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Effect of corticosterone injection on carbohydrate metabolism and tenderness of broiler breast muscle Whalley, Linda Louise 1974

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a  EFFECT OF CORTICOSTEROID INJECTION ON CARBOHYDRATE  METABOLISM  AND TENDERNESS OF BROILER BREAST MUSCLE  BY  LINDA LOUISE  B.Sc.  (Agr.),  University  WHALLEY  of B r i t i s h  Columbia,  1972  A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE  REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in  t h e Department o f Food  We a c c e p t required  THE  this  Science  thesis  as c o n f o r m i n g  to the  standard  UNIVERSITY OF BRITISH COLUMBIA APRIL, 1974.  In p r e s e n t i n g  t h i s t h e s i s i n p a r t i a l f u l f i l m e n t of the requirements f o r  an advanced degree at the U n i v e r s i t y o f B r i t i s h Columbia, I agree t h a t the L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e  and  study.  I f u r t h e r agree t h a t p e r m i s s i o n f o r e x t e n s i v e copying of t h i s t h e s i s f o r s c h o l a r l y purposes may by h i s r e p r e s e n t a t i v e s .  be granted by  the Head of my  Department or  I t i s understood t h a t copying or  publication  of t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l not be allowed without written  permission.  Department o f  o  .5r„/ /- /v  The U n i v e r s i t y o f B r i t i s h Columbia Vancouver 8, Canada  my  ABSTRACT  The  effect  of intramuscular  on p o s t mortem m u s c l e m e t a b o l i s m P e e t o r a l i s Major muscle  was  until  showed no  shear r e s i s t a n c e  c o n d u c t e d o v e r an significant  6 hours post i n j e c t i o n  increased  a t which  from 6 week-old had r e c e i v e d  alteration  broilers  due  8 hour p o s t of  injection  Blood glucose l e v e l s injection.  o f s h e a r v a l u e s o f P. M a j o r and  corticosterone  d i f f e r e n c e was  effect  from 2 to 8 hours p o s t  A comparison  of  time shear v a l u e s  from those o f the c o n t r o l s .  rose s i g n i f i c a n t l y  injection  studied.  Short term s t u d i e s , injection period,  and  corticosterone  8 week-old injections  broilers  muscle after  showed t h a t  both  the  only  t o t h e d i f f e r e n c e o f age n o t t o any  i n the p h y s i o l o g i c a l  handling  of the  injected  corticosterone. The  greatest difference  f o u n d between b i r d s was  subjectively  of d i f f e r e n t  t h e r a n g e h o u s e and  level.  Stress  to the b i r d s to the  was level  reaction  birds  to handling. Long  illustrated injections Shear  stress  evaluated with respect  to a person e n t e r i n g response  i n treatment e f f e c t  t e r m s t u d i e s c o n d u c t e d o v e r 5 and  homeostatic adjustments to d a i l y and  recovery after  t e s t i n g was  cessation of  15  days  corticosterone  injections.  p e r f o r m e d on P. M a j o r m u s c l e  which  was  chilled  on  Excised  muscle d i s p l a y e d  and  cessation  a lack on  the  of  carcass  of  and  on  muscle e x c i s e d  a greater  initial  i n j e c t i o n and  injection.  r e a c t i o n to i n j e c t i o n  i n j e c t i o n than d i d  skeletal restriction.  Shear v a l u e s  increased  s t e a d i l y on rapidly  B l o o d g l u c o s e and  i n j e c t i o n of  at cessation.  As  during  required.  i n j e c t i o n than  t i s s u e glycogen and  expected ultimate increase.  i n j e c t i o n period  terone production,  values  to  t o d i s p l a y a much more  corticosterone  decreased with muscle glycogen increased  of  times.  r e a c t i o n to c o r t i c o s t e r o n e  shear t e s t i n g .  to  decreased  upon c e s s a t i o n  C h e m i c a l t e s t s were f o u n d definite  i n t a c t m u s c l e due  Homeostatic adjustments returned  n e a r - n o r m a l between t h e s e  at death.  f o r which  as  did increased  decreased muscle  Blood  pH  cholesterol  endogenous c o r t i c o s -  i t i s a precursor,  was  not  ACKNOWLEVGEMENTS The authon ootAheA to thank heA adviAon, VA. J.F. RtchaAdA, AAAoclate  PAO&ZAAOA,  6uggeAtlonA  VepanMnent o^ Food Science,  ^OA  'hlA kind  duAlng the couAAe o& thJj, Atudy. She  OIAO  uxliheA to thank the. membeAA ofa heA Graduate  Committee:  • VA. P.M. TownAley, Ve.pantme.nt ofi food VA. J. [/andeAAtoep, VepaAtment oi food  Science Science  Vh.. R.C. FXXzAlmmonA, VepaAtment o£ Foult/iy ^OA fievtew ofa thtA Financial  theA-iA. AuppoAt &Aom the UntveAA-lty o& BAltiAh  thn.ou.gh a Post-GAaduate Council  Science  FeJLlomhlp and finom the National  o/) Canada IA gnate^ully  appreciated.  Columbia ReAeaAch  iv TABLE OF CONTENTS PAGE ABSTRACT  i  ACKNOWLEDGEMENTS  i  i  i  TABLE OF CONTENTS  iv  L I S T OF TABLES  VI  L I S T OF FIGURES  v i i  INTRODUCTION  1  REVIEW OF LITERATURE  3  a) b)  Alarm Reaction R e s i s t a n c e o r A d a p t a t i o n Stage  3 3  c)  Stage  5  of Exhaustion  PHYSIOLOGICAL RESPONSE TO HYPERCORTICALISM  8  PHYSIOLOGICAL RESPONSE TO CORTICOID  9  ADRENAL CORTICAL INDUCED ALTERATIONS  INJECTION I N MUSCLE  ADRENAL CORTICAL CONTROL OF A V I A N METABOLISM  9 12  MATERIALS AND METHODS P o u l t r y Source Slaughter Procedure I n j e c t i o n Procedure Tenderness 1) I n t a c t m u s c l e 2) E x c i s e d m u s c l e Chemical Tests 1) B l o o d g l u c o s e 2) T i s s u e g l y c o g e n 3) B l o o d c h o l e s t e r o l 4) pH  15 15 15 15 16 16 16 17 17 17 17 18  RESULTS AND DISCUSSION  19  E f f e c t o f I n j e c t i o n Method S h o r t Term S t u d i e s L o n g Term S t u d i e s 1) 5 Day s t u d y 2) 15 Day s t u d y  19 21 27 27 30  TABLE OF CONTENTS CONTINUED Shear t e s t i n g Chemical t e s t s Blood glucose Tissue glycogen H P  Blood SUMMARY AND CONCLUSIONS BIBLIOGRAPHY APPENDIX  cholesterol  vi.  L I S T OF  TABLES  PAGE  TABLE 1  2  3  4  5  6  7  8  9  10  11  C o m p a r i s o n o f s h e a r v a l u e s o f P. M a j o r m u s c l e when c o r t i c o s t e r o n e was d i l u t e d w i t h d i s t i l l e d water o r p h y s i o l o g i c a l saline solution  19  C o m p a r i s o n o f s h e a r v a l u e s o f P. M a j o r m u s c l e f r o m u n i n j e c t e d b i r d s and b i r d s i n j e c t e d w i t h d i s t i l l e d water  20  E f f e c t o f c o r t i c o s t e r o n e i n j e c t i o n on s h e a r v a l u e o f P. M a j o r m u s c l e a t several i n t e r v a l s post i n j e c t i o n  22  E f f e c t o f c o r t i c o s t e r o n e i n j e c t i o n on blood g l u c o s e l e v e l s a t s e v e r a l time i n t e r v a l s post i n j e c t i o n  22  C o m p a r i s o n o f s h e a r v a l u e s o f P. M a j o r m u s c l e f r o m 6 and 8 week o l d b i r d s a t s e v e r a l time i n t e r v a l s post i n j e c t i o n of c o r t i c o s t e r o n e  24  C o m p a r i s o n o f s h e a r v a l u e s o f P. M a j o r muscle w i t h s u b j e c t i v e e v a l u a t i o n o f s t r e s s l e v e l of the b i r d  24  S h e a r v a l u e s o f P. M a j o r m u s c l e f r o m p l a c i d b i r d s o v e r an 8 h o u r p e r i o d p o s t i n j e c t i o n of corticosterone  25  S h e a r v a l u e s o f P. M a j o r m u s c l e f r o m m i l d l y e x c i t a b l e b i r d s o v e r an 8 h o u r p e r i o d p o s t injection of corticosterone  25  S h e a r v a l u e s o f P. M a j o r m u s c l e f r o m e x t r e m e l y e x c i t a b l e b i r d s o v e r an 8 h o u r period post i n j e c t i o n of corticosterone  26  U l t i m a t e pH o f P. M a j o r m u s c l e f r o m broilers receiving daily corticosterone i n j e c t i o n s f o r 8 d a y s and o b s e r v e d f o r a f u r t h e r 7 day r e c o v e r y p e r i o d  35  Blood c h o l e s t e r o l l e v e l s o f b r o i l e r s receiving daily corticosterone injections f o r 8 d a y s and o b s e r v e d f o r a f u r t h e r 7 day r e c o v e r y p e r i o d  36  vii.  