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Qualitative and quantitative aspects of the protein nutrition of juvenile chinook salmon (Oncorhynchus… McCallum, Ian 1985

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QUALITATIVE AND QUANTITATIVE OF THE PROTEIN  NUTRITION  ASPECTS  OF JUVENILE  CHINOOK SALMON ( O n c o r h y n c h u s  ts hawytscha)  by  IAN McCALLUM  B. M.  Sc. Sc.,  A THESIS  (Agr.).  McGill  University SUBMITTED  of  University, Saskatchewan,  1973 1978  IN PARTIAL FULFILLMENT OF  THE REQUIREMENTS  FOR THE DEGREE OF  DOCTOR OF PHILOSOPHY  in THE FACULTY OF GRADUATE STUDIES Department  We a c c e p t  of  this  Poultry  thesis  to_tiie_required  THE UN/IVERSI^KOF July  as  Science  conforming  standard  BRITISH 1985  Ian M c C a l l u m ,  1985  COLUMBIA  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  requirements f o r an advanced degree at the  the  University  of 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 it  f r e e l y a v a i l a b l e f o r reference  and  study.  I  further  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  be granted by the head of  department or by h i s or her  representatives.  my  It i s  understood t h a t copying or p u b l i c a t i o n of t h i s t h e s i s f o r f i n a n c i a l gain  s h a l l not be  allowed without my  permission.  Department o f  9ou  LTfi.Y  The U n i v e r s i t y of B r i t i s h 1956 Main Mall Vancouver, Canada V6T 1Y3  DE-6  (3/81)  ^(HEMcg Columbia  written  ABSTRACT  Two  experiments  protein  source  (Oncorhynchus protein  and l e v e l  mix  differently  from the  protein  duplicate replaced energy  growth be  slight High  of  of  best  in  temperature  depressed  quality growth  of  intake  whole  of  were  fish  for  water  at  fed  The  muscle  and  processed  and D L - m e t h i o n i n e  three  to  levels  satiation  period.  relative  on body  d r y i n g of  protein  of  to  Protein  to  was  metabolizable evaluate  protein  to  FPE.  In  protein  gain,  FPE was  h e r r i n g meal  quality  compared  d r i e d h e r r i n g meal  protein  fish  fed  nitrogen  excretion  tanks.  terms  of found  source.  (75°C)  into  arginine  methods employed  based  salmon  pollock  on an e s t i m a t e d  values  of  chinook  h e r r i n g meals  a 42-day  and g l u c o s e  effect  raw h e r r i n g and a  diets  source.  were o b t a i n e d  in  of  exogenous  the  juvenile  fresh  tested  quality  loss  and  in  sources  (150°C)  endogenous protein  of  with  protein  poor  study  supplemented  and a s s a y s  reduction  protein  same l o t  different  temperature  extremely  three  The v a r i o u s  yielded  the  diet  reared  isocaloric  by d e x t r i n  rate  Low  in  groups  basis.  quality  to  mix  The p r o t e i n  dietary  the  to  compared were a f r e e z e - d r i e d  (9:1)(FPE),  casein-gelatin (CS).  in  tshawytscha)  sources  euphausid  were c o n d u c t e d  the for  for  amounts each  -  enabled  used  _  found  food  to  for  the  intake of  the  partitioning  growth,  source.  be an  estimates  The d e t e r m i n a t i o n  protein  11  was  C S , lower  a  freeze-drying .  Although high  CS d i e t s . from f i s h  to  caused  maintenance  In E x p e r i m e n t containing  17  FPE  levels  at  two  + 0.25398x = protein the  to  2,  two  47% p r o t e i n , of  dietary  8.37872x  energy:total  dietary  series  protein  in  increments  for  105-day  period.  Maximum growth  of  0.55.  However,  for  The  was  range  efficiency  found  permits in diet  to the  practical lie fish  in  the  purposes range to  the  provided  growth  rate, to  chinook  achieved  at  by  and x  quantify salmon  a PE:TE  PE:TE  between 0.35 consider  tested  y = -0.50699  derived  juvenile  was  culturist  formulation.  was  were  10%,  The e q u a t i o n  (PE:TE))  requirement  diets  of  y = specific  a  required  isocaloric  energy.  ,(where energy  of  over  ratio  ratio and  0.55.  economic  Table  of  Contents  Section  Page CHAPTER 1  1.0  INTRODUCTION  1 CHAPTER 2  2.0  REVIEW OF LITERATURE  2.1  The u t i l i z a t i o n  of  dietary  protein  3  2.2  General  of  protein  metabolism  5  2.3  Nitrogen  2.4  Endogenous  2.5  Methods used  2.6  The p r o t e i n r e q u i r e m e n t s of s a l m o n i d s to d i e t a r y e n e r g y CHAPTER 3  3.0  aspects  3  balance  8  nitrogen to  EXPERIMENT 1. measurement of salmon f r y  excretion  determine  Protein protein  12  protein  quality with  u t i l i z a t i o n and the q u a l i t y in diets for  15 respect 22  chinook 28  3.1  Introduction  28  3.2 3.2. 3.2.2 3.2.3 3.2.4 3.2.5 3 . 2 .'6 3.2.7 3.2.8  M a t e r i a l s and methods Test p r o t e i n sources Diets Aquarium f a c i l i t y Protocol Measurement of growth C h e m i c a l a n a l y s i s of f i s h Available lysine Data a n a l y s i s  32 32 37 42 42 42 43 43 43  3.3  Results  3.3.1  The e f f e c t of d i e t a r y p r o t e i n s o u r c e and l e v e l on •body w e i g h t g a i n The e f f e c t of d i e t a r y p r o t e i n s o u r c e and l e v e l on f o o d i n t a k e and g r o s s f o o d c o n v e r s i o n efficiency The e f f e c t of d i e t a r y p r o t e i n s o u r c e and l e v e l on energy i n t a k e and g r o s s energy u t i l i z a t i o n The e s t i m a t i o n of endogenous n i t r o g e n l o s s by j u v e n i l e c h i n o o k salmon The r e l a t i o n s h i p between p r o t e i n i n t a k e and protein u t i l i z a t i o n — iv —  3.3.2  3.3.3 3.3.4 3.3.5  and d i e t s  :  .  46  46  52 57 59 65  Table 3.3.6 3.3.7 3.3.8 3.3.9 3.4 3.4.1  The measurement efficiency The measurement productive The measurement Available lysine  of  Contents  (cont'd)  of p r o t e i n q u a l i t y by p r o t e i n r a t i o and net p r o t e i n r a t i o 70 of p r o t e i n q u a l i t y by p r o t e i n v a l u e and net p r o t e i n u t i l i z a t i o n . . . . 80 of p r o t e i n q u a l i t y by s l o p e ratio....88 c o n t e n t of the p r o t e i n s o u r c e s 93  3.4.6  Discussion 95 The e f f e c t of d i e t a r y p r o t e i n s o u r c e and l e v e l on body w e i g h t g a i n 95 The e f f e c t of d i e t a r y p r o t e i n s o u r c e and l e v e l on f o o d i n t a k e and g r o s s f o o d c o n v e r s i o n efficiency 97 The e f f e c t of d i e t a r y p r o t e i n s o u r c e and l e v e l on energy i n t a k e and g r o s s e n e r g y u t i l i z a t i o n . . . 98 The e s t i m a t i o n of m a i n t e n a n c e r e q u i r e m e n t s f o r protein 100 The e f f e c t of d i e t a r y p r o t e i n s o u r c e on p r o t e i n utilization 105 The d e t e r m i n a t i o n of p r o t e i n q u a l i t y by b i o a s s a y . . . . 112  3.5  Summary of  3.4.2  3.4.3 3.4.4 3.4.5  Experiment  1  127  CHAPTER 4 4.0  EXPERIMENT 2. P r o t e i n r e q u i r e m e n t s c h i n o o k salmon i n r e l a t i o n to content  of j u v e n i l e d i e t a r y energy 129  4.1  Introduction  129  4.2 4.2.1 4.2.2 4.2.3  M a t e r i a l s and methods Protocol Diets Data a n a l y s i s  130 130 130 13,1  4.3 4.3.1  Results E s t i m a t i o n of the p r o t e i n r e q u i r e m e n t of j u v e n i l e c h i n o o k salmon from growth d a t a The e f f e c t of d i e t a r y p r o t e i n and e n e r g y l e v e l on food c o n v e r s i o n and g r o s s e n e r g y utilization The e f f e c t of d i e t a r y p r o t e i n and energy l e v e l on p r o t e i n u t i l i z a t i o n The e f f e c t of d i e t a r y p r o t e i n and energy l e v e l on the p r o x i m a t e body c o m p o s i t i o n of fish  135  4.3.2  4.3.3 4.3.4  v  135  147 148  150  Table  of  Contents  (cont'd)  4.4  Discussion  A.4.1  The d i e t a r y p r o t e i n r e q u i r e m e n t of j u v e n i l e c h i n o o k salmon The e f f e c t of d i e t a r y e n e r g y on the efficiency of p r o t e i n u t i l i z a t i o n The e f f e c t of the s o u r c e and l e v e l of d i e t a r y e n e r g y on the p r o x i m a t e body c o m p o s i t i o n of f i s h  A.4.2 4.4.3  4.5  Summary of  155  Experiment 2  155 162  165 168  CHAPTER 5 5.0  Conclusions  170  6.0  Bibliography  174  7.0  Appendices  189  vi  List  of  Tables  Table  Page  1.  Amino a c i d r e q u i r e m e n t s f o r c h i n o o k s a l m o n , c o m p o s i t i o n of eggs and the a n a l y z e d e s s e n t i a l amino a c i d c o m p o s i t i o n of f r e e z e - d r i e d p o l l o c k m u s c l e , freeze-dried whole e u p h a u s i d s and v i t a m i n - f r e e casein 30  2.  C o m p o s i t i o n of  protein  3.  C o m p o s i t i o n of  diets  4.  Proximate  5.  Summary of  6.  F i n a l mean wet body w e i g h t s and s p e c i f i c growth r a t e s of f i s h f e d the v a r i o u s p r o t e i n s o u r c e s a t each d i e t a r y p r o t e i n c o n c e n t r a t i o n  7.  8.  9.  10.  11.  12.  13.  14.  composition dietary  sources  35-36 of  diets  treatments  S l o p e s of body w e i g h t a g a i n s t various p r o t e i n sources Total gross  34  38-39 and codes  protein  intake  for  50 the 53  d r y food i n t a k e , mean d a i l y food i n t a k e food c o n v e r s i o n of the v a r i o u s d i e t s  Gross food c o n v e r s i o n ( G F C ) , of d i f f e r e n t p r o t e i n s o u r c e s concentration  41  and 56  r e l a t i v e GFC and r a n k i n g at d i f f e r e n t d i e t a r y 58  Mean d a i l y g r o s s energy i n t a k e , mean d a i l y m e t a b o l i z a b l e e n e r g y i n t a k e and g r o s s e n e r g y u t i l i z a t i o n of f i s h fed the v a r i o u s d i e t s  60  G r o s s energy u t i l i z a t i o n ( G E U ) , r e l a t i v e GEU and r a n k i n g of d i f f e r e n t p r o t e i n s o u r c e s at d i f f e r e n t dietary concentrations  61  Wet body w e i g h t and p e r c e n t of i n i t i a l body w e i g h t f i s h f e d p r o t e i n - f r e e and a m a i n t e n a n c e d i e t E s t i m a t i o n of mean d a i l y c a r c a s s a n a l y s i s of f i s h low p r o t e i n d i e t  of 63  endogenous n i t r o g e n l o s s by f e d a p r o t e i n - f r e e and a 63  The u t i l i z a t i o n of d i e t a r y p r o t e i n on a d a i l y b a s i s by f i s h f e d the v a r i o u s e x p e r i m e n t a l d i e t s d u r i n g the 42 day p e r i o d  VII  66  List 15.  of  Tables  (cont'd)  P r o t e i n e f f i c i e n c y r a t i o c a l c u l a t e d on a d r y body w e i g h t b a s i s ( P E R ) , r e l a t i v e PER and r a n k i n g of d i f f e r e n t p r o t e i n s o u r c e s at d i f f e r e n t dietary concentrations  72  16.  Net p r o t e i n r a t i o c a l c u l a t e d on a d r y body w e i g h t b a s i s ( N P R ) , r e l a t i v e NPR and r a n k i n g of different p r o t e i n s o u r c e s a t d i f f e r e n t d i e t a r y c o n c e n t r a t i o n s . . . . 75  17.  P r o t e i n e f f i c i e n c y r a t i o ( P E R ) , r e l a t i v e PER, net p r o t e i n u t i l i z a t i o n ( N P U ) , and r e l a t i v e NPU f o r v a r i o u s p r o t e i n s when f e d as the s o l e s o u r c e of p r o t e i n to d i f f e r e n t f i s h s p e c i e s h e l d under dissimilar conditions  18.  19.  20.  21.  22.  23.  24.  25.  26.  Protein ranking dietary  76-79  p r o d u c t i v e v a l u e ( P P V ) , r e l a t i v e PPV and of d i f f e r e n t p r o t e i n s o u r c e s at different concentrations  81  Net p r o t e i n u t i l i z a t i o n c a l c u l a t e d by the method of Bender and M i l l e r (1953) ( N P U - 1 ) , r e l a t i v e NPU-1 and r a n k i n g of d i f f e r e n t p r o t e i n s o u r c e s a t different dietary concentrations  82  Net p r o t e i n u t i l i z a t i o n c a l c u l a t e d by the method of Ogino et a l . ( 1 9 8 0 ) ( N P U - 2 ) , r e l a t i v e NPU-2 and r a n k i n g of d i f f e r e n t p r o t e i n s o u r c e s at different dietary concentrations  83  The r e l a t i o n s h i p between p r o t e i n i n t a k e and net p r o t e i n u t i l i z a t i o n c a l c u l a t e d by the Bender and M i l l e r ( 1953) f o r m u l a  84  The r e l a t i o n s h i p between p r o t e i n i n t a k e and net p r o t e i n u t i l i z a t i o n c a l c u l a t e d by the method of Ogino et a l . ( 1980)  85  S l o p e r a t i o s of intake for f i s h  dry body w e i g h t f e d the v a r i o u s  89  S l o p e r a t i o s of intake for f i s h  body p r o t e i n g a i n s on p r o t e i n f e d the v a r i o u s p r o t e i n s o u r c e s  Available lysine, lysine salmon and p e r c e n t a g e of p r o t e i n source  g a i n s on p r o t e i n protein sources  requirements requirements  for chinook s u p p l i e d by  each 94  Estimated absorbed n i t r o g e n requirement for m a i n t e n a n c e of f i s h , d e t e r m i n e d by f e e d i n g and carcass a n a l y s i s experiments  —  90  vm  -  104  List 27.  28.  29.  of  Tables  (cont'd)  Summary of the r e l a t i v e e s t i m a t e s of the n u t r i t i v e v a l u e of the t e s t p r o t e i n s o u r c e s employed i n t h i s s t u d y d e t e r m i n e d by d i f f e r e n t methods  119  Correlation coefficients and l e v e l of significance between the d i f f e r e n t p a r a m e t e r s used to e s t i m a t e the n u t r i t i v e v a l u e of d i e t s c o n t a i n i n g the v a r i o u s p r o t e i n sources  119  C o m p o s i t i o n of d i e t s c o n t a i n i n g v a r i o u s p r o t e i n and e n e r g y ( E x p e r i m e n t 2)  132  levels  30.  P r o x i m a t e c o m p o s i t i o n and c a l c u l a t e d d i e t s employed i n E x p e r i m e n t 2  31.  Wet f i s h body w e i g h t a t day 43 and day 105, and s p e c i f i c growth r a t e s of f i s h f e d d i e t s c o n t a i n i n g d i f f e r e n t l e v e l s of p r o t e i n and e n e r g y , and OMP  138  G r o s s f o o d c o n v e r s i o n e f f i c i e n c y and g r o s s e n e r g y u t i l i z a t i o n of d i e t s c o n t a i n i n g d i f f e r e n t l e v e l s of p r o t e i n and energy  149  32.  energy  of  contents  of 133  33.  Protein e f f i c i e n c y r a t i o , protein productive value and net p r o t e i n u t i l i z a t i o n of d i e t s c o n t a i n i n g d i f f e r e n t l e v e l s of p r o t e i n and energy 151-152  34.  Whole body p r o x i m a t e c o m p o s i t i o n a t day 42 of f i s h f e d the d i e t s c o n t a i n i n g d i f f e r e n t l e v e l s of p r o t e i n and e n e r g y  153  Whole body p r o x i m a t e c o m p o s i t i o n at day 105 of f i s h f e d the d i e t s c o n t a i n i n g d i f f e r e n t l e v e l s of p r o t e i n and e n e r g y  154  35.  ix  List  of  Figures  Figure 1.  2.  Page The u t i l i z a t i o n fish . Stylized  of  food  nitrogen  i n growing  juvenile 6  representation  of  nitrogen  balance  10  3A.  Growth of c h i n o o k salmon f e d the v a r i o u s p r o t e i n s o u r c e s i n d i e t s c o n t a i n i n g a p p r o x i m a t e l y 17% p r o t e i n . . 4 7  3B.  Growth of c h i n o o k salmon f e d the v a r i o u s p r o t e i n s o u r c e s i n d i e t s c o n t a i n i n g a p p r o x i m a t e l y 27% p r o t e i n . . 4 8  3C.  Growth of c h i n o o k salmon f e d the v a r i o u s p r o t e i n s o u r c e s i n d i e t s c o n t a i n i n g a p p r o x i m a t e l y 37% p r o t e i n . . 4 9  4.  S p e c i f i c growth r a t e s s o u r c e s of p r o t e i n a t  5.  S l o p e s of w e i g h t g a i n a g a i n s t p r o t e i n i n t a k e c h i n o o k salmon f e d d i e t s c o n t a i n i n g the t e s t sources  of c h i n o o k salmon different dietary  fed v a r i o u s concentrations.52 of protein 55  6A-E  P e r c e n t u t i l i z a t i o n of p r o t e i n f e d f o r m a i n t e n a n c e and g r o w t h , and p e r c e n t e x c r e t e d by f i s h f e d the t e s t protein sources 68-69  7.  E f f e c t of d i e t a r y p r o t e i n l e v e l on p r o t e i n efficiency r a t i o c a l c u l a t e d on a wet body w e i g h t b a s i s f o r c h i n o o k salmon f r y f e d f r e e z e - d r i e d p o l l o c k and e u p h a u s i d s , for p l a i c e f e d f r e e z e - d r i e d cod muscle and f o r r a t s f e d casein containing diets 71  8.  The r e l a t i o n s h i p efficiency ratio diets containing  9.  The r e l a t i o n s h i p between p r o t e i n i n t a k e u t i l i z a t i o n of the t e s t p r o t e i n s o u r c e s  10.  between p r o t e i n i n t a k e and p r o t e i n and net p r o t e i n r a t i o of f i s h f e d f r e e z e - d r i e d p o l l o c k and euphausids...73 protein  S l o p e s of d r y body w e i g h t g a i n on p r o t e i n i n t a k e of c h i n o o k salmon f e d d i e t s c o n t a i n i n g the t e s t p r o t e i n sources  11.  Slopes salmon  12.  Weight g a i n of c h i n o o k salmon from stage fed d i e t s c o n t a i n i n g various and e n e r g y , and f e d OMP  13.  and net  of body p r o t e i n g a i n fed d i e t s c o n t a i n i n g  91  on p r o t e i n i n t a k e of c h i n o o k the t e s t p r o t e i n s o u r c e s . . . 9 2 the f r y - to s m o l t l e v e l s of p r o t e i n  S p e c i f i c growth r a t e of j u v e n i l e c h i n o o k salmon f e d d i e t s c o n t a i n i n g d i f f e r e n t l e v e l s of p r o t e i n and energy x  86  137  139  List  14.  15.  of  Figures  (cont'd)  S p e c i f i c growth r a t e of j u v e n i l e c h i n o o k salmon diets containing different protein energyrtotal energy r a t i o s  fed  The second o r d e r p o l y n o m i a l c u r v e f i t t e d to specific growth r a t e of j u v e n i l e c h i n o o k salmon f e d d i e t s w i t h d i f f e r e n t p r o t e i n e n e r g y : t o t a l energy r a t i o s  xi  142  145  List  of  Appendices  Tables  Table 1.  2.  3.  4.  5.  6.  7.  8.  9.  10.  Page A n a l y s i s of v a r i a n c e Experiment 1  of  day  42  body w e i g h t s  in 191  A n a l y s i s of v a r i a n c e and c o v a r i a n c e f o r s l o p e (GR%) t e s t of l o g body w e i g h t s i n E x p e r i m e n t A n a l y s i s of c o v a r i a n c e of body w e i g h t p r o t e i n i n t a k e ( E x p e r i m e n t 1)  gain  1  192  against 193  A n a l y s i s of v a r i a n c e of g r o s s f o o d c o n v e r s i o n and g r o s s e n e r g y u t i l i z a t i o n of d i e t s f e d i n E x p e r i m e n t A n a l y s i s of net p r o t e i n  variance r a t i o of  of p r o t e i n e f f i c i e n c y r a t i o d i e t s fed i n Experiment 1  A n a l y s i s of v a r i a n c e of d i e t s fed i n Experiment  protein 1  A n a l y s i s of net p r o t e i n  net p r o t e i n -2 of d i e t s  v a r i a n c e of utilization  productive  and 194  value  of 194  u t i l i z a t i o n -1 fed i n Exp. 1  Summary of s t a t i s t i c a l a n a l y s i s f o r the s l o p e s of d r y body w e i g h t and body p r o t e i n g a i n ( E x p e r i m e n t A n a l y s i s of v a r i a n c e Experiment 2  of  A n a l y s i s of v a r i a n c e Experiment 2  of  of  day 42  1.193  body w e i g h t s  and 195  1)..195  in 196  day  105  body w e i g h t s  in 196  11.  Analysis  covariance  for  body w e i g h t s  i n Exp. 2  12.  S t a t i s t i c a l a n a l y s i s f o r the second o r d e r p o l y n o m i a l model to e s t i m a t e p r o t e i n r e q u i r e m e n t s i n E x p . 2  197  198  13.  S t a t i s t i c a l a n a l y s i s f o r the second o r d e r p o l y n o m i a l model to e s t i m a t e the p r o t e i n e n e r g y : t o t a l energy i n Experiment 2 ..198  14.  A n a l y s i s of v a r i a n c e of g r o s s f o o d c o n v e r s i o n g r o s s energy u t i l i z a t i o n i n Experiment 2  15.  16.  A n a l y s i s of and p r o t e i n  v a r i a n c e of p r o t e i n e f f i c i e n c y p r o d u c t i v e value i n Experiment  A n a l y s i s of v a r i a n c e in Experiment 2  of  net  protein  and 199  ratio 2  utilization  199 -2 200  _  xii  —  List 17.  18.  of  Appendices  Tables  (cont'd)  T a b l e of mean s q u a r e s f o r f i s h body m o i s t u r e , l i p i d and p r o t e i n a t day 42 ( E x p e r i m e n t 2)  ash,  T a b l e of mean s q u a r e s f o r f i s h body m o i s t u r e , l i p i d and p r o t e i n a t day 105 ( E x p e r i m e n t 2)  ash,  xiii  _  201  201  ACKNOWLEDGEMENTS  The  assistance  completion indebted  of  to  this  of  study.  guidance.  Professor  Beryl  supervision.  who was  instrumental author  Fisheries  Oceans, West  to  the  been most  would  analyses advice  In  Dianne  like  provided  is  processing Very  their  would  the  assuming  the  author  very  in  indebted fish  to  much and  to  studies  and  D r . B r u c e Owen  study. D r . Edward D o n a l d s o n  for of  is  encouragement  due  this  the  of  Fisheries  the  the  and  facilities  student Fish  use.  of  of  the  Being  Culture  Section  the  staff  sincere  provided by H e l e n  of  the  West  assistance,  Nancy R i c h a r d s o n ,  technical  Helen  good  Bakhshish Dye,  Andy Lamb  assistance.  Crepeau with  the  humor  The  statistical  D r . Bob McKay p r o v i d e d  valuable  inference. like  to  thank  Debra Wadsworth  for  manuscript.  special  for  are  Jack M a r k e r t ,  acknowledged.  author  thank  particular  on s t a t i s t i c a l  The  to  Plotnikoff,  Edgell a l l  assistance  interest  i n making  activities  author  greatly  (Department  support  Laboratory for  Phill  also  in  rewarding.  friendship.  Dosanjh,  the  patience,  acknowledges  Branch  his  instrumental  thanks  initiating  research  author  Vancouver  and  in  is  Vancouver L a b o r a t o r y a v a i l a b l e  The  and  his  keen  Special  Research for  for  her  gratefully  Canada)  exposed has  for  was  p a r t i c u l ar  The a u t h o r  March  skilled  the  In  D r . David Higgs  dedicated  The  many p e o p l e  the  thanks  are  burden of  completed  this  due extra  to  the  work  study. —  xiv  —  author's on the  wife,  family  Lesley,  farm  while  CHAPTER 1  1.0  INTRODUCTION  The  relationship  important  aspects  of  of  hatchery-raised  of  their  1984).  nutritional  salmonids  have  predominantly component  of  high  fish  fishmeal  in  does  goal  al . ,  1982,  1983).  fish  nutritionists  all  This  could  nutrients  protein  source  1979;  in  in  to  the  terms  diet. of  the  fish  and the  total  in  the  diet  would  be e x p e c t e d  Further,  there  is  a need  measurement  of  the  energy  (Cowey,  1979).  for  the  biological  in  f ish.  1  ratio  of  Higgs  objective  among  of  to  of  the  amino  of  to  a major the  value  the  acid  energy  develop  nutritive  balance  quality  play  et  dietary  a correct  protein to  sources  1981a,b;  essential  the  expensive  replacement  utilization  of  is  readily  1980,  requirements  al.,  (Oncorhynchus  be a  In p a r t i c u l a r ,  et  becoming  by o b t a i n i n g  meeting  function  protein  salmon  an i m p o r t a n t  be a c h i e v e d  a  most  Total  to  ability  Protein  the  are  most  species,  quality  Fowler,  the  the  Plotnikoff  and i s  fishery  maximize  of  may be  animal  high  appear  Consequently, is  1969;  chinook  however,  (Westgate,  wild  round b a s i s .  for  one  requirements.  meal,  Columbia  on a y e a r  not,  attainable  protein  Suitable  diets  the  cultured  by f i s h  British  in  is  F u r t h e r m o r e , the  (Burrows,  other  dietary  practical  tshawytscha)  proteins.  most  and growth  survive  history  diets.  scarce  food  production.  to  furnished  from the  increasingly  of  fish  fish  Compared to  derived  between  of  role  criteria protein  The o v e r a l l were  twofold.  source to  objectives First,  and l e v e l  determine  requirements  to  of  evaluate  on p r o t e i n  the  effect  this  of  study the  effect  utilization dietary  and u t i l i z a t i o n  of  on c h i n o o k  dietary  (Experiment  energy  protein  of  salmon  level  for  on  growth  1).  fry  protein Second,  the (Experiment  2). The the  following  reader  with  understanding  of  review  of  literature  a background i n the  protein  experimental  2  goals  is  intended  nutrition  to to  and p r o c e d u r e s  provide facilitate employed.  CHAPTER 2  2.0  REVIEW  2.1  The u t i l i z a t i o n The  OF LITERATURE  utilization  dietary  ability  and  i n d i v i d u a l amino a c i d content  genetic  Most  fish  carnivorous fish  amino been  of  and have  for  Shimeno  Compared  to  metabolize (Walton  et  most  a greater  of  lower  than  energy  poikilothermic. in  the  to  synthesize  (Smith  form of  et  al.,  to  ammonia, acid  energy,  conditions  play  body of  while  same ten  the  3  and  dietary have  1982).  (Phillips,  salmonids for  energy  requirements a  of  considerable Fish  excreted  are by f i s h  and mammals r e q u i r e  respectively  1979).  These  protein  energy  the  1982).  food,  nitrogen  the  essential  poorly  require  The  (Cowey,  (Ketola,  temperature.  birds  1978) .  for  protein  are  animals.  species  upon  role.  culturists  dietary  the  and u r e a  a major  requirement  mammals which  most  dietary  raised  However,  maintain  total  and A t k i n s o n ,  p r o p o r t i o n of  for  the  animal.  carbohydrates Hilton  dependent  the  fish  the  fish  species  1982).  of  by most o t h e r  1979;  Also,  uric  to  require  a dozen  animal  of  protein  utilize  al.,  and Cowey,  are  amount  fish  mainly  fulfill  animal also  needed  over  is  environmental  a high  are  to  source  concern  studied  that  reviewed  the of  protein  requirements  the  of  protein  source  diet,  species  Carnivorous  fish  the  state  species  acids  1969;  protein  predisposition,  physiological  All  the  of  dietary  the  energy  of  of  for  is  energy  excretion  Unlike acids if  carbohydrates  derived  fed  i n excess  considerable in  the  from  globules.  them,  of  deposits  body  For t h i s  reason  is  dependent  amino a c i d s  absorbed  sources  utilization  is  subject  the  to  vary  three  food  nitrogen  in  Fig . 1 .  catabolic via  the  retained  nitrogen they  and used  fractions  play  a role  for  in  the  are  they  are  wastes,  they  represent  by the  protein of  1959).  