t o the s m a l l e r number of b i r d s i n c l u d e d i n t h e m i l d l y d e f i c i e n t group and/or t o the s i g n i f i c a n t t r e n d of  i n c r e a s i n g glycogen  content with increasing  49 age e x h i b i t e d by c o n t r o l b i r d s .  T h i s t r e n d was  not e v i d e n t  f o r the c o n t r o l  group i n Experiment 2 where s a m p l i n g began when the b i r d s were 6 weeks o l d compared t o the 9 week-old b i r d s i n i t i a l l y sampled i n Experiment 3. These r e s u l t s are reported  i n accord w i t h N o c k e l s and  a n o n s i g n i f i c a n t t r e n d of  P h i l l i p s (1971a)  i n c r e a s i n g muscle glycogen w i t h  v i t a m i n A d e f i c i e n c y f o r m i l d l y d e f i c i e n t c h i c k e n s a t 8 and  increasing  16 weeks of  The  age-dependent i n c r e a s e noted f o r c o n t r o l s s u p p o r t s Rosebrough and  who  found t h a t age  c a r b o h y d r a t e and The  significantly  who  age.  Begin (1975)  i n f l u e n c e d muscle glycogen c o n t e n t i n both  fat diets.  elevated  in Experiment 2 may  glycogen l e v e l s i n the f i r s t two  weeks of  r e f l e c t the s t r e s s of Newcastle d i s e a s e  drop i n the t h i r d week may  be due  deficiency  w h i l e the  t o the a d d i t i o n a l s t r e s s of the  slight  vaccination  procedure. The  P. major samples e x c i s e d  from t r e a t m e n t b i r d s e x h i b i t e d a normal  p a t t e r n of postmortem i s o m e t r i c t e n s i o n development and required ciency, The  longer t o develop maximum t e n s i o n and,  p r e v i o u s l y observed d e l a y  s i g n i f i c a n t l y higher  l a t e r s t a g e s of  defi-  In response t o the r e f e e d i n g of a v v i t a m i n  samples. A  - both i s o m e t r i c t e n s i o n parameters are  f o r t r e a t e d b i r d s i n the f i r s t week of r e p l e t i o n but  s i m i l a r f o r the c o n t r o l and There are s e v e r a l  are  t r e a t e d b i r d s of the second week. possible explanations  i s o m e t r i c t e n s i o n development.  The  f o r t h i s e f f e c t of v i t a m i n  f i r s t would c o n s i d e r  v i t a m i n A d e f i c i e n c y on the s t r u c t u r a l and samples.  generally  i n d i c a t e d g r e a t e r t e n s i o n development than c o r r e s p o n d i n g c o n t r o l  adequate r a t i o n i s a g a i n e v i d e n t  on  i n the  d e c l i n e but  cellular  T h i s cannot be c o m p l e t e l y d i s r e g a r d e d  the e f f e c t of  i n t e g r i t y of the muscle  as h i s t o l o g i c a l  were not performed on any of the samples; however, a s ( 1 )  examinations  a normal p a t t e r n  of  A  50  development and d e c l i n e was shown by t h e t r e a t m e n t b i r d s and (2) some o f t h e d e f i c i e n t b i r d s were c a p a b l e o f d e v e l o p i n g  s i g n i f i c a n t l y greater tension  than  c o n t r o l s , t h i s p o s s i b i l i t y may not deserve g r e a t emphasis. A second e x p l a n a t i o n  c o n s i d e r s t h e e f f e c t o f a v i t a m i n o s i s A on t h e  a v a i l a b i l i t y and/or c o n c e n t r a t i o n in postmortem s h o r t e n i n g :  o f t h e major m e t a b o l i t e s  ATP and g l y c o g e n .  which have a r o l e  There a r e no r e p o r t s which d i r e c t l y  i n d i c a t e t h a t muscle ATP i s not r e a d i l y a v a i l a b l e f o r c o n t r a c t i o n but N o c k e l s and  P h i l l i p s (1971a) found t h a t v i t a m i n A d e f i c i e n c y impared l a c t i c a c i d  t h e s i s from glycogen i n w h i t e muscle. overall  If t h i s  impairment i s i n d i c a t i v e o f an  s l o w e r r a t e o f a n a e r o b i c g l y c o l y s i s - an e s s e n t i a l , though  source o f new ATP postmortem - an e x t e n s i o n maximum i s o m e t r i c t e n s i o n would be e x p e c t e d .  syn-  inefficient  i n t h e t i m e r e q u i r e d t o reach On t h e o t h e r hand, i f t h e r a t e  of a n a e r o b i c g l y c o l y s i s i s n o t markedly a f f e c t e d by a v i t a m i n A d e f i c i e n c y , actual metabolite ATP  concentrations  must be examined.  