LIST OF FIGURES  FIGURE  PAGE B a s i c response to s t r e s s  6  Blood glucose l e v e l s o f b r o i l e r s r e c e i v i n g d a i l y c o r t i c o s t e r o n e i n j e c t i o n s over a 5 day p e r i o d  28  Shear v a l u e s o f i n t a c t and e x c i s e d P. Major muscle o f b r o i l e r s r e c e i v i n g d a i l y c o r t i c o s t e r o n e i n j e c t i o n s over a 5 day period  29  Shear value o f e x c i s e d and i n t a c t P. Major muscle of b r o i l e r s r e c e i v i n g d a i l y c o r t i c o s t e r o n e i n j e c t i o n s f o r 8 days and observed f o r a f u r t h e r 7 days without injection  31  Blood glucose l e v e l o f b r o i l e r s r e c e i v i n g d a i l y c o r t i c o s t e r o n e i n j e c t i o n s f o r 8 days and observed f o r a f u r t h e r 7 days r e c o v e r y period  33  T i s s u e glycogen l e v e l s of P. Major muscle from b r o i l e r s r e c e i v i n g d a i l y c o r t i c o s t e r o n e i n j e c t i o n s f o r 8 days and observed f o r a f u r t h e r 7 day recovery p e r i o d  34  1.  INTRODUCTION  The  degree  o f p o s t mortem t e n d e r i z a t i o n  i n chicken  m u s c l e h a s b e e n shown t o be a f f e c t e d  by the ante  physiological  Stressful  conditions  i n muscle chemistry  w h i c h have  preslaughter deleterious The been by  cause changes effect  stress  a d r e n a l m e d u l l a r y hormone,  metabolic muscle glucose  effect  and l i v e r levels.  (deFremery,  tissues  1966).  This  resulting  If  tends  Should  slaughter  to deplete  the r e l e a s e  important  occur  of blood  the at  tissue  this  point,  t o a minimum e x t e n t a c o n d i t i o n known t o  tenderization.  or u t i l i z a t i o n of epinephrine could tissue  g l y c o g e n o l y s i s w o u l d be  and p o s t mortem g l y c o l y s i s w o u l d p r o c e e d  •normal o r e l e v a t e d tating  of r i g o r ,  i n p o s t mortem  be m i n i m i z e d p r e s l a u g h t e r , decreased  caused  tissue,  I t s most  i n elevation  action  i n a r a p i d onset  cause a decrease  the s t i m u l a t i o n  i n the muscle  mortem g l y c o l y s i s w o u l d p r o c e e d  resulting  extent,  reducing ultimate  to a  pH and f a c i l i -  tenderization. . The  the  1963,  e p i n e p h r i n e , has  i s the promotion of g l y c o g e n o l y s i s i n  g l y c o g e n and A T P .  post  between  and t h e c h e m i c a l r e s p o n s e 1968),  a  o n meat q u a l i t y o n p r o c e s s i n g .  i m p l i c a t e d as a m e d i a t o r  (Aberle,  of  s t a t e of the c h i c k e n .  mortem  metabolic  effect  hormones o f t h e a d r e n a l  o f e p i n e p h r i n e i s m o d i f i e d by  cortex,  (Zimmerman,  1972),  (Brown, 1 9 5 9 ) . g l u c o s e and  The  glucocorticoids  act  t i s s u e glycogen l e v e l s .  An  glucocorticoid deposits  hormones has  i n the  liver  and  (Greenman e t a l . 1 9 6 1 ) . ticoid  i n j e c t i o n on  This  However, t h e  was  has  not  liver  modification  of  t i s s u e s and the  blood  of glycogen  glucose,  e f f e c t of  been  conducted  o f g l u c o c o r t i c o i d i n j e c t i o n on m u s c l e and  blood  m u s c l e g l y c o g e n and  research  overdose  b e e n shown t o p r o d u c e  to elevate  p o s t mortem t e n d e r i z a t i o n  to m a i n t a i n  glucocor-  i t s r e l a t i o n to elucidated.  to determine the  carbohydrate metabolism  t o o b s e r v e any  s t r e s s r e a c t i o n by  the  inhibition  effect of or  glucocorticoids.  REVIEW OF LITERATURE  In 1936 Seyle proposed the General A d a p t a t i o n Syndrome (GAS)  as a theory on s t r e s s r e s i s t a n c e and s u s c e p t i b i l i t y i n  animals.  Since t h i s time much r e s e a r c h has been  conducted  to e l u c i d a t e a p h y s i o l o g i c a l and b i o c h e m i c a l b a s i s f o r the r e a c t i o n to, c o n t r o l o f and a d a p t a t i o n t o s t r e s s .  Von Faber  (1964) d e f i n e d the g e n e r a l a d a p t a t i o n syndrome as "the sum o f a l l n o n - s p e c i f i c responses  to s y s t e m a t i c s t r e s s " and i n d i c a t e d  that i t evolves i n three d i s t i n c t phases: a)  Alarm  Reaction When the body i s exposed to damaging s t r e s s o r s , a  s e r i e s o f p h y s i o l o g i c a l a l t e r a t i o n s occur which r e p r e s e n t e i t h e r shock o r attempts o f the organisms t o defend a g a i n s t shock.  itself  The p h y s i o l o g i c a l m a n i f e s t a t i o n o f shock i s  the r e l e a s e of epinephrine from the a d r e n a l medulla  which  mediates the breakdown o f l i v e r and muscle glycogen  to glucose  and  lactic acid.  counter  The p r i n c i p l e a l t e r a t i o n s t h a t occur d u r i n g  shock are, i n c r e a s e d r e l e a s e o f a d r e n o c o r t i c o t r o p i c  hormone (ACTH) from the a n t e r i o r p i t u i t a r y and c o n s e q u e n t i a l release of c o r t i c a l  steroids,  s t e r o i d s a f f e c t carbohydrate,  ( B r i s k e y , 1966).  f a t and p r o t e i n metabolism  along w i t h water and e l e c t r o l y t e balance b)  Resistance or Adaptation  Cortical  (Tepperman, 1968).  Stage  Adjustments made i n counter  shock g a i n supremacy and  the  capacity  internal  o f the organism  environment  (Sayers,  to  be  of  agents  early  as  normal  and  Further this.  either  extracts  experimentation  in increasing  the s u r v i v a l  could  replace-  restore  Anoxia  t o be o f  that  among  Thorn  rats  exposed  cortical low  first the  (1948). i f stress  showed t h a t  i s frequent there i s a  i n the requirement of a d r e n a l e c t o m i z e d  s t e r o i d s d u r i n g the i n i t i a l  atmospheric  steroid  follow  L a n g l e y e t a l . (1942),  marked i n c r e a s e  therapeutic  t o low b a r o m e t r i c p r e s s u r e .  o b s e r v a t i o n s were c o n f i r m e d by K o t t k e  prolonged.  (1945)  stresses  an e x t r a c t o f b e e f a d r e n a l s r e d u c e d  of intact  period  of  pressure but d e s p i t e continued  requirement  no  to a hot e n v i r o n -  i s unique  extracts.  rats.  not confirm  found  t h e r e s i s t a n c e o f man 1945).  that  of i r r a d i a t e d  e x t r a c t s were a l s o  A d a p t a t i o n t o s t r e s s may  the  observed  variety  (1947) r e p o r t e d t h a t a d r e n a l  ( S t r a u b e , 1949)  of adrenal c o r t i c a l  mortality  to  He  of a  r e g a r d t o t h e number o f c o n f i r m e d r e p o r t s o f t h e  demonstrated  for  increases  found a d r e n a l e c t o m i z e d animals  completely or p a r t i a l l y  increased  (Moriera e t a l . ,  benefit  or  functions  t o t h e damaging e f f e c t s  Ellinger  Adrenal c o r t i c a l  benefit  Swingle  e n v i r o n m e n t a l changes.  resistance.  cortical  His  1944,  extremely s e n s i t i v e  ment t h e r a p y w o u l d  in  normality of bodily  the  1950). As  ment  and  to maintain constancy bf  r e t u r n s to pre-exposure  animals  exposure exposure  levels.  c.  Stage of  Exhaustion  In t h i s similar  stage  to those  ensues u n l e s s  acquired  i n the  alarm  Figure  adrenal  1 o u t l i n e s the  complex  forms an  the response  to s t r e s s .  