metabolic  of  intake,  dietary  growing  utilized give  utilization  fraction  to  of  of  the  of  4  from  the  depicted  nitrogen  and  losses  intake  The endogenous because of  nitrogen  losses  exogenous  which have  and o t h e r  of  anabolic  some e s t i m a t o r s  fractions  These  is  primarily  endogenous  in  utilization  nitrogen  and m a i n t e n a n c e . interest  the  through  rise  protein  digestibility  fish  the  indistinguishable  animal.  and  metabolic  calculation  although  utilized  fat  and O r t o ,  to  of  is  been  (Albanese  which  quality.  nitrogen  found  and p r o p o r t i o n s  of  are  protein that  are  or  dietary  representation  growth  The f e a t u r e  animal  no  granules of  amino  by the  or amino a c i d s  utilization  Another  (Fig.l)  such  quantities  and the  acids  and k i d n e y .  the  species. of  processes  or  means t h a t  digestibility  applies  amino  as  glycogen  meal  A general it  This  factors:  tract,  enzymatic  gills  stored  efficiency  animal  general  as  Once a b s o r b e d ,  to  each  their  proteins,  protein  the  or e f f i c i e n c y  products.  of  is  in  gastrointestinal  digestion  be  upon the  with  by v a r i o u s  The q u a l i t y  of  comparable  utilization  Protein  cannot  requirements.  inert  animal  and l i p i d s ,  aspects  actually of  protein  utilization based  on the  '2 .2  General In  are as  scheme  fish,  of  undergo  animals  deamination  conversion  to  polypeptides, synthesis protein  broken that,  lipid  followed  and a r e are  in a continual  only  does each  different  rate  of  1980). are  For example,  primarily  mass. to  be  In the  regulated  in  for  on the  other  turnover,  studies  have  protein  the  have  mechanisms  rates  of  regulation  a  for  hand,  protein of  being  confirmed  p r o t e i n may d i f f e r  the  the  of  (Garlick, synthesis  muscle  protein  p r o t e i n mass i s  thought  results  from a b a l a n c e  between  growing  rate  state  maintenance  the  of  dietary  adults,  synthesis  pools  protein  In  rate  and breakdown ( M i l l w a r d  animal p r o t e i n  In s t u d i e s  to  of  breakdown.  the  synthesis  various  protein  muscle  protein  protein  possibly  of  sensitive  are  of  non-growing of  tissue  changes  by the  and  and  reactions  These  The r a t e s  and each  but a l s o  each  responsible liver,  1982).  Recent  tissue  turnover,  mass of  biochemcial  acids.  extremely  pathways  under h o r m o n a l c o n t r o l .  not  the  same  and s y n t h e s i s  down and r e s y n t h e s i z e d .  regulating  the  by o x i d a t i o n  and n u c l e i c  and breakdown a r e  proteins  complex  and g l u c o s e ,  purines  intake  Body  same  enter  ( W a l t o n and Cowey,  synthesis,  thesis  metabolism  amino a c i d s the  this  in F i g . l .  protein  assimilated  presumed to i n other  depicted  aspects  discussed in  w i l l be  and e v a l u a t i o n  exceeds  concerned  amino a c i d s  are  growth  results  from  et  al.,  the  1978).  fact  In  that  p r o t e i n breakdown. w i t h whole  body p r o t e i n m e t a b o l i s m  conceptualized  5  (Garlick,  1980).  two  One  PROTEIN INTAKE DIGESTION  FECAL NITROGEN (FN) (Undigested protein)  ABSORBED PROTEIN METABOLISM  BRANCHIAL ( B N ) - » and URINARY (UN) NITROGEN ( Protein utilized for energy)  (Anabolic-catabolic processes)  PROTEIN RETAINED  (NET PROTEIN RATIO)  GROWTH  MAINTENANCE ENDOGENOUS PROTEIN LOSS  ENDOGENOUS: BRANCHIAL (EBN) and URINARY (EUN) NITROGEN  PROTEIN PRODUCTIVE VALUE (PPV)  METABOLIC FECAL NITROGEN (MFN)  INSENSIBLE NITROGEN LOSS (SN) (Mucus, scales, etc.)  Fig.  1.  Simplified  scheme  protein  j u v e n i l e growing  in  for  6  the  -  utilization fish.  of  food  represents  the  "metabolic  p o o l " and the  These  exist  metabolic  total  in  are  to  compartments The  rate  shown  to  salinity  teleosts  be  dependent  on water  ( S m i t h and T h o r p e ,  of  replenish  the  This  body  the  that  body  were d e p l e t e d  may i n d i c a t e  that  the  rather acids  as  from one  The above provide  tissue  concepts  the  of  background f o r  synthesis  excretion.  from s t a r v e d the  S m i t h and T h o r p e , stores  fish  above  in  fish  (Savitz,  1976).  have  been  1971)  of  and  be h i g h  starvation, in  order  probably r e f i l l e d  are  tissue  pool  protein  of  last  for  container the  7  balance  first.  more has  of  the of  and F i s h e r ,  anabolic/catabolic  the  led  to  is  not  body,  but  The p o o l  transfer  (Shapiro  nitrogen  was  metabolism  amino a c i d s .  fluid  to  regeneration,  depleted  of  another  the  of  a period  will  mechanism  to  curves  temperature  be a d i s o r g a n i z e d  an i n t e g r a t e d  from p r o t e i n  last  of to  breakdown.  During  the  a metabolic  protein  nitrogen  pool. the  stores.  The dynamic n a t u r e  considered  protein  into  the  1976).  essential. concept  are  1970;  protein  protein  and body  existence  provided after  dietary  body known as  body p r o t e i n  subsequent  (Iwata,  of  is  pool  the  depletion  the  The i n p u t s  excretion  of  When p r o t e i n  tissues  nitrogen  of  whole  intake  with  demonstrate  in  retention  the  metabolic  oxidation  Characteristic served  other  from f o o d  from the  and amino a c i d  have  amino a c i d s  dynamic e q u i l i b r i u m .  pool  The o u t p u t s  free  amino 1962).  interrelationships studies  of  animals.  2.3  Nitrogen The  balance  p r i n c i p l e of  identical  to  input  versus  purpose  is  protein  utilization.  quantity  to  of  measure  nitrogen  Many e x p e r i m e n t a l devised  for  nutrient  nitrogen  the  output  net  gain  i n any o t h e r  simply  1951)  as  can be d e f i n e d  as  that  measurement  has  based of  on n i t r o g e n  the  nutritive  either  repletion  of  balance  normal  depleted  are  state  this of  balance  value  of  body.  have foods  been and  protein  metabolism  equation  (Allison,  others  the  new t i s s u e s  are  nitrogen  UN = u r i n a r y  in  nitrogen  because  some t i s s u e s  found  to  be  human s u b j e c t s  have  demonstrated  8  mean t h a t  under animals,  all  tissues the  1977).  totally  correct.  -  are,  equilibrium.  f o r m u l a of  significant  in perspiration.  -  the  Adult  balance  that  or  may be m a i n t a i n e d a t  (Young and S c r i m s h a w , nitrogen  gaining  animals  balance.  e q u i l i b r i u m does not  eliminated  intake,  animal is  A l l growing  been  are  (UN + FN)  growth of  not  nitrogen  balance  positive,  stores.  The a f o r e m e n t i o n e d has  is  conditions, nitrogen  (1951)  of  NI = n i t r o g e n  in positive  Maintaining  expense  by the  nitrogen.  t h r o u g h the  conditions,  in  state  nitrogen  balance,  and FN = f e c a l  nitrogen  protein,  under  retained  follows:  nitrogen,  normal  been  the  animals.  by the  NB = n i t r o g e n  If  its  Nitrogen balance  NB = NI where  since dietary  procedures  described  science,  is  o c c u r r i n g with  of  or  The dynamic a n a b o l i c - c a t a b o l i c is  in n u t r i t i o n a l studies  loss  intake  requirements  balance  Also,  Allison Studies  with  amounts  of  losses  from  skin  -  and h a i r must be a c c o u n t e d studies. losses  These (SN)  factorial  procedure for  consider the  in  all  the  mucus  of  as  for  1977).  of  and r a t e  of  nitrogen loss.  balance  nitrogen for  when  the  protein (Maynard and  for  fish  amounts  of  expected  are  form a c u r v e  1976). in  equilibrium  As i n t a k e  the is  infection  recognized  negative  reached.  1980)  or h a n d l i n g ,  that  are  and  1973). (Ingram,  is  known  ( P i c k e r i n g and  consideration secretion  Insensible where  (BN)  Also f i s h  (Ingram,  mucus n i t r o g e n  fish  nitrogen are  for  would loss  crowded and  an a p p r o p r i a t e n i t r o g e n  balance  (FN + UN + BN + SN) of  energy, of  excretion  fish.  fish  must  be:  a series  metabolizable  to  reduction  fed  fish  and c a r b o y d r a t e  production in  of  to  (Cameron and E n d e a n ,  protein  Therefore, would  in  protect  in aquaculture  NB = NI When f i s h  loss  to  balance  Branchial  p r o t e i n metabolism.  handled.  relationship  of  (1969)  significance  frequently  both  mucus  Savitz  estimates  al.,  accounted  calculate  nitrogen  a result  an i n c r e a s e d  may be of  et  insensible  farm a n i m a l s  nitrogen  contains  unknown amount  bias  in  application  sources  Stress,  cause  the  to  swimming and o s m o r e g u l a t i o n  1980).  Macey,  term n i t r o g e n  termed  i n mucus w h i c h f u n c t i o n s  Epithelial  to  maintenance  long  They a r e  employed  major mechanism  covered aid  been  1977).  is  in  1969).  Similarly,  is  have  (Bressani,  requirements Loosli,  losses  for  the  diets their  type  increases balance  -  9  the which  equal  balances  in F i g .  from z e r o  to  supply  nitrogen  shown  until  The e x t e n t  which  be  2 (McDonald  there  point  can  of  further  is  a gradual  exact nitrogen  Fig. (2).  Stylized representation of nitrogen balance. Taken from McDonald et al.(1976).  -  10  -  Figure  2 intake  potential  of  supply  other  of  further  promotes  the  tissue  animal,  the  nutrients.  nitrogen  intake  growth  quality  of  The c u r v e  fails  to  depends on the the  protein  growth  and  the  becomes h o r i z o n t a l  promote  additional  when  nitrogen  retention. The  information derived  been used  since  calculation  of  Value  (BV) i s  which  is  1909 the  (Thomas,  Biological  defined  retained  from n i t r o g e n  as  the  by the  cited  Absorbed A b s o r b e d food Fecal  With  food  fecal  = Fecal N -  Excreted  food  N = Urine  value  to  fish  reported (FN -  N -  (UN -  EBN = Endogenous  Branchial  EUN, and EBN a r e  nitrogen  (1979)  (BN -  EBN)  for  was:  x  100  Nitrogen fractions  (Fig.l).  the  nitrogen  excreted  diet.  formula  Nitrogen  nitrogen  protein-free  the  and T i e w s  food  by  urinary N  Nitrogen  Urinary  demonstrated  E x c r e t e d food N  EUN) -  EUN = Endogenous  of  feces  (FN - MFN)  MFN = M e t a b o l i c F e c a l  independent  food N  Endogenous  by C a s t e l l  NI -  nitrogen  ioo  nutrition research,  MFN) -  the  1924):  M e t a b o l i c N of  N = Absorbed N -  respect  x  in  food-N  food  BV = NI -  MFN,  N  Retained  biological  where:  food  has  Biological  absorbed  (Mitchell,  N = Food N -  N  of  studies 1962)  proteins.  percentage  BV = R e t a i n e d  where:  by M i t c h e l l ,  V a l u e of  organism  balance  They r e p r e s e n t  11  Their  whose e x c r e t i o n existence  when an a n i m a l i s  fractions  which have  is  is given  a  actually  been  u t i l i z e d - by the  excretions.  The n u m e r a t o r i n  total  nitrogen  It  noteworthy  is  accounted  2 .4  a n i m a l even  for  utilized that  Endogenous  Castell  nitrogen nitrogen  catabolism  represents  metabolism  and i s  urea,  and o t h e r  nitrogenous  ammonia.  The major end fish  is  amino a c i d s ,  (Watts  and W a t t s ,  breakdown of  of  creatine  have  fraction  dietary of  from  p r o d u c t of though  creatinine  purine metabolism  and t i s s u e  (1979)  endogenous the  growth.  not  acid,  and c r e a t i n i n e  protein to  also  formed from  (Forster  yield  in  trimethylamine excreted  u r e a and u r i c  is  action  are  In  nitrogen  metabolism  and c r e a t i n e  The o r i g i n of  intake.  catabolized  uric  acid  nitrogen  breakdown of  nitrogen  urea,  i n muscle  of  protein  compounds w h i c h a r e  1974).  represents  o r i g i n a t i n g from amino  maintenance  which a r i s e s  oxide,  from  excretion  ammonia,  as  loss.  constituent  is  equation  and T i e w s  independent  principal  above  appear  excretion  the  excretion  teleost  the  they  both m a i n t e n a n c e  mucus n i t r o g e n  Endogenous  mammals the  for  though  acid  is  the  and G o l d s t e i n ,  1969) . Endogenous u r i n a r y homeothermous similar  to  animals  caloric  and endogenous both a r e metabolic  and i s  metabolism  nitrogen  a function rate  nitrogen  is  of  excretion  related (Brody,  excretion metabolic  p r o p o r t i o n a l to  are  to  varies  body w e i g h t  1945).  Basal  related  to  body s i z e . the  (W  )  (Maynard  three-fourths  and L o o s l i ,  12  one  i n a manner metabolism another  In mammals  0.75 body w e i g h t  in  1969,).  the  power  of  and  Gerking in  the  (1955)  bluegill  reported  sunfish  that  endogenous n i t r o g e n  (Lepomis m a c r o c h i r u s )  excretion  decreased  with  0.54 increasing to  fish  size  has  Groves,  been  1979).  calorimetry in to  four 4.0g  from  by W  .  reported S m i t h et  to  Since  study  the  to  to  the  exponent  v a r y from 0.34  al.(1978)  employed  effect  fish  s p e c i e s of s a l m o n i d s . f i s h have a m e t a b o l i c  4.0g  then,  50.Og i n w e i g h t  of  to  1.0  applicable (Brett  a method  size  of  direct  on m e t a b o l i c  Their results indicated r a t e p r o p o r t i o n a l to W have  and  a metabolic  rate  rate  that 1.0 . Fish  proportional  0.63 to W McDonald mg of  et  al.(1976)  endogenous u r i n a r y  metabolism value  of  for  warm b l o o d e d  7 mg of  endogenous  sunfish.  This  obtained  by Bonnet  (1933)  that  fish  more endogenous animals.  This  excretion  is  a large from  carbohydrate The d a i l y shown  to  bluegill At  low  nitrogen  excreted  with  per  that  characteristic  of  and f a t rates  increase sunfish  of  with  rising  7.2  rate  in  results  and a h a l f  times  do warm b l o o d e d  poikilothermous  animals.  of  fish  large  nitrogen  are  Hence, derived  proportions  excretion  environmental and c a r p  excretion  13  have  temperature  (Ogino  rates  a  mg/kcal  nitrogen  can use  2  the  of  purpose.  1971)  nitrogen  of  than  obtained  per k c a l  three  requirements  this  (1955)  of  basal  of  endogenous  (Savitz,  temperatures  a high  homeotherms for  kcal  value at  The f o r e g o i n g  excrete  basal  energy  a value  frogs.  established  whereas  per k c a l  Gerking  agrees with  nitrogen  the  excreted  accepted  animals.  and a m p h i b i a n s  p o r t i o n of  protein,  a generally  nitrogen  bluegill  suggest  state  et  were  al., found  been  in 1973). not  to  fluctuate,  suggesting  adaptation  in cold  Smith the  process  in  increase water.  of  the  onset  By c o n t r a s t , excretion  indicate  that  the  may be g r e a t e r Another  does not arises  higher  enzyme  MFN o u t p u t the  intake,  Mitchell  the  the  O.lg/lOOg  dry matter  varying excreted  of  steers  at  any  extrapolation  to  in  fresh  food  residues with  (1954)  the  for  nitrogen  intake  of  a fixed  point  of  of  zero  14  of  MFN i s pigs  the  (Fig.l). the  trout 1973).  that It  digestive  because  digestive  a  the juices  nitrogen-free  almost  totally  approximately  total  Titus  nitrogen  with  diets  of  e s t i m a t e d MFN  by s t r a i g h t  protein  -  of  metabolic  and man.  ration level  intake  results  nitrogen  off  sea  cells.  against  at  is  to  of  (Zeitoun,  intake  levels  rats,  He a r r i v e d food  food  that  consumed  at  protein  in  These  loss  d e t e r m i n e d MFN w i t h low  rate  fecal  secretion  They e s t i m a t e d  level  water  epithelial  containing  found  maintenance  sloughed  bulk  was  water.  for  study  smoltification  their  p r o p o r t i o n of  greater  content.  the  in fresh  capacity  to  and c o n t i n u e  decreased  undigested  fed  the  endogenous n i t r o g e n  intestinal  total  protein  than  for  balance  excretion  requirements  increases  protein.  excretion  nitrogen  smolts  and c e l l  rations  plotted  nitrogen  that  from  digestible  (1927)  of  and B e r t  or w i t h  employed  smoltification  water  (MFN) or  and a b r a s i o n of  ration  sea  ability  1971).  when r e t a i n e d  originate  from  tract.  post  component  nitrogen  of  protein  in  the  (Savitz,  Endogenous  nitrogen  fecal  (1971)  have  p r o t e i n metabolism with  trout.  with  sunfish  climates  and T h o r p e  association  that  intake.  line Mitchell  and  Bert  (1954)  by d i r e c t  et  al.(1973)  determine  between  in  different  the  water  feces,  values  Ogino  et  al.(1973)  matter  2.5  showed  purpose  of  efficacy  of  Then  economic  the  quality  of  there  and b a l a n c e  in a feedstuff  subject  to  is  the  S k r e d e et 180  were  was  in  of  nitrogen  within  the  was  employed.  carp  increases  of  obtained  dietary  quality to  is  to  support  the  constituent a given  species.  an e x p e n s i v e  a  dry  quality  how w e l l  of  at  fed.  protein  its  above  values  al.(1980)  mg/lOOg  sources  to  of  evaluate  rapid  the  growth. The  available essential  species.  amino  As m e n t i o n e d  differences  in  The d e t e r m i n a t i o n  the  amino  of  amino  p r o c e d u r e which i s  often  inter-laboratory variation.  reliance  (NRC, 1 9 7 3 , 1 9 8 1 )  ( P a y n e , 1972) .  values  the  a p r o t e i n can be e s t i m a t e d .  needs  fish  considerable  Alternatively, tables  of  amount  MFN e x c r e t i o n  of  method  relationship  MFN e x c r e t i o n  a r e major q u a n t i t a t i v e  requirements  acids  refers  the  and the  protein  protein  r e t u r n of  c o r r e s p o n d to  From the  These  diet  measuring  a protein  concentration  earlier,  diet  that  (1927)  species.  obtained  determine  different  Titus  a s i m i l a r a p p r o a c h to  increases.  to  the  when a n o n - p r o t e i n d i e t  when a p r o t e i n - f r e e  The  acid  the  they  with  carp.  MFN i n r a i n b o w t r o u t  Methods used  acids  of  obtained  temperature  for  in  temperatures.  of  value  several  followed  MFN e x c r e t i o n  range  water  agreement  for  p r o t e i n content  excreted  as  good  determination  Ogino to  obtained  on t a b u l a t e d  often  leads  to  F u r t h e r m o r e , amino a c i d  15  amino a c i d  erroneous  composition  estimates  composition  data  give  no  indication the  of  amino a c i d  availability  a n i m a l when i n c l u d e d i n Methods  divided animal  into  of  experiments  final  assess  reviewed  and W i t t y  Studies  (1978)  with  with  fish  protein  These methods ( 1977),  usually  analysis  feedstuffs.  The use  of  et  who found no c o n s i s t e n t  presumed  gain  casein  or  (1979)  demonstrated  corn g l u t e n .  amino a c i d s  pattern.  However,  they  can be  (PER).  protein  This  intake.  determinations highest allow  value  for  growth. similar  i n the  is  of  protein  weight  as  several the  protein  i n concept  to  et  al.  growth  trials  attempted  relationship  fed  (1977),  (1983).  was  hepatic  diets  of  by Cowey  between  the  activities  containing  between the  and the  quality  enzyme  and K a u s h i k  fishmeal,  and L u q u e t proportions  plasma f r e e  require further  quality of  that  amino  of acid  refinement  levels  that  (NPR) (Bender  16  the  efficiency gram of  out PER  and a c c e p t e d  protein.  for  protein  carried  and assume  PER e x c e p t  is  an a n i m a l per  al.(1919)  PER f o r  ratio  gain  protein  needs  been  successfully.  Osborne et at  have  indices  (1972)  diet  metabolic  protein  techniques  employed  maintenance Net  Nose  systems,  the  opposing  by t r o u t  these  A common e s t i m a t e ratio  of  may be  evaluate  a correspondence  essential  before  metabolic  e q u i l i b r i u m point  and maximum w e i g h t  to  on  Satterlee  employed  carcass  al.(1981)  quality  based  and Von der Decken have  to  diet.  and p r o c e d u r e s  by B o d w e l l  usefulness  m i c r o b i o l o g i c a l assay  protein metabolism.  critically  coupled  to  c h e m i c a l methods,  feeding  indices  Evans  commonly used  the  and hence  the  PER e s t i m a t e s all  food  and D o e l l , inclusion  is  do  not  used  for  1957) of  is  a group  fed  a diet  devoid  of  protein.  NPR i s  defined  by the  following  equation: gain i n weight of t e s t group  NPR =  w e i g h t l o s s of group fed a n o n - p r o t e i n d i e t  + protein  Investigators Sargent,  1972;  because  of  protein  free  estimated  the  species  at  of  quality  because  it  has  gain  been  Diet  factors  in  also  level  as  of of  fat  cannot  sources,  growth  rate  when an a t t e m p t  protein. for  digestion  Techniques variation  than  is  and a b s o r p t i o n  to  to  the  made to  protein  acid  al.,  measure  sources,  1978).  17  given  1969)  i n w e i g h t may  profiles (Evans  of  of  of  new alters these  and W i t t y ,  anti-nutritional of  plant  origin,  1979).  Fish  feeding  assessment  nitrogen  or a l t e r e d  a  animal d i e t s  p a r t i c u l a r l y those  utilizing in  gain  elaboration  presence  et  of  and L o o s l i ,  be d i s c o u n t e d  (Higgs  the  estabilishing  since  sources  and the  may be more b e n e f i c i a l  quality  (Maynard  rather  at  history.  growth,  protein  acceptability  alter  life  on a  PER i s  usually  a c r i t e r i o n for  true  fish  maximum growth  v i t a m i n and f a t t y  effects  protein  of  not NPR  Usually  level  for  but  maintaining  1973).  criticized  reflect  mineral,  and t h e s e  causes  protein  The a d d i t i o n  overall  trials  protein  weight  may not  1978).  will  one  from d e p o s i t i o n  tissue.  diets,  al.,  PER (Cowey and  1982),  with  ( Z e i t o u n et dietary  report  Pfeffer,  associated  a p a r t i c u l a r stage  protein  the  at  routinely  1981;  problems  dietary  The use  result  fish  Steffens,  diet  only  recommended  with  intake  the  retention  protein  fate  balance for  of  of  depict  example (Evans  ingested the  impaired and W i t t y ,  Some i n v e s t i g a t o r s formula  for  BV f o r  fish  efficiency  of  1970;Brett  and Z a l a ,  nitrogen of  cite  Storer  nitrogen  of  nitrogen  excretion  controls  fed  considerable mammals,  the  1955).  relied  Under  requirement  composition, is  that  Savitz  that  this  Nutrient feedstuffs  balance  for  domestic  Quantitative  collection  fraught  problems  with  separated using  from  large  an i n d i g e s t i b l e  extensively  to  in  the  in  that  with  the  the  the  onset  found  sunfish  in  fish,  levels  is  of  was  rate  as  the  rate  higher  it  from  of  in  of  body  starvation  when  fat  requirements.  and e x c r e t o r y because  in  they  1969). are  must  be  measurement  chromic oxide of  evaluating  products  Indirect  digestibility  18  does  acid  (Maynard and L o o s l i ,  as  present  protein  extensively  water.  of  than  a major p o r t i o n  by a n a l y s i s in  of  of  that  amino  derived  used  marker such  determine  reported  studies of  (1979)  endogenous  energy  feces  volumes  and G r o v e s  of  changes  are  animals  fish  excretion  a measure  showed  satisfy  of  Endogenous  nitrogen  conditions  fishes  studies  1976).  nitrogen  starvation  (1971)  to  Iwata,  W a l t o n and Cowey (1982)  situation  upon h e a v i l y  1969;  of  bluegill  normal  of  gross  rate  (1969,1971)  alone.  increases  enzymes.  caloric  (Gerking  to  the  increase  starved  glucose  the  Mitchell  the  and B r e t t  that  (1967)  Savitz  evidence  leads  degrading  d i d not  in  from  provides  Stover  However,  it  (Birkett,  (1970)  evidence fish  excretion.  simpler  S m i t h and T h o r p e ,  Iwata  as  the  and c a l l e d  determined  fish.  catabolism  studies  1975;  starved  modified  utilization  been  (1967)  starvation.  in  has  starved  excretion  protein  nitrogen  loss  rate  have  has  been  feedstuffs  for  used  fish  (Nose,  Windell  et  1960; al.,  1978;  W i l s o n et  the  ratio  of  the  food  determining of  a nutrient  resulting be  in  from the  estimated  output.  S m i t h and L o v e l l ,  food,  without  Urinary  digestibility  sample  of  (Windell  et  al.,  1978;  problems  of  this  method.  collection  facilitates  the  water  or  for  Rychly  in  (1973) to  chambers  the  hydroxide  effluent  collection fish  for  is  water  employed of  feces,  were r e s t r a i n e d  al.,  of  the  methods fish  al., all  that  and p r e s e r v e Nitrogen an i o n  an a p p a r a t u s  and g i l l  in  a way  19  Iwata,  of  with  the  1970;  automated With  these  matter  is Ogino  tank  water  cupric  are  recovered  column.  excretions. the  balance  and a n a l y s i n g  permitted  that  major  excreted.  excretions  urinary  a  simply  effluent  exchange  only  leaching  nitrogen  1969;  feces  that  the  1975).  filters  and  excretions  tank  is  feces  nutrient  insoluble  material  can  obtaining  are  development  (Cho et  or  v a r y from  Birkett,  the  nutrient  in  that  feces  collected  1979)  By  marker to  intake  not  1979;  1982).  the  digestibility,  made t h a t  such  of  and n i t r o g e n o u s  to  with  in  Difficulties  1955;  an a p p a r a t u s  the  and i n a v o i d i n g  soluble  separate  of  food  are  from the  fish  and c h l o r o f o r m .  (1971,1976)  measure  These  1979),  o r i g i n and t h a t  developed  the  feces  feces  assumption  precipitate,  from  feces  (Gerking,  and S p a n h o f f ,  methods fecal  siphoning  nitrogen  metabolism  to  excretions  energy.  Pfeffer,  same r a t i o  C h o u b e r t et  of  1981;  digestibility  determination  and m e t a b o l i z a b l e scooping  and the  having  Cho and S l i n g e r ,  concentration  the  and g i l l  al.,  can be c a l c u l a t e d .  representative  Total  the  1973;  the  Smith separate  However,  determination  of  the the  nutritive  value  considerable Data  on the  foodstuffs of  of  feedstuffs  stress  determined  of  The u t i l i z a t i o n  utilization  as  of  well  (NPU) i s  for  fishes  its  the  efficiency method  is  animals  and Bender  product  of less  than  excretory nitrogen  nitrogen  is  the  products. content  resulting  over  It  is  resulting  the  in  of  tables  two  depend  Net  on  its  protein  values  :  digestibility)  the  procedure  contents  tabulated  value.  