Though s l i g h t l y  elevated  l e v e l s were observed (Experiment 2), t h i s e f f e c t was n o t as c o n s i s t e n t  as t h a t observed f o r muscle glycogen c o n t e n t where m i l d h y p o v i t a m i n o s i s  A  induced g l y c o g e n d e p o s i t i o n . Bate-Smith and B e n d a l l  (1949) d e s c r i b e d one t y p e o f r i g o r m o r t i s  t h a t was c h a r a c t e r i z e d by a long d e l a y t o i t s o n s e t and o c c u r r e d a n i m a l s whose muscles had a high t o e x p e c t an i n c r e a s e levels  level of glycogen.  in well-fed  I t i s not unreasonable  i n t h e time t o maximum t e n s i o n where e l e v a t e d  glycogen  occur. The  concentrations  i n t e r r e I a t e d n e s s o f i s o m e t r i c t e n s i o n parameters w i t h  metabolite  can be seen when t h e c o r r e l a t i o n c o e f f i c i e n t s determined f o r  t h i s study a r e examined.  F o r c o n t r o l b i r d s (Experiment 2 ) , a s i g n i f i c a n t  p o s i t i v e a s s o c i a t i o n e x i s t s between t h e time t o maximum t e n s i o n and ATP; a  51  finding  in accord  w i t h Vanderstoep and  Richards  (1974) who  reported  that  a p p r o x i m a t e l y f i f t y p e r c e n t of the observed v a r i a t i o n s i n the t i m e v t o maximum t e n s i o n c o u l d be e x p l a i n e d p e c t o r a I i s major.  by d i f f e r e n c e s i n the  T h i s a s s o c i a t i o n was  Experiment 2, t o g e t h e r  i n i t i a l ATP  c o n t e n t of  turkey  a l s o found f o r the d e f i c i e n t b i r d s of  w i t h a s i g n i f i c a n t dependence on muscle g l y c o g e n  For the t r e a t e d b i r d s of Experiment 3, o n l y the glycogen c o n t e n t was  content.  significantly  c o r r e l a t e d w i t h the t i m e t o maximum t e n s i o n . The and  the  e f f e c t of v i t a m i n A d e f i c i e n c y on P. major m e t a b o l i t e  i n f l u e n c e of t h e s e changes on the parameters of  development should  be e v i d e n t  i n shear v a l u e s .  isometric  Avitaminosis  concentrations tension  A does not  appear t o s u f f i c i e n t l y a l t e r any of the f a c t o r s c o n t r i b u t i n g t o shear u n t i l the  value  l a t e r s t a g e s of d e f i c i e n c y , when the t r e a t e d b i r d s were found t o  be s i g n i f i c a n t l y tougher than c o n t r o l s . a t t r i b u t e d t o e i t h e r the tend t o r e q u i r e  This  length of the a g i n g  increased period  s h e a r v a l u e may  (as the t r e a t e d  longer t o reach maximum t e n s i o n ) o r an  increased  myofibrillar contraction.  The  The  f o r t h e d e f i c i e n t b i r d s of the f i f t h  maximum t e n s i o n v a l u e s  l a t t e r seems the most l i k e l y  be  birds  degree of  possibility. depletion  w e e k e n d the m i l d l y d e f i c i e n t b i r d s i n the f i r s t week of r e p l e t i o n were s i g n i f i c a n t l y g r e a t e r than c o n t r o l v a l u e s , and c o r r e s p o n d i n g l y  greater  shear  v a l u e s were observed f o r t r e a t e d b i r d s . Nakamura a t aj_. (1975) r e p o r t e d s h e a r v a l u e and age  c o u l d be found i n c h i c k e n s ,  gested t h a t t h i s age r e s i s t a n c e of the  and  r e l a t e d t e n d e r n e s s r e f l e c t e d an  Marsh (1977) l a t e r sugincrease  in the thermal  i n t e r m o l e c u l a r c r o s s l i n k s of the muscle c o l l a g e n .  changes i n the c o l l a g e n m o l e c u l e may dependent i n c r e a s e  t h a t a very high c o r r e l a t i o n between  These  have c o n t r i b u t e d somewhat t o the  i n shear v a l u e noted i n Experiment 2 but a more  age-  likely  52 explanation  c o n s i d e r s t h e developmental s t a g e :  t h e b i r d s a r e a c t i v e l y growing  between 6 and 10 weeks o f age and t h e o v e r a l l i n c r e a s e  i n muscle mass may be  the f a c t o r p r i m a r i l y r e s p o n s i b l e f o r t h e t r e n d . The  n a t u r e o f t h e r e l a t i o n s h i p s t h a t e x i s t between  concentrations,  i s o m e t r i c t e n s i o n parameters and shear v a l u e  wide v a r i a t i o n .  