stimuli  the  of the  activity.  or  important The  then stimulates  hypothalamic-pituitary-  neuro-endocrine chain  c o n c e p t has an  area  interbrain,  These n e u r a l  lying  are  a t the  v i a general  (ACTH).  This  circulation  and  steroid  floor  to a  is  an  to secrete  which  the  hormone  i n t u r n reaches the  adrenal  c a u s e s an  increase  the  immediate r e s p o n s e o f  t o combat t h e o f f e n d i n g  t o accommodate o r a c c l i m a t e m e d i a t e d by  immediate  epinephrine  from m e d u l l a r y  provide  for increased  in  the  hormones.  proportions,  effort  the  neural-  While t h i s c h a i n of events r e q u i r e s time to detectable  in  cardiovascular  converted  the. a n t e r i o r p i t u i t a r y  in  neural  which i s important  a p p e t i t e and  stimuli  been t h a t  c o r t i c o t r o p i n - r e l e a s i n g f a c t o r --  adrenocorticotropin  secretion of  b i r d s the  temperature,  humoral f a c t o r —  gland  death  b a s i c p h y s i o l o g i c a l response  hypothalamus,  diencephalon  r e g u l a t i o n of  r e a c t i o n a p p e a r and  Symptoms  STRESS  I n mammals and  reach  is lost.  s t r e s s i s withdrawn.  PHYSIOLOGICAL RESPONSE TO  to s t r e s s .  adaptation  to i t .  body  environment r a t h e r The  responses  secretions of epinephrine t i s s u e and  r e l e a s e of  the  reach  or  than  are nor-  a r e mechanisms w h i c h  energy.  I n mammals  the  6.  STRESS  ^ HYPOTHALAMUS  STIMULI  NERVE STIMULI  ADRENAL CORTEX  ADRENAL MEDULLA  CHANGES I N CORTEX 1) i n c r e a s e d  weight  2) c h o l e s t e r o l d e p l e t i o n 3) s u d a n o p h i l i a ^EPINEPHRINE  CORTICOIDS ADAPTATION REACTION 1) l y m p h a t i c 2) b l o o d c e l l  FIGHT OR FLIGHT MECHANISM  .  involution  4) g a s t r o - i n t e s t i n a l  sugar  2) v a s o c o n s t r i c t i o n  count  3) c h a n g e s i n b l o o d ions cholesterol nitrogenous sugar  1) i n c r e a s e d b l o o d  chemistry  3) i n c r e a s e d r e s p i r a t o r y 4) i n c r e a s e d , m u s c l e  products  5) i n c r e a s e d n e r v e  rate  tone sensitivity  ulceration  5) a n t i - i n f l a m m a t o r y a c t i o n 6) a n t i b o d y  activity  A f t e r H.S.. S i e g e l (1971) A d r e n a l , S t r e s s a n d t h e E n v i r o n m e n t . World's P o u l t r y S c i . J . 27(4) 327. FIGURE 1.  B a s i c Response t o S t r e s s  release a  o f ACTH f r o m t h e  feedback  system.  particularly elevated, process  the  that  activity  (Frankel,  When t h e  leads to  secretion  adrenal cortex apart  to  Tissues  to  processes are  a l o s s of  shrink  t h r i v e on  to  are  and  they are  required  (Brown, 1 9 5 9 ) . homeostatic  of  and  control,  to  the for  cortical  accompaniment o f  of  organisms to the  the  hormones a r e  an  liver  parallel  which  only  the stressors stress,  accompanies  s t r e s s may increase  be in  adrenal cortex  is  at the  an  stress, certain alterations  c h a r a c t e r i s t i c s of  Evans  the  hormones.  metabolic pattern also  against  activity  the  cortical  doubt t h a t  protect  cellular  responsible of  stress  i n h i g h e r amounts d u r i n g  Since hyperactivity invariable  during  a p p e a r s t o be  T h e r e i s no  necessary  Increased  utilization  s u g a r and  some measure  substance throughout  e n l a r g e and  adjustments of  least partially  accelerated  living  stress.  steroids  rate  maintains  the  1971).  i n weight except f o r adrenal  cortical that  of birds  is  deletes  (Siegel,  from p i t u i t a r y s t i m u l a t i o n  t i s s u e which continues tissue  concentration,  c i r c u l a t i n g blood  o f ACTH,  by  1970)'.  thus l e a d  body.  i n the  steroid  c e n t r a l nervous system i n h i b i t s or  Catabolic and  pituitary is.controlled  cortical  glucocorticoids,  However, t h e of  anterior  the  action  of  i n v a r i a b l e accompaniment o f  (1935) showed t h a t  the  increase  in  in  cortical  stress. blood  glycogen, which n o r m a l l y o c c u r s a f t e r  stress,  does n o t o c c u r  i n the absence o f a d r e n a l s .  continued  this  study  of  t h e r e appears  stress  tion  and  confirming that d u r i n g the i n i t i a l  of carbohydrate;  store of l i v e r liver  as c o r t i c a l Laiwrie,  i s maintained  i s reduced;  with  normal.  glucocorticoids,  glycogen  The h y p e r g l y c e m i c  express  and reduces  reaction  blood  sugar  o f trauma f a i l s  TO  A diabetic-like  corticoid  state  i s to i n h i b i t  results  glucose  overdose,: r e s u l t i n g  an u n s t r e s s e d o r g a n i s m , glycogen,  these  cortical  cortical  non-traumatized  trauma i n an  actions of the gluco-  (Sayer,  symptoms a p p e a r  by d i m i n i s h i n g 1950).  adrenal c o r t i c a l  Glucohypertrophy  can cause e x c e s s i v e d e p o s i t i o n o f  blood glucose elevation,  and i n s u l i n  levels.  from  from h y p e r c o r t i c a l i s m ;  utilization  u p t a k e by p e r i p h e r a l t i s s u e s  alterations  to occur  HYPERCORTICALISM  s i n c e one o f t h e most c h a r a c t e r i s t i c  glucose  of  below  r a t (Seyle, 1941).  PHYSIOLOGICAL RESPONSE  corticoids  sugar  r e s t o r e s the hyperglycemic  adrenectomized  liver  glucose  themselves.  the absence o f a d r e n a l s , b u t a dose o f a d r e n a l  normal r a t s ,  of  blood  t o g r e a t e r than normal c o n c e n t r a t i o n s ,  e x t r a c t , w h i c h does n o t e l e v a t e b l o o d  in  time  while the  (1966) s t a l e s , t h a t i n t h e a b s e n c e o f a d r e n a l s , m i l d  stress depletes l i v e r  in  blood glucose  rise  steroids,  stage  t o be an i n c r e a s e d r a t e o f u t i l i z a -  glycogen  glycogen  L e w i s e t a l . (1942)  sensitivity,  protein  metabolism  (Tepperman, 1 9 6 8 ) .  i n a s t r e s s e d animal  with  None  elevated  9.  PHYSIOLOGICAL RESPONSE TO Cortical  CORTICOID INJECTION  s t e r o i d s may  glucose u t i l i z a t i o n  and  produce anabolism  d e p e n d i n g upon t h e d o s e o f t h e Von  Faber,  i n experimental  changes i n c l u d i n g  leucocyte  steroid  (1964) s t a t e s t h a t r e p e a t e d  injections gical  p r o m o t e as w e l l as  count,  animals  lymphatic  adrenal weight.  administration hypertension Green,  produced  possibility removing  observed  the  of a r r e s t i n g  the  steroid  changes i n  acid,  retardation  hyperglycemia  (1959) o b s e r v e d  that  p i g , dog  and  same i n humans and  o r by  nullifying  of  and the  produced monkey.  suggested  hypertensive disease either  adrenal gland  needs.  numerous p h y s i o l o -  involution,  r a t , c a t , guinea  catabolism  tissue  of large' doses of g l u c o c o r t i c o i d s  i n the  (194 8)  Selye  the  cortical  growth, h y p e r l i p e m i a , h y p e r c h o l e s t e r e m i a , decreased  a s w e l l as  and  increased blood c i t r i c  inhibit  the  by  i t s activity.  ADRENAL CORTICAL INDUCED ALTERATIONS IN MUSCLE Research be  related  tenderness  an  (TSH,  shown t h a t a d r e n a l g l a n d  to l a c t i c a c i d  accumulation  ( A d d i s e t a l . , 1965)  p o r c i n e muscle, that  has  (Cassen  imbalance  ACTH, STH)  (Ludvigsen,  of trophic  Ludvigsen  hormones o f t h e  i s r e s p o n s i b l e f o r the  The  data  may  have d e f i c i e n c i e s  Exudative  t h a t muscular animals i n adrenal  steroid  in  suggested  pituitary  association (PSE)  may  1957),  p o s t mortem e v e n t s  e t a l . , 1965).  