these  devised  for  balance.  (NRC,1981).  of  utilization  cumbersome  were under  nitrogen  energy  are  biological  (1953)  fish  by an a n i m a l w i l l  NPU = BV x (% Miller  the  negative  methods  a protein as  of  metabolizable  by t h e s e  requirements  digestibility  made when  and i n a s t a t e  digestibility  nutrient  was  NPU to  of  a test  the  the  protein.  determination  involving  based  assess  of  NPU i n  collection  on a c o m p a r i s o n of  from a t e s t  same p e r i o d of  protein  time  Their  from a  of  the  source  small  body  with  that  nitrogen-free  diet: body N of t e s t group  NPU =  _  N consumed Ogino suited  to  et  al  fish  (1980) studies.  by c a r c a s s  during  experimental  the  endogenous  body w e i g h t / d a y  loss  of  by t e s t  described  determined the  body N of group f e d a non-protein diet  a method  after  period. nitrogen  and changed  They in  that  loss  feeding found  is  of  particularly  nitrogen  was  a protein-free  diet  that  the  value  rainbow t r o u t  was  9.5  mg/lOOg  body  weight.  in direct  20  100  group  The endogenous  analysis  x  p r o p o r t i o n to  for  Accordingly,  NPU f o r  rainbow t r o u t  Body N g a i n of t e s t group (g)  W 1  +  Nitrogen Where:  =  ^2  intake i  n  i  t  i  a  = final  d = days By  this  readily  adapted  difficult can  task  of  of  efficiency gained  to  studies  for  protein  1979;  protein  in  fed  utilization  with  fish  g  of  nitrogen  method  sufficiently proteins  has  the  would  when c o n d u c t e d  conditions  (Cowey  make a l l o w a n c e s  from the  determination  be more a c c u r a t e  losses  be  analysis.  wastes  The water of  because  i n c l u d i n g the  n u t r i t i o n have by the  1971;  This  protein  has  to  of  provide  under  been  maintenance  al.,  difficult  being  1973; Higgs  et net  (PPV): x  100  and  ratings  to  various  experimental  However,  requirements  21  protein  apparent  simple  relative  body  1977;  value  the  al.,  termed  gained  standardized 1979).  of  Z e i t o u n et  productive  advantage  estimated  ratio  H i g u e r a et  and S a r g e n t ,  for  can  losses.  De l a  reproducible  balance  by c a r c a s s  (%)PPV = c a r c a s s p r o t e i n p r o t e i n fed This  100  i ht  nitrogenous  (Gerking,  1982).  x  weight  fish  utilization  or  w e  xd  feeding  nitrogen  1975;  Pfeffer,  y  follows:  -3  F u r t h e r m o r e , the  all  protein  Rumsey and K e t o l a , al . ,  of  insensible  workers of  d  body n i t r o g e n  measuring  Several  o  body  recovering  and f i n a l  b  as  x 9.5 x 10 100 group(g)  test  principles  avoided.  would a c c o u n t  task  the  to  be e n t i r e l y  initial it  method  l  calculated  W 2 (g)  + 2 of  NPU =  was  PPV does  and assumes  not that  all  food  growth  the  protein  is  used  demand p r o t e i n , (1959)  proposed  rate  change  of  to  of  compare  an a s s a y  with  gain  on n i t r o g e n  with  rats  nitrogen These  fed  intake  are  the  Slope-ratio  assay  without  of  1965a;  2.6  use  Samonds  The  protein  in  fish  protein  is  chiefly  have fed  the  respect  of  of  undertaken  been  to  with  1965b).  critically  linearity  and  curves.  proposed  both w i t h  (Hegsted  and  with  and  Chang,  M c L a u g h l a n and K e i t h ,  salmonids  weight  obtained  and Chang  and have  be  described  slope  also  measuring  intake  regression  of  group  line  (1965a)  (Hegsted  been  1977;  nutrition.  equal  of  the  dietary  70% of  the  energy  form of  protein  by i t s  the  importance  are  has  been  As m e n t i o n e d  determined  Perhaps  lipid  the  dose-response  requirement  requirements  the  gain  the  protein  amino a c i d  About  the  priority.  protein  method was  with  requirements  studies  programs  of  and  respect  1977).  to  energy.  gross  of  of  of  and Chang  assays  (1979)  and H e g s t e d ,  The p r o t e i n dietary  This  first  with  a percentage  protocols  a zero  slope  balance  weight  intercepts  the  slope  slope-ratio  by M c L a u g h l a n  of  the as  replacing  methods  origin  using  As b o t h m a i n t e n a n c e  receives  Hegsted  lactalbumin.  gain  reviewed  that  nitrogen  proteins.  rats  growth.  maintenance  Allison  used  in  for  in  a n i m a l under the  in natural  design  ( G u l b r a n d s e n and U t n e ,  of  1977).  22  the  of  to  the  of  many  value  satisfy  of  the  consideration.  of  of  foods  subject  earlier,  ability  concentrations  and most  the  economic  protein salmon  remainder  is  and is  feeding energy. provided  supplied  Most c o m m e r c i a l  salmon  by  a  diets, and  however,  lipid.  energy  contain  There i s  less  of  considerable  the  total  reliance  energy  as  protein  on c a r b o h y d r a t e as  an  source.  The o b j e c t i v e minimum amount Delong with  far  et  of  was  protein  protein  al.(1958)  chinook  protein  of  supplied  composition  simulating  composition  was  (dextrin) isocaloric period  formulated protein  et  that to of  diets  40 and 55% when water  the  to  temperature  was  8.3  levels  of  carbohydrate  the  diets  a ten-week gains  feeding  of  protein  and  This  indispensable  maintain  contained  acid  protein.  of  weight  the  and  different  After  studies  amino  egg  the  growth.  i n which  digestible  that  diet  find  these  essential  containing  found  to  gelatin,  an e x c e s s  basis.  of  a diet  chicken  by s u b s t i t u t i n g  when  first  casein,  whole  contain  al.(1958)  salmon were h i g h e s t  the  of  is  p r o d u c e maximal  an o v e r a l l of  energy  studies  were f e d  i n an a t t e m p t  on a g r o s s  Delong  with  thought  A series  for  of  by a m i x t u r e  amino a c i d s  was  one  The f i s h  crystalline  protein  r e q u i r e d to  conducted  salmon.  amino a c i d s .  requirement  chinook  at  levels  of  found  that  the  14.5°C  respectively. Employing  similar  minimum p r o t e i n 10 or  20 ppt  protein directly as  1973).  requirement  salinity  requirement related  salinity  was  Satia  requirements  diets,  to  of  of  coho  al.(1974)  salmon m a i n t a i n e d  approximately  juvenile  water  raised  (1974) of  was  Z e i t o u n et  rainbow t r o u t  salinity  from 10 to  conducted  40%.  ppt  a study  a p a r t i c u l a r genetic  23  to  the  was  be  (Zeitoun  found  of  to  from 40% to et  determine  strain  either  By c o n t r a s t ,  and i n c r e a s e d 20  at  45%  al., the  rainbow  protein trout.  This  strain  Olson, to  1957).  27°C  from  is  30% to  over  data, of  20g  to  of  the  the  the  to  for  lost  that  value  for  kcal/g excrete  ingested  of  all  values  of  lipid  16  by  on a  The f i s h  the  of  be i s o c a l o r i c  Based  the  were  fed  on food *•  protein A0% as  1979).  in  these  is  than  the  fish  The m e t a b o l i z a b l e  of  each  protein  should for  are  case  and the  a more a p p r o p r i a t e  the  frequently  This  is  because  for  ammonia r a t h e r  carbohydrate  the  -  2A  than  picture  -  which  is  Smith  a lack  then for 1969; of  employed (1971)  has  metabolizable used fish  that  value  of  of  energy 3.9  primarily  energy  by the  is  is  (Phillips,  values  non-metabolizable  be a c c o u n t e d  estimated  protein,  there  For e x a m p l e ,  more  of  products  energy  usually  component  the  procedures  the  studies  by i n d i v i d u a l  Unfortunately,  (1969). Thus,  foregoing  and c a r b o h y d r a t e  content  ammonia.  to  to  and  content  from f i s h m e a l  day.  assigned  feedstuffs.  A.5 k c a l / g  respect  protein  d r o p p e d from 50% to  excretory  value  in  1969).  per  nitrogenous  Phillips  combustion  protein  and i n  protein  of  temperatures  showed t h a t  between i n v e s t i g a t o r s .  shown  high  formulated  digestibility  in  (Donaldson  ranging  (Phillips,  (197A)  trout  energy  standardization  with  to  body w e i g h t  the  and G r o v e s ,  differ  were  at  rate  weight.  protein,  corrected  diets  basis  caloric  gross  growth  were r e a r e d  uncertainty  investigators  Brett  fast  relation  Satia  in  The major relates  in  A.5% of  requirements  energy  its  The d i e t s  energy  of  conversion  from  fish  dextrin  a rate  values  These  50%.  metabolizable  grew  for  and they were f e d  adjusting  at  noted  heat  fraction  of  of  urea.  complicated  Also, further  because the  for  diet  Hence,  example,  result  the  lowered  metabolizable  components  metabolizable  Jobling energy  composition,  dry  efficiency diet  Zeitoun  has  (Delong, et  al.,  also  1976).  the  promote  protein  the  sparing  amino a c i d s  to  to  eat  1973;  satisfy  Brett  (1979)  have  et  utilization  and of  for  energy  and G r o v e s ,  1979).  stated  since  studies  is  relation  that  energy,  best  be e x p r e s s e d  of in  to  terms  as  of  energy  1974;  upon  protein both  as  the  have  p r o p o r t i o n of  Lee  the  diet  a of  energy shown  Sargent of  protein  content  the  and Putman,  objective  dietary  protein  of  been  Cowey and  ultimate  of  al.(1975)  a source  (Lee  or  1975;  (1971),  and have  Fish  gain  Cowey,  components  Recently,  the  only  a percentage  Ringrose  requirements  the  dietary  not  and Cowey et  dietary  estimate  dietary  estimated  depend  Satia,  (1969),  the  the  1967).  optimum w e i g h t  synthesis.  their  of  in  level.  stated  Protein acts  requirement to  of  and N o s e ,  various  that  for  1971;  or d e x t r i n  level  will  al.(1975)  tissue  that  out  level  been  the  the  upon r a t i o n  non-protein  effect.  of  a diet  Phillips  have can  of  Luquet,  Takeda  that  pointed  starch (Singh  vary with  generally  1958;  of  values  protein  and Putman ( 1 9 7 3 ) , reported  to  (1983)  The minimum d i e t a r y feed  energy  content  but  amounts  digestibility  can be e x p e c t e d  carbohydrate.  diet  in  increased  of  protein level  in  the  diet  can  energy  that  it  contributes. Previous best  ratio  diet  is  of  0.50  Gulbrandsen  work w i t h  chinook  salmon  protein  energy  to  (Combs et  al.,  1962;  and Utne  (1977)  total  indicated  energy  Fowler  reported  25  has  that  et  that  (PE:TE) al.,  dietary  in  the the  1964). PE:TE  ratios  varying  between 0.37  rainbow  trout.  feeds  for  content  the  approximately oil,  higher other  employed  20%. for  The l i p i d 40% to  this  studies  digestible in  caused  on f i s h ,  component,  42% of  the  diet  maximal  practical  high PE:TE  the  (1977)  The  level  of  (dextrin)  supplied  in  and c o n s e q u e n t l y  kept  the  lipid  energy levels  U n l i k e most  furnished  by  at  the  20% of  energy  was  capelin  lipid  growth.  energy  was  that  of  dry  (1977)  by  dietary  found  growth  Norwegian  ratios.  total  a depression  carbohydrate the  that  for  by G u l b r a n d s e n and Utne  G u l b r a n d s e n and Utne than  energy  noted  had e x c e s s i v e l y diet  accounted  content.  were o p t i m a l  They f u r t h e r  trout  of  and 0.41  supplied  total by i  protein  was  balanced  Whether  protein  carbohydrate protein 1979;  could  be  is of  replaced great  (1969)  to  carbohydrate former  the  omnivorous  al.,  to  1979).  juvenile was  increased  salmon  decreased 43%.  were i n c r e a s e d  high  from l i p i d to  They found  by s u b s t i t u t i n g  outcome  (Shimeno  not  able  are  14% of  that  the  of  of  to  have  diets  protein  dextrin  for  from of et  a al,  of tolerate  diet. fish  are  utilize shown  carbohydrate (1961)  i n which  40% and l e v e l s  that  to  carnivorous  ability  levels  26  or  fish  B u h l e r and H a l v e r  from 71% to  the  evidence  some e x p e r i m e n t s  a series  oil.  statement  their  dietary  fish  the  above  in  For example,  chinook  to  fish  of  supports  indicating  effectively,  do a d a p t  1982)  levels  reports  by energy  Recent  carnivorous  carbohydrate  Despite inferior  that  levels  significance  experiment.  H i l t o n and A t k i n s o n ,  digestible  level  energy  requirement  Phillips  et  by v a r y i n g d i e t a r y  of  the  the  (Shimeno fed protein  dextrin  efficiency protein  that  were  ratios  whereas  growth  rate  dietary that  remained s i m i l a r  treatments.  elevating  while  S i m i l a r l y , Bergot  the  dietary  demonstrated  action  in  fish,  optimal 1971;  fish  also  carbohydrate  spares  in  difficult occur.  owing  gained  ratio  the  reducing  studies (Lee  1977; concept  to  the  energy  different  Therefore,  the  protein  fish  trout  improved  these  in  experiments  a protein extent to  of  determine  1973; al.,  sparing  Ringrose,  1953  some e x t e n t  the  in  diets  dietary  protein.  the  in  in  resulted  directed  to  found  different  30%  and Putman,  that  the  from 15% to  the  and Cowey et  to  be c o n t i n u o u s l y  intake  Therefore,  proper d e f i n i t i o n  relation  (1979a)  glucose  protein  through  Additional  support  receiving  c a r b o h y d r a t e may have  and W i l s o n ,  In summary, protein  that  protein/energy  Garling  of  utilization.  perhaps  gluconeogenesis.  of  level  maintaining equivalent  growth and p r o t e i n have  among f i s h  re-evaluated  as  of  the  content nutrient  correct of  nutrition.  27  a salmonid  interactions  requirements more  level  of  fish  fundamental  of diet  is  which  can  may have  knowledge  is  to  CHAPTER 3  EXPERIMENT 1  3.0  Protein in  3.1  utilization  diets  for  evidence  hatchery-reared their  1980;  during best  size  Bilton  juvenile  salmon  the  at et  of  protein  quality  fry.  critical  fish  culture.  understanding nutritive  value  enhance  Fishmeal salmonid essential  is  diets,  for or  minerals.  entry  (Fowler  their their it  The p r o t e i n  potential through  recognized  fish  the  the the  Columbia  foods  of  an  highest conditions  quality. protein  protein  of  to  in  commercial  supply  lipids,  fishmeal  is  salmon  was  and how p r o c e s s i n g  of  is  In  salmonid  objective  have  that  feeds.  effectiveness  contributes  28  that  in B r i t i s h  first  provides  quality  essential  comprising  source  also  al.,  history  profitable  related  et  growth  well  sources  protein  only but  is  the  salmon  is  life  farming  of  protein  it full  and c o s t  principal  amino a c i d s ,  seawater  proteins  chinook  and not  directly  study,  diminish  the  is  salmon  of  this  which  ocean  of  performance  of  the  in  supply  quality  in  of  Also,  of  on the  Hence,  survival  realize  nutrition.  the  the  Therefore  stage  development  to  of  1982).  salmon  dependent  that  salmon  time  al . ,  particular,  either  the  freshwater  possible  partly  suggests  chinook  chinook  successful  and  chinook  measurement  Introduction Recent  to  and the  vitamins,  presently  available This  in  British  v a r i a b i l i t y is  (origin,  species,  conditions),  fishmeal  Both  of  these  with  balance.  acid  profile  compared  In of  requirements noteworthy, in  a protein  of  conditions.  would  contribute  with  The q u a l i t y compared  to  that  certain  of of  this  drying,  appetite observed muscle  a l .( 1971 , 1972)  and  d r i e d under was  be m a n i f e s t e d  of  the  a mix of  29  marine amino  amino  it  acid  is  excellent cod muscle  as  different a common b a t c h  of  different  made t h a t by l o s s e s  damage in  due  to  the  lysine,  and  sources  was  this  quality.  foregoing casein  of  especially  protein  the  amino  obtained  quality,  of  euphausids  freeze-dried  amino a c i d s ,  a  was  and i m p r o v e  Also,  effects  assumption  pollock  that  1).  on p r o t e i n  for  to  a species  (Table  the  final  into  selected  freeze-dried  salmon  decreased all  were  euphausids,  using  the  processed  essential  and t h e n  of  M o r e o v e r , the  pollock  storage  grinding,  conditions  were  was  raw m a t e r i a l  study.  promote  plaice  the  scraps,  stated  alone,  cooked  to  fish  of  A tentative  of  or  considerably.  of  commonly  muscle  to  that  d u r i n g d r y i n g would  availability  in  To e v a l u a t e  methods  raw h e r r i n g was  to  Cowey et  source.  are  regard i t  chinook  studies  processing  heat  value  freeze-dried  that  nature  (cooking,  p o r t i o n of  this  favourably  vary  h e r r i n g and p o l l o c k  i n an a t t e m p t  acid  fish  availability.  a small  zooplankton,  the  and s t o r a g e  species  nutritive  amino a c i d  of  methods  h e r r i n g and p o l l o c k  maintain  results  fish  known to  whole  stabilization)  and of  comparison  blended  season,  Several  is  a function  processing  antioxidant product.  Columbia  protein  and g e l a t i n  fortified  with  Table  1. A m i n o a c i d amino a c i d casein.  requirements f o r chinook salmon (NRC.1981), composition of f r e e z e - d r i e d p o l l o c k muscle,  Amino  acid  Requirements (NRC, 1981)  R  amino  composition o f e g g s and t h e a n a l y z e d essential f r e e z e - d r i e d whole euphausids and v i t a m i n - f r e e  acid  / 16g n i t r o g e n "  1 Freeze-dried eyed chinook eggs  3 Freeze-dried p o l l o c k muscle  3 Freeze-dried whole euphausids  Vitamin-free casein  Arginine  6 .0  6 .39  9,.30  5 .09  3 .83  Histidine  1..8  2,.88  2. .15  2 .00 ,  2 .95  Isoleucine  2, .2  5..94  .34 4 .  3. .65  4 .10  Leuc i n e  3, ,9  9..74  8..61  5,.81  7 .60  Lysine  5,.0  8,.65  12 .,92  6,.17  7,.39  2, .51  4,.10  4..27  3, .01  Methionine Cystine2  4,.0  Phenylalanine  1 .56 ,  1 .,68  0,.99  0,.74  5..39  4.,39  3, .74  4,.91  Tyrosine2  5.. 1  4,.62  4.,13  3. .60  5,.59  Threonine  2 .2  5,.02  5.,13  3. ,55  3, .83  Tryptophan  0 .5  1 .42  1 . .22  1 , .00  1 .23  Valine  3 .2  7 .32  4,.26  3, .75  5 .39  1. 2. 3.  D e t e r m i n e d by A . A . A . L a b o r a t o r y , S e a t t l e , W a s h . C y s t i n e and t r y o s i n e a r e d i s p e n s a b l e , but spare r e q u i r e m e n t s f o r methionine and p h e n y l a l a n i n e r e s p e c t i v e l y . D e t e r m i n e d a t t h e Department o f A n i m a l and P o u l t r y S c i e n c e , U n i v e r s i t y o f S a s k a t c h e w a n , S a s k a t o o n , Canada.  arginine  and m e t h i o n i n e  requirements because  of  chinook  a blend  of  source  fish  in  based the  case  casein  and g e l a t i n  permit  direct  and T i e w s , Owing  the  of  the  goal  second of  assessment  food  been  juvenile  been fish  salmon and  extensively  (Cowey,  1976;  results  has  be  diets to  study  Halver,  diet  been  to  other  and i n d e e d ,  used  source  done  considered  a standard reference  of  this  several of  the  containing  proposed  among l a b o r a t o r i e s  nutritive the  estimates  were  made of  groups  fish  during  of  protein  approach  enabled  utilized  for  for  in  was  to  (Castell  to  value  protein.  endogenous the  protein  of  growth  ingesting  31  the  quality this  loss  versus used  lost  of  in  diets. at  several  goal,  by t h e  different  permit  growth.  amounts  of  1979),  which c o n s i d e r  as  and t h a t  proteins  fish  p e r i o d to  well the  merits  in  As p a r t of nitrogen  as  for  procedures  protein  by methods  maintenance  the  bioassay  experimental  quality  maintenance,  assess  of of  techniques  (Cowey and S a r g e n t ,  quality  determination  by f i s h  fish  evaluation  dietary  protein  standardized  study  protein  of  measurement  of  proteins  concentrations  excretions  of  a protein  paucity  necessitated  dietary  has  was  1980) . to  of  for  have  c o m p a r i s o n of  evaluation  demerits  as  for  amino a c i d This  nutrition studies,  and g e l a t i n  the  essential  and g e l a t i n  n u t r i t i o n a l requirements Also,  the  (NRC, 1 9 8 1 ) .  protein  previous  on c a s e i n  1982).  This  of  meet  salmon  casein  an e x c e l l e n t species  to  use  This  protein  from  exogenous  varying quality.  of  3.2  M a t e r i a l s and  3.2.1  Test Frozen  partially grinder  protein pollock  plate),  (Fitzmill, U.S.  sources. ( T h e r a g r a chalcogramma)  thawed,  comminuted  loosely  freeze-dried.  size  methods  model 20  J T ) so  sieve.  pacifica)  were p r e p a r e d  compositions  of  respectively:  that  of  16.45%;  of  the  on a d r y w e i g h t  muscle  and whole  ( C l u p e a harangus  (-20°C)  month w h i l e  being  (FRH) except of  was  ethoxyquin  0.005% b e f o r e  fish  meal  Organization,  prevent  The  proximate were  4.97%  and  89.74% and  with  one  60.01%.  part  freeze-dried  12.10%;  of  the  pollock  caught  on J a n u a r y 8,  transported plastic  bags  Freeze-dried  a similar  manner  added  to  to  the  batch  on i c e for  whole the  mixed  of 1981,  and  not  off  kept  more  than a  raw h e r r i n g  pollock  muscle  h e r r i n g at  a  level  freeze-drying.  The h e a t - d r i e d pilot  to  p r e p a r e d from a s i n g l e  pallasi)  was  100% t h r o u g h a  (FPE).  in a i r t i g h t  in  ground  (Euphasia  ash,  protein,  produce  meal  processed.  prepared  that  to  The h e r r i n g was  deep f r o z e n  pass  mm  and  and e u p h a u s i d s  were combined  basis  euphausid  B.C.  was  manner.  and 3.40%;  and c r u d e  The h e r r i n g meals were  Ladysmith,  would  euphausids  pollock  1.80%  former  latter  herring  particles  (6.35  refrozen  muscle  i n an i d e n t i c a l  freeze-dried  and  trays,  pollock  Whole f r o z e n  moisture,  2.01%  parts  over  were  a meat g r i n d e r  E t h o x y q u i n (,025%)was added  rancidity.  Nine  spread  The f r e e z e - d r i e d  oxidative  lipid,  through  fillets  h e r r i n g meals were  m a n u f a c t u r i n g machine  Esbjerg,  Denmark).  32  prepared  in a  (Chemical  The machine  continous  Research  consists  of  a  steam-jacketed dryer.  cooker,  The t e m p e r a t u r e  controlled. pressing, aqueous  Minced,  the  oil  fraction  steam-jacketed to  the  at  (LTH).  The h i g h in  150°C  for  extract  one  solvent  in to  wire  times. mesh  remove added  any to  through  0.57%  of  hexane  a size  it  was  the  30% s o l i d s  solubles  were set  in a  added to  d r i e d h e r r i n g meal  and f u r t h e r the  with  U.S. of  (HTH) was oven-dried  mainly at  LTH and HTH p r o d u c t s necessary  hexane  stand  filtered  paper.  (5:1  residual  for  to  partially  v/w)  to  of  30 m i n u t e s ,  t h r o u g h a Buchner  This  h e r r i n g meal  permit  p r o c e d u r e was products  ambient  air  were for  the funnel  repeated placed  five  E t h o x y q u i n (0.025%)  extracted  products  w h i c h were  20  screen  with  the  fish  a hammer m i l l .  protein  to  was  ground  T a b l e 2 shows  sources.  mix s u p p l e m e n t e d a modified  by NRC ( 1 9 7 3 ) ( S w a r o k  33  then  in  hours  hexane.  D L - m e t h i o n i n e (CS) was outlined  to  a current  casein-gelatin  Diet  both  After  and  The d r y e r was  temperature  120°C  meals  filter  traces  the  approximately  be  75°C.  decanted  d r i e d h e r r i n g meal  lipid,  s l u r r y was  over  compositions The  low  at  rotary  formulation.  The f i n a l  trays  to  was  rotary dryer.  m i x i n g and a l l o w i n g  and meal  cooked  The condensed  Because of  and d r y e r can  liquid  condensed  from the  and d i e t  cooker  press  produce  hour.  and s t e a m - j a c k e t e d  h e r r i n g was  temperature  Whatman N o . l  three  the  the  an e x c e s s  After  whole  the  r o t a r y d r y e r at  lipid  grinding  Test  75°C  the  contained  the  was  presscake  dried  in  of  press  bowl c o o k e r .  operate  over  screw  with  1.4%  version  of  and H i g g s ,  L - a r g i n i n e and the  Oregon  1981).  Table  2.  Composition of protein sources (values i n parenthesis and l y s i n e e x p r e s s e d on a d r y m a t t e r b a s i s ) .  show  percentages  % Code  % Moisture  % Protein  % Lipid  %  Ash  f o r each  Available lysine  proximate  constituent  Available lysine (% o f c r u d e protein)  Freeze-dried pollock muscle & freeze-dried euphausids  FPE  1 .96  8 6 .72 ( 8 8 .45)  3,.43 ( 3 ,. 5 0 )  5.69 (5.80)  7 .42 (7 .57)  8 .56  Freeze-dried raw h e r r i n g (hexane extracted)  FRH  8 .11  7 0 .56 ( 7 6 ,.79)  1 0 ,.48 ( 1 1 ..40)  11 .49 (12.50)  3 .86 (4 .20)  5,.47  Low t e m p e r a t u r e d r i e d h e r r i n g meal (hexane extracted)  LTH  7 .77  6 3 , .95 ( 6 9 ,.34)  1 6 . .98 (18. 41)  10.21 (11.07)  3 .45 ( 3 ,. 7 4 )  5,.39  H i g h t e m p e r a t u r e d r i e d h e r r i n g HTH meal (hexane extracted)  3,.83  7 2 . .13 ( 7 5 . ,00)  1 3 .,12 ( 1 3 . ,64)  11.15 ( 1 1 .59)  3,.28 ( 3 ,. 4 1 )  4..55  2.,61  9 0 . ,82 (93.. 2 5 )  6.,69 ( 6 ., 8 7 )  7 . ,37  2 Casein  T. 2.  mix  90T" f r e e z e - d r i e d p o l l o c k C o n t a i n s 8 8 % c a s e i n , 10%  CS  a n d 10% fr e e z e - d r i e d euphausids g e l a t i n , 1.4% L - a r g i n i n e , 0.57%  on a d r y w e i g h t basis. DL-methionine (I.C.N., St. Louis,  HO.)  Table  3.  Composition  of d i e t s  Diet  Ingredients  PF 0  Code  FPE 7  79. 14  Freeze-dried pollock-euphausid  (g/kg d i e t  (Protein  FPE 17  on  source  FRH 17  221 . .38  HTH 17  protein)  CS 17  FPE 27  FRH 27  305 . .26  351 .61  -  -  H i g h temp d r i e d h e r r i n g meal Casein-gelatin, a r g i n i n e , and m e t h i o n i n e mix  Mineral  LTH 17  % dietary  245 . 17  Low temp d r i e d h e r r i n g meal  Herring o i l ^  and  basis).  -  192. 20  Freeze-dried raw h e r r i n g  a dry matter  226 .67  182 .31  -  127 . 22  123 .22  104, .76  83. 7  79 .6  71 . 1  50, .4  50 .4  85 .5  69 .24  85 .5  91 . 66  87 . 77 92 . 05 105 .,49  130 .0  mix^  Ground cellulose  84 .186  99 .08 50 .4  130 .0 83 .7  119 ,, 32  89, .92  62 , 7  28 , . 1  127 . 52 110 .57  Dex t r i n  346 .0  307 ..0  250 .5  250 .5  250 .5 250 . 