Busch e t aj_. (1967) r e p o r t e d  metabolite i s subject t o  t h a t no d i r e c t r e l a t i o n s h i p ,  e x i s t e d between ATP and shear r e s i s t a n c e . Lee et_ aj_. (1976) found t h a t  initial  glycogen l e v e l s and t h e postmortem g l y c o l y s i s r a t e were s i g n i f i c a n t l y c o r r e l a t e d w i t h t h e shear v a l u e o f ground muscle.  Other workers have i n d i c a t e d a s i m i l a r  a s s o c i a t i o n between shear and t h e r a t e o f postmortem g l y c o l y s i s (deFremery, 1966;  Khan and Kim, 1975).  Khan (1974) r e p o r t e d  t h a t t h e u l t i m a t e shear f o r c e  appears t o be d i r e c t l y p r o p o r t i o n a l t o t h e maximum i s o m e t r i c t e n s i o n whereas Wood (1973) found t h a t shear had a p o s i t i v e r e l a t i o n s h i p w i t h  developed, both  t e n s i o n and time. The  r e s u l t s o f t h i s study i n d i c a t e s i g n i f i c a n t p o s i t i v e a s s o c i a t i o n s  between t h e time t o maximum t e n s i o n w i t h ATP and/or glycogen c o n t e n t . the n e g a t i v e ATP  However,  c o r r e l a t i o n s found f o r t h e maximum t e n s i o n and shear and f o r  and shear r e q u i r e f u r t h e r s t u d y .  53  SUMMARY AND CONCLUSIONS A v i t a m i n o s i s A had l i t t l e o r no e f f e c t on e i t h e r body weight g a i n or  l i v e r weight d u r i n g i t s e a r l y s t a g e s but depressed both as t h e d e f i c i e n c y  progressed.  A s i g n i f i c a n t decrease  i n feed e f f i c i e n c y was a l s o noted  i n the  later deficiency stages. V i t a m i n A d e f i c i e n c y d i d e x p r e s s an e f f e c t on both h e p a t i c and muscle c a r b o h y d r a t e metabolism and t h e natu're o f t h e response appears t o t h e p r o g r e s s of d e f i c i e n c y .  M i l d h y p o v i t a m i n o s i s A induced an i n c r e a s e i n  glycogen d e p o s i t i o n whereas a s e v e r e d e f i c i e n c y elevated  related  led t o a reduction of these  stores. The a b i l i t y o f P. major s t r i p s t o develop i s o m e t r i c t e n s i o n postmortem  was n o t i n f l u e n c e d by d i e t a r y v i t a m i n A though some changes i n t h e measured parameters were noted. deficiency required  The P. major from c o c k e r e l s i n t h e l a t e r s t a g e s o f  l o n g e r t o reach maximum t e n s i o n and developed  g r e a t e r maximum t e n s i o n than t h o s e from c o n t r o l s . e l e v a t e d muscle glycogen c o n t e n t .  significantly  The extended t i m e s r e f l e c t e d  A s i g n i f i c a n t i n c r e a s e i n shear v a l u e was  observed when t h e i n c r e a s e i n maximum t e n s i o n o c c u r r e d . Although t h e r e was a delayed response t o t h e r e f e e d i n g o f a v i t a m i n A adequate r a t i o n , v i t a m i n A r e p l e t i o n a l s o i n f l u e n c e d t h e s t u d i e d  parameters.  Muscle g l y c o g e n c o n t e n t , i s o m e t r i c t e n s i o n parameters and s h e a r r e s i s t a n c e r e t u r n e d t o v a l u e s s i m i l a r t o those o f c o n t r o l s w i t h i n a two week p e r i o d . On the^ b a s i s o f t h i s data i t can be concluded t h a t t h e n u t r i t i o n a l s t a t u s o f v i t a m i n A i n f l u e n c e s both t h e c a r b o h y d r a t e metabolism and t h e i s o m e t r i c t e n s i o n parameters of muscle t i s s u e , w i t h a c o n c o m i t a n t e f f e c t on t h e u l t i m a t e muscle t e n d e r n e s s .  deleterious  54  LITERATURE CITED  Bate-Smith, E.C. 1948. The p h y s i o l o g y and c h e m i s t r y of r i g o r m o r t i s , w i t h s p e c i a l r e f e r e n c e t o the a g i n g of beef. Adv. Food Res. 1: 1. Bate-Smith, E.C. and B e n d a l l , J.R. 1949. F a c t o r s d e t e r m i n i n g the time of r i g o r m o r t i s . J . P h y s i o l . ' 1 1 0 : " 4 7 . B e n d a l l , J.R. 1973. 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