muscularity with Pale, Soft, indicate  and  function  of  muscle c o n d i t i o n .  w i t h PSE  musculature  production i n  10.  comparison with  l e s s muscular animals w i t h q u a l i t a t i v e l y normal  muscles.  further  lated not  After  that  of  glucocorticoids  sufficient  function. levels their  urine  M a r p l e and  Cassen,  1 9 7 1 ) , and  glucocorticoids  that  an  by  white muscle f i b r e s .  lactate after  Creatine  Phosphate  muscle b i o p s y established larger  (CP)  samples  that  f i b r e s and  m u s c l e m e t a b o l i s m due  f i b r e s and  larger  increased  s u p p o r t e d by  (Lister,  1970).  most l i k e l y  t o p r o d u c e ATP  by  of of  i n plasma  consumption  Marple  a greater  of  accumulation  1 9 7 0 ) , and  a more r a p i d  to  potential  Adenosine Triphosphate  (Lister,  size  reports  (ATP)  of in  e t a l . , (1973a)  muscle from s t r e s s - s u s c e p t i b l e  similar capacity  1969),  s u c h a n i m a l s were shown  f i b r e / u n i t mass w i t h a l o w e r c a p a c i t y but  Cassen,  a more r a p i d  samples  and  (Topel,  of  suggestion of  and  Further  aberrant muscle metabolism  Muscles of  1969)  pigs.  of  swine h a v i n g  (Topel,  metabolic  1973a)  white muscle  i n muscle b i o p s y  stress  ( M a r p l e and  glycolysis i s also  stress-susceptible  increased  production  for aerobic  The  in  i n s t r e s s - r e s i s t a n t animals.  to  swine.  low  catecholamines excreted  increased  which lead  a h i g h percentage of  a n i m a l s were  were shown t o have  stress-susceptible  t o have a l o w e r c a p a c i t y  anaerobic  and  utilization  stress-susceptible  for  pigs  (1973b) o b s e r v e d an  Cortisol  indicated  to  postu-  to ensure adequate c i r c u l a t o r y  steroids  compared  c l e a r a n c e of  Weiss,  quantity  adrenal  (1957)  in stress-susceptible  Stress-susceptible  of  evidence  i n v e s t i g a t i o n Ludvigsen  swine  mass o f w h i t e  for oxidative oxidative  have anaerobic  metabolism  phosphorylation.  Eikelenboom and Van Den Bergh,  (1971) suggested  t h a t muscle  g l y c o g e n o l y s i s w i l l be enhanced i n s t r e s s - s u s c e p t i b l e swine i n an e f f o r t to produce ATP d u r i n g p e r i o d s o f i n c r e a s e d energy  demand such as the s t r e s s response.  s t r e s s response by Topel  During such a  the r a p i d u t i l i z a t i o n o f e p i n e p h r i n e  noted  (1972) would a l s o s t i m u l a t e the r a p i d muscle  g l y c o g e n o l y s i s and g l y c o l y s i s noted by S a i r  (1970).  Similarly  l i v e r glycogen would be converted t o blood glucose by the a c t i o n o f e p i n e p h r i n e thus producing the i n c r e a s e d blood . glucose c h a r a c t e r i s t i c o f s t r e s s - s u s c e p t i b l e p i g s . a t e l y C o r t i s o l would be expected neogenesis  Ultim-  to stimulate l i v e r gluco-  t o convert h i g h l e v e l s o f l a c t a t e produced  by  anaerobic muscle f i b r e mass t o g l u c l o s e and t o muscle and l i v e r glycogen, with the high r a t e of C o r t i s o l metabolism in stress-susceptible pigs; (Marple and Cassen,  t h i s c o n v e r s i o n i s slow,  1973a).  Russian workers  (Nestorov e t a l . ,  1972) found an  i n c r e a s e i n ACTH and 17-hydroxy c o r t i c o s t e r o i d s the b l o o d o f c a l v e s d u r i n g t r a n s p o r t .  (17-HCS) i n  The l e v e l s o f 17-HCS  d u r i n g an 8-hour r e s t p e r i o d f o l l o w i n g t r a n s p o r a t i o n was maint a i n e d a t h i g h e r than normal v a l u e s , while the content o f glycogen i n the l i v e r and muscle a t t a i n e d and surpassed the i n i t i a l quantity.  Analogous data was o b t a i n e d f o r p i g s .  Effects of stress  (simulated by an e p i n e p h r i n e  t i o n ) as l o s s o f muscle e x t e n s i b i l i t y ,  injec-  low pH and ATP i n  muscle,  i n rabbits  overcome  by B e n d a l l  by an i n j e c t i o n o f c o r t i s o n e  epinephrine cattle  was d e m o n s t r a t e d  injection.  T h i s was a l s o  (Hendrik et a l . , 1957).  shown t o o c c u r  However, P u r c h a s ,  may be d e t r i m e n t a l  quality,  i n high  ADRENAL CORTICAL The of  the b i r d  1958), in  and d e a t h  adrenocortical  after  for continued s u r v i v a l  extracts  or pure c r y s t a l l i n e cortical  in control,  adaptive  hormones w h i c h t e n d t o c o u n t e r a c t  the  glands of b i r d s ,  More s p e c i f i c a l l y ,  are countered  (Brown,  present,  concluded  that  release  s h o c k p r o d u c e d by  the shock  symptoms i n d u c e d  release of adaptive  hormones  from  cortex.  to i d e n t i f i c a t i o n of c o r t i c o s t e r o n e ,  tentative  1958).  a s w i t h mammals,  I n c u b a t i o n o f White Leghorn c o c k e r e l lead  typical  by t h e r e l e a s e o f ACTH f r o m t h e  and t h e c o n s e q u e n t i a l  adrenal  prevented  Brown ( 1 9 5 8 ) ,  the adrenal  pituitary  (Brown,  corticosteroids.  steroids  normally  stress  is rapid,  n o r m a l c o n d i t i o n by t h e a d m i n i s t r a t i o n o f  From t h e s e e x p e r i m e n t s ,  by  states  t o p o s t mortem m u s c l e  adrenalectomy  s h o c k symptoms w h i c h a p p e a r e d  stress.  (1970)  A d r e n a l e c t o m i z e d b i r d s c a n be m a i n t a i n e d  Administration of adrenal  severe  i n beef  doses.  gland i s necessary  1954).  an a p p a r e n t l y  the  CONTROL OF AVIAN METABOLISM  adrenal  (Leroy,  t o be  2 0 hours before  that glucocorticoids i f injected  (1962)  1960).  cortex  a l d o s t e r o n e and  i d e n t i f i c a t i o n o f s m a l l amount o f (de R o o s ,  adrenal  hydrocortisone  A n a l y s i s o f f o w l p l a s m a by P h i l l i p s  e t a l . (1957) i n d i c a t e d the primary adrenal  s t e r o i d to  c o r t i c o s t e r o n e followed by s m a l l e r amounts of c o r t i s o n e and  aldosterone.  In v i v o and  be  hydrocortisone,  i n vitro investiga-  t i o n s have shown t h a t c o r t i c o s t e r o n e i s the p r i n c i p l e f r e e cortical Nagra,  s t e r o i d i n chickens i960).  (Sandor, 1963;  Corticosterone  was  a l s o e s t a b l i s h e d to be  p r i n c i p l e f r e e c o r t i c o s t e r o i d i n turkey and  the adrenal  e f f l u e n t blood  de Roos, I960,;  plasma  of the capon  (Brown,  the  1961)  ( P h i l l i p s et a l .  1957) . Greenman sterone  (1961), found h y d r o c o r t i s o n e  to be of high g l u c o c o r t i c o i d a c t i v i t y  demonstrated t h a t blood  glucose and  Cortisone,  a c t i v i t y i n the b i r d .  i n chickens  and  t i s s u e glycogen showed  marked i n c r e a s e s w i t h d a i l y h y d r o c o r t i s o n e injections.  and c o r t i c o -  or  corticosterone  possesses l i t t l e g l u c o c o r t i c o i d  Stamler  (1952) r e p o r t e d  a failure  to  induce hyperglycemia i n the c h i c k w i t h c o r t i s o n e at doses which were e f f e c t i v e w i t h  hydrocortisone.  Brown (1961) suggested t h a t i t i s v a l i d to use corticosterone concentrations of s t r e s s c o n d i t i o n s s t r e s s may  plasma  as a measure of the s e v e r i t y  i n the b i r d s .  