5  250 .5  194 ,.5  194 .5  Glucose  346 .0  307 .0  250. 5  250, .5  250 . 5 250 . 5  250 . 5  194 , 5  194 ,.5  C a r boxy-me t h y 1 cellulose ... Vitamin  .3 mix  Choline chloride ( 50%) 1. 2.  3.  20  20  20  20  20  20  20  20  20  7 .3  7 .3  7 . .3  7 .3  7 .3  7. 3  7 .3 .  7 .3 ,  7. 3  3. 5  3 .5  3. 5  3 .5  3 .5  3 .5  3 .5  3 .5 ,  3,.5  S t a b l i z e d w i t h BHA-BHT ( 1 : 1 ) 0 . 3 3 % . M i n e r a l mix f o r m u l a t e d so t h a t e a c h d i e t c o n t a i n e d ( g / k g d r y d i e t ) : Ca 12.6, P 9.0, Mg 1.8, Fe 0.2, Na 2.4, A l 0.05, Cu 0.01, Mn 0.03, Co 0.01, Zn 0.08, K 8.3, I 0.004, Na:K 0.29, Ca:Mg 7, P:Mg 5, Ca:P 1.4. C a l c i u m l e v e l s i n FRH, LTH and HTH c o n t a i n i n g d i e t s w e r e c a l c u l a t e d t o be 13.9 and 19.6 g / k g f o r d i e t s c o n t a i n i n g 27 and 3 7 % p r o t e i n . Similarly, p h o s p h o r u s l e v e l s were 10.0 and 14.0 r e s p e c t i v e l y t o m a i n t a i n equivalent Ca:P i n a l l d i e t s . S o u r c e s o f m i n e r a l s were p o t a s s i u m phosphate monobasic, potassium phosphate d i b a s i c , sodium phosphate monobasic, c a l c i u m phosphate d i b a s i c , c a l c i u m c a r b o n a t e , magnesium s u l p h a t e , f e r r i c o x i d e , z i n c o x i d e , sodium c h l o r i d e , manganese s u l p h a t e , c o b a l t c h l o r i d e , c u p r o u s c h l o r i d e , potassium i o d i d e and a l u m i n u m c h l o r h y d r a t e . V i t a m i n mix t o s u p p l y (mg/kg d r y d i e t u n l e s s o t h e r w i s e i n d i c a t e d ) : a s c o r b i c a c i d 1200, i n o s i t o l 400, n i a c i n 300, C a - p a n t o t h e n a t e 150, r i b o f l a v i n 6 0 , m e n a d i o n e 8 0 , p y r i d o x i n e HCL 30, t h i a m i n HCL 30, f o l i c a c i d 20, b i o t i n 3, v i t a m i n A 10,000 I U , v i t a m i n D3 1,000 I U , v i t a m i n E 600 I U .  35  Table  3.  cont'd  Diet  Ingredients  LTH 27  Code  HTH 27  (Protein  CS 27  Freeze-dried raw h e r r i n g  FRH 37  LTH 37  389 .39  533 .60  -  -  58 .31  80.90  CS 37  —  360.0  Casein-gelatin, a r g i n i n e , and m e t h i o n i n e mix  protein)  HTH 37  481 . 83  -  H i g h temp d r i e d h e r r i n g meal  Herring o i l '  FPE 37  and % d i e t a r y  418 . 32  Freeze-dried pollock-euphausid  Low temp d r i e d h e r r i n g meal  source  531.37  -  -  -  -  493 . 33  .54 289 .  130  396 .78  115. 36  75 .07  FPE 47  31 . 76  •  .  83 .7  71  130  -  Ill.40  83, . 7  56 .00  17 .04  17, .4  17 .4  Ground cellulose  104 .40  11 .20 132 , .56  142 . 12  118. 90  .44 110 ,  119 . 76  149 .02  Dextrin  194 . 5  194 .5  194 ..5  138  138  138  138  138  82  Glucose  194 .5  194 . 5 194 ,.5  138  138  138  1 38  138  82  20  20  20  20  20  Mineral  mix^  Carboxy-methyl cellulose  28 . 1  20  28 . 1  20  20  20  47.4 115.43  3 7 .3 ,  7.3  7 .3 .  7 .3  7 .3  7 . .3  7. 3  7 .3  7.3  C h o l i n e c h l o r i d e 3,.5 ( 50%)  3.5  3 .5 .  3 .5  3. 5  3 ,5 .  3 .5.  3 .5  3.5  Vitamin  1. 2.  mix  S t a b l i z e d w i t h BHA-BHT ( 1 : 1 ) 0 . 3 3 % . M i n e r a l mix f o r m u l a t e d s o t h a t e a c h d i e t c o n t a i n e d ( g / k g d r y d i e t ) : Ca 1 2 . 6 , P 9.0, Mg 1.8, Fe 0.2, Na 2.4, A l 0.05, Cu 0 . 0 1 , Mn 0.03, Co 0 . 0 1 , Zn 0.08, K 8.3, I 0.004, Na:K 0.29, Ca:Mg 7, P:Mg 5, Ca:P 1.4. C a l c i u m l e v e l s i n FRH, LTH a n d HTH c o n t a i n i n g d i e t s were c a l c u l a t e d t o be 13.9 a n d 19.6 g / k g f o r d i e t s c o n t a i n i n g 27 and 3 7 % p r o t e i n . Similarly, p h o s p h o r u s l e v e l s were 10.0 a n d 14.0 r e s p e c t i v e l y t o m a i n t a i n equivalent Ca:P i n a l l d i e t s . S o u r c e s o f m i n e r a l s were p o t a s s i u m phosphate monobasic potassium phosphate d i b a s i c , sodium phosphate monobasic, c a l c i u m phosphate d i b a s i c , c a l c i u m c a r b o n a t e , magnesium s u l p h a t e , f e r r i c o x i d e , z i n c o x i d e , sodium c h l o r i d e , manganese s u l p h a t e , c o b a l t c h l o r i d e , c u p r o u s c h l o r i d e , potassium i o d i d e and aluminum c h l o r h y d r a t e . V i t a m i n mix t o s u p p l y (mg/kg d r y d i e t u n l e s s o t h e r w i s e i n d i c a t e d ) : a s c o r b i c a c i d 1 2 0 0 , i n o s i t o l 4 0 0 , n i a c i n 3 0 0 , C a - p a n t o t h e n a t e 150, r i b o f l a v i n 6 0 , m e n a d i o n e 80, p y r i d o x i n e HCL 30, t h i a m i n HCL 3 0 , f o l i c acid 20, b i o t i n 3, v i t a m i n A 10,000 I U , v i t a m i n D3 1,000 I U , v i t a m i n E 600 I U .  t  3.  36  3.2.2  Diets  The 3.  Each  source and  formulae  of  protein  of  the  source  protein.  37% of  two  diets  protein  groups  of  fish.  to  diets.  all  mineral  kg  glucose. glucose  a non-protein  diets  diets  were  metabolizable  energy  employed  considered  of  in  Lastly,  95% f o r energy  gross  for  to  to  fish  each  diets  was  derived  for  values  of  the  (Cho and  protein  (Smith,  diets  common  4000  4.5  the  dextrin  (5.66  to  be  Slinger, the  kcal/g  heat  of  kcal/g)(Brett coefficient  1971)  based  and  and  found  by d e d u c t i n g (0.95  kcal/  kcal/g  dextrin  been  and a p p l y i n g a d i g e s t i b i l i t y casein-gelatin  of  oil,  has  ammonia n i t r o g e n of  two  equalize  contain  kcal/g  trout  energy  of  3).  and 4 . 0  for  and F P E .  on a n a l y s i s  The are  4. groups  of  fish  received  Oregon M o i s t P e l l e t s  (0MP) which  for  salmon  chinook  offered  (ME) v a l u e s  because  protein  excreted  1979)  two  valid  rainbow  for  from the  in Table  was  formulated  values  calculated  levels  endogenous and  diet  (Table  The h i g h  of  FPE at  p r e p a r e d to  all  27%,  An  were  of  sole  17%,  supplements  lipid  and G r o v e s ,  (N x 6 . 2 5 ) .  estimate  kcal/g  combustion  contain  prepared containing to  the  Mineral  The v a l u e  shown  protein  to  as  proportionately  1979).  protein  crude  diets  in Table  were  digestible  protein)  were f o r m u l a t e d  9.5  are  the  shown  ingredients  highly  of  loss  experimental  protein,  as  are  Basal  by a s c r i b i n g  crude  in  included  In o r d e r  composition  All  was  were  7% and 47% p r o t e i n . metabolic  diets  Diets  dry m a t t e r  additional  experimental  and coho  in  is  the  British  37  a commercial  diet,  standard hatchery Columbia.  Values  diet of  4.2  Table  k.  Proximate c o m p o s i t i o n of d i e t s l y s i n e and e n e r g y v a l u e s b a s e d  Analysis  PF 0  Crude p r o t e i n (Z N x 6.25)  1 .07 .  Available Total  lysine  crude  lipid  14 .08  Lipid contributed by f i s h m e a l Ash  7 .46  Digestible carbohydrate  69. .2  Moisture  9 .17  FPE 7  (% on  d r y - m a t t e r b a s i s ) and the a n a l y s e s .  calculated  FPE 17  FRH 17  LTH 17  HTH 17  16 .94  18 .01  16, .78  17 .00  0 .58  1 .45  0..99  0 .90  13..22  .40 12 ,  12 .46  13 , .33  11 .95 ,  (2. .52)  (4, .51)  (3, .09)  6 .67  7 .13 ,  7 .06 ,  6.. 73  (•.28)  (..67)  7..38  7 . 15  .4 61 . 10..44  50, . 1 7,.89  50, . 1 9 .20  50 , .1 8,.12  0,. 77  50 , .1 6,.10  CS 17  available  FPE 27  FRH 27  18 , .55  27 .24  27 . .59  1 .37 ,  2 .33  11 .61 ,  12..31  12 . .81  ( 1 , .07)  (4, .01:  7,.37  7 .34 ,  7 .73 ,  50 .. 1 .88 8 ,  38, .9 7,.72  1 ., 51  38. ,9 10, .95  Energy (kcal/kg) Protein: ME ( A . 5 k c a l / g )  48  303  762  811  755  765  835  1126  1242  GE  61  384  966  1027  956  969  1057  1553  1573  kcal/g)  1338  1256  1178  1184  1266  1135  1103  1169  1217  Carbohydrate: ME & GE ( 4 . 0 k c a l / g )  2768  2456  2004  2004  2004  2004  2004  1556  1556  Total  (ME)  4154  4015  3495  3998  4026  3904  3942  3951  4015  Total  (GE)  4167  4096  4148  4215  4226  4108  4164  4278  4346  (5.7  Lipid: ME S GE  mg  kcal/g)  (9.5  p r o t e i n / k c a l (ME)  Protein energy  energy:total (ME)  2 .58  0..01  16,,8  0..08  42, .9  0 , .19  45, .0  0 , .20  41 , 7  0..19  43, .5  0.,20  47 ., 1  0 . ,21  68, .9  0,.28  68. . 7  0.,31  Table  4.  cont'd.  Analysis  Crude P r o t e i n (% N x 6.25) Available Total  lysine  crude  lipid  Lipid contributed by f i s h m e a l Ash Digestible carbohydrate  LTH 27  HTH 27  28, .45  27 , .38  1 .53  1 . 25  11 .43 ,  .60 12 ,  (7, .17)  (4, .91)  7. , 78  7 .11 ,  38, .9  Moisture  38 .9  8,.36  6 . 28  CS 27  26 .40  FPE 37  FRH 37  LTH 37  HTH 37  CS 37  FPE 47  OMP  35 .96  39, .58  38, .42  39 .33  35, .99  46 .15  1 .95  3 .08  2 .17  2 .07  1 .79  2 .65  3 .95  12 .60  11 .79  12, .20  11 .81 ,  11 .29  11 .52 ,  12 .95  (1 .46)  (5, .49)  (9 .82)  ( 6 •73)  7,.61  8,.07  7 .50 ,  7,.84  7,.21  38 .9 8 .32  27 .6 6 . 72  27 , .6 9 .12  27 .6 8 . 19  27 .6 6 .73  4 9 . 40  14 . 53  (1 .86) 7 .30 ,  8 .30  27 , .6  16 .4  8 . 59  11 .92  16. 4  6.03  24. 62  Energy ( k c a l / k g ) Protein: ME ( 4 . 5 k c a l / g )  1281  1232  1188  1618  1781  1729  1797  1619  2077  GE  kcal/g)  1622  1561  1595  2050  2256  2190  2276  2051  2631  2816  (9.5 k c a l / g )  1086  1197  1146  1120  1159  1122  1073  1094  1230  2380  Carbohydrate: ME & GE ( 4 . 0 k c a l / g )  1556  1556  1556  1104  1104  1104  1104  1104  656  2662  Total  (ME)  3922  3985  3890  4044  3955  3974  3818  3818  3963  3721  Total  (GE)  4264  4314  4207  4274  4519  4416  4453  4249  4517  4462  (5.7  Lipid: ME S GE  mg  protein/cal  Protein energy 1. 2.  (ME)  energy:total (ME)  2075  1  2  72. 5  0. 33  Based on 4.2 k c a l / g p r o t e i n . Based on 1.6 X c a l / g carbohydrate.  68. 7  0. 31  67 .9  0. 31  93. 6  0. 42  97 . 9  0. 44  97. 1  0. 44  100 .5  0. 45  94. 3  0- 42  116..5  0.,52  132 . 8  0. 56  kcal/g  of  employed  p r o t e i n and 1.6 to  adequately in  this  estimate reflect  diet  the  the  (Table  ME c o n t e n t  feeds  were  mixer.  The mix was  CL-type  2 laboratory pellet the  hand  to  screened  remainder  of  then  were  the  herring  (needle  mill  oil  size  containers  during  the  respective  p r o t e i n sources  summarized  in Table  results  this  natural lights time  more  for  the  20 m i n u t e s  i n a Hobart  in a California 1.59  mm d i e .  sprayed,  into  the  by means  of  crumbles.  refrigerated  The d i e t a r y  code  in  to  The  a hand  The  diets  airtight  treatment  may be used  p i n and  crumbles.  codes,  and p r o t e i n c o n c e n t r a t i o n s  This  model  are  interpret  the  study.  facility  The e x p e r i m e n t a l  10.5°C  18),  study.  5.  oil  with  was  mixed t h o r o u g h l y and s t o r e d  fiberglass  were  p r o t e i n and raw s t a r c h  appropriate sized  were  Aquarium  of  they  crumbled with a r o l l i n g  syringe  3.2.3  OMP b e c a u s e  pelleted  held  of  of  extract  p r e p a r e d by m i x i n g  the  cold  pellets obtain  the  nitrogen-free  digestibility  w i t h a p o r t i o n of  Subsequently,  of  4).  The e x p e r i m e n t a l ingredients  kcal/g  tanks.  well-water.  aquarium  Each  (Vitalite,  tank c o n t a i n e d  Flow r a t e  p h o t o p e r i o d was  facility  was  4 to  contained 150  litres  40w)  of  rows  of  aerated  6 litres/minute/tank.  p r o v i d e d by a s e r i e s  Durotest  two  of  fluorescent  c o n t r o l l e d by an a s t r o n o m i c a l  clock.  40  A  Table  5.  Dietary  Summary of d i e t a r y  protein  source  Protein-free  treatments  Formulated protein concentration  0  and c o d e s .  Dietary treatment code  PF  Freeze-dried pollock muscle & f r e e z e - d r i e d euphausids  7 17 27 37 47  FPE -7 FPE--17 FPE -27 FPE -37 FPE -47  F r e e z e - d r i e d raw whole herring (hexane-extracted)  17 27 37  FRH -17 FRH -27 FRH -37  Low t e m p e r a t u r e d r i e d whole h e r r i n g meal (hexane-extracted)  17 27 37  LTH -17 LTH -27 LTH -37  High temperature d r i e d whole h e r r i n g meal (hexane-extracted)  17 27 37  HTH -17 HTH -27 HTH -37  C a s e i n - g e l a t i n mix, plus L - a r g i n i n e & DL-methionine  17 27 37  C S - 17 C S - 27 C S - 37 OMP  Oregon m o i s t p e l l e t s ( # 2 ) (complete diet)  41  3.2.4  Protocol  Chinook  salmon  fry  Qualicum Hatchery, for  uniform s i z e  the  selected  150  in  in  each  the  B.C.  with  of  38  the  During excess  with  a diet  except  that  protein  matter. 1.56 fed  of  by hand t h r e e of  feed  satiation  and second  in  case.  3.2.5  to  one  food  21  was  towel.  period  was  + 0.24.  selected Fish  into  from  groups  constituted  the  at  to  at  the  start  FPE-47 10% of  of  experimental  when  test  each  fed  the  feeding  were  daily  group.  The  feeding  ceased  The i n t e r v a l  between  the  feedings  was  two and one  size  adjusted  to  dry  was  fish  and a  3)  diet  "active"  was  to  (Table  period  satiated for  were  point over  60  the  and  half  hours  suit  fish  size  (1971).  fish  i n 0.5  feeding  were  trial,  following  randomly removed  ml 2 - p h e n o x y e t h a n o l / l  Individual fish  recorded.  42  weights  (to  from  16  g)  hrs  each  and p l a c e d 0.01  a  first  growth of  of  a block  were a l l o t e d  fish  50% and l i p i d  maintained  hour.  and 42  anaesthesized  fish  treatments  composition  F o w l e r and Burrows  of  1.03g  tanks  day u n t i l  particle  deprivation,  an a b s o r b e n t  per  and t h i r d  Food  Measurement  On day 0,  tank,  of  1900  from  row.  the  determined  second,  according  fish  During  times  period  each  in  comprised  intake  was  The 19 each  similar  of  of  row of  acclimation  Mean w e i g h t  feeding  of  within  +_ 0 . 0 2 4 g ( S E M ) .  record  Each  design.  a 14-day  of  1981  were r a n d o m l y d i s t r i b u t e d  tanks.  tanks  i n March,  A population  a mean w e i g h t  population  experimental  random to  were o b t a i n e d  on  were  3.2.6  Chemical a n a l y s i s  Four  samples  common to of  the  were  treatment  groups,  trial,  following  48  from each  analysis  at  Prior  -20°C  to  homogenized  used  to  1959)  nitrogen Diets  was  total  methods  Available  lysine  values  reported  a percentage  3.2.8  Data  The  of  in  crude  weight  data  conducted  as  fixed  fish  2 ml of  and s t o r e d  for  fish  of  homogenat'e  the  1975),  (Technicon  to  estimate  similar  protein  as  in Table  The a n a l y s i s  source  in  12  samples  lipid  were  (Bligh  Instrument  Co.  Percent  protein  content.  procedures.  sources  modified 2 as  protein,  was  by Booth  determined (1971).  a percentage i.e.  g/16g  of  by  The  the  meal  and  N.  analysis  body  dietary  the  method  (ANOVA).  The  of  end  lysine  (1960)  as  samples  thawed  (AOAC,  nitrogen  employing  Carpenter's were  At the  bags.  and ash  by 6.25  each,  day 0.  369-75 A / A and 334-74 W / B ) .  multiplied  Available  fish  were k i l l e d  Aliquots  moisture  at  d r y on a t o w e l  heat-sealed  of  starvation,  in a blender.  were a n a l y z e d  3.2.7  of  25g  taken  partially  and  industrial  were  analysis  determine  Dyer,  diets  The f i s h  blotted  chemical  were  hrs  group.  in  and  approximately  all  taken  Ltd.  fish  containing  2-phenoxyethanol/l,  and  of  a mixed  effects  of  model  treatments  and d i e t a r y  were the  subjected randomized  analysis  block  tank  two-way ANOVA w i t h o u t  were not  protein  to  replicated  concentration  and row ( b l o c k )  effects  43  of  variance  means  was  replication.  within  rows.  were assumed  were random.  to  The  Protein be  appropriate  interaction  mean s q u a r e  significant  differences  due  Zar  (1974)  effects  there  indued  drawn.  tanks was  1 error  The  reduced.  (1955) data  Specific  growth  covariate  percent  to  variance weight general compared Food  (g))  inference  individual  fish  the  means  then  weights  in  this  of  a Type  subjected  body w e i g h t  day as  the  and  subjected  covariate.  slope;  GR% = ( e s l o p e -  i)  100.  GR% e x p r e s s e s  in  body w e i g h t  were s u b j e c t e d significant  against  linear  per  using  intake  procedure  intake  were used  energy  weight  to  44  These  least  1979).  calculate  from  The  test  (P =  analyses  were  squares  analysis  equations  were computed 1982).  derived  (1959)  Regression  gain(g)  utilization  (Brett,  Scheffe's  (SAS,  (P = 0 . 0 5 ) .  (GFC),(dry  day  a general  by DMR t e s t data  x  differences.  GENLIN) .  protein  models  to  to  (DMR)(P = 0 . 0 5 ) .  log  with  were  was  body  probability  fish  was  program ( U . B . C .  and g r o s s  between i n d i v i d u a l  Range T e s t to  residual  1979)  by computer  conversion  the  to  block  al . ,  detect  gain  Nevertheless,  for  (GR%)(Higgs et  slopes  performed  test  that  The t r e a t m e n t  covariance  According  The a p p r o a c h t a k e n  assure  transformed  for  rate  increase  covariate 0.05)  of  the  test  effects.  no major  replicates. to  to  to  variance  However,  New M u l t i p l e  were  fixed  model.  the  used  term a v a i l a b l e  viewpoint  were not  an a n a l y s i s  the  ANOVA.  conservative  is  Duncan's  to  the  from  From a s t r i c t  within study  derived  in  the  no c o r r e c t  in a non-replicated  mean s q u a r e s is  is  to  was  of  by means  The s l o p e s  gross  body of  were  food  x 100 -f- dry f o o d  ( G E U ) , ( g r o s s energy  intake gain  of  a  (kcal) ratio  x 100 -r (PER),  weight et  net  gain)  al.,  protein  protein  value  gain)  (PPV),  designated  (NPU-1),  by the  a l . ( 1980)(NPU-2) . previous  The above  above  estimate was  derived  These  are  shown  describe. differences shown w i t h  formulas  carcass  Net  protein  method for  of  basis in  (Higgs  the  protein  and SR  (for  utilization  (1953) Ogino  these  (SR),(for  was  method, et  methods  is  described  chapter. were  subjected  randomized block  the  to  the  table  ANOVA.  model w i t h o u t  tabulated  respective  on each  derived  are  variance  Superscripts  each  described  utilization  efficiency  ratio  composition,  protein  The p r o c e d u r e  sample from  Protein  and s l o p e  Bender and M i l l e r  The a n a l y s e s of  the  and by the  indices  a two-way  assumed.  (NPR),  were c a l c u l a t e d .  both  the  net  (kcal)).  on a dry body w e i g h t  From f i s h  calculated  in  intake  ratio  a c c o r d i n g to  chapter.  productive  energy  were c a l c u l a t e d  1979)  previous  gross  in  the  standard error  with  were used  from m u l t i p l e  45  -  of  the of  value  designate  comparison  table.  appendices.  error  first  described  replication  mean s q u a r e  the to  As  As an  mean the  was  (SE)  ANOVA.  they  significant  tests.  These  are  3.3  RESULTS  3.3.1  The e f f e c t weight  day  in  dietary  Figs.  in  body w e i g h t  3A, 3B,  p e r i o d was  meals.  17% p r o t e i n to the  42  compared to  PF d i e t The  the  lost  growth  dietary  growth  to  to  response as  from the  diets  groups  increase dietary  of in  diets  proximate  (Fig.  3A).  d u r i n g the  on  not  all  growth  rate  body  was  to  in  over  42  dried  containing  7 and  from day  21  fish  fed  period. to  the  gains  hand summarize of  various  and 3C a r e  body w e i g h t  terms  the  temperature  Predictably,  3A,  are  containing  growth  dietary  formulated  over  FPE as  to  showed  the  in  over  the  specific  concentration  at  the  the  each range  sole  as  (Fig. clear  46  contain  that  of  4). for  the  exact  levels  compositions  prescribed  protein  of  levels  source  diminished with  not  diets  sources.  concentration  returns  of  entire  Figs.  other  equivalent  with  diets  3B,  analysis  fish  fed  in  response  However,  response  fish  treatments  were  various  experimental  form showing  4).  protein  diminishing  in  the  and t h o s e  rate  the  different  the  were  (Table  and l e v e l  were h i g h  the  depicted  dietary  fed  fish  diets,  in  21  of  nutritional  nutrients,  level  these  day 0 to  elicited  Although  the  fed  body w e i g h t  the  rates  protein  of  for  T a b l e 6 and F i g . 4 on the  responses  fish  The growth  linear  a decline  treatments  classical  time.  source  The e x c e p t i o n s  Fish  showed  of  and 3 C .  generally  27% and 37% p r o t e i n . herring  protein  gain  The changes shown  of  each  protein of  protein  increment  The p a t t e r n the  other  the  of  of  protein  of  fed  DAYS  Fig.  3A.  protein  Growth of sources  17% p r o t e i n . free are  (PF)  diets  Growth of  diets  mean wet  in  chinook  are  fish  containing fish  47  fed  shown.  weights,  (SE).  -  also  salmon  fed  the  various  approximately  FPE-7 Values  +_ 2 s t a n d a r d  and  protein  plotted errors  DAYS  Fig.  3B.  protein  Growth of sources  27% p r o t e i n . free are  (PF)  diets  Growth  diets  mean wet  in  chinook  are  fish  of also  -  containing fish  48  -  fed  shown.  weights,  (SE).  salmon  fed  the  various  approximately  OMP and Values  +_ 2 s t a n d a r d  protein plotted errors  DAYS  Fig.  3C.  Growth of chinook  salmon fed the  p r o t e i n sources i n d i e t s c o n t a i n i n g 37% p r o t e i n . p r o t e i n free plotted  Growth of  fish  approximately  fed FPE-47, OMP, and  (PF) d i e t s are a l s o shown.  are mean wet  fish  e r r o r s (SE) .  -  49  -  various  Values  weights, + 2 standard  Table  6.  F i n a l ( d a y 4 2 ) mean wet body w e i g h t s p e r c e n t body w e i g h t p e r d a y ) o f f i s h dietary protein concentration.  Protein Source  FPE  ( g ) and s p e c i f i c g r o w t h fed the various protein  Z 17  Final G.R.  wt. ( + S E ) ( I ) (+SE)  cl +0.06 Cl 1.04 +0.06  2.33  Protein 27  i n Diet 37  Mean  3.94i  3.45 DE 1.86  r a t e s (G.R.I; s o u r c e s a t each  3.24  +0.03  EF  Y  2.26  1.72  +0.03  de FRH  Final G.R.  wt. (Z)  2.43  3.02  4.25  3.23  1.04  1.52  2.38  1.64  2.99  3.90  M  be LTH  Final  wt.  2.16 BC  G.R.  (Z)  3.02  X  EF  0.74  1.52  1.59  1.85  0.06  2.20  1 .48  2.34  1.92  0.39  0.89  0.44  3.06  3.34  2.85  1.52  1.76  1.33  +0.02  2.87  3.55  +0.03  1.36  1.90  ab HTH  Final  wt.  v  AB G.R.  (Z)  V  Final  wt.  2.16  v  de  be CS  BC G.R.  Mean  (Z)  Final  wt.  0.70  2.13  h  P G.R.  OMP  (Z)  Final  0.71  wt.  3.59  G.R.(Z)  2.08  1. V a l u e s w i t h t h e same s u p e r s c r i p t f o r e a c h p a r a m e t e r w i t h r e s p e c t t o t h e s o u r c e x l e v e l ( a - f ) p r o t e i n s o u r c e ( v - y ) , a n d p r o t e i n l e v e l ( p - r ) e f f e c t s do n o t d i f f e r significantly. S u p e r s c r i p t s f o r G.R.(I) a r e c a p i t a l i z e d (DMR t e s t f o r body w e i g h t , P • 0.05); S c h e f f e ' a t e s t f o r G.R. ( Z ) , P - 0 . 0 5 ) . 2. OMP d a t a waa n o t a n a l y z e d statistically. t  - 50 -  sources  as  was  inadequate drying  protein  LTH at  found the  salmon  protein  compared  protein  source  (Table  of  g  at  three  that  of  the  test  protein  each  6).  B o t h the that  protein  mean f i n a l  in  were m a n i f e s t e d  at  of  severe  of  fish  fed  F P E , FRH and  a popular  fish  widely  containing  concentrations  in  analysis  of  level  fed  different  to  were  covariance due  to  (P < 0 . 0 5 ) .  slopes)  of  groups  were compared  showed  of  (Table  growth  growth,  proteins  of  due  effect  and s p e c i f i c  results  was  (P < 0 . 0 1 )  fastest  the  analyses  body w e i g h t  level  levels  the  The above  performance  diets  protein  body w e i g h t s  HTH.  OMP i s  from the  all  FPE and FRH promoted  CS and l a s t l y ,  differences  (derived at  of  fed  a significant  and d i e t a r y  source  fish  factorial  and p r o t e i n  indicated  rates  effect  an  49% p r o t e i n .  randomized block  (P < 0 . 0 1 )  HTH was  The p e r f o r m a n c e  Similarly, a factorial  growth  fed  quality  fed  also  the  protein.  fish  (P < 0 . 0 0 1 )  source  fish  LTH,  to  containing  of  Clearly,  demonstrating  dietary  sources  6).  specific  showed  feed  body w e i g h t s  protein  FPE.  At day 42 marked d i f f e r e n c e s  found  The  of  by a two-way  variance.  fed  be c o m p a r a b l e  performance  various  fish  on h e r r i n g m e a l .  37% l e v e l  commercial The  to  for  source,  temperature  OMP was  log  noted  rates  followed  by  that  of  varying  dietary  concentrations. Since  protein  palatibility against slopes gain  of  protein of "the  relative  intake  may have  particular diets, intake  for  each  restricted  weight protein  regression  equations  to  intake.  protein  been  51  gain source  indicate These  was  are  the  due  to  poor  regressed  (Table rate  depicted  7). of  The  weight  graphically  *  0  5  10  15  20  25  30  35  40  45  50  PERCENT PROTEIN IN DIET  Fig. fed  4.  Specific  rates  of  of  protein  at  concentration.  Values  various  dietary  growth  sources  specific  growth  rate  standard  errors  (SE).  52  (% wet  chinook  salmon  different  plotted  are  body w e i g h t s / d a y ) ,  + 2  Table  7.  Protein source  S l o p e s of body w e i g h t g a i n a g a i n s t p r o t e i n the v a r i o u s p r o t e i n s o u r c e s .  Equation  SE o f  intake  for  slope  a FPE  y = - 0 . 2 6 3 + 3.33x (n=8,  FRH  r=0.9969,  y = -0.2483 (n=8,  0.28  P<0.01) a  + 3.23x  r=0.9957,  0.28  P<0.01) b  LTH  y = -0.1522 (n=8,  + 2.56x  r=0.9843,  0.27  P<0.01) c  HTH  y = -0.3842 (n=8,  + 1.78x  r=0.9966,  0.33  P<0.01) a  CS 1.  y = - 0 . 4 2 7 1 + 3.52x 0.24 S l o p e s w i t h the same s u p e r s c r i p t do not d i f f e r (DMR t e s t , P = 0 .r 0= 50 ) . 9 8 4 8 , P<0.01) (n=8,  -  53  -  significantly  (Fig.  5).  protein  The a n a l y s i s  source  significant  the  intake  protein-free  diet  sources.  Relative  supported  the  highest  results  suggests  that  above food  diets.  Similar  meals.  The s l o p e  (LTH)  was  meal  slope  The e f f e c t food  Food  of  intake  (P < 0 . 0 0 1 )  was  rate  weight  gain.  of  where  intake was  that  was  with  the  obtained  obtained  not  the  This  in  fish  the  the  weight data  diets  fed  (CS) the  This the CS  freeze-dried dried  meal  freeze-dried  high  for  protein  contrasts  temperature  with  with  all  considered.  