Degree of r e s i s t a n c e to  be measured i n a number of ways, i n c l u d i n g  m o r t a l i t y , maintenance o f normal c o n c e n t r a t i o n s  of  metabolites  i n the body f l u i d s and maintenance or r a t e of r e s t o r a t i o n of individual functions. to cause an i n c r e a s e  Surgically inflicted  s t r e s s was  i n corticosterone concentration  in  shown  14 .  a d r e n a l venous blood o f c o c k e r e l s 1967), pheasants  (Nagra,' 1963)  (Nagra, 1963),  and ducks  (Frankel,  (Macchi, 1967).  Cold s t r e s s i n c r e a s e d c o r t i c o s t e r o n e i n c o c k e r e l s 1963), ducks  ( B o i s i n , 1967)  and t u r k e y s (Brown, 1961).  However, continued exposure to h i g h ambient for  (Nagra,  temperatures  4 weeks had l i t t l e e f f e c t on c o r t i c o s t e r o n e c o n c e n t r a -  tions.  Chickens s u b j e c t to s t r e s s imposed  (Garren e t a l . 1952), c o l d  (Wolford, 1962), heat  1961a,b,c) l i m i t a t i o n o f feed and water handling 1960)  by muscle  (Conner,  ( B e l l o f f e t a l . 1963), i m m o b i l i z a t i o n  and crowding  ( S i e g e l , 1960)  (Hill, 1959),  (Newcomer,  have been shown to e l i c i t  a s t r e s s response and an a d r e n a l c o r t i c a l i n h y p e r a c t i v i t y of a d r e n a l s .  fatigue  reaction  resulting  I f s t r e s s i s not r e l i e v e d  and  h y p e r a c t i v i t y o f a d r e n a l t i s s u e i s prolonged, death w i l l result.  Terminal symptoms are muscular weakness, decreased  blood g l u c o s e , decreased body temperature and c o n v u l s i o n s (Brown, 1959).  Brown (1973) demonstrated t h a t t u r k e y s which  can adapt to s t r e s s c o n d i t i o n s without i n i t i a t i n g the g e n e r a l a d a p t a t i o n syndrome o r which a t l e a s t respond by a low r e l e a s e o f a d a p t i v e hormones grow f a s t e r , c o n v e r t feed more efficiently,  reproduce more e f f i c i e n t l y  and are more  r e s i s t a n t to a wide v a r i e t y of s t r e s s c o n d i t i o n s .  MATERIALS AND METHODS P o u l t r y Source The c h i c k e n s used i n t h i s study were commercial b r o i l e r s o b t a i n e d from a l o c a l p r o c e s s i n g p l a n t and maintained i n range houses on the U.B.C. P o u l t r y Farm.  The b i r d s were  6 o r 8 weeks o f age when o b t a i n e d . The b i r d s were s u b j e c t e d t o experimental treatments e i t h e r w i t h i n 48 hours o f t r a n s p o r t o r a f t e r 5-7 days a c c l i m a t i z a t i o n depending upon the requirements o f the experiment. Slaughter Procedure B i r d s were p l a c e d i n a metal f u n n e l , which  restricted  wing and l e g movement, and were exsanguinated by an o u t s i d e neck c u t and allowed t o b l e e d f r e e l y . t h i s time. In j ect ion  Carcasses were then p l a c e d i n i c e t o c h i l l . Pro c edure  In llB,  Blood was c o l l e c t e d a t  an i n i t i a l  experiment c o r t i c o s t e r o n e  (4-pregnen-  21 D i o l - 3 , 20-dione*) was d i s s o l v e d i n p h y s i o l o g i c a l  s a l i n e and i n d i s t i l l e d water t o determine optimum  solubility  and t o determine i f any s i g n i f i c a n t change i n 'muscle shear value was caused by e i t h e r d i l u e n t .  In subsequent t e s t s 2 ml o f  a 5 mg/ml c o r t i c o s t e r o n e i n d i s t i l l e d water s l u r r y was B i r d s were i n j e c t e d i n t r a m u s c u l a r l y i n the majorum  * Schwartz/Mann, Orangeburg,  N.Y.  injected.  peronaeus  longus.  When repeated  i n j e c t i o n s were g i v e n ,  i n j e c t i o n s were given i n a l t e r n a t e l e g s .  consecutive  Some c o n t r o l b i r d s  were i n j e c t e d with 2 ml d i s t i l l e d water t o compare with uninjected controls.  A l l subsequent c o n t r o l b i r d s were  uninjected. Tenderness 1)  I n t a c t muscle For shear t e s t i n g carcasses'.• were c h i l l e d  i n i c e s l u s h p l a c e d i n a c o l d room a t 4°C.  overnight  The P e c t o r a l i s  Major was then e x c i s e d and p l a c e d between metal p l a t e s f o r cooking.  The p l a t e s h e l d the muscle a t a c o n s t a n t  thickness  of 6 mm.  The muscles were then immersed i n b o i l i n g water  f o r 10 minutes, c o o l e d i n c o l d water f o r 5 minutes and r e l e a s e d from between the p l a t e s .  The cooked muscle was then  cut i n t o s t r i p s 4 cm wide a t random o r i e n t a t i o n t o the muscle fibre.  Shear was measured on an Allo-Kramer  shear  with a 250 pound p r o v i n g r i n g and a s i n g l e blade attachment. 2)  press  shear  Range was s e t a t 20.  E x c i s e d muscle Muscle was e x c i s e d from the b i r d immediately  death,  t o allow s h o r t e n i n g without  skeletal  after  restrictions,  and c h i l l e d o v e r n i g h t i n an i c e s l u r r y p l a c e d i n a c o l d room a t 4°C.  Cooking procedure and shear measurement was as  p r e v i o u s l y d e s c r i b e d f o r i n t a c t muscle.  Chemical 1)  Tests  Blood  glucose  B l o o d was in a test  tube  method tion  i n blood  exsanguination, o x a l a t e and  prevent  (Washko, 1 9 6 1 ) , an e n z y m a t i c  Tissue  c o a g u l a t i o n and  samples.  Glucose  method  was  glucostat*  f o r the  determina-  solution  d e n s i t y was  m e a s u r e d a t 420  nm  with  a  glycogen  immediately  and  after  liver death.  t i s s u e s were removed  from  the  Samples were p l a c e d i n an  bird  alkaline  i n a b o i l i n g w a t e r b a t h , w i t h i n 20 m i n u t e s o f  death,  digest. Liver  and  m u s c l e g l y c o g e n were d e t e r m i n e d  method o f Roe  and  Dailey  carbohydrate  (1966),  Beckman DB Blood  an  anthrone  by  the  method f o r  determination.  O p t i c a l d e n s i t y was  r e c o r d e d a t 62 0 nm  with  a  spectrophotometer. cholesterol  Blood  *  mg  spectrophotometer.  Muscle  3)  25  of glucose.  Beckman DB  to  of  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 the  Optical  2)  activity  time  potassium  These c h e m i c a l s  glycolytic  measured  a t the  c o n t a i n i n g 20 mg  sodium f l u o r i d e . inhibit  collected  c h o l e s t e r o l was  Worthington  determined  B i o c h e m i c a l Corp.,  •9  by  Freehold,  t h e method  N.J.  of  Zlatkis  e t a l . (1953) . Optical  d e n s i t y was  spectrophotometer 4)  a t 560  measured  with  a Beckman  DB  nm.  pH The  and  chilled  with  40 m l  P e c t o r a l i s M a j o r m u s c l e was overnight.  20g o f muscle  excised  tissue  at  was  of water i n a Waring Blender f o r 5 min.  determined with  a Fisher Accumet,  Model  230  pH  death  homogenized pH  Meter.  was  RESULTS AND  DISCUSSION  E f f e c t of I n j e c t i o n Method When an  i n j e c t i o n i s made i n t o a l i v i n g animal, i t  should be made i n such a way a minimum and  t h a t t i s s u e trauma i s kept to  t h a t c e l l u l a r environment i s not d r a s t i c a l l y  altered. A preliminary  study was  c a r r i e d out  to determine  a f a v o u r a b l e d i l u e n t f o r the c o r t i c o s t e r o n e to s o l u b i l i t y  i n i n j e c t i o n s o l u t i o n and  muscle t i s s u e when i n j e c t e d . Greenman  saline.  respect  absorption i n  the  (1961) r e p o r t e d  f a v o u r a b l e r e s u l t s when c o r t i c o s t e r o n e s l u r r y with p h y s i o l o g i c a l  both w i t h  was  Saline  i n j e c t e d as s o l u t i o n was  a used  to prevent osmotic shock i n the muscle t i s s u e which may promoted i f d i s t i l l e d water was Table 1  used as a d i l u e n t .  