with low  between  casein-based  restricted  obtained  were  and the  common f o r the  the  differ  relationships  intake  intake  feeding  of  are  The l a t t e r  corrects  for  decreased  the  shown  concentration. containing  protein food  temperature  groups both  basis  of  size  fish  on a per  a wide  100  most  range  54  and l e v e l  meals. dried  of  on  efficiency  by hand to  satiety  fish  basis  (TFI),  al.  1979,  1982)  et  more m e a n i n g f u l content.  body w e i g h t  The t r e n d was  FPE o v e r  fed  (DFI)(Higgs  and m o i s t u r e  increasing  source  conversion  may be c o n s i d e r e d  fish  with  dietary  and g r o s s  on a d r y body w e i g h t 8).  d i d not  of  (HTH).  3.3.2  each  groups  obtained  was  the  effects intake)  protein  intake  than  the  (protein  were l i n e a r  s l o p e s were  lower  The l o w e s t  since  fed  to  that  The i n t e r c e p t s  (P > 0 . 0 5 )  and p r o t e i n  growth  covariate  (P < 0 . 0 0 1 ) .  significantly gain  and the  indicated  Generally,  and d i e t a r y  evident dietary  since  for  and (Table  it DFI  protein  fish  protein  fed  at  diets  levels.  0.50  025  0.76  LOO  gain  against  PROTEN NTAKE (g/FRH )  Fig.  5.  intake test  Slopes of  of  chinook  protein  weight salmon  sources.  -  55  -  fed  diets  protein  containing  the  Table  8.  T o t a l d r y food i n t a k e w e i g h t / d a y ) and g r o s s experimental period.  ( T F l ) ( g / 1 0 0 f i s h ) , mean d a i l y f o o d i n t a k e ( D F I ) ( g / 1 0 0 g mean d r y body food c o n v e r s i o n ( G F C ) ( P e r c e n t ) of the v a r i o u s d i e t s during the  %  Protein Source  Crude  7  233 8,.86  247 8,.21  .50 11 ,  2 1 . .15  27 .11  2 9 ,.86  TFI DFI  196 11 .,01  180 8.,49  233 8..04  GFC  12 ,,68  2 0 .,69  31 ,.12  LTH  TFI DFI GFC  173 10 . 18 9,.79  187 8 .85 1 9 ,.84  271 1 0 .48 22 .67  HTH  TFI DFI  133 9 , .83  140 9..54  156 8..88  GFC  1 .26 ,  4 , .99  12 ,.60  TFI DFI  178 10 .98  186 8 .95  176 7 .77  1 9 .67  24 .72  FRH  On O  CS  GFC OMP  ( + SE)  162 12 .,13  47  234 9..89  GFC  123 11 . 0 0  37  188 1 0 ,.91  FPE  TFI DFI  Protein i n Diets 17 27  0.,44  +0.74  7 . 19  TFI DFI  203 8.70  GFC  25.28  Gross percent al.,  food  conversion  conversion  1982).  protein  A rapid  level  increase  to  was  increase  increased  at  source  protein of  the  diets  with  3.3.3  (MEI)  size  was  basis  fish  to  of  intake  in Table  that  9).  of  poorly  dietary  a means  relative (9).  each of  source  energy  energy  protein  content.  level  and P < across  ranking  fish  by t h o s e  fed for  fish  ingesting  diets  shows t h a t  and l e v e l  the  basis  an a t t e m p t  Food  increased.  57  on  utilization by the  intake  Food energy in  protein  utilized.  (Table  basis  for  proteins  ( G E I ) and  body w e i g h t  of  source,  fish  compared on both a g r o s s 10).  way  evaluating  Values  other  a two  GFC and the  followed  protein  and g r o s s  for  protein  as  (P < 0.001  The GFC of  the  on a d i e t a r y  and m o i s t u r e  dietary  as  regard,  containing  containing  The e f f e c t s  obtained  were used  shown  diets  were s i g n i f i c a n t  GFC v a l u e s  were e x t r e m e l y  The e f f e c t  fish  dry  is  test  experiment.  LTH and CS ( T a b l e  Food i n t a k e  diets  The  small.  FRH and FPE were h i g h e s t ,  energy  of  level  In t h i s  HTH r e l a t i v e  HTH d i e t s  FPE i n  27% and 37% were a n a l y z e d  protein,  proteins  37% ( T a b l e 8) .  fed  et  dietary  17%,  factorial  the  as  of  quality.  receiving with  of  relatively  found  as  Higgs  fed  The a v e r a g e  levels  groups  1971;  fish  and p r o t e i n  0.05). all  block  7% to  calculated  for  levels  randomized  (Brett,  i n GFC was  from  fish  37% and 47% p r o t e i n was  sources  (GFC) was  d r y body w e i g h t  i n GFC between  The GFC d a t a  efficiency  energy This  to  various  metabolizable  was  reported  correct  intake  groups  for  on a fish  decreased  t r e n d was  noted  to  as be  the  Table  9.  G r o s s f o o d c o n v e r s i o n c a l c u l a t e d on a d r y b o d y w e i g h t b a s i s ( G F C ) ; r e l a t i v e GFC ranking of d i f f e r e n t protein sources at d i f f e r e n t dietary concentration.  %  Protein Source  17  FPE  GFC ( + S E ) R e l a t i v e GFC Ranking  1 cd 11.50 +1..55 (100) 2  FRH  GFC Relative Ranking  GFC  12.68 (110) 1  GFC  9.79 (85) 3  HTH  GFC Relative Ranking  GFC  GFC Relative Ranking  GFC  1. did  GFC  21 .50 (108) 1  22 .67 (84) 4  17 .43 (88) 3  w  b  e  6.28 (32) 5 w  gh  19.67 (93) 4  17.19 (86) 4  24.72 (91) 3 r  q +0..70  V  d 12.60 (46) 5  4.99 (24) 5  7.19 (63) 4  23.64  17 .27  V a l u e s w i t h t h e same s u p e r s c r i p t w i t h r e s p e c t t o s o u r c e n o t d i f f e r s i g n i f i c a n t l y (DMR t e s t P = 0 . 0 5 ) .  wx +0.89  X  31 .12 (115) 1 e  a 1 .26 (11) 5  8.48  19.92 (100) 2 j  19.84 (94) 3  P Mean  Mean  h  ef  b CS  i n Diets 37  27.11 (100) 2  20.69 (98) 2 c  GFC Relative Ranking  Protein  ef 21 . 1 5 (100) 1  d  LTH  Crude  27  ( i n p a r e n t h e s i s ) and  x level  ( a - i ) and  protein  source  (v - x)  effects  lower fish  for fed  protein  GEI t h a n  OMP and f i s h and g r o s s  supported  a lower  Gross directly dietary  energy  protein  also  fed  gross  energy  was  noted to  for  meals  ( F P E , FRH) and the dietary  energy  produced s i m i l a r  extremely  3.3.4  37% no f u r t h e r  the  diets  quality  fish  fed  diets  the  At in  dried  F P E . GEU  proteins  the  is  freeze-dried  meal (LTH)  The c a s e i n  The GEU of  energy  protein  various  containing  be  10).  increase  of  effectively. LTH.  to  containing  the  low-temperature most  noted  FPE-47.  the  based  HTH d i e t s  diet was  poor.  requirements) Two d i e t s  of by  salmon  satiation)  i n which a l l be used  Therefore, available  tissue for  to  maintenance conditions  endogenous n i t r o g e n juvenile  were employed  by c h i n o o k  designed  (Table  fed  GEU to  The e s t i m a t i o n  possibly  level  fed  However, OMP  (GEU) t h a n  protein  between  a similar  10).  was  fish  Fish  (Table  diets  i n GEU of  (11).  similar  the  compare  in Table  was  utilization  of  above  shown  (CS)  concentration  dietary  levels  The t r e n d  utilized  Energy i n t a k e  F P E - 4 7 which c o n t a i n e d  energy  to  employed  sources.  MEI.  utilization  related  utilization was  for  fry.  of  protein  maintenance  the  than a l l o w e d  dietary  would  endogenous n i t r o g e n  energy  purposes.  the  only  that  metabolizable  protein-free  59  source  The second  under more normal by the  diet  c a r b o h y d r a t e and be  (maintenance  salmon  a protein-free  from  sufficient  requirements  measure  One was  originated  provide  to  chinook  loss  (fed  to  could lipid. of  diet  energy  nitrogen was  to  satisfy  physiological diet.  loss  Exactly  Table  1 0 . Mean d a i l y g r o s s e n e r g y i n t a k e ( G E I ) ( k c a l / l O O g mean d r y b o d y w e i g h t / d a y ) , mean d a i l y e n e r g y i n t a k e ( M E 1 ) ( k c a l / l O O g mean d r y b o d y w e i g h t / d a y ) a n d g r o s s e n e r g y u t i l i z a t i o n various diets during the experimental period.  Protein Source  FPE  GEI MEI  O o  LTH  HTH  CS  OMP  7  4 5 , .87 4 5 , .65  4 9 . .73 4 8 . ,76  % Crude P r o t e i n 17  i n Diets 27  37  47  4 5 . , 28 4 3 . 09  4 2 , .33 3 9 . .56  3 7 . .83 3 5 . .71  3 7 . .11 32 ..51  1 6 . ,41  3 1 ,.86  3 9 , .66  4 1 ..71  GEI MEI  4 6 , .46 4 4 ,.04  3 9 .93 3 4 .13  3 6 .34 3 5 .79  GEU  1 7 . .08  2 9 , .43  4 0 , .93  GEI MEI  4 3 . .06 41 ,.03  3 7 ,.70 3 4 ,.69  4 6 . ,32 4 1 ,.50  GEU  1 4 ,.42  2 9 .11  3 2 . 14  GEI MEI  4 0 , .40 3 8 , .34  4 1 .12 3 8 .06  3 9 , .52 3 5 , .25  GEU  0,.96  6 .38  1 5 , .54  GEI MEI  4 5 , .68 4 3 , .26  3 7 .68 3 4 ,.82  3 3 , .02 2 9 , .60  GEU  8..20  2 6 , .09  3 3 , .49  GEU FRH  0  metabolizable (GEU) of t h e  ( + SE)  - 0 . .29  +1.37  GEI MEI  3 8 , .80 3 2 , .36  GEU  3 3 , .88  Table  11.  G r o s s e n e r g y u t i l i z a t i o n ( G E U ) , r e l a t i v e GEU sources at different dietary concentrations.  Protein Source  % 17  ( i nparenthesis)  Crude 27  Protein  in  and  ranking  of d i f f e r e n t  Diets 37  protein  Mean  1 c FPE  GEU ( + S E ) Relative GEU Ranking  16.41 (100) 2  FRH  GEU Relative Ranking  GEU  17.08 (104) 1  GEU Relative Ranking  GEU  14.42 (88) 3  GEU Relative Ranking  GEU  0.96 (6) 5  GEU  8. 20 (50) 4  +1.48  31 .86 (100) 1  c  de  Mean  GEU Relative Ranking  GEU  11.41  e  h  b CS  wx 25.22 (86)  c 15.54 (39) 5  d  +0.66  29.15 (99) 2  3 2 .14 (81) 4  6.38 (20) 5  ±0.85  x  40.93 (103) 1  39.11 (91) 3 a  HTH  29.31 (100) 1  f  de 29.43 (92) 2  c LTH  39.66 (100) 2  7.62 (26) e  26.09 (82) 4  33.49 (84) 3  24.57  32.35  1. V a l u e s w i t h t h e same s u p e r s c r i p t with respect to source x level (a p r o t e i n l e v e l ( p - r ) e f f e c t s d i d n o t d i f f e r s i g n i f i c a n t l y (DMR t e s t P =  f ) , protein 0.05).  22.59 (77) 4  source  ( v - x ) , and  sufficient were f e d than  42 In  (97%) and  to  the  over  fish  by c h i n o o k  was  in  salmon  FPE-7  postulated  growth  and e x a c t  to  fish  maintained  experimental  Body w e i g h t diets  for  63  days  deprivation. diet  lost  42  Fish  FPE-7  fed  to  body w e i g h t no f u r t h e r to  have  fish  fed  63  gain  the  similar  nitrogen  loss  are  totally  absorbed  intake  digestible  of  protein  that  r e q u i r e d to  As m e n t i o n e d  previously,  this  diet  loss  during  fish.  support  the  protein-nitrogen  the  by the  represented  was  their  fed  the  of  a group  the  42  the of  effects  fish  their  day  21.  offered  showed  By day  diet  weight.  and the  day  42  and the  the  It  body  is  content.  estimates  observed  62  body  Endogenous  weight.  increase  were  there  in was  observed  noteworthy  7% p r o t e i n  body w e i g h t s  13.  in  groups  protein  weight.  however,  energy  in Table  FPE-7  protein-free  a slight  42,  the  and  severe  reduction  and by day 63 initial  of  initial  no f u r t h e r  metabolizable  shown  protein-free  show the  7% p r o t e i n  respective  employed  highly  balance  22.5%  protein-free  rations  be  (more  weight.  assumed  12)  i n weight  their  body  reasonably  fish  day 0 to  maintained  The d a t a  of  there  maintained  of  of  containing  from  time  may be  protein  From day 0 to  From day  to  fish  period.  (Table  an a v e r a g e  their  shown  on a p r o t e i n - f r e e  data  p e r i o d of  the  levels  endogenous  feeding  when  varying in  nitrogen  corresponds  that  diets  almost  the  requirement. the  FPE was  it  was  that  an e x t e n d e d  fed  Therefore, diet  p r o v i d e d so  would m a i n t a i n  maintenance  of  was  preliminary studies  energy.  zero  energy  satiation,  days),  protein It  protein  that  diet  to  day 63  for  on  endogenous  nitrogen  loss  Table  12.  Wet body w e i g h t (BW) and p e r c e n t w e i g h t (%BW) of f i s h f e d p r o t e i n maintenance d i e t ( F P E - 7 ) .  of i n i t i a l body f r e e ( P F ) and a  _Day_ 21  Diet  PF  BW g / f i s h (+SE)  1.56 (+0.04)  1 .44  % BW  FPE-7  BW g / f i s h (+SE)  Diet  13.  FPE-7  FPE-7  91.93  77.51  77.80  1.67  1 .55  1 .52  99.33  97 .09  E s t i m a t i o n of mean d a i l y endogenouse n i t r o g e n l o s s by c a r c a s s a n a l y s i s of f i s h f e d a p r o t e i n - f r e e ( P F ) and a low p r o t e i n ( F P E - 7 ) d i e t f o r the 42 day experimental period.  Nitrogen gain  Nitrogen intake  on a wet  Endogenous nitrogen loss  body w e i g h t  (SE)  b a s i s (mg/lOOgBW/day))  -17.09  3.63  20.73  (2.77)  -1.15  26.72  25.52  (0.18)  (Calculated PF  1.21  07.04  (Calculated PF  63  1.21  1.56 (+0.04)  % BW  Table  42  on a d r y body w e i g h t  b a s i s (mg/1OOgDBW/day))  -88.48  18.80  107.28  (15.60)  6.16  130.64  124.96  (0.16)  63  -  was  calculated  low  nitrogen  derived  by a d d i n g n i t r o g e n  content  from p r o t e i n  determination.  fish  fed  (SE = 0 . 1 8 ) (FPE-7).  the  Because  have  an e l e v a t e d  also  calculated  fish  fed  dry  (SE = 1 5 . 6 0 )  loss  diet  group.  The above  maintenance fish  The v a l u e s 13)  can  to  nitrogen  Since,  endogenous  to  fish  body w e i g h t  the  of  Ogino (1980)  ENL (g)  =  EPL (g)  =  W  W  was  1 + 2  1 + 2  W  W  nitrogen  be  the  the  the  the  protein  loss  adapted  level  diet  found  to  loss  the  was  FPE-7  intake  loss  7% p r o t e i n  that  (Table for  (EPL)(ENL x in  direct  and G r o v e s ,  follows: -5  x 25.52 x  10  2  x  10  x  diet  estimates  loss  (Brett  159.5  BW/day  diet.  changes  as  from  protein  for  protein  2  64  be  case,  zero  protein  quality  protein  (size)  to  107.28 mg N / l O O g  (SE = 0 . 1 6 )  fed  nitrogen  were  In t h i s  fed  a non-protein  (ENL) or  proportion method  groups  show t h a t  correct  loss  to  be  the  protein  diet  basis.  estimated  from f i s h  in  to  (BW)/day (SE = 2 . 7 7 )  endogenous  approximates  The  assumed  calculated  maintenance  d r y BW/day  fed  obtained  be employed  endogenous 6.25).  were  loss.  and 25.52 mg N / l O O g  content,  values  was  protein-free  from the  and 124.96 mg N / l O O g  when  the  was  diet  occurred  the  was  included  body w e i g h t diet  nitrogen  diet  loss  on a d r y body w e i g h t  nitrogen  for  nitrogen  fed  moisture  endogenous  required  therefore  protein-free fish  to  protein-free  and was  (N)/100g  from the  BW/day  the  Endogenous  2 0 . 7 3 mg n i t r o g e n the  of  intake  -5  x d  x d  1979)  DW ENL (g)  =  _ x 124.96  DW DW 1• + 2 2  x d  _ x 10  ENL = endogenous  nitrogen  loss  EPL = endogenous  protein  l = initial  W  W  2 = final  D  w  D W  body w e i g h t  body w e i g h t  l = initial 2 = final  d  = days  The r e l a t i o n s h i p  between  protein  x d  (g)  (g)  dry body w e i g h t feeding  5  loss  dry body w e i g h t  of  5  x 10  x 781.0  EPL (g)  3.3.5  DW 2  1 + 2  (g)  (g)  intake  and  protein  utilization To i l l u s t r a t e protein the  quality,  various  intake  was  this  energy protein  relationship way  intake  intake  were e x p r e s s e d  was  and, 1)  the  nitrogen  the  groups  was  amounts tissues  and m e t a b o l i c  digested  obtained  endogenous plus  body  of  fish  protein  and the  refer plus  by the  to  the  utilized  protein  gain.  The per  14).  In  fraction  amount  used  of  for  between to  quantities day  for  lost  (Table  (equal  body w e i g h t  Protein  amount  the  and  utilized  difference  loss  65  of  investigated.  protein  lOOg wet  balance  excretion  excretions  not  was  i n mg per  of  sources  body  exogenous  This  Fig.  of  fecal  that  (Fig.l).  maintenance,  into  and growth  exogenous  i n which  protein  partitioned  discussion,  protein  the  dietary  maintenance through  the  T a b l e 14.  Diet  The u t i l i z a t i o n o f d i e t a r y p r o t e i n on a d a i l y b a s i s by f i s h f e d the v a r i o u s experimental d i e t s d u r i n g t h e 42 day p e r i o d .  Amount o f P r o t e i n U t i l i z e d by lOOg f i s h f o r : Excretion P r o t e i n Intake Maintenance Growth (mg/lOOgBW/d) (mg/lOOgBW/d)  Percent o f P r o t e i n Intake used f o r : Excretion Maintenance Growth  FPE-7  167.0  160  7.1  0  96.0  4.3  0  FPE-17  395.3  160  146.6  88.7  40.5  37.0  22.5  FPE-27  604.1  160  298.2  145.9  26.6  49.4  24.1  FPE-37  728.9  160  351.4  217.5  22.0  48.2  29.9  FPE-47  898.6  160  378.8  359.8  17.6  42.2  40.3  OMP  941.4  160  312.7  468.7  17.0  32.3  49.8  38.3  40.5  21.3  FRH-17  422.8  160  154.0  108.9  FRH-27  511.8  160  227.5  111.7  31.3  44.5  24.2  FRH-37  749.9  160  321.4  268.6  21.3  41.8  36.9  LTH-17  366.4  160  131.0  75.5  43.7  35.8  20.5  LTH-27  554.5  160  232.6  161.9  28.9  42.0  29.1  UTH-27  914.3  160  340.5  413.8  17.5  37.3  45.2  HIH-17  341.7  160  -16.2  197.8  46.9  -4.1  57.2  HIH-27  534.8  160  53.3  321.6  30.0  10.0  60.1  HIH-37  754.8  160  121.9  473.0  21.2  16.2  62.6  CS-17  417.7  160  88.6  169.2  38.4  21.4  40.2  CS-27  500.9  160  238.6  102.5  32.0  43.1  25.0  CS-37  609.6  160  257.6  191.9  26.3  42.3  31.5  -  66  -  °  (mg/lOOg  BW/day).  Next,  the  utilization  maintenance,  growth  and the  quantity  a percentage  of  protein  fed  graphically  (Figs.  In r e g a r d for  the  to  maintenance  6A -  fish was  all  maintenance  and none  6A). the  percentage  (Fig.  the  6A).  protein intake  was of  protein  was  way all  diets  protein  protein growth  diets  with  with  meet  above  for  protein  the  growth lost for  fish  growth  rose  (Fig. (Op),  sharply, rapidly  optimum  resulted  of  excreta.  growth  at  a  In c o n t r a s t  largely  for  dropped  because  in  fed  intake  an FPE  protein to  fish  OMP u t i l i z e d because  of  increased  6A). processing  (Fig. fed of  a protein protein  HTH r e l a t i v e  their  intake  protein  utilization  maintenance  F P E , the  for  16.5%  p r o p o r t i o n of  depicted  growth  growth  600mg/100g BW/day.  chinook  Only  for  utilized  utilized  groups,  in diets  of  h e r r i n g meal  was  poorly.  A higher  LTH to  of  as  level  for  for  containing  (Fig.  been  and u t i l i z e d  or a v a i l a b l e  utilization  effectively  absorbed  used  of  At t h i s  intake  efficiently  losses  other  was  level  protein  protein  of  expressed  and has  and o p t i m a l  percentage  most  14)  F P E , the  wasted  protein  rate  the  The method  fed  protein  approximately  less  excretory  to  the  Additional  in  receiving  the  ingesting  p r o p o r t i o n employed  a decreased  increase  of  (Table  was  for  6E).  Between m a i n t e n a n c e  whereas  in  of  protein  excreted,  160mg/100g BW/day.  practically  of  to  6B,  diets  the  markedly  6C, 6D).  fed  concentration intake those  maintenance  67  In  containing  protein  was fed  of  influenced  was  comparison  HTH u t i l i z e d utilized  37% ( T a b l e  r e q u i r e d by the diets  containing  for 14). fish  FRH or  Fig.  6A -  E.  maintenance (•)  when of  intake  that  growth (see is  test  intake.  promotes is  (A) i s  F i g . 1).  shown  f e d the t e s t  protein  It  utilization  (O) and growth  fish  levels  growth.  Percent  proteins.  ( A ) , and p e r c e n t protein  optimal  that  percent  The p r o t e i n  for comparison  The dashed  portions  extrapolated.  -  68  excreted at  of  of  various protein  protein  utilization  productive  utilization  fed f o r  f o r the  utilization  the same as p r o t e i n  symbols  sources  (Op) s t a n d s  noteworthy  i n dark  of p r o t e i n  of  for  v a l u e (PPV)  fish  with  for  f e d OMP  each  the c u r v e s  of  the  were  PROTEIN UTILIZATION  I X Of PROTEIN  P(0|  PROTEIN  UTILIZATION  I Z OP PROTEIN FED )  8  3  8  PROTEIN UTILIZATION  8  o  o  o  o  o  (X OP PROTEIN PED|  S  o  o  2* 2  M  m\  \  •  \  M  « Tl  8 »  I ? PROTEIN  V UTILIZATION  ! • (Z OP PROTEIN  FPE.  §  requirements protein  for  intake  protein.  over  Further,  60% of  the  at  the  highest  protein  fed  was  level  excreted  of (Fig.  6D) . Because CS a t  low  of  levels,  ingesting  high  utilization groups of  an i n c o n s i s t e n c y  (Fig.  protein  meeting  terms  in  3.3.6  The measurement  Protein protein assess  Predictably  it  for  noteworthy  extent  effect  rats  to  at  protein  which  problem noted  in  fish  of  that  CS was  fed  protein  and i n t e r p r e t  quality  for  the CS  a good  source  of  chinook  ( P E R ) ( O s b o r n e et  Figure  1957)  and PER i n trend  by o t h e r species  intake  efficiency  dependent was  commonly  fish  fed  similar  are  the  partially  allowance  was  made  for  maintenance  between  the  PER and NPR c u r v e  by the  to  that The  intake.  overcome  different  net to  between  F i g u r e 8 shows  NPR method. at  used  protein  on p r o t e i n  protein  and  FPE d i e t s .  investigators.  low  70  1919)  relationship  maximum PER o c c u r r e d .  PER was  al.,  are  7 shows the  a classical  among  by p r o t e i n  ratio  of  difference  pattern  requirements  and D o e l l ,  and p l a i c e  which  fish  utilization.  concentration followed  of  C S , the  indicated  ratio  quality.  performance  construct  protein  differences  concentrations the  protein  of  of  protein  (NPR)(Bender  protein  reported  the  efficiency  protein  dietary  of  and net  ratio  to  The d a t a  salmon  ratio  levels  difficult  6E).  for  the  and a p a l a t a b i l i t y  dietary  was  in  The  once an The protein  -0-5  1  0  • 5  • 10  1  • 15  • 25  20  PERCENT  Fig.  7.  protein  Effect  for et  plaice al.,  Hegsted  PROTEIN  of  ratio  basis fed  1972)  • 35  for  and C h a n g ,  71  protein  chinook  rats  1965)  1  40  ' 45  ' 50  55  level  ' 60  on  (PER) c a l c u l a t e d  freeze-dried and f o r  1  I N DIET  dietary  efficiency  body w e i g h t  1  30  salmon  fry  cod muscle fed  casein  containing  on a  wet  fed F P E ,  (from (from  diets.  Cowey  Table  15.  Protein efficiency ratio and r a n k i n g of d i f f e r e n t  Protein Source  c a l c u l a t e d on a d r y body w e i g h t b a s i s protein sources at d i f f e r e n t dietary  % 17  FPE  PER ( + S E ) R e l a t i v e PER Ranking  1 def 0.679 +0.010 (100) 2  FRH  PER Relative Ranking  PER  0.704 (103) 1  PER  0.583 (85) 3  PER  0.074 (10) 5  PER  0.387 (57) 4  PER Relative Ranking  PER Relative Ranking  Mean 1. did  PER Relative Ranking PER  0.486  de  b  w 0.623 (84) 3  c  v  0.315 (43) 5 f  V a l u e s w i t h t h e same s u p e r s c r i p t w i t h r e s p e c t n o t d i f f e r s i g n i f i c a n t l y (DMR t e s t P = 0 . 0 5 ) .  0.747 (101) 1  0.590 (79) 4  0.182 (23) 5  0.687 (92) 3  0.630  0.626  to source  0.191 (26) 5 def  0.745 (96) 3  x level  (a -  +0.06  X  f  ef  +0.055  0.736 (100) 2  0.786 (105) 1  0.697 (90) 4  c CS  X  f  f  a HTH  ( i n parenthesis)  Mean  0.754 (100) 2  0.750 (96) 2 d  LTH  i n Diets 37  f 0.776 (100) 1  f  VI  Crude P r o t e i n  27  ( P E R ) , r e l a t i v e PER concentrations.  f ) and  w 0.606 (82) 4  protein  source  (v - x)  effects  100  200  300  PROTEIN INTAKE  Fig. and  8.  for  FPE.  (NPR)(A) Values  comparison.  73  600  700  800  900  (mg/IOOg BV^doy)  efficiency  ration  containing symbols  500  The r e l a t i o n s h i p  protein  protein  400  between  ratio of for  protein  (PER)(0)  fish  and  fed  diets  OMP a r e  shown  intake net  in  dark  intakes latter  is  method  Those 17,  due  27,  effect  to  the  (Fig.  diets  f o r m u l a t e d to  dietary  (P < 0 . 0 0 5 ) . methods  employed  protein  Despite  distinguished  an a l m o s t  identical  different  quality  was  found  protein  concentrations. meals PER a t  the  quality on the  of  other  protein  values  CS ( i . e .  PER of  an e x t r e m e l y were  (Table  when t e s t e d  However,  at  17)  were  those  for  for  meal  the  low  reports  74  in  the  above  the  protein levels.  ranking  NPR  dietary of  the  PER and NPR by  other PER r e l a t i v e  indicated the  protein  different  ( F P E = 100)  p r o t e i n whereas  the CS  freeze-dried  The  an i d e n t i c a l  The r e s u l t s  the  th~e poor  dietary  at  in  (LTH) show maximum  made by t a b u l a t i n g  reported  for  Excluding  lower  noted  Comparisons with  CS = 1 0 0 ) .  to  noted  high d i e t a r y  values  mean r e l a t i v e  poor q u a l i t y  comparable  were  PER u n d e r e s t i m a t e s  hand p r o v i d e d s i m i l a r  obtained.  investigators  that  sources  Inconsistant  protein concentration.  proteins  a c c o r d i n g to  was  PER was  Low-temperature dried  concentrations.  proteins  15 and 1 6 ) .  not  The  significant  protein  concentrations.  was  of  i n PER and NPR b o t h  low compared to  may s u g g e s t  poor  the  protein quality.  different  relative  effect  intermediate  observations  by  concentration  on PER and NPR was  (Tables  dietary  that  This  evaluate  PER and NPR v a l u e s  (HTH) at  (FPE, FRH).  maintenance  a protein  differences  manner  proteins it  to  between the  in r e l a t i v e  diets,  have  source  the  differences at  made f o r  8).  and 37% were of  allowance  that  other  literature.  to  HTH was  fishmeals  Table  16.  Net p r o t e i n r a t i o c a l c u l a t e d ranking of d i f f e r e n t protein  Protein Source  on a d r y body v e i g h t sources at d i f f e r e n t  b a s i s ( N P R ) , r e l a t i v e NPR dietary concentrations.  % Crude 27  17  FPE  NPR ( + S E ) R e l a t i v e NPR Ranking  1 ef 0.974 +0.009 (100) 2  FRH  NPR Relative Ranking  NPR  0.993 (102) 1  NPR Relative Ranking  NPR  0.901 (93) 3  NPR Relative Ranking  NPR  NPR  Protein  i n  Diets 37  cdef 0.921 (100) 3  0.934 (101) 1  cd 0.920 (100) 2 cde  +0.005  xy  0.884 (102) 1  0.937 (102) 1  0.872 (95) 4  0.679 (79) 4  0.817 (89) 3  0.481 (49) 5  0.422 (46) 5  0.464 (53) 5  0.456 (50) 5  0.667 (69) 4  0.934 (101) 1  0.833 (97) 3  0.811 (88) 4  0.816  0.745  cdef  and  Mean  0.864 (100) 2 def  LTH  ( i nparenthesis)  cd  a HTH  def CS  Mean 1. did  NPR Relative Ranking  NPR  0.803 +0.004  V a l u e s w i t h t h e same s u p e r s c r i p t w i t h r e s p e c t n o t d i f f e r s i g n i f i c a n t l y (DMR t e s t P = 0 . 0 5 ) .  to source  x level  (a -  f ) and  protein  source  (v -  y)  effects  T a b l e 17.  P r o t e i n e f f i c i e n c y r a t i o (PER), r e l a t i v e PER, n e t p r o t e i n u t i l i z a t i o n (NPU), and r e l a t i v e NPU f o r v a r i o u s p r o t e i n s , when f e d a s the s o l e source o f p r o t e i n t o d i f f e r e n t f i s h species h e l d under d i s s i m i l a r experimental c o n d i t i o n s .  