i n d i c a t e s t h a t t h e r e was  no  i n shear values of p e c t o r a l i s major, was  be  However,  significant difference when  i n j e c t e d as a s l u r r y i n p h y s i o l o g i c a l  corticosterone s a l i n e or  distilled  water TABLE 1.  COMPARISON OF SHEAR VALUES OF P. MAJOR MUSCLE WHEN CORTICOSTERONE WAS DILUTED WITH DISTILLED WATER OR PHYSIOLOGICAL SALINE SOLUTION  Hours*  D i s t i l l e d Water Saline Shear~Value  0 4 8 * **  15.88 14.67 11.10 Hours post i n j e c t i o n P < .05  16.00 14.14 10.52  T Value .521 .511 .592  df 90 83 54  T-calc.** 1.99 1.99 2.02  20, T e s t s were c a r r i e d o u t injected with physiological as  well  that  as  at  received  osmotic  subsequent  of  the  birds,  not  studies  an  d e p e n d e n t on  that  only,  birds  was  of  the  environment.  i n j e c t e d as  water,  that  absorption  a saline  corticosterone  validity  e x p e r i m e n t was  due  to  a simplicity  of  using  control  compare s h e a r  values  b i r d s with those of  birds  injected with  distilled  2 indicates  significant difference  Table  found.  Therefore,  c a r r i e d out  uninjected  to  water.  that  control  no  birds  i n a l l subsequent  was  studies  uninjected. In  these preliminary  in  s h e a r v a l u e was  in  the  muscle  i n d i c a t e only reflect  an  TABLE 2.  Treatment Injected Uninjected * **  water  i n j e c t i o n , on  and  slurry in distilled  t e s t the  uninjected  were  distilled  These r e s u l t s i n d i c a t e  non-significant  was  b i r d s w h i c h were  method. To  of  s a l i n e or  corticosterone.  t h o r o u g h l y mixed  control  time i n t e r v a l s p o s t  s h o c k was  corticosterone In  two  on  u s e d as  i n d u c e d by very  an  the  studies  a significant difference  i n d i c a t i o n of treatment.  s p e c i f i c metabolic  general  alterations  Chemical t e s t s  c h a n g e s and  would  would not  o v e r a l l a l t e r a t i o n i n muscle c h a r a c t e r i s t i c s . COMPARISON OF SHEAR VALUES OF P. MAJOR MUSCLE FROM UNINJECTED BIRDS AND BIRDS INJECTED WITH D I S T I L L E D WATER Shear  Value**  15.88 16.56  T  Value  .455  •  df  90  T-calculated for a two-tailed T-test at P < .05 S h e a r v a l u e s a r e an a v e r a g e o f s e v e r a l values.  T-calc.*  1.99  Short Term S t u d i e s G l u c o c o r t i c o i d s are not r e l e a s e d immediately upon s t r e s s but a c t to i n c r e a s e r e s i s t a n c e by i n d u c i n g a l t e r a t i o n s only a f t e r s t r e s s has  continued  the event t h a t g l u c o c o r t i c o i d s were present  f o r a time.  In  a t the onset  of  s t r e s s , as when i n j e c t e d , they would i n i t i a t e the adaptation  modified  on glycogen  and ATP  reserves.  a l t e r a t i o n s of s t r e s s should be decreased  or  i f g l u c o c o r t i c o i d i n j e c t i o n were used to i n c r e a s e  the r e s i s t a n c e to the s t r e s s of s l a u g h t e r and e f f e c t on meat q u a l i t y should Table  a  3 i n d i c a t e s the e f f e c t of c o r t i c o s t e r o n e  post i n j e c t i o n . noted u n t i l  favourable  result.  i n j e c t i o n s on muscle tenderness at s e v e r a l time No  intervals  s i g n i f i c a n t e f f e c t of the i n j e c t i o n i s  6 hours a t which time the shear v a l u e s  from those of the c o n t r o l . ous  general  syndrome.and thus modify the c a t a b o l i c e f f e c t of  s t r e s s - i n d u c e d epinephrine Metabolic  metabolic  T h i s i s a r e s u l t of the  increase continu-  a b s o r p t i o n of the c o r t i c o s t e r o n e from the i n j e c t e d  muscle.  In an e x c i t e d b i r d , as one  i n j e c t i o n i n the p r e v i o u s utilized  t h a t has r e c e i v e d  an  few hours, the c o r t i c o s t e r o n e i s  to overcome the s t r e s s of h a n d l i n g  and  injection,  and  i s i n c o r p o r a t e d i n t o the body metabolism, i n i t i a t i n g  GAS  to r e s t o r e homeostatic c o n d i t i o n s .  As the s t r e s s  the  subsides  the c o n t i n u i n g a b s o r p t i o n of c o r t i c o s t e r o n e from the i n j e c t e d muscle c r e a t e s an overdose i n the b i r d .  In an  unstressed  TABLE 3.  EFFECT OF CORTICOSTERONE INJECTION ON SHEAR VALUE OF P. MAJOR MUSCLE AT SEVERAL INTERVALS POST INJECTION  Hours*  Shear Value  0 2 4 6 8  12.25 11.66 12.81 16.69 16.27  T Value  df  .588 .575 4.68 3.66  70 70 76 70  T-calc.**  1.99 1.99 1.99 1.99  * Hours post i n j e c t i o n ** P < .05 animal c o r t i c o s t e r o n e overdose i s r e p o r t e d induce at  increased  c e l l u l a r a c t i v i t y and  by L i s t e r  (1970) to  d e p l e t i o n of ATP  which  s l a u g h t e r w i l l u l t i m a t e l y l e a d to a r a p i d onset o f r i g o r  mortis  and post mortem toughening of muscle t i s s u e . R e s u l t s of blood  confirm  glucose t e s t s i n d i c a t e d i n Table 4  t h a t c o r t i c o s t e r o n e has been absorbed and  an e f f e c t a t 2 hours post  injection.  is eliciting  T h i s i s f u r t h e r evidence  t h a t the l a c k of a s i g n i f i c a n t d i f f e r e n c e i n shear values 2 and  4 hours post  corticosterone TABLE 4.  Hours*  0 2 4 6 8 * **  i n j e c t i o n i s not merely due  at  to. a l a g i n  absorption.  EFFECT OF CORTICOSTERONE INJECTION ON BLOOD GLUCOSE LEVELS AT SEVERAL TIME INTERVALS POST INJECTION Blood Glucose mg % 268 325 367 373 337  Hours post i n j e c t i o n P < .05  T Value  2.87 4.67 2.89 3.14  df  10 10 14 14  T-calc.**  2.23 2.23 2.15 2.15  Blood glucose l e v e l s increase hour p o s t i n j e c t i o n p e r i o d .  This  s t e a d i l y over the 8  i s consistent with the  r e s u l t s o f S i e g e l e t a l . (1960) who i n j e c t e d ACTH, a corticosterone potentiator, into chickens, action of corticosterone u t i l i z a t i o n of  and r e f l e c t s t h e  i n i n h i b i t i n g c e l l u l a r u p t a k e and  glucose.  Table 5 i n d i c a t e s a comparison o f shear values  of  P. M a j o r m u s c l e f r o m 6 w e e k - o l d b r o i l e r s a n d 8 w e e k - o l d b r o i l e r s a f t e r b o t h have r e c e i v e d  corticosterone injections.  A c o n s i s t e n t d i f f e r e n c e i n shear value t h a t the shear values  i s observed.  The f a c t  were c o n s i s t e n t l y d i f f e r e n t i n d i c a t e s  t h a t t h e d i f f e r e n c e was due t o an age and w e i g h t e f f e c t r a t h e r t h a n t o an e f f e c t o f t h e d i f f e r e n c e i n t h e p h y s i o l o g i c a l handling  and e f f e c t o f t h e c o r t i c o s t e r o n e  injection.  The  e f f e c t o f age o n t e n d e r n e s s o f p o u l t r y m u s c l e h a s b e e n w e l l e s t a b l i s h e d b y May e t a l . ( 1 9 6 2 ) . increase  i n shear value  It i s likely  that there  with  They f o u n d a c o r r e s p o n d i n g  increased  is little  age o f t h e c h i c k e n .  significant physiological  c h a n g e i n t h e m e t a b o l i s m o f c o r t i c o s t e r o n e w i t h i n t h e two week  period. The  greatest  d i f f e r e n c e i n t r e a t m e n t e f f e c t was  between b i r d s o f d i f f e r e n t s t r e s s l e v e l . subjectively evaluated a person entering handling.  with respect  Stress  level  found was  to the birds reaction to  t h e r a n g e h o u s e , and t h e i r r e s p o n s e t o  The s h e a r v a l u e s  o f t h e P. M a j o r f r o m t h e c o n t r o l  24 . TABLE 5.  