P r o t e i n Source of Diet  Freeze d r i e d cod muscle  Species  (g)  Plaice (Pleuronectes platessa  Single c e l l protein p l u s methionine White f i s h meal  Size  M  Temperature degree C  Protein level % of dry diet  50  1.78  42  50  1.23  38  13  15  satiation  14  15  it  15  15  PER wet body weight basis  Relative PER (casein=100)  1  Ration  NPU '  2  Relative NPU (casein=100)  Cowey e t a l . (1974)  50  1.29  35  50  0.89  34  15  50  0.60  23  13  15  50  0.84  30  14  15  50  0.98  33  2.5  25  41  1.78  68  34  74  Casein  2.5  25  43  2.60  100  46  100  Leaf p r o t e i n concentrate  2.5  25  30  2.09  59  38  72  Casein  2.5  25  27  3.55  100  53  100  M  Soy p r o t e i n p l u s methionine Fish protein concentrate L i p i d extracted h e r r i n g muscle  "  II  L i p i d extracted sprats  Leaf protein concentrate  cont'd.. ./2  Carp (cyprinos carpio)  14  15  13  "  not  stated  Reference  Ogino e t a l . (1978)  T a b l e 17. cont'd..(2)  P r o t e i n Source of Diet  Leaf p r o t e i n concentrate  meal  Rainbow t r o u t (salmo g a i r d n e r i )  Rainbow t r o u t (salmo g a i r d n e r i )  htethanophilic bacteria S p i r u l i n a algae Petroyeast  n »  Soybean Brewers y e a s t  "  Casein  Herring  meal  Methanophilic bacteria  Carp (cyprinus carpio) "  S p i r u l i n a algae Petroyeast Soybean cont'd.../3  Size (g)  ii  Casein  Herring  Species  "  Temperature degree C  3.5  19  3.5  19  30  12  30  12  30  12  30  12  30  12  Ration  satiation  ••  II  «  Protein level % o f dry diet  PER wet body weight basis  Relative PER (casein=100)  1  '  2  Relative NPU (casein=>100)  41  2.03  70  38  69  39  2.91  100  55  100  30  1.91  97  38  95  30  1.62  82  37  93  30  1.33  68  32  80  30  2.01  102  42  105  18  45  30 II  NPU  30  1.17  59  30  75  30  1.97  100  40  100  4% o f body weight  25  2.82  114  64  131  "  35  2.54  102  49  100  25  30  1.15  42  36  73  30  25  31  2.08  84  47  97  30  25  29  1.35  54  42  86  30  12  30  12  30  25  30  25  30  Reference  Ogino e t a l . (1978)  Atadc and Matty (1978)  Atadc e t a l . (1979)  T a b l e 17. cont'd..(3)  P r o t e i n Source of Diet  Species  Size (g)  Temperature degree C  Ration  Protein level % of dry diet  PER wet body weight basis  Relative PER (casein=100)  NPU  1 2 '  Relative NPU (casein=100)  Reference  Casein  Carp (cyprinus carpioT  30  25  4% o f body weight  30  2.48  100  49  100  Atack e t a l (1979)  White f i s h meal  Rainbow t r o u t (saliro g a i r d n e r i )  34  10.7  not s t a t e d  31  3.2  89  63  95  Watanabe e t a l . (1983)  "  31  3.7  103  61  92  Peruvian anchovy meal  "  34  10.7  S a r d i n e and p o l l o c k scrap meal  "  34  10.7  32  3.2  89  63  95  G r e e n l i n g meal  "  34  10.7  30  3.6  100  66  100  Casein  "  34  10.7  33  3.6  100  66  100  2.0  17  30  3.8  109  60  107  Whole egg  2.0  17  30  3.8  109  61  109  Egg albumin  2.0  17  30  3.9  111  62  111  Trout muscle protein  2.0  17  30  3.8  109  62  111  Squid muscle protein  2.0  17  30  3.9  111  62  IU  2.0  17  30  3.5  100  56  100  Egg y o l k  cont'd.../4  Rainbow t r o u t (salmo g a i r d n e r i )  not s t a t e d  Ogino and Nanri (1980)  T a b l e 17. cont'd..(4)  -o  P r o t e i n Source of Diet  Species  Casein & squid muscle (1:1)  Rainbow t r o u t (salrtD g a i r d n e r i )  Size (g)  Temperature degree C  4.0  17  C a s e i n & whole egg (1:1)  4.0  Casein  FPE  Chinock salmon (oncorhynchus tshawytscha)  FRH LTH HTH  II  CS (casein-gelatin, + a r g i n i n e & inetJuonine)  »  T. 2.  Ration  PER wet body weight basis  Relative PER (casein=100)  NPU  1  ,  2  Relative NPU (casein=100)  30  3.9  108  61  105  17  30  3.8  106  63  109  4.0  17  30  3.6  100  58  100  1.50  11  27  2.94  98  79  103  1.50  11  28  2.87  96  78  101  1.50  11  "  28  2.65  88  73  95  1.50  11  »  27  0.76  25  40  52  1.50  11  26  3.00  100  77  100  Determined by method o f Bender and M i l l e r (1953). rjetermined by method o f Ogino e t al.(1980).  not s t a t e d  Protein level % o f dry diet  satiation  Reference  Ogino and N a n r i (1980)  Present study  3.3.7  The measurement value  and net  Protein (NPU)  to  value  and B e n d e r ,  retention  to  in  differentiate  body w e i g h t  but  PPV and NPU would provide  for  1955;  protein  The v a l u e s depicted 6E).  intake.  protein  fed  parameter respect  protein  graphically  to  and  for  was  the  the  the  1980)  relate  methods,  analogous  however,  that  promote  body  protein.  effectiveness  to  shown  various the  intake  mode of  utilization  PPV and NPU a r e  in  PPV a r e  similar  gains  Therefore,  of  in Table  protein  estimate  retained.  protein  productive  a protein  to  synthesis.  for  identical  that  gains  protein  al.,  The l a t t e r  measure  obtained  PPV i s  Ogino et  between p r o t e i n s  best  by p r o t e i n  (PPV) and net  different  muscle  quality  utilization  PER and NPR r e s p e c t i v e l y .  cannot  -  protein  protein  productive  (Miller  protein  of  of  sources the  protein  previously  utilization  in  are  Fig.(6A  between  described for  and  percentage  The r e l a t i o n s h i p  was  (18)  of this  with  growth  (section  3.3.5) . A progressive source  order:  FPE > FRH > LTH > CS > HTH ( T a b l e the  quality  Low t e m p e r a t u r e statistically  chinook  i n mean PPV was  protein  previously  in  decrease  salmon  of  the  d r i e d meal  significant  herring  meal  (FRH).  similar  quality  to  for  the  protein  sources  19,  20).  was  found  P > 0.05)  to  based  A l t h o u g h NPU was  obtained calculated  80  in  18).  as  have  As  protein  highest. (not  freeze-dried  source  ranking (of  following  stated  a lower than  the  the  meals was  quality  An i d e n t i c a l was  changed  freeze-dried  The c a s e i n LTH.  diets  observed  (CS)  mean  was  values)  by PPV and NPU ( T a b l e s by two  methods,  of  the  18,  Table  18.  Protein sources  Protein Source  p r o d u c t i v e v a l u e s ( P P V ) , r e l a t i v e PPV at different dietary concentrations.  ( i n parenthesis)  % Crude 27  17  Protein  and  ranking  i n Diets 37  of d i f f e r e n t  protein  Mean  1 d FPE  PPV ( + S E ) R e l a t i v e PPV Ranking  36.87 (100) 2  FRH  PPV Relative Ranking  PPV  40.45 (110) 1  PPV  3 4 .23 (93) 3  PPV  -4.81 (-13) 5  PPV  21.50 (58) 4  f +3.23  def  PPV Relative Ranking  PPV Relative Ranking  Mean 1. did  PPV Relative Ranking  PPV  25.65  41 . 7 8 (87) 3 def 37.53 (78) 4  V a l u e s w i t h t h e same s u p e r s c r i p t with respect n o t d i f f e r s i g n i f i c a n t l y (DMR t e s t P - 0 . 0 5 ) .  to  v 7.07 (16) 5  def  43.06 (87) 3  42.23 (88) 2  37.81  3 7 . 18  source  wx 37.95 (85) 3  16.23 (34) 5 def  +1.45  wx  be  b 9.81 (20) 5  x  level  +1.87  42.26 (94) 2 de  42.08 (85) 4  c CS  44.84 (100) 1 def  def  a HTH  48.11 (100) 1  44.56 (90) 2 d  LTH  X  ef  49.55 (100) 1  ( a •- f ) a n d  w 35.59 (79) 4  protein source  (v -  x)  effects  Table  19.  N e t p r o t e i n u t i l i z a t i o n c a l c u l a t e d by t h e m e t h o d o f B e n d e r a n d M i l l e r ( 1 9 5 3 ) ( N P U - 1 ) , ( i n p a r e n t h e s i s ) and r a n k i n g of d i f f e r e n t p r o t e i n sources a t d i f f e r e n t d i e t a r y concentrations.  Protein Source  % Crude 27  17  Protein in  Diets 37  relative  Mean  1 FPE  NPU-1 ( + S E ) R e l a t i v e NPU Ranking  56.88 (100) 2  FRH  NPU-1 Relative Ranking  NPU  60.02 (105) 1  NPU  55.74 (98) 3  NPU  22 .69 (40) 5  NPU  40.41 (71) 4  de +0.90  e  e  NPU-1 Relative Ranking  de  NPU-1 Relative Ranking  Mean  NPU-1 Relative Ranking  NPU-1  47.11  1. V a l u e s w i t h t h e same s u p e r s c r i p t e f f e c t s d i d not d i f f e r s i g n i f i c a n t l y  be  a  with (DMR  w 51.10 (89) 3 V  a 25.04 (44) 5  26.32 (47) 5 de  +0.40  55.14 (96) 2  43.5 (78) 4  26.12 (44) 5  52.27 (94) 2  50.41  45.23  respect to source test P = 0.05).  w  cde  55.78 (94) 3  x level  +0.52  V  bed  de  b CS  57.32 (100) 1  48.42 (87) 3  53.86 (91) 4 a  HTH  w  55.65 (100) 1  57 .00 (96) 2 de  LTH  de  59.31 (100) 1  ( a - e) and  49.51 (86) 4  protein  source  (v -  x)  NPU-1  Table  20.  N e t p r o t e i n u t i l i z a t i o n c a l c u l a t e d by t h e m e t h o d o f O g i n o e t a l . ( 1 9 8 0 ) ( N P U - 2 ) , r e l a t i v e NPU-2(in p a r e n t h e s i s ) and r a n k i n g of d i f f e r e n t protein sources a t d i f f e r e n t d i e t a r y concentrations.  Protein Sou r c e  00 CO  FPE  NPU-2 ( + S E ) R e l a t i v e NPU-2 Rank i n g  FRH  NPU-2 Relative Ranking  LTH  HTH  % Crude 27  17  NPU-2  1 ef 79.06 +3.46 (100) 3 f 84.05 (106) 1 ef 79.57 (101) 2  NPU-2 Relative Ranking  NPU-2  NPU-2 Relative Ranking  NPU-2  41 .85 (53) 5  NPU-2 Relative Ranking  NPU-2  59 .82 (76) 4  Mean  NPU-2  68.87  1. V a l u e s w i t h t h e same s u p e r s c r i p t e f f e c t s d i d not d i f f e r s i g n i f i c a n t l y  Diets 37  Mean  def 72.56 (100) 1  ef  76.74 (100) 1 bed  77.69 (99) 2 def  75.84 (99) 2 b  wx  73.07 (93) 4  56.99 (79) 4  69.88 (91)  39.70 (51) 5  38.24 (53) 5  39 .93 (52) cde  76.56 (97) 3  70.10 (97) 2  69.16  60.74  respect to source t e s t P = 0.05).  +2.00  wx  65.78 (91) 3  def  +1.55 with (DMR  i n  ef 78.69 (100) 1  be CS  Protein  x level  ( a - f ) and  68.89 (90) 4  protein  source  (v - x)  Table  21.  The r e l a t i o n s h i p protein  utilization  and M i l l e r  (1953)  Chinook  Protein  Source  between p r o t e i n (NPU)(y)  intake  calculated  (x)  and net  by the  Bender  formula.  Salmon  (present  study)  Equation  FPE  y == 65 .01 -- 0 .0161x  (n=10,  FRH  y == 73 .81 -- 0 .0333x  (n=6,  r = -0.8848,  P<0 .05)  LTH  y == 65 .28 -- 0 .0233x  (n = 6,  r = -0.8358,  P<0.05)  HTH  y == 20 .38 -- 0 .0085x  (n=6,  r = 0.3732,  NS)  CS  y == 22 .17 -- 0 .0536x  (M=6,  r = 0.6036,  NS)  -  84  -  r = - 0 . 5 6 1 1 , NS)  Table  22.  The r e l a t i o n s h i p protein of  utilization  Ogino  et  al.  Chinook Protein  between p r o t e i n  Source  (NPU)(y)  intake  calculated  (x)  and n e t  by the  method  (1980)  Salmon  (present  study)  Equation  FPE  y = 104.76 -  0.0476x  (n=10,  FRH  y = 106.45  -  0.0545x  (n=6,  r= - 0 . 9 5 9 4 ,  P<0.01)  LTH  y =  95.69  -  0.0422x  (n=6,  r= - 0 . 8 5 9 5 ,  P<0.05)  HTH  y =  45.15  -  0.0096x  (N=6,  r= - 0 . 6 5 9 9 , NS)  CS  y =  4 7 . 9 3 + 0.0412x  (n=6,  r= 0 . 4 0 7 6 ,  Rainbow (calculated Egg  yolk  y =  97.9  P<0.01)  NS)  Trout  from Ogino -  r= - 0 . 9 1 2 0 ,  0.0458x  et  al.,  (n=5,  1980)  r= - 0 . 9 6 6 4 ,  P<0.01)  Plaice (cal culated Freeze-dried cod m u s c l e  y =  75.8  from Cowey  -  0.0579x  85  -  et  al.,  (n=6,  1972)  r= - 0 . 9 8 2 3 ,  P<0.01)  IU  Z  10 . 0 0  100  200  PROTEIN  Fig.  9.  300  400  INTAKE  500  600  700  The r e l a t i o n s h i p  between  protein  utilization  protein  sources  determined  by the  Slopes  rainbow  al .  yolk fed  (1980).  (taken  containing  for  from Ogino et  cod muscle  (taken  diets  The NPU v a l u e  for  900  1000  (mg/100g BW/day)  and net  et  800  are  (NPU) of  al.,  also fed  shown  trout  1980)  for  OMP i s  the  method  from Cowey et  fish  86  protein  and al.,  intake test of  Ogino  fed  egg  plaice 1972)  comparison.  also  shown.  Bender  and M i l l e r  respectively, obtained  very  by b o t h  allows  for  = 100)  values  those  reported  The  (1953) close  maintenance obtained  in  by o t h e r  method  O g i n o et  0.01) method  of  protein  equations  al.(1980)  was  NPU on p r o t e i n  FRH (P < 0 . 0 1 ) ,  present  (Tables  intake  was  (1953)  was  intake  was  21).  (1972)  state  and S a r g e n t  nutritive provide  value  the  species.  protein was  study that  for  method the  explained.A  in  sources, measure are  of  O g i n o et  greater  assay.  of  been  used,  87  quality 9)  egg  of  the  for  may fish  f o r NPU The l i n e cannot  fish  lines  (Table  absence  relationship for  the  equations  obtained  The r e g r e s s i o n  9).  when  less  the  (Fig.  expected  fed  in  inverse  correlation  al.(1980).  inverse  rainbow t r o u t  the  the  FPE (P <  determining  F P E , F R H , and LTH compare w e l l with  for  regression  slope  When  However,  their  below  comparison.  a significant  that  depicted  positive  would have  this  obtained  1980)(Fig.  equations  because  intake  found  protein  for  22).  evident  for  NPU (CS  and NPU was  considerably  concentrations  relative  by the  omitted  adequately  of  best  These  determined was  dietary  NPU-2  tabulated  17)  21,  employed  between NPU and p r o t e i n  standardized  intake  were  Relative  are  (Table  and LTH (P < 0 . 0 5 ) .  Bender and M i l l e r  Co wey  basis.  (NPU-2)  values  previously,  study  investigators  between  by r e g r e s s i o n  of  As o u t l i n e d  the  al.(1980)  NPU ( F P E = 100)  on a body w e i g h t  described  dependence  relative  procedures.  relationship  of  (NPU-1) and Ogino et  fed  of  f o r CS  be NPU on  HTH than  obtained  with  one  yolk  ( O g i n o et  in  another  this and  al.,  3.3.8  The measurement  The  slope  includes protein  (Hegsted the  calculating 1977).  the  zero  the  gain  or  test The  compared  body the  as  covariate. fed  Chang,  1965)(Tables  were  included  obtained the  for  present  protein  1979).  Samonds  and  reference  protein  of  linear  the  and meet at  various  of  proteins  covariance  protein  gain  of  the the  of data  the  intake  assay of  free  diet  FPE,  LTH and HTH from one  body  between  were c o m p a r e d , another  88  the  the  (Table  were  gain intake  zero  The  standard  (Hegsted  and  as  reduced diet  the body  the  and C h a n g ,  protein  fry  fed  groups  coefficients  under  satisfied  (Hegsted  ( F P E ) and  protein  were free  the  intercept.  11.  The c o r r e l a t i o n  body In  considered  These  that  of  percentages  protein  relationship  slopes  groups)  assay.  indicated  and p r o t e i n  as  10,  rat  Hegsted,  weight  with  of  when  chinook  including  and may be  for  assay.  the  in a slope  the  to  24)  Figs.  were c a l c u l a t e d 23 and 24)  a common  d r y body  data  in  (slope)  fed  (Table  for  depicted  study,  fed  1974;  some  been o m i t t e d  intake.  slope  When the  group has  which  levels  in  on p r o t e i n  each  gain  linearity  in  Alternatively,  gain  slopes  when  or more d i e t a r y  protein  precision  considerably  procedure  response  are  of  of  fed  The s l o p e s  errors  estimates  the  1965).  ratio  on the  the  groups  a multi-dose  and t h r e e  slopes  and body  is  by s l o p e  based  by a n a l y s i s  23)  protein  diet  quality  (McLaughlan,  are  for  (Table the  is  proteins slopes  assay  protein  slope  SR a s s a y ,  protein  and C h a n g ,  The a s s a y  weight  (SR)  a non-protein  bioassays,  ideal  ratio  of  conditions weight  (including assay 24).  or  requirements 1965;  of  for  McLaughlan,  the  protein  distinguished With  the  T a b l e 23.  Slope r a t i o s o f d r y body weight gains on p r o t e i n i n t a k e f o r f i s h f e d the v a r i o u s p r o t e i n sources. C a l c u l a t e d both by i n c l u d i n g and e x c l u d i n g t h e p r o t e i n f r e e d i e t (PF) f e d groups.  Protein Source  Slope r a t i o s o f d r y body weight gains e x c l u d i n g PF groups i n c l u d i n g PF groups  ,z  1  0.8817 +0.0432 4.90 -0.0852 0.9957 (100) 3  8  0  Slope SE o f s l o p e SE o f s l o p e as % Intercept Correlation coefficient Relative slope Ranking  0.8048 +0.0623 ~7.74 -0.0329 0.9937 (100) 3  Slope SE o f s l o p e SE o f s l o p s a s % Intercept Correlation coefficient Relative slope Ranking  0.8382 +0.0517 6.17 -0.0445 0.9973 (104) 2  Slope SE o f s l o p e SE o f s l o p e as % Intercept Correlation coefficient Relative slope Ranking  0.5752* +0.0431 7.49 0.0275 0.9858 (71) 4  Slope SE o f slope SE o f s l o p e as % Intercept Correlation coefficient Relative slope Ranking  0.4618 +0.0826 T7.89 -0.0943 0.9873 (57) 5  3  0.4675 +0.0611 T3.07 -0.0972 0.9928 (53) 5  Slope SE o f s l o p e SE o f s l o p e a s % Intercept Correlation coefficient R e l a t i v e slope Ranking  1.0202 +0.1093 TO. 71 -0.1859 0.9566 (127) 1  b  0.8875 +0.0591 6.66 -0.1168 0.9835 (101) 2  m  0.8940 +0.0405 4.53 -0.0840 0.9970 (101) 1  b  C  0.6769™ +0.0361 5.33 -0.0520 0.9827 (77) 4  8  A  0  ! • A n a n a l y s i s o f oovariance o f d r y body weights w i t h the o o v a r i a t e ( p r o t e i n intake) h a v i n g d i f f e r e n t s l o p e s f o r d i f f e r e n t p r o t e i n sources i n d i c a t e d P > 0.001, when c a l c u l a t e d both by i n c l u d i n g and e x c l u d i n g PF groups. 2.  Slopes w i t h t h e same s u p e r s c r i p t do n o t d i f f e r s i g n i f i c a n t l y ( S c h e f f e s ' t e s t P = 0.05).  89  T a b l e 24.  Slope r a t i o s o f body p r o t e i n g a i n s on p r o t e i n i n t a k e f o r f i s h f e d t h e v a r i o u s p r o t e i n sources. C a l c u l a t e d both by i n c l u d i n g and e x c l u d i n g t h e p r o t e i n f r e e (PF) f e d groups.  Protein Source  Slope r a t i o s o f body p r o t e i n gains e x c l u d i n g PF groups i n c l u d i n g PF groups  c  FPE  Slope SE o f slope SE o f s l o p e a s % Intercept Correlation coefficient Relative slope Ranking  0.5588"+0.0422 7.55 -0.0541 0.9849 (100) 2  0.5756 +0.0286 4.97 -0.0652 0.9943 (100) 1  FRH  Slope SE o f slope SE o f slope a s % Intercept Correlation coefficient R e l a t i v e slope Ranking  ab 0.4255" +0.0350 8.23 0.0000 0.9986 (76) 3  BC 0.4896 +O.0268 5.47 -0.0454 0.9887 (85) 3  LTH  Slope SE o f slope SE o f slope as % Intercept Correlation coefficient Relative slope Ranking  0.3785 +0.0292 7.71 -0.0029 0.9818 (71) 4  HTH  Slope SE o f s l o p e SE o f slope as % Intercept Correlation coefficient R e l a t i v e slope Ranking  0.2807 +0.0560 19.95 -O.0715 0.9807 (57) 5  CS  Slope SE o f s l o p e SE o f s l o p e as % Intercept Correlation coefficient Relative slope Ranking  0.6489 +0.0740 Tl.40 . -0.1327 0.9661 (116) 1  ab  3  1  •B 0.4345' +0.0238 5.48 -0.0405 0.9839 (76) 4 0.2711* +0.0404 14.90 -0.0671 0.9881 (47) 5 BC 0.5483 +0.0390 7.11 -0.0790 0.9838 (95) 2  T~. An a n a l y s i s o f c o v a r i a n c e o f body p r o t e i n gains w i t h the c o v a r i a t e ( p r o t e i n i n t a k e ) having d i f f e r e n t s l o p e s f o r d i f f e r e n t p r o t e i n sources i n d i c a t e d P > 0.001, when c a l c u l a t e d both by i n c l u d i n g and e x c l u d i n g PF groups. 2.  Slopes w i t h t h e same s u p e r s c r i p t do not d i f f e r s i g n i f i c a n t l y ( S c h e f f e s ' t e s t P = 0.05).  -  90  -0-20 0  0 10  0 20  0 30  0-40  0-50  PROTEIN INTAKE  Fig.  10.  intake protein points free  Slopes  of  chinook  sources. obtained  diets  of  (see  0-60  070  0-80  0-90  100  MO  (g/ FISH)  dry body w e i g h t  salmon  fed  The s l o p e s from g r o u p s Table  23).  91  diets  gain  on  containing  were c a l c u l a t e d of  fish  protein  fed  the  the  test  including protein  -0-10  .  1  0  0 10  .  .  0-20  0-30  .  . 0 40  . 0-50  PROTEIN INTAKE  Fig. of  11.  Slopes  chinook  salmon  of  fed  The s l o p e s  obtained  from g r o u p s  (see  protein  diets  .  .  0-80  0-90  . 1 00  gain  1 10  on p r o t e i n  containing  the  test  were c a l c u l a t e d  including  of  protein  T a b l e 24) .  -  . 070  (g/ FISH)  body  sources.  . 0-60  92  fish  fed  the  intake protein  points  free  diets  exception  of  FRH s i m i l a r  either  body w e i g h t  3.3.9  Available  damage  in  values  are  available  in  shown  al.(1958) is  5% of  dietary  dietary  Ogino of  al.,  (Table  25).  egg  available by K e t o l a  almost  sufficient  (1982)  with  the  suggest  that  a certain  in  protein the  and a r e  for  (Table  the  (1982) lysine  of  The  to  calculate  25). for  to  chinook  containing  5.3%  the the  Halver  showed t h a t was  lysine  compared  diet  was  protein  used.  requirement  2% of  sources  extent  method  requirement 1958)  only  However,  on the  et salmon 40%  the  minimum  and 6.1%  of  source  of  due  the  to  to  fish  amount  meal.  93  to  lysine  the  lysine  dietary  meet  the  requirement  is  sufficient requirements  LTH and FRH c o n t a i n e d  lysine  of  5% of  FPE c o n t a i n e d  25).  available  performance  the  to  lysine  of  HTH f a i l e d  only  (Table  value  if  relative  available  coupled  of  or  trout  lysine  Based  drying  lysine  by  sources  protein  the  salmonids  composition,  1982).  this  the  diets  and K e t o l a  lysine  et  stated  protein  They were used  the  of  were o b t a i n e d  respectively.  the  (Halver  on f i s h  surplus  of  protein  rainbow  of  evaluate  (2).  the  (1980)  protein  requirement based  that  the  processing  requirements  Employing protein  content  stated  requirement  the  in Table  lysine  the  protein.  to  to  values  gain.  of  content  an a t t e m p t  slope  protein  content  lysine  relation  known l y s i n e  body  lysine  The a v a i l a b l e determined  or  relative  meet r e q u i r e m e n t s content fed  lysine severe  obtained  HTH, the was  f o r HTH  results  rendered  heating  (Ketola,  may  unavailable  during  the  Table 25.  A v a i l a b l e l y s i n e , l y s i n e requirements f o r chinook salmon and percentages s u p p l i e d by each p r o t e i n source ( i n p a r e n t h e s i s ) .  of  requirements  % Crude P r o t e i n i n D i e t s 7  17 % of Req  27 % of Req  37 % of Req  47 % of Req  % of Req  FPE Available lysine  % of dry d i e t  0.59  Halver et al.(1958)5% of p r o t e i n K e t o l a (1982) 8.8% p r o t e i n FRH  0.34 0.59  1.45  2.33  3.08  3.95  Requirement:  Available lysine  (171) (97)  0.85 1.49  % of dry d i e t  0.99  Halver et al.(1958)52 of p r o t e i n K e t o l a (1982) 8.8% p r o t e i n LTH  0.90 1.58  Available lysine  % of dry d i e t  0.90  Halver et al.(1958)5% of p r o t e i n K e t o l a (1982) 8.8% p r o t e i n HTH  0.84 1.48  Available lysine  % of dry d i e t  0.77  Halver et al.(1958)5% of p r o t e i n K e t o l a (1982) 8.8% p r o t e i n CS  0.85 1.50  Available lysine  % of dry d i e t  1.37  H a l v e r et al.(1958)5% of p r o t e i n K e t o l a (1982) 8.8% p r o t e i n  0.93 1.63  (171) (97)  1.36 2.40  (171) (97)  1.51  1.80 3.16  (171) (97)  2.17  Requirement: (109) (62)  1.38 2.43  (109) (62)  1.53  1.98 3.48  (109) (62)  2.07  Requirement: (108) (61)  1.42 2.50  (108) (61)  1.25  1.92 3.38  (108) (61)  1.79  Requirement: (91) (52)  1.27 2.41  (91) (52)  1.95  2.00 3.51  (91) (52)  2.65  Requirement: (148) (84)  1.32 2.32  (148) (84)  1.80 3.17  (148) (84)  2.31 4.06  (171) (97)  3.A  DISCUSSION  3.4.1  The e f f e c t weight  of  growth  protein  supply  influence  studies  of  p r o t e i n would  on g r o w t h .  calculate  Various  growth  Zeitoun  (1973)  compared a v e r a g e  initial  weight  and s p e c i f i c  weight 1969)  per  day).  concluded  compare growth gain  of  describe  and L l o y d ,  1959).  inferences  could  results source  salmon.  With  HTH ( F i g s .  of  nutrient Daily  of  been  in  any  growth  for  fish  3B,  3C)  for is  in  rates  do not  a lag  differing  rate  fed  the  likely  due  to  adjust  effect  nutrient  in  the  -  it  to  daily  does  not  (Crampton  similar  the of  final  body  effect  that  chinook  the  protein  influence  acclimation to  has  dietary  increase  levels  day 21  protein  metabolic  composition  95  way  average  by b o t h  from  low  instantly the  al.,  age  study,  dietary  growth  during  et  HTH body w e i g h t  fish  accumulated  body  6).  fed  fed  in  because  performance  fish  of  useful  of  demonstrated  had on the  linear  21  (Table  study  of  most  growth  a  fish  gain  (Brett  specific  present  have  methods.  (% i n c r e a s e  The use  to in  percent  critized  at  the  rate  The d e c l i n e  of  rate  fish.  change  exception  reserves  The e x i s t e n c e diets  the  day 0 to  growth  has  growth  and l e v e l the  3A,  gain,  be made by e x p r e s s i n g  27% and h i g h e r . to  investigators  p r o v i d e d the  size  However,  approximately  compared  on body  be e x p e c t e d  investigators  latter  weight  and s p e c i f i c  protein  was  the  different  accurately  The  and l e v e l  by d i f f e r e n t  daily  He and o t h e r that  rate  growth  i n animal experiments  weight  source  gain  An i n a d e q u a t e profound  dietary  to  42  diets  or  of  period. change.  adaptation  been  well  body  to  of  established fish  (Maynard  n u t r i t i o n by S t e f f e n s  indication would  of  protein  be r e a c h e d  The e f f e c t  of to  weight  was  gain  dietary  juvenile weight the  time  study,  1983)  had the  diet  growth  response  followed  a smooth  increase  (Fig.  plaice  fed  smooth  c u r v e was Since  intakes,  the  sources intake  was  unit was  of  the  not  this  than  to  casein  protein  conjectured  based  actual  this  of  was  the to  of  protein  diets  (CS) weight  due  96  adaptation. similar  in  fish  for  feeding  most  trials  reported  by H i g g s  et  experimental influence  of with  that  In c o n t r a s t  experimental protein  to  al. period,  of  the  period  the  to  supplied  decreasing was  (Cowey  fish  slopes  dietary  intake, that  the  tests  period.  observation  with  rate  that  1983)  the  a steadily  considered  compared by the  a longer  levels  cod muscle  of  of  reported  a 42-day  obtained was  reason  in  reliable  exponentially  masking  trial  A similar  performance  rather  although  the  of  curve with  freeze-dried  tested.  during  a  growth  exponential.  rates  increasing  4).  likely  al.(1982,  effect  that  w h i c h was  Higgs  following to  diet,  increase  growth  during  Nevertheless,  most  nearly  recognized  2 week p e r i o d of  period  et  was  of  results  was  the  by means  From the  were measured  perhaps  acclimation  to  This  who s t a t e d  a 1 to  the  noted  salmon,  over  present  which  is  treatments.  gain  (1982,  after  FPE-47,  chinook  (1981)  acclimation  not  1969).  utilization  only  the  composition  the  and L o o s l i ,  et  other be due  fed  the  weight  rate  reported al.,  for A  sources  differing  food  various  protein  gain  protein  to  concentration. supported gain  of  1972).  protein to  by FPE  was  rapid not  a palatability  Thus, growth  realized.  per It  problem with  the  CS d i e t s .  