COMPARISON OF SHEAR VALUES OF P. MAJOR MUSCLE FROM 6 AND 8 WEEK OLD BIRDS AT SEVERAL TIME INTERVALS POST INJECTION OF CORTICOSTERONE. SHEAR VALUE  Hours*  6 Weeks o l d  0 2 4 6 8 *  8 weeks o l d  14.93 13.98 13.67 12.61 12.29  Hours post  16.16 15.56 15.72 15.68 15.36  injection  b i r d s confirmed  the s u b j e c t i v e o b s e r v a t i o n s .  Table 6 i n d i c a t e s  an i n c r e a s e i n shear v a l u e o f the P. Major muscle from the c o n t r o l b i r d s w i t h a corresponding i n c r e a s e i n the l e v e l o f stress. Table 7 i n d i c a t e s treatment birds.  r e s u l t s o f very p l a c i d  The b i r d s d i d not bunch up i n a corner o f the house,  d i d not f l a p about when handled and showed no s i g n o f s t r e s s on i n j e c t i o n .  Shear v a l u e s i n c r e a s e d immediately  and remained e l e v a t e d throughout period.  the 8 hour p o s t  injection  I t may be e x p l a i n e d t h a t , as no s t r e s s was  i n the b i r d s , c o r t i c o s t e r o n e absorbed i n j e c t i o n produced  elicited  even a t 2 hours post  a m e t a b o l i c overdose  t e n s i o n and a p o s t mortem toughening TABLE 6.  on i n j e c t i o n  r e s u l t i n g i n hyper-  o f muscle t i s s u e .  COMPARISON OF SHEAR VALUES OF P. MAJOR MUSCLE WITH SUBJECTIVE EVALUATION OF STRESS LEVEL OF THE BIRD. Stress Level Placid Mildly excitable Extremely e x c i t a b l e  Shear Value 10.88 16.16 16.20  25. TABLE 7.  SHEAR VALUES OF P. MAJOR MUSCLE FROM PLACID BIRDS OVER AN 8 HOUR PERIOD POST INJECTION OF CORTICOSTERONE .  Hours*  Shear Value  T Value  df  0 2 4 6 8  10.88 13.43 14 .58 13.06 11.86  3.63 6 .20 3.00 1.60  158 147 155 167  * **  T-calc.**  1.96 1.96 1.96 1.96  Hours post i n j e c t i o n P < .05 Table 8 i n d i c a t e s shear v a l u e s o f P. Major muscle  from m i l d l y e x c i t a b l e b i r d s , f o r an 8 hour p e r i o d injection of corticosterone. observed i n shear v a l u e .  post  No s i g n i f i c a n t changes were  B i r d s were not s t r e s s e d t o t h e  e x t e n t t h a t the c o r t i c o s t e r o n e would be immediately and r a p i d l y u t i l i z e d a f f e c t i n g a t e n d e r i z a t i o n o f muscle t i s s u e when compared t o t h e c o n t r o l , nor were they so p l a c i d t h a t c o r t i c o s terone was never r e q u i r e d .  The i n j e c t e d c o r t i c o s t e r o n e  would  e n t e r i n t o the metabolism, be u t i l i z e d and never c o n s t i t u t e an overdose, nor produce d e l e t e r i o u s e f f e c t s on muscle q u a l i t y . TABLE 8.  SHEAR VALUES OF P. MAJOR MUSCLE FROM MILDLY EXCITABLE BIRDS OVER AN 8 HOUR PERIOD POST INJECTION OF CORTICOSTERONE.  Hours*  Shear Value  0 2 4 6 8  16.16 15.56 15.72 15.68 15.36  * **  Hours post i n j e c t i o n P < .05  T Value  df  1.240 1.23 1.29 1.57  79 67 70 68  T-calc.**  1.99 1.99 1.99 1.99  26,  In obtained was  comparison,  from m i l d l y  excitable  a significant difference  post  injection.  corticosterone excitable in  Variability  due  birds.  shear  In t h i s case  i n the  excitable  birds  T h e r e was  4  6 8 * **  stress  homeostatic  0 2  a  corti-  a s i g n i f i c a n t decrease i n  i n j e c t i o n appears  *  error  i n j e c t i o n s , marked c h a n g e s i n s h e a r v a l u e s o c c u r r e d ,  injection.  Hours  and  received  post  TABLE 9.  by  mildly  treated  hydrate  hours  i n s h e a r v a l u e s as a f f e c t e d  shear v a l u e s f o r c o r t i c o s t e r o n e  to  8  judgement.  shown i n T a b l e 9.  relieved  values  there  i n s h e a r v a l u e a t 6 and  to p h y s i o l o g i c a l v a r i a b i l i t y  When e x t r e m e l y costerone  indicates  i n j e c t i o n i s the g r e a t e s t  birds  subjective  as  Table 3 also  Corticosterone symptoms and conditions  initiated  of c e l l u l a r  birds  a t a l l times t o have  a more r a p i d activity  and  return carbo-  metabolism.. SHEAR VALUES OF P. MAJOR MUSCLE FROM EXTREMELY EXCITABLE BIRDS OVER AN 8 HOUR PERIOD POST INJECTION OF CORTICOSTERONE.  Shear V a l u e  T Value  16.20 13.69 14.45 14.77 10.77  2.41 1.98 2.08 5.91  df  139 174 174 145  T-calc.**  1.96 1.96 1.96 1.96  Hours p o s t i n j e c t i o n P < .05 When t r e a t m e n t  e f f e c t was  were o b t a i n e d a t t h e same t i m e  t o be  compared a l l b i r d s  as t h i s s t u d y was  conducted  over  an  range  18 month p e r i o d  house v a r i e d .  ranged  from  4 5°F  comparisons  and  The  m a i n v a r i a b l e was  t o 85°F.  beyond  environmental  Due  to this  c o n d i t i o n s i n the temperature  variability  t h o s e t h a t a r e made may  n o t be  which  further valid.  Long Term S t u d i e s On  a l o n g term b a s i s ,  carbohydrate metabolism, and  glycogen  1)  5 Day  deposition  particularly i n the  corticosterone  increase blood glucose l e v e l s the r e s u l t s  corticosterone To over  of Siegel  to i n h i b i t  indicate  t h e 5 day  and  period  intact  E x c i s e d muscle treatment is  on  i n j e c t i o n s were f o u n d (Figure  2).  (1962) and cellular  uptake  conducted.  Major  muscles  o r m o d i f i e d by  hours  24  apparent  i n the i n t a c t  s h e a r v a l u e s may  skeletal  a dramatic decrease  recorded  after  be  the f i r s t tissue.  the r e s u l t  ATP  level,  by  a  study  values f o r both i n Figure  effect  of  attachments.  i n shear v a l u e  injection.  In was  T h i s i s not  T h i s marked d e c r e a s e of the m o d i f i c a t i o n  the i n j e c t e d  the  muscle s h o r t e n i n g  i n d u c e d p o s t mortem c h a n g e s i n m u s c l e t i s s u e , and  metabolism  are d i s p l a y e d  shear v a l u e s p r o b a b l y because  of  of glucose.  showed a much more p r o n o u n c e d  not c o n t r o l l e d  the a b i l i t y  injection,  Shear  to  This i s consistent  reflects  of corticosterone  P.  the e x c i s e d t i s s u e  pH  mobilization  tissue.  g r o s s changes i n muscle  o f m u s c l e t e n d e r n e s s was excised  glucose  study Daily  with  hypercorticalism modifies  as  corticosterone.  of  in stress  decreased Following  3.  FIGURE 3.  Shear v a l u e s  of i n t a c t  muscle of b r o i l e r s injections  and  excised  receiving daily  o v e r a 5 day  period.  P.  Major  corticosterone  this the  initial  24 h o u r p e r i o d ,  homeostatic  s i g n i f i c a n c e of the c o r t i c o s t e r o n e  proceeding t o the c o n t r o l l e v e l . the  modifying  tuates  shear  The i n t a c t m u s c l e  values with  i n f l u e n c e of s k e l e t a l r e s t r i c t i o n merely  fluc-  during  the  study A 15 day s t u d y was  effect  of d a i l y  c a r r i e d out to confirm  corticosterone  i n j e c t i o n found  study and t o o b s e r v e changes d u r i n g following  cessation  of daily  is  seen i n e x c i s e d  by  a return  B i r d s were i n j e c t e d  f o r a f u r t h e r 7 day  A marked d e c r e a s e  muscle,  after i n i t i a l  t o more n o r m a l s h e a r v a l u e s  e f f e c t was a l s o s e e n i n F i g u r e again  3.  injection,  intact  Shear v a l u e s  of i n j e c t i o n .  followed This  for intact  A l a g i n homeostatic  as an i n c r e a s e increase  tissue.  This  i n s t r e s s symptoms  i n shear values i s i n accord  levels of corticosterone  adjust-  a f t e r d a y 8, and an  ultimate  i n both e x c i s e d  and  with the f i n d i n g s of  Zimmerman e t a l . , (1972a, 1972b) who logical  value  (Figure 4 ) .  ment p r o d u c e s a t e m p o r a r y h y p o c o r t i c a l i s m  significant  i n shear  f l u c t u a t e d n o n - s i g n i f i c a n t l y about the c o n t r o l  l i n e on c e s s a t i o n  detected  period  period. Shear t e s t i n g ;  tissue  the  i n t h e 5 day  the recovery  injection.  f o r 8 d a y s and were o b s e r v e d  recovery  stress  decrease  period.  15 Day  daily  effect,  n o n - s i g n i f i c a n t l y about t h e c o n t r o l v a l u e  5 day 2)  adjustments  reported  that  physio-  suppressed non-stress,  induced p i t u i t a r y - a d r e n a l f u n c t i o n  in rats.  and  A f t e r the  FIGURE 4.  S h e a r v a l u e s o f e x c i s e d and i n t a c t  P. M a j o r  muscle of b r o i l e r s  corticosterone  injections  receiving daily  f o r 8 d a y s and o b s e r v e d f o r a f u r t h e r  7 days, without i n j e c t i o n .  o— •  excised intact  i n i t i a l r e a c t i o n to c e s s a t i o n o f c o r t i c o s t e r o n e i n j e c t i o n s , homeostasis  ensures  t h a t metabolism r e t u r n s to a normal  c o n d i t i o n and shear v a l u e s proceed  to t h a t o f the c o n t r o l  birds. A l t e r a t i o n s o f shear v a l u e s as a r e s u l t o f c o r t i costerone i n j e c t i o n s a r e not as d e f i n i t i v e as are the r e s u l t s o f s p e c i f i c chemical t e s t s .  Shear v a l u e r e f l e c t s the sum o f  a l l changes i n muscle metabolism induced by c o r t i c o s t e r o n e and not the r e s u l t o f any one d e t e c t a b l e change. Conclusions made on the b a s i s o f a l t e r a t i o n s i n shear value a r e o n l y p o s s i b l e w i t h r e f e r e n c e t o r e s u l t s o f s p e c i f i c chemical Chemical  tests. tests:  Blood g l u c o s e .  F i g u r e 5 i n d i c a t e s blood glucose  l e v e l s d u r i n g the 15 day study: i n j e c t i o n s caused  As p r e v i o u s l y noted,  an e l e v a t i o n i n blood g l u c o s e  levels.  When i n j e c t i o n s were d i s c o n t i n u e d g l u c o s e r a p i d l y as would be expected  i n a hypocortical condition.  d e p r i v a t i o n o f glucose d u r i n g c o r t i c a l  steroid  daily  decreased Cellular  injection  produces a marked c e l l u l a r glucose d e f i c i e n c y t h a t i s r a p i d l y c o r r e c t e d on d i s c o n t i n u i n g i n j e c t i o n s , producing a c o n d i t i o n i n the blood.  hypoglycemic  Progress t o a normal glycemic  c o n d i t i o n f o l l o w e d as expected. T i s s u e glycogen.  Figure 6 indicates tissue  l e v e l s d u r i n g t h i s 15 day study.  Glycogen  glycogen  content i n c r e a s e d  w i t h c o r t i c o s t e r o n e i n j e c t i o n as c o r t i c o s t e r o n e acted t o  FIGURE 5..  Blood glucose l e v e l of b r o i l e r s corticosterone injections for  a further  7 day  receiving  f o r 0 d a y s and  recovery  period.  daily observed  3.5.  modify The  the t i s s u e  glycogen  depletion effect  of epinephrine.  gluconeogenic  activity  of corticosterone i s well  docu-  mented. Gluconeogenesis  i s an i n v o l v e d m e t a b o l i c  i n c l u d i n g many i n t e r m e d i a t e s t e p s . cessation of i n j e c t i o n pH. a decrease  to  lactic  i n u l t i m a t e pH i s e x p e c t e d .  At death  acid, this  Blood  acts to metabolize  cholesterol.  Cholesterol  TABLE 10.  Values  this  anaerobic, glycogen  T a b l e 10  increasing  i s the basic  steroids.  pre-  With  daily  the adrenal cortex i s spared  them, w i t h a r e s u l t i n g  cholesterol,  illustrates  glycogen  pH d e c l i n e .  of corticosteroids  of producing  tissue  d e c r e a s i n g t h e u l t i m a t e pH.  f o r the biosynthesis of c o r t i c a l  injections  blood  i s n o t immediate. i n muscle  illustrates  cursor  Therefore, response a t  With the i n c r e a s e noted  p o s t mortem g l y c o l y s i s  process,  decrease  circulating  i n uptake o f  levels.  T a b l e 11  effect.  ULTIMATE pH OF P. MAJOR MUSCLE FROM BROILERS RECEIVING DAILY CORTICOSTERONE INJECTIONS FOR 8 DAYS AND OBSERVED FOR A FURTHER 7 DAY RECOVERY PERIOD. Day  pH*  0 8 10  5.79 5.57 5.40  a r e an a v e r a g e  o f those  from  4 birds.  . .  TABLE 11.  BLOOD CHOLESTEROL LEVELS OF BROILERS RECEIVING DAILY CORTICOSTERONE INJECTIONS FOR 8 DAYS AND OBSERVED FOR A FURTHER 7 DAY RECOVERY PERIOD.  Day  0 8 15 *  V a l u e s a r e an a v e r a g e  Blood  Cholesterol* mg % 115 275 120  o f those determined  for 4 birds.  37.  SUMMARY AND  Injected  corticosterone  carbohydrate metabolism Short indicated  term  d i d express  and m u s c l e  studies  an e l e v a t i o n  injection.  CONCLUSIONS  tenderness o f b r o i l e r s .  conducted  higher  than the c o n t r o l  excitable value. in  immediately  birds  over  displayed  Extremely  s h e a r v a l u e s a t 2-8 h o u r s  v a l u e upon i n i t i a l return  muscle  injections cessation blood tion  studies  and a marked  cholesterol  t h e 8 hour  birds  corticosterone  period.  indicated  indicated  Mildly  a marked  a decrease  decrease  i n shear  adjustments  metabolic condition  during  to continued  i n shear values a t the  B l o o d g l u c o s e , t i s s u e g l y c o g e n and  l e v e l s s i m i l a r l y increased  o f i n j e c t i o n s and t h e n d e c r e a s e d  returning  of the  post-injection.  increase  of injections.  found  and r e m a i n e d s i g n i f i c a n t l y  i n j e c t i o n , homeostatic  t o a normal  level  post-  no s i g n i f i c a n t c h a n g e i n s h e a r  excitable  Long t e r m  period  2-8 h o u r s  stress  When p l a c i d b i r d s were i n j e c t e d w i t h  shear v a l u e s i n c r e a s e d  an 8 h o u r  m u s c l e t e n d e r n e s s was  t o be d e p e n d e n t on t h e p r e - i n j e c t i o n bird.  over  i n b l o o d g l u c o s e from  E f f e c t on P. M a j o r  an e f f e c t on  t o those l e v e l s found  to the cessa-  rapidly,  i n the c o n t r o l  ultimately birds.  U l t i m a t e pH o f t h e p o s t mortem t i s s u e d e c r e a s e d w i t h d a i l y corticosterone As the s t r e s s metabolic  injections.  proposed,  reaction  corticosterone  i n that  injections did offset  i t modified stress  c h a n g e s i n t h e m u s c l e and r e s u l t e d  induced  i n a decrease  38,  i n shear value o f some post mortem muscle. I t appears from the r e s u l t s o f the s h o r t term t h a t the e f f e c t o f i n t r a m u s c u l a r c o r t i c o s t e r o n e dependent  studies,  injection i s  on the p h y s i o l o g i c a l s t a t e o f the b i r d and i t s s t a t e  of r e s i s t a n c e  t o environmental s t r e s s o r s a t the time o f  injection. 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Med. 41: 486.  APPENDIX I Parameters of S u b j e c t i v e E v a l u a t i o n of S t r e s s L e v e l o f B i r d s  S T R E S S  STRESS Placid  L E V E L  Mildly Excitable  Extremely Excitable  A person entering range house  Head movement remained sitting  Walked away from person  Rapidly moved to f u r t h e s t corner o f house  Continued presence o f person  Got up and walked about  Bunched i n the corner  Fought t o g e t i n corner-most position  Person h a n d l i n g other b i r d s  Walked  Moved i n t o t i g h t e r bunch some f i g h t i n g  Fought w i l d l y sat on each other i n c o r n e r  Being handled  Flapped wings a few times  Flapped wings continuously and o c c a s i o n a l l y pecked  Flapped w i l d l y , w r i t h i n g body and p e c k i n g ,  about  

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