sources  The n u t r i t i o n a l q u a l i t y  will  be  The growth fish  fed  or  low  to  rear  rates  diets  juvenile of  of  lowering  this  study  growth  rate  of  3.4.2  The e f f e c t intake  in  fish  is  and Putman,  assuming  that  all  report  salmon was the  largest  level  of  tables  fish  (1971)  influenced relative  dietary  (Fowler  Food  other  standardized.  by B r e t t  represents  that  the  1973),  biotic  is  and B u r r o w s ,  of OMP  OMP c o n t a i n s origin,  impair  of  caloric  this food  the  for  the  on  food  agree  intake  of  in  1971);  (Brett,  study  smaller  of  of fish  fish  hatchery  1979)  with  of  a major i n f l u e n c e  weight  consuming  size  on  feeding  on the  -  97  particular interest gain  profit  to  food  a  sockeye  1971). of  a  and  content  (Brett,  factors  The i n f l u e n c e  a parameter  The r a t i o  used  The  content  plant  size  and a b i o t i c  accounted  most  efficiency  and f i s h  weight,  of  freeze-dried  and l e v e l  by the  that  those  survival.  source  The r e s u l t s  to  diet  However,  conversion  amount.  is  protein  hatcheries.  ocean  regulated  by  the  OMP would p e r h a p s  protein  by f i s h  similar  protein  50%.  who showed  intake  conversion  producers.  of  food  (Lee  been  local  i n c l u d i n g some of  dietary  diet  have  OMP i s  and c o n s e q u e n t l y  of  various  supplied  at  present  level  and g r o s s  Food i n t a k e the  suggest  sources  fish  OMP were  meals.  salmon  from the  protein  fed  37% p r o t e i n  dried fish  protein  the  chinook  chinook  be r e d u c e d  variety  of  the  later.  containing  temperature  findings could  discussed  of  to  intake  m a r g i n i n an a n i m a l  production of  enterprise.  increase  i n GFC to  The t r e n d increments  concentration  are  consistent  investigators  who have  For e x a m p l e ,  Gulbrandsen  and Utne  maximum GFC when and 45%.  More  adequately conditions. is  under They  similar obtained  series  of  protein.  because  of  GFC v a l u e s  test  level  in  without  suppressing  protein  is  by GFC i n  with  ranging  less  the  protein.  sources  study  reports  cannot  experimental et  al.(1982)  same s p e c i e s present  50%  this This  study  the  the  with  suggests  diets  could  quality to  and  study.  approximately  judged  were  between 35%  a  salmon  GFC, provided that  lay  20% and 25% f o r  containing in  of  obtained  by H i g g s  on the  between  were o b t a i n e d  present  diets  these  as  other  (1974),  differing  conditions  The p r o t e i n the  Satia  the  conducted  p r a c t i c a l chinook  high.  of  widely  practical diets  substantially  of  requirements  an e x p e r i m e n t  was  experimental  protein  value  it  Higher values  containing  level  Comparison with  more v a l i d  protein  rate  protein  results  (1973),  comparisons  be made b e c a u s e  dietary  decreasing  and K o s h i i s h i (1980)  protein  precise  a steadily  the  the  Zeitoun  (1977)  the  in  with  studied  salmonids.  of  diets  that be  the  reduced  of  dietary  be of  highest  freeze-dried  fish  meals.  3.4.3  The e f f e c t intake  (Lee  dietary  and g r o s s  As m e n t i o n e d needs  of  energy  previously,  and Putman,  protein  1973).  source  and l e v e l  on  energy  utilization  fish In  98  eat this  to  satisfy  study  their  an a t t e m p t  energy was  made  to  formulate  the  diets  (except  metabolizable  energy  this  practice  in  been  reviewed  and h a r s h l y  main o b j e c t i o n s measure  fish  pertain  energy  inaccurate  caloric  In  study  this  replaced  dietary  to  satiety  one  to  be  similar  at  however,  suggests too  kcal/g;  low  that or  each  of  was k e p t  protein  level.  energy  and l e s s  c a r b o h y d r a t e was  on t h e i r  fish  However,  value  concentration  the  on d i e t a r y  while  was  shown was  too  energy  of  the  to  decrease  In the  Thus  an e r r o r  introduced  by not  in  correcting  the  high,  diet  the  consuming  99  This was  causing  an  carbohydrate  since  and  study  was equal  to  as  Nose,  may have  ascribed  is  their  progressively  study  ME v a l u e  trend  polysaccharides  (Singh  present  fish  intakes  protein  monosaccharides  present  the  energy  to  high  in  of  basis  monohydrate and p a r t i a l l y h y d r o l y z e d d e x t r i n  employed.  of  carbohydrate.  the  increased  absorption level.  to  protein  Since  by f i s h  assigned  receiving  metabolizable  trout  used  The g e n e r a l  intake  for  MEI i n  His  of  energy  metabolizable  value  the  use  constant  energy  of  recently  estimates  and the  of  coefficients.  more p r o t e i n  concentration  glucose  feeding,  the  the  dietary  of  dependency  may e x p e c t  their  depend  of  has  methods  carbohydrate = 4 k c a l / g ) .  in  to  the  lipid  digestibility  1967).  u n r e l i a b i l i t y of  either  Actually,  dependent  the  a decreased  containing  overestimation diets.  was  on a  by J o b l i n g ( 1 9 8 3 ) .  by c a r b o h y d r a t e on a m e t a b o l i z a b l e  fed  diets  to  conversion  were  the  criticized  on l e v e l  = 4.5  noted,  isocaloric  The i n a c c u r a c y and i n c o n s i s t e n c y  energy,  (protein  (MEI)  be  growth and n u t r i t i o n s t u d i e s  metabolizable  metabolizable  was  basis.  OMP) to  not  found  portions were been  carbohydrate Higgs  et  with  al.(1983)  ascribed  to  the  fiber  high  They  also  study) of  their  was  of  the  in  as  proteins  diet.  also  in  of  the  diet. they  and energy  due  protein  diets.  low  (cellulose  in  the  and i n c r e a s e s  interesting  also  the  dilemma which  protein  fiber  is  diet  showed  appeared  of  to  study,  to to  for  fish  Pieper  eat  rate  that  to  to  present  the  note  relative  from  lipid.  The r e l a t i o n s h i p states  that  fats,  recorded  Perhaps  a source  of  that  the  the  satisfy  when n i t r o g e n  energy  requirements  between n i t r o g e n  100  of  due not  available  maintenance  value  (1980)  diets  (N)  intake  intake is  and  p a r a m e t e r may of  found  GEU to  various  to  extremely  fed  was  diet.  nutritive  and p r o t e i n s  fish  utilization  level  this  utilization  this  of  the  protein  HTH was  for  energy  of  and P f e f f e r  receiving  sources. HTH as  ratio  dietary  evaluating  carbohydrates,  of  dietary  on GEU, s u g g e s t i n g  diets.  values  that  protein:calorie  present  The e s t i m a t i o n  retention  of  contents  It  c o m p a r i n g the  protein  inadequacy  3.4.4  the  The GEU of  comparison other  by the  fish  in  supplements  a similar  dietary  digesta.  a criteria  for  be u s e f u l  level  mineral b i o a v a i l a b i l i t y  had an e f f e c t  used  its  requirements.  influenced  source  but  the  that  and Putnam (1973)  Similarly,  be  and d e x t r i n  protein-free  energy  Lee  for  impaired d i g e s t i b i l i t y  depresses  fed  encountered  suggested  transit  fish  consideration  a  basal  poor  in  containing only  to  from  for and  zero,  the  the  protein  protein nitrogen  nitrogen  retention  is,  nitrogen  excretion  retention  is  nitrogen 1955a;  and must  consider  nitrogen  Several  Birkett,  equal  (Bressani,  zero,  loss.  be,  1969;  to  1977).  intake  studies  Savitz,  endogenous n i t r o g e n  the  total  Conversely,  should  discussed  1971; loss  endogenous  Brett  to  equal  when  nitrogen  endogenous  earlier  (Gerking,  and G r o v e s ,  represent  1979)  the  protein-nitrogen  requirement  for  maintenance  in  fish  1).  requirement  for  maintenance  of  chinook  The p r o t e i n  fry  was  estimated  containing was  made t h a t  FPE was  is  97% d i g e s t e d  valid and i s  et  al.,  The m a i n t e n a n c e contrasted fishes  above  with  reported . 22.1mg larger present  by B r e t t  fish  by the  1974)  al . ,  in  This  that  FPE was  digestibility  and r a i n b o w  present  endogenous  1979).  assumption  trout  1984).  the  daily  on  diets  fish.  found  reports  salmon  fed  The  Fig.  study  may be  nitrogen  The p r o t e i n  excretion  requirement  an endogenous  nitrogen  excretion  rate  of  This  in  agreement  with  that  value  and Z a l a  is  (1975)  although  a higher  for  their  water  close  sockeye  salmon,  estimate  was  temperature  than  namely  obtained in  using  the  study.  Gerking  (1955a,b)  requirements days  to  BW/day,  and at  et  requirement  BW/day.  N/lOOg  Opstvedt  of  with  al.,  fish  dry f o o d .  assimilated  et  and G r o v e s ,  corresponds  for  a p r e l i m i n a r y study  (Cowey  estimates  (Brett  25.5mg/100g  totally  i n agreement  1980;  BW/day  3950 k c a l / k g  since  raw cod by p l a i c e  (Skrede  in  be 0 . 1 6 0 g / 1 0 0 g  approximately  assumption  of  to  (see  and t h e n  of  was  fish.  they  were  the  first  Bluegill fed  to  sunfish  daily  101  study  maintenance  were s t a r v e d  an amount  of  for  glucose  protein three  via  stomach  tube  Gerking  (1955a)  nitrogen this  equivalent  daily  initial  the  were  decline until  placed  the  rate  reserve  ensues at previous  (1918)  is  the  stabilization Body w e i g h t  of  data  stabilization protein-free Gerking  nitrogen  (1955b)  fish  difference  beginning  mealworms. N/day  was  the  in  42  the  in  the  of  necessary  to  Morgulis and f o u n d  a four  suggest  fish  time  period  level  excretion.  12)  adaptive  excretion  brook t r o u t  (Table  an  this  time  following  reserves.  is  nitrogen  have  endogenous  minimum amount  nitrogen in  and the  mealworms to  regarded  to  from  shown  in  since  a p e r i o d of  The h i g h e r  excretion study  there  that  fish  excretion  over  of  unlikely  protein  p e r i o d of  balance  5.84mg  is  During  nitrogen  fish.  week  no  fast.  that  fed  the  diet.  the  were f e d  this  the  of  of  diet  established.  longer  it  nitrogen  and u r i n a r y  the  day f a s t e d  excretion  had o c c u r r e d by day  equivalent  the  the  in  However,  level.  of  excreted  depleted  nitrogen  nitrogen  maintain  were  fully  minimum l e v e l  determined  BW) .  endogenous  depleted  nutrition,  sunfish  on a p r o t e i n - f r e e of  rate  upon a t h r e e  a minimal constant  establish  the  of  a new e q u i l i b r i u m i s  protein  being  based  measure  metabolic  a 29.7g  p r o b a b l y not  are  in  that  estimate  a good  When f i s h  the  (19.7mg N / l O O g  provided fish  found  to  at  body end of  nitrogen  rates  98% of  the  maintain a 29.7g  of  as  required  to  was fish  in  estimated  sunfish  determined sampled  period.  intake.  protein  (1955b)  102  This  feeding of  excretion  protein  content  a 30 day  A c c o r d i n g l y , Gerking r e q u i r e d to  of  equilibrium.  different  absorbed  nitrogen  in  The  The  fish  sunfish  the that  7.2mg  nitrogen  at  equilibrium  at  amounted  24.24mg  this  to  value  nitrogen  to  25°C.  be  Ogino  to  for et  19 d a y s .  feeding  analysis  than  Other  estimates  obtained protein  the  from diets,  that  salmon 23.5%  the  to  either  the  maintenance  the  casein-gelatin method  of  growth was In a s t u d y  maintenance  in was  These  protein fish  from  in  in  for  protein  study,  poorly of  it  by  this  fishes,  maintenance  of  low  salmon  effects  of  or more  dietary  for  diets  would a p p e a r  percentage  26.  young  rats,  protein  level  was  Henry  about  103  (1965)  2% f o r  found  egg  coho  was low  that likely,  protein  inaccurate. with  fed  a l . ( 1973,1974)  based  assimilated,  protein  in Table  requirement  the  and  study.  in  chinook  F P E , Z e i t o u n et  14  considerably  either  shown  of  be 8.89  are  a series  casein-gelatin  their  and  periods  to  values  are  Discounting  extrapolation  reported  endogenously  were f e d  feeding  dietary fed  for  salmon  estimate  from  diet  requirements  analysis,  present  maintenance  glucose  carcasses  excreted  chinook  where  7% p r o t e i n  from  oral  estimate  trout  lost  nitrogen  for  carcass  regarded  obtained  following the  rainbow  nitrogen  obtained  water  that  them a p r o t e i n - f r e e  and 15% r e s p e c t i v e l y . of  with  (1955b)  requirement  fry.  analysed  and r a i n b o w t r o u t ,  salinity  zero  salmon  respectively.  and c o n d u c t i n g  found  agrees  or e s t i m a t e d  containing  their  of  with  a q u a r i u m water  experiments  In c o n t r a s t diet  of  nitrogen  Gerking  agreement  and by t o t a l  one  maintenance  BW/day.  estimated  BW/day,  lower  the  value  chinook  They  9.96mg/100g  diets  This  al.(1980)  after  carcass  close  determination  study  excreta  N/lOOg  in  administration. this  Thus,  that  the  a l b u m i n and  at  a  Table 26. Estimated absorbed nitrogen requirement f o r maintenance of f i s h , determined by feeding and carcass a n a l y s i s experiments.  Size (grams)  Species  B l u e g i l l sunfish (Lepomls macrochlrus)  Water Temperature (degrees C)  11 - 34  17  0.2 - 1.8  Reference  Gerking (1955b)  lugwonns  20.3  B i r k e t t (1969)  17  whiteworms  24.4  B i r k e t t (1969)  17  lugwonns  38.1  B i r k e t t (1969)  100 - 150  17  earthworms  17.3  B i r k e t t (1969)  Carp (Carassius auratus)  2.5 - 4.5  20  chironomid larvae  22  Iwata (1970)  Rainbow trout (Salmo g a i r d n e r i )  0.9 - 26  12 - 19  Carp (Carassius auratus)  1.5 - 12  19 - 25  Y e a r l i n g sole (Solea v u l g a r i s )  25  Requirement (mg/lOOgBW/d) 24.24  Yearling plaice (Pleuronectes platessa)  30  Diet  Y e a r l i n g sole (Solea v u l g a r i s ) Perch (Perca  5-53  fluvlalis)  Chinook salmon (Oncorhynchus tshawytscha)  1.56  Chinook salmon (Oncorhynchus  1.56  Ogino (1980) protein-free  protein-free  protein-free  13.9  Ogino (1980)  20.7  Present study  Present study  11  FPE-7  t&hawytscha)  -  10A  -  herring  meal  increased were  and between  when  poor  quality  fed.  Similarly,  for  maintenance  level  dietary  protein  required amount  for  of  approximately  note  should  dangerous  requirements  of  protein  organ w i t h i n maintenance  since of  they  the  do not  consider  growing  animal.  of  protein  intake  by F i g .  1.  The e f f e c t  of  dietary  or  to  for  are  6).  in  on t o t a l  nitrogen  body  balance  a steady the  the  (Fig.  protein  based  state  does  level  nutritional  of  status  The p r a c t i c a l  the  they  according  source  to  fish  culture  physiological  Rather  protein  was  the  requirements  the  partitioning  of  body.  protein  protein  Whereas a  employed  respect  Maintenance  metabolism  dietary  zein  FRH and L T H , d o u b l e  26)  maintenance  and  180mg/100g BW/day  requirements  (Table  by f i s h .  tissue  application  values  reflect  a particular  be added w i t h  maintenance  turnover  individual  containing  requirement  gluten  source.  r e q u i r e d when HTH was  necessarily  3.4.5  protein  The  the  was  fish.  described  that  protein  These  the  observed  as  diets  growing  is  of  such  fed  of  of  was  varied with  intake  interpretation  not  it  proteins  casein.  fish  A final  nitrogen  2% and 3% f o r  to  on  serve the  to  aid  in  scheme  protein  utilization The effect  results that  utilization protein  of  this  proteins of  of  dietary  utilization  is  study  demonstrated  different protein also  in  nutritive growing  a function  105  the  of  magnitude  value  had on  salmon.  Since,  protein  intake,  of  the  the  the  protein  sources  were compared at  different  dietary  concentrations. The of  relationship  protein  for  between p r o t e i n  maintenance  described  in  described  by O g i n o et  protein  consumed  efficiency, did  section  the  the  and  (3.3.5) al.  followed  (1973)  used  p r o p o r t i o n used  however,  concomitantly  economic  loss,  for  nitrogenous  dietary  most it  between value.  desired  is  mode of  entirely  The amino a c i d compares chinook has  been  amply  patterned produced Ketola,  after the  known to  cause  availability. plaice  with  and the  the  FPE was any  stated  that  composition in  of  fish  detrimental  of  fish  fish fed  muscle  poor and  amino  by Cowey et  since  muscle  (Table  diet  the  proteins.  requirements  on amino  rats  fish.  growth  1).  for It  acids  eggs or whole  with  of  vary  and  by f r e e z e - d r y i n g  (1953)  form  an  surprising  eggs  as  loss,  to  pollock  a basal  effects  106  gradually  confined  not  skeletal  of  f o r maximum  the  found  supplemental  processed  and Bender  in  amino a c i d  Experiments conducted  and M i l l e r  is  that  representing  excellent  freeze-dried  composition  gain  needed  excellent  utilization  of  the  from  to  amount  Exogenous  was  of  teleostean  demonstrated  highest  1982).  protein  composition  that  pose a h a z a r d to  sources  pattern  As the  accumulation  protein  utilization excreted  decreased  Apart  FPE s u p p o r t e d  made up of  favourably salmon  protein That  carp.  growth  waste  The u t i l i z a t i o n  of  a similar  above  increased.  and the  and amount  maintenance.  feces  nutritional  for  for  ammonia and d e c o m p o s i n g  considerably  growth  by c a r p i n c r e a s e d protein  intake  fry  (Arai, which  1981; is  not  acid al.(1971)  have  shown  with  excellent Cowey et was  protein al.(1972)  completely  over  FRH was  and  reported  digested  90% d i g e s t e d  both  utilization  at  some of  compared  the  and anchovy  level  almost  as  to  of  white  fish  comparable  the  source.  meal  fishmeal  Cowey et  et  meal  protein  al.(1971)  in  Watanabe  rainbow  utilization obtained  dried in  observed (30°C)  their  different  different  drying conditions  growth  biological  The f i n d i n g s  presumption  FPE.  may not  diet.  Although  have  et  fish  been  al.  trout  in  (1983)  diets.  trout  with  respectively,  supported  acids.  i n the  sardine They  regardless  of  FPE and FRH, a w h i t e  support  the  observations  al.(1983).  indistinguishable  the  and  p r e p a r e d from whole  proteins  The r e s u l t s  temperature  affected  levels  inclusion as  protein  brown meals made from g r e e n l i n g ,  and a brown meal  Watanabe  of  muscle.  cod muscle  dietary  efficiently  Recently,  kinds  lower  FRH was  integumental  reported  low  highest  assimilated.  three  freeze-dried at  meals were f r e e z e - d r i e d ,  completely  of  that  v a c u u m - d r i e d cod  by p l a i c e  the  utilized  with  although  that  although  cod meal  essential rates. for  this  the  study  effect  content,  They presumed  the  cod meal  of  of  that  had to  certain  would  they  the  some  extent  essential  corroborate  a mild  and  almost  amino a c i d  availability  in  were  freeze-dried  the  amino same  d r y i n g temperature  was  minimal. LTH was ideal  processed  commercial  depletion attempt  of  the  should  in  a laboratory  facility  to  h e r r i n g meal.  C o n s i d e r i n g past  herring stocks  in B r i t i s h  be made  towards  and  Columbia,  i m p r o v i n g the  107  simulate  an  present every  utilization  of  the  rendered that  product.  whole  The p e r f o r m a n c e of  h e r r i n g can be c o o k e d ,  product  suitable  adverse  effect  herring  meal  included  to  been  of  have  utilization probably It  severely  low  rate  (1954) of  steam  dried  from  their  formation (Bender,  of  of  was  destruction  was  of  are as  lysine  amino a c i d s  i n heated  destroyed  heat  present  the  amino a c i d s  between the  during not  digestion,  due  can  (Tarr  extremely  as  and et  biological compared  to  HTH may have  oxidation, amino a c i d s and,  so  the digestion  by odor and d a r k  color, minor  to  1)  methionine  temperature  suffered  less  (Table  more damage. al.  and l y s i n e  somewhat  108  drying  Although a r e l a t i v e l y  may have  rendered  and  to  diet.  Tarr  hydrolyzed during  evidenced  lower  digestive  study.  by  likely  value  by the  damage  h e r r i n g meal C a r p e n t e r et  but  this  during  drying  the  from LTH i s  and b i o l o g i c a l  flame  HTH had o c c u r r e d .  available  in  digestibility  to  that  appears  protein  present  the  linkages  delayed that  noteworthy study  produce a  little  growth and h i g h  processing  tryptophan, arginine,  totally  in  subjected  the  with  amply d e m o n s t r a t e d  protein  to  The s l i g h t l y  excessive  HTH f o r  Certainly,  of  other  of  some of  1972).  observed,  that  a r e d u c t i o n of  linkages  of  LTH.  ingredients  During  release  destruction  that  this  the  some c h a r r i n g  example,  of  meal.  modification that  This  menhaden meal  resulted  to  is  LTH d e m o n s t r a t e s  and d r i e d  in this  protein d i g e s t i b i l i t y  observed  value  other  utilization  loss  It  fed  diets  i n OMP compared t o  established  1972).  of  quality.  i n OMP t e s t e d  protein the  pressed  salmon f r y  similar quality  impair  metabolic al.  of  well  and B i e l y ,  chinook  on p r o t e i n  due to  is  for  fish  (1962) were  was For  found  not  available.  Opstvedt  et  al.  (1984)  pollock  reported a depression  muscle  investigators heat  (above  bonds.  noted  95°C)  protein  Several quality  animal  coincided via  with  and the  available  Ketola salmon  the  lysine in  have  Hence,  to  the  lysine  should  foods.  He r e p o r t e d  8.8%  of  p r o t e i n which agrees w i t h  from  the  analysis  requirement  CS c o n t a i n e d  as  that  of  on a f e e d i n g  lysine and  be used  eyed  trial, was  5% of  lysine  FRH and LTH a p p e a r e d  Halver  to  be p r o t e i n  lysine  supplementation  chinook  salmon  according  et  March values  et  the  of  diet  fish  have  to  be  composition  egg  amino  acid  fish  chinook  of  8.65%  obtained  this  study  (Table  Table  stated  25  eggs  that  is  1). the  shows t h a t FPE  stated  sources  that  requirements. would  practical diets  lysine  the  of  ( 1958)  most  of  of  requirement  for  not juvenile  proposed  by  ( 1958) . al.(1966)  were not  biological in  al.  to  in  in  al.  protein  formulating  value  protein.  warrant  Halver  et  to  to  determination  the  content  eggs  i n excess  relate  the  for  the  chinook  found  amino a c i d  that  lysine  due  disulfide  requirement  of  the  a guideline the  to  lysine  eggs and p r o p o s e d  of  were not  chemical  published  of  study.  content  pattern  Based  residues  the  trout  in d i g e s t i b i l i t y  formation  their  in  treatments.These  attempted  both  (1982)  heat  depression  biologically  lysine.  chinook  the  investigators  considered.  various  digestibility  measured  available  to  that  Cross-linkages  affect  of  subjected  in d i g e s t i b i l i t y  found  assays,  combination with  reported to  be  that  significantly  supplementation arginine  although  of  the  lysine  correlated  with  the  fish  with  lysine  and m e t h i o n i n e  109  available  meals  significantly  increased  chick  the  lysine  total  temperature available  weight  content  content  former.  The low  somewhat  (sic)  products  contained  requirements  meals  reduced  for  based  amino  distinctly  inferior  differentiate  of  relied  inclusion The  trials  in  have meal.  to  at  43°C  to  et  1971).  of  lysine  for  it  al.(1954) effects  of  diets.  this  drum.  meal  for  attributed  occurred after  heat  prolonged  to  of  fish  meals  estimates  of failed  to  for that  protein  value  of  of  the  sources proteins  salmon. diets to  c o n t a i n i n g HTH  other  nutrients  a number of  in  feeding  on the  nutritive  value  They compared  presscake  dried  heated  2,  They n o t e d  heating  done  conducted of  et  availability  various  fed  a  the  quality  nutritive  by f i s h  the  both  may be c o n c l u d e d  chinook  the  supported  improved q u a l i t y  the  the  the  the  low  in  March  detecting  or  in  juvenile  also  that  But,  than  of  protein  in  study,  protein  chick  after  value  for  the  in a rotating  nutritive  acids  in  al.,  average  predict  and  However,  excess  i m p a i r e d by damage  portions  were d e s t r o y e d in  of  determine  h e r r i n g meal  149°C  diets  et  lower  product  However,  this  upon to  Tarr  of  in  was  plaice.  analyses  of  cod m u s c l e  in  were u s e f u l  available  been  lysine  quality.  utilization  may a l s o herring  acids  of  estimates  among meals  determination  for  that  latter  rate  (Cowey  observed  indistinguishable.  air-dried  available  Similarly,  be  the  upon l a b o r a t o r y  individual  cannot  of  growth  plaice  al.(1971)  freeze-dried  temperature  concluded  poultry.  in  Cowey et  d r i e d cod were a l m o s t  lysine  al.(1961)  gains.  for  that  1 hour,  1,  110  -  3 hours  heat-labile  whereas  availability  heating  or  (Bisset  of  the  at  vitamins impairment  essential  and T a r r ,  amino  1954).  By c o m p a r i n g h e x a n e - e x t r a c t e d heat  damage,  growth  that  heating  could  was  extraction  Biely  et  al.(1955)  meals  heating  extracted  and c o m p l e t e of  fresh  made to  extract  levels  However, of  HTH i n  utilization Casein diets  for  due has  in  and K e t o l a , al.(1958)  a higher with  lipid  been  used  certain  the was  that  synthesis level this  salmon  These  it  of  in  CS i n  study  was  relative d i d not  findings  to  both the  damage  an a t t e m p t  h e r r i n g meals  with  growth  and  may have  to  as  alone  diet  has  been  for  in  its  The major  apparent  high  intake  low  quality  support  a growth  concur with  reports  111  Halver  rate  was  was  of  depressed  meal  for  protein.  comparable  by o t h e r  at  encountered  palatability  rate  et  deficiency  study  problem  fish  Rumsey  results  an e f f i c i e n t  food  test  be  1980;  this  The  in  to  an a r g i n i n e  and m e t h i o n i n e .  diet.  shown  salmonids,  mix employed  tissues,  source  and N a n r i ,  for  correct  protein  occurred. a protein  (Ogino  to  was  some i m p a i r m e n t of  test  the  of  higher  CS s u p p o r t e d  fish  However,  severe  study  hexane  and at  amino a c i d s  although  test not  casein  arginine  this  when  i m p r o v e d by  that  to  chick  except  149°C.  not  in  complete  The c a s e i n - g e l a t i n with  in  subjected  i m p r o v e d by  quality  was  extensively  However,  was  suggesting  damage  gelatin  Consequently FPE.  to  employed  CS i n  chinook  in  diets  In the  supplemented  protein  This  Similarly,  1975).  casein.  indicated  meals.  extraction  quality  at  i m p a i r e d the  lipids  the  fish.  deficient  in  the  lipid  2 hours  herring o i l ,  had o c c u r r e d .  to  meals  no d i f f e r e n c e s  performance  for  severely  protein  hexane.  Chick  heated  prolonged  addition  found  be a t t r i b u t e d  excessive. of  and u n e x t r a c t e d  to  investigators  who c o n c l u d e d  that  preferable  casein  fish,  and  various 1980;  to to  fish  species  McLaughlan, there  is  that  the  as  1979).  an u r g e n t diets  or  quantity  nutrition  quality  There the  First, mixture the  were  protein the of  can  and o t h e r energy  with  for  be  method  than  Samonds  three  (Walton  use.  have  of  has  been  and  bird  protein  been  that quality  quality  as  no c o n s e n s u s  The same p r o b l e m e x i s t s 1972,  attempted  1979) to  and v e r y  evaluate  on in  few  protein  method.  estimates  function amino  of  of  consider  determined  in  the  1982).  112  the  application  this is  study.  to  provide  proportions  a  for  proteins.  In  this  animals.  In  salmonids  also  This  in  right  other  are  in  protein  tissue  than  proteins  and Cowey,  to  dietary  acids  no d i f f e r e n t  species,  al.,  agreed  protein  has  et  1977;  been g e n e r a l l y  there  of  bioassay  and H e g s t e d ,  in  in  1983).  animal  which  and m a i n t e n a n c e  fish  human,  measures  to  al.,  evaluation  vary  growth  Pfeffer  by  adequate  main p o i n t s  quality  are  has  and S a r g e n t ,  one  et  quality  to  compared,  fish  essential  fish  respect  1959;  need  Watanabe  quality  for  requirement  1970,1972;  protein  were  potential  protein  al.,  protein  marine o r i g i n  genetic  1981;  ingredients  primary  synthesis  respect  et  Although i t  (Cowey  by more  the  of  dietary  of  with  (Allison,  investigations  of  for  most a p p r o p r i a t e  fish  the  (Cowey  frequently  applications  well  meet  The d e t e r m i n a t i o n  discussed  sources  and S t i c k n e y ,  The m e t h o d o l o g y  so  to  determine  Winfree  3.4.6  protein  major  contrasts  sources with  of  omnivorous  mammals where supplying  protein  energy.  The q u a l i t y  measurable  biological  satisfying  the  acids,  growth  for  the  amino  is  of  response,  of  protein  and b a l a n c e  acid  needs  for  for  significance as  dependent  and m a i n t e n a n c e , of  less  a protein,  is  required levels  unknown q u a n t i t y that  catabolism  as  well  energy.  determined  on the  by a  protein  of  essential  amino  as  supplying  an  Rat  maintenance  in  studies  differ  have  shown  both  a  quantitatively (Maynard that  and L o o s l i ,  protein  growth  quality  advantage  of  utilization intake  1969).  sources  and p r o v i d e  protein  other  and q u a l i t a t i v e l y  would  for  at  hand,  as  be  protein  metabolizable  intake  excess  (Phillips, estimates The quality  1969; is  assay  in the  of  as  desirable  to  fully  aspect  biologically a protein  that  fish  et  make  concerns  identified  standard.  classified  relative  of  support  determine have  the  protein  levels  of  protein  1979).  On the  effect  tolerable  on p r o t e i n  purpose  This  of  source  the of  was  FPE ( T a b l e  113  to  protein  study  of  on an  of  carbohydrate  by  salmonids  quality  in  protein  amino a c i d  in a diet.  relation  protein  27).  the  classify  essential  classification In t h i s  to  be  1982)  the  studies.  sole  this  to  by c a r b o h y d r a t e  to  expected  intake  McLaughlan,  the  growth  understood.  suitability the  protein  single  basis,  al.,  be  ability  picture at  replaced  believed  rapid  would  their  of  1977;  energy  Hilton  not  second  obtained  is  it  for  Methods w h i c h  levels  and H e g s t e d ,  of  in  a more c o m p l e t e  estimated in  differ  maintenance.  t h a n would  (Samonds  Therefore,  different  giving  from t h o s e  to  sources  Ultimately,  It a  balance is  well  were it  was  most  Table  27.  Summary sources  o f r e l a t i v e ( F P E = 100) e s t i m a t e s o f t h e n u t r i t i v e v a l u e employed i n t h i s study d e t e r m i n e d by d i f f e r e n t methods.  Protein source  GR  GFC  GEU  PER  NPR  FPE  100  100  100  100  100  FRH  95  108  99  101  LTH  86  88  86  HTH  26  32  CS  77  86  1.  PPV  of the test  NPU-1  NPU-2  100  100  100  100  100  102  94  96  99  101  85  84  89  85  89  91  77  76  26  26  50  16  44  52  53  47  77  82  88  79  86  90  101  95  NPU-1 a n d NPU-2 r e f e r s t o t h e m e t h o d s o f B e n d e r a n d M i l l e r respectively employed to c a l c u l a t e this parameter.  Slope (weight gain)  protein  (1953)  and O g i n o  Slope (protein gain)  e t a l . (1980)  desired in  to  identify  quality,  content  in  to  content (1972)  the  of  measure  the  of  feedstuff  proteins  as  the  chemical  determination  the  amount of  diet.  In t h i s  c o n f i r m the  states  should  as  indicates  included served  just  the  what  some f a c t o r x f  Therefore,  nutritionist protein  growth  is (f),  to  1.0 this  formulate  available  to  the  the  of  the  of  fish.  such  that  that  is  of  essential  and be  the  to  the  to  a  amino  fish  rather  acid Hegsted is  p r o t e i n content animal.  p r o p o r t i o n a l to  based  be  bioassay  For example,  scheme would e n a b l e diets  poor  protein  results  of  or  r e q u i r e d by human d i e t i c i a n s  = protein available  v a r y from 0 to  quality.  for  intermediate  ingredient  sense,  suitability  protein that  the  good,  on the  Such a the  the  the of  the  factor  true  fish  biological  t h a n on p r o t e i n  value  of  content  alone. Thus, several  the  biological  potential  applications  i)  The measurement  meet  specific  and S a r g e n t , ii)  of  the  of  protein  which are  as  effectiveness  performance requirements  of  in  salmonids  has  follows:  of  a protein  source  cultured  fish  (Cowey  that  used  in  1972).  The m o n i t o r i n g of  manufacture  evaluation  of  fish  processing  food  methods  ingredients  (i.e.  are  fishmeal)  and  the fish  f eed . iii)  The d e t e r m i n a t i o n  acids iv)  and p r o t e i n s  by o t h e r  the  (Zeitoun,  The c o m p a r i s o n of  sources  of  minimum r e q u i r e m e n t s  amino  1973).  results  investigators  for  obtained  with  when e x p r e s s e d  basis.  115  -  novel on a  protein relative  to  v) for  The p r e d i c t i o n  of  the  other  animals,  The  third  aspect  protein  quality  the  nutritive  that  should  parameters  affected  the  protein  intake  PPV,  NPU and s l o p e  usually based  upon body  present gain  study,  protein  Protein  gain al.  retention  whether of  slope  the  present  study  ranking  of  good  that  modifications  The mean v a l u e s  concentrations this  for  all  was  protein terms are  of  effort.  sources  in  -  for  protein  each  partly  values  obtained  the  overcome  since  were c o n s i d e r e d .  116  could  The  important  the  protein it of  relative at  at  all  In  be  the  gain  for  the and  levels. suggests  protocol.  dietary  the  was  values  different  experimental  in  weight  weight  and p r e c i s i o n  source.  the  0.0096.  PER, NPR, PPV, and NPU r e s p e c t i v e l y values  In  former  Clearly  tested  the  are  those  with  conditions  sensitivity to  and  percent  determining  found  necessary  gain  correlated.  the  rats  equation:  that  experimental  from the  p r o b l e m was  regression  in  and the  example, PER, NPR,  1979).  correlated  reported  worth  disagreement  the  taken  is  For  gain  the  are  same manner  following  work i n  Under the  shortcoming  were  extra  all  methods w i t h  = 0.9888)  also  that  study  on w e i g h t  highly  by the  is  (McLaughlan,  were h i g h l y  carcass  This  27)  (1976)  the  assay.  the  proteins  species.  same f a c t o r s . in  based  was  this  trial  retention  gain  in  = 0.6172 x dry w e i g h t  protein  content  those  latter  et  is  affects  coefficient  Zeitoun  point  the  into  from the  dietary  recognized  by the  Feeding  nitrogen  (correlation  predicted  and  assay.  subdivided  be  determined  and a r e  of  of  p a r t i c u l a r l y carnivorous  interrelated level  value  slope  (Table protein assays  analysis  of  Although there  is  linear  disagreement group  in  with  protein  was  noted  the  shortcomings  be  clear  to  respect quality  to  the  assays  be a d v a n t a g e o u s  that  proteins  of  a protein-free  (Samonds  in  in estimating  use  the  and H e g s t e d ,  present  protein  study.  quality  were m e a s u r a b l y  diet  it  different  1977)  was in  found  support  growth  and meet m a i n t e n a n c e  needs.  inference  is  in  a poor  quality  as  suffice  if  nutritive NPR, of  fed  at  value  50% of  NPU, and s l o p e  that  25% of  FPE as  performances conversion  quality the  Ultimately,  and d i e t a r y protein  the  efficiency. should  diet case  consider  the  quality  effectiveness the  of  above  to  state  be  recommended  the  of  then  diets.  of  HTH l i e s  diet  protein  routine  a  double  the  in  the  on  amount  The v a l u e  will  region  of  similar of  growth,  were o b t a i n e d  content  a protein a whole  raised  of  117  the  ocean  accepted,  feeding  the  feed  with  lies  final  be g i v e n  the  7,  in  with 9).  its  cost  feed food,  one  product. to  For  the  survival. then  quality  trials  for  diet  5,  growth and  should  protein  of  (Table  source  for  fish  promote are  which  HTH c o u l d  ( F P E ) based  terms  doubled  and y i e l d  to  in  The  HTH has  In f a c t  utilization  farmed  statements  for  27),  salmon  consideration  categorically  protein  to  ( F P E , FRH, LTH and CS) and a poor  of as  If  fish  approximately  the  fish  in  value  the  In  hatchery-reared  If  the  (Table  energy  usefulness  in  quality  chinook  sources  was  level.  by PER, and PPV.  (HTH) when  source  effectiveness  best  relative  juvenile  protein  latter  the  high  be needed  estimated in  quality  the  estimates  p r o t e i n would  be q u a d r u p l e d i f  good  a sufficiently  such  of  their  to  protein  it  In s p i t e  abilities  that  fed  it  assay  with  is  difficult  method  salmonids.  should When  the  relative  (Table  27),  biological rate,  scores  for  was  a general  assays  for  evaluating  gross  food  conversion  and M c L a u g h l a n  chicks  were c o r r e l a t e d  proteins.  NPR.  (casein  the  these  (1965)  = 100)  other  for  other,  protein  methods  was  gain  (Higgs  obtained chinook herein  are  for  et  (Table  et  fry. 15)  1979)  Morrison  the  protein,  duration  with  of  (Abernathy)  sex,  of  other of  test.  genetics, dietary  byproducts  variety relative  values  confirm  both  and NPR and NPU  (i.e.  between  weight these  on a dry f i s h  gain  two  differing (1960) the  (Higgs  et  of  et  al.,  118  -  with  reported  should  be  experimental  quantity  for  values  al.(1983)  results  stated  components,  modifications  weight  comparison with  investigators  The PER v a l u e s  and v a r i o u s  rapeseed  hand,  same a s s a y  and  by PER, NPU and  study  and P l o t n i k o f f  and C a m p b e l l  and o t h e r the  this  facilitate  because  age,  a  much l o w e r  C o m p a r i s o n s between the  conditions.  rats  28).  al.(1982)  cautiously  only  of  agreement  to  and t h o s e  in  similar  proteins  PER and PPV on one  (Table  and Coon  PER v a l u e s  PER y i e l d s  the  growth  energy  Johnston  numerically  The r e s u l t s  and  various  NPU and NPR f o r  quality  related),  interpreted  not  for  PER were c a l c u l a t e d  al.,  by H i g g s salmon  high  the  and g r o s s  that  obtained  essentially  excellent  The v a l u e s ba s i s  Since  are  those  proteins,  methods.  findings.  on the  from  with  quality  (1962),  found  were compared  between  protein  efficiency  (1979)  F o r poor q u a l i t y  than  and  Henry  proteins  agreement  Henry and T o o t h i l l  (1979)  values  various  there  utilization.  of  the  but  that  PER v a r i e d  and q u a l i t y also  with  of  the  a commercial dry  diet  this  diet  with  protein  1982)  were  determined  T a b l e 28.  C o r r e l a t i o n c o e f f i c i e n t s and l e v e l of s i g n i f i c a n c e between the d i f f e r e n t parameters used to e s t i m a t e the n u t r i t i v e value of d i e t s c o n t a i n i n g the v a r i o u s p r o t e i n sources.  GFC  0.9668 P - 0.000  GEU  0.9658 P = 0.000  0.9936 P = 0.000  PER  0.7172 P = 0.000  0.4635 P = 0.002  0.8096 P = 0.000  NPR  0.2360 P = 0.083  0.2790 P = 0.50  0.3126 P = 0.032  0.6765 P = 0.000  PPV  0.7391 P = 0.000  0.8056 P = 0.000  0.8308 P = 0.000  0.9765 P = 0.000  0.6519 P = 0.000  NPU-1  0.2525 P = 0.069  0.2863 P = 0.045  0.3271 P = 0.026  0.6319 P = 0.000  0.9620 P = 0.000  0.6669 P = 0.000  NPU-2  0.0213 P = 0.451  0.0396 P = 0.409  0.0727 P = 0.337  0.4037 P = 0.007  0.8896 P = 0.000  0.4374 P = 0.004  0.9266 P = 0.000  LYS  0.8503 P = 0.000  0.8410 P = 0.000  0.8379 P = 0.000  0.5349 P = 0.001  0.0497 P = 0.390  0.5776 P = 0.000  -0.1220 0.0955 P = 0.295 P = 0.246  GR  GFC  GEU  PER  NPR  PPV  NPU-1  NPU-2  under  similar  study  except  experimental that  a 69  conditions  day  trial  for  a commercial  diet  order  the  PER r e p o r t e d  by H i g g s  practical obtained  diets. with  diets.  It  source  such  of  FPE at  may be as  evaluation, group  However,  in  the  FPE i s  et  present  protein  employed  as  to  value  15)  the  same  is  of  for  various  a higher  to  if  a control  present The  study  that  the  employed.  level  comparison  performing closer  of  al.(1982)  therefore  a more m e a n i n g f u l  fish  (OMP) ( T a b l e  a similar  suggested  those  p e r i o d was  obtained as  to  all  of  PER was  the  a premium  in  above  protein  practical  diet  may be made w i t h  their  genetic  a  potential  for  growth. Higgs  et  evaluated  al.(1982)  practical  management Hatchery)  the  higher  laboratory.  This  stocks  temperatures may be  or  fed  values  only to  may be  Higgs  et  significantly PER w i t h present  probably  genetic  that  if  Robertson  al.(1983)  locality  determined due  to  as  chinook  West  the  a sole  salmon,  Vancouver  between  elevated  at  h e r r i n g meal  Creek  PER v a l u e s  the  and to  strategy  FPE were used  Creek  (Robertson  differences  constitution feeding  at  also  hatchery  They r e p o r t e d  premium q u a l i t y  the  water  hatchery. protein  were employed  in  o u t s t a n d i n g PER  obtained. al.(1983)  showed t h a t  influenced  chinook study  is  those  and d i f f e r e n t  postulated  source, diets  in  study.  than  et  i n c l u d i n g OMP under  in a different  present  considerably  chinook  diets,  conditions to  and P l o t n i k o f f  for  fry. diets  protein  dietary  protein  utilization  The p r o t e i n containing  120  levels  as  determined  employed  FPE c o v e r e d  content  in  a wider  by  the range  It  than  those  diets  of  Higgs  containing  consistent 1983),  with  that  and  carp f r y  grass of  similar  al,  pattern  difference which  to  dietary  and  other  In the  quality,  official  method  a readily  For  as  curves  is  may be  for due  previously  Over  diets  species  mentioned  1980)  a wider  The  fed  casein  noteworthy  of  dietary  protein  levels  this  protein  the  Differences  of  16% i n  between  study  differences, factors  a  containing  Maximum PER w i t h  plaice.  trout  and r a t s  level  dietary  al . ,  PER f o l l o w e d  fed  maximum PER i n to  et  rainbow  with  7).  is  (Koshiishi,  1972)  the  obtained.  level  of  the  are  and C a n a d a ,  Association  a control  for  casein  comparisons when  protein.  pure  the  protein. are  PER i s  (27%)  although  affecting  of  rats.  the  121  measure  is  customary  used  because relative  i n an a t t e m p t  casein  and i s  among was  an  Analytical  PER e s t i m a t e s  reported  for  It  Casein is  to  purposes,  Official  i n PER measurements PER v a l u e  used  regulatory  performed with  obtainable  example,  (Fig.  p a r t i c u l a r l y for  Tests  determined  facilitate  1965)  al . ,  F i g . 7)  (Higgs  1979).  plaice  i n PER i n  of PER.  protein  casein  et  and 40% i n  United States  Chemists.  juvenile  relationship in  15,  salmon  chum salmon  o c c u r r e d at  protein  determination  1970),  (Cowey  these  chinook,  by the  chinook  observed  and C h a n g ,  studies  confounded  that  the  that  proteins  the  feed  levels  maximum PER was  27% i n  (Table  ( D a b r o w s k i and K o s a k ,  between  respective rats,  with  juvenile  cod m u s c l e  (Hegsted  The downward t r e n d  27% p r o t e i n  found  1978),  protein  freeze-dried  at  (1983).  c a r p . ( O g i n o and S a i t o , et  diets  al.  more t h a n  (Takeuchi  range  et  set  to it  is to  to  investigators. at  100,  for  each  assay,  beef  88,  1979).  soy  relative  78,  mind  the  that  casein  in  the  PER v a l u e s  obtained  proteins)  those  for  for  proteins  those  of  lower  digestibility  animal or  fish  fed  protein  bacteria  of  plant  while  algal  studies  deposition  to  and S a i t o ,  1970;  Higgs  the  value (the  former  gain  to  were  highly  do not in  measure  et  fish the  1979;  discussion Since  relates  protein  were  of  petroyeast inclusion  17).  report  the  al . ,  Pfeffer, this  is  body  intake)  correlated  protein the  termed  (Table  28).  122  al. ,  than  have  relative  contains  1983).  chinook  of  to  one  Single  protein  obtained  of  Higuera  Clarke  and the  salmon  et  diets  protein  a feedstuff  essentially  gain  results  source  De l a  1982;  PPV and PER a r e  lower  balance  to  The PER  proteins  efficiency  value  1973;  meals,  and m e t h a n o p h i l i c  into  (Table  relative  (fish  study.  plant  et  bearing  a r g i n i n e and  consistently  (Higgs  be  a s i m i l a r order  this  protein  nutritive  Z e i t o u n et  al.,  present (PPV).  as  promising for  proteins  Several  1977;  such  can  Generally,  and a p o o r e r amino a c i d plant  (McLaughlan,  study  animal p r o t e i n s  o r i g i n are  each  6  gelatin,  17).  p r o b a b l y because  Also,  defatted  investigators;  F P E , F R H , and LTH i n  sources  look  this  with  (Table  more a n t i - n u t r i t i o n a l f a c t o r s  cell  In  study  109,  gluten  in  other  investigators  animal o r i g i n ,  proteins.  of  supplemented  with  meal was  obtained  those  present  by o t h e r  determined  values  was  fish  23 and wheat  values  compared w i t h  methionine  egg  PER f o r  pea p r o t e i n  Similarly,  cautiously in  the  et  al.,  (Ogino al . , 1982).  productive the  latter by the  same  assay  body two  weight  methods  Under Higgs  et  similar al.(1982)  commercial with  experimental  diet  reported  stock  of  al.(1982) and P l o t n i k o f f of  approximately  practical reported  dry d i e t s . for  the  present  al.,  1973;  trout that  Higgs  (Zeitoun chinook  t h a n coho former  fish  fed  salmon  in  et  al. ,  al . ,  1973;  those  for  Clarke  Pfeffer,  salmon  species  for  body  protein  a value  coho  f o r PPV  and may  p r o t e i n more  synthesis than  and  et  rainbow suggest  effectively  and t h e r e f o r e the  in  (Zeitoun  1982) This  et  been  salmon  al.,  and  test  PPV have  1982).  may be more p r e f e r a b l e  another  investigations  et  dietary  for  Creek) Higgs  OMP and v a r i o u s for  study,  temperatures  reported  above  salmon can u t i l i z e  present  PPV (32%)  water  the  1979;  the  (Robertson  al.(1983)  compared to  et  fry  Higher estimates  chinook study  At h i g h e r  chinook et  40% w i t h  to  an e q u i v a l e n t  (Abernathy).  a different  conditions  latter  the  for  fish  farming. In  fish  measuring Bender close  nutrition  NPU ( T a b l e  and M i l l e r relative  procedures. the  latter  17).  (1953)  corrects  for  avoids  overestimating  achieving et  protein  and Ogino et values  Theoretically  a higher  al.(1980) intake  that  al.  been  (1980)  were o b t a i n e d  more a c c u r a t e  quality  of  protein  exactly  met  obtained  maintenance  9).  123  the very  both-  since  sources  With  for  is  that  on a body  is  100% was  by  two methods  requirements  mean body w e i g h t .  by  methods,  this  a NPU of  adopted  calculated  between the  maintenance  the  have  A l t h o u g h NPU was  The main d i f f e r e n c e  basis.  Ogino  two methods  NPU ( F P E = 100)  weight  fish  studies  the  with  it  measured method  a  of  dietary  requirements  (Fig.  in  In  this  dietary et  study  protein.  al.(1979)  with  diets  17).  The l e v e l  the at  contained  for  as  has  evaluation one  level  quality  of  fish  it  of  quality  carp  fed  of  et  (Cowey  al.,  The this  study similar  al.,  1978;  1983).  may not  one  level  1978)  Ogino,  likely  due  can o n l y as  to  be m e a n i n g f u l they  were  for  dietary  at  intake  arbitrary  rats.  However,  also  1970), fed  since  estimate protein  previously,  penalized  been  trout  an  true  (McLaughlan,  has  (Table  protein  one  protein  both  1979).  A  reported  fed  for  casein  freeze-dried  reported  to  those  for  in  by a s i m i l a r  values  between  differences  results if  in  regimen  PER ( T a b l e  1980;  cod  muscle  the  chinook rainbow  in trout  (Takeuchi  Watanabe et studies  species,  by o t h e r  However,  et al.,  (Table  strain,  17)  fish  ingredients  NPU ( c a s e i n  17).  124  with  method  and o t h e r  obtained  relative  juvenile  obtained  Ogino and N a n r i ,  feeding  C o m p a r i s o n of  evaluating  As m e n t i o n e d  proteins  protein  1980;  respectively  dose d e p e n d e n t ,  1977).  because  9).  comparable of  for  Atack  obtained  and h e r r i n g meal  p r o p o r t i o n a l to  and p l a i c e  NPU v a l u e s  types  by  is  and S a i t o ,  1972)(Fig.  are  problem encountered  employing  be  of  quality  (Ogino  of  of  standardized  quality  and H e g s t e d ,  temperature,  diets.  used  protein  The d i f f e r e n c e s  most  size,  methods  al.,  range  for  are  for  NPU on p r o t e i n  (Watanabe et  protein  been  intake  casein  levels  bacteria  been  and low  several  35% and 25% p r o t e i n ,  dietary  at  at  c o m p a r i n g NPU v a l u e s  not  (Samonds  dependence  the  has  of  standardization high  avoided  methanophilic  their  sources  This  determined  who had d i f f i c u l t y  carp for  level  NPU was  in  the  investigators = 100)  values  are  since  casein  was  not  used  as  meaningful  the  at  few  method  se  evaluate  discussed balance, (Iwata, the  (Hegsted  level  of  protein  for  between  intake  assumed  (De Long  1971).  The q u e s t i o n  group  of  with  et  excessive which  of  caused  catabolism  availability intakes,  of  (see  the  not  (weight  1958;  that  the  in  use  fed  lysine  this  on the  other  of  were  nitrogen  Rychly,  1980).  Up to  slope  rat  some  this  to  often  diet  fed  the is at  wheat  due  to  very  gluten  1979).  upon  The  At low  resulting  in  lysine the  Threonine  upward c u r v a t u r e  have  no  rate  the  amino a c i d .  to  Gerking,  proteins.  1977).  show  was  has  slopes  such as  retarded,  amino a c i d a p p e a r s  1965;  assays  the  dependent  hand,  gain)  assay  Objections  of  relationship  a protein-free  with  125  on  per  intake  (Bodwell,  is  adopted  protein  group i n  is  slope  against  and C h a n g ,  proteins  in rats  protein  studies  or p r o t e i n  the  1977).  amino a c i d of  the  been  a linear  gain  Hegsted  linearity to  4)  a  subjective.  minimum r e q u i r e m e n t  Chap.  and r e - u t i l i z a t i o n of  as  1971;  study,  measure  Several  regressed  encountered  lysine  proteins,  low i n t a k e s  has  fish.  to  intake,  downward c u r v a t u r e of  is  highly  p r o b l e m (Yanez and M c L a u g h l a n ,  catabolism  conservation deficient  of  1965a)  in  lysine-deficient  this  protein  where  a zero-protein  Specifically, have  of  is  present  attempted  Nose,  (McLaughlan,  reports  intakes,  met  respect  of  low  intake  al.,  employment  of  the  reporting observations  and r e s p o n s e  animals  of  in  1971;  growth was  raised  level  in  studies  have  g a i n was  Gerking,  protein  been  other  quality  where,  protein  1970;  protein  and C h a n g ,  protein  earlier body  of  investigators  more t h a n one  ratio to  source  comparison with  Because quality  sole  at  conservation  mechanism  (McLaughlan  and M c L a u g h l a n includes  the  (1979)  than  former  situation  the  led  dose  data  the  to  same time  number of  three,  the  have  been  each  diet.  zero  protein  meaningful able  to  diet  of  In the  When the  both  comparisons  diet  for  groups were  between  the  was  study,  groups  found. test  126  the  because  the  lines  intercepts when  for  the At  the  high.  precision  reduced  of  the  used.  from  six  protein  fed  including  (Table  By t h i s  were  could  had been  gain  to  groups  and s e n s i t i v i t y  protein  When  by  duplicate  fish  slope  rats  increased  of  which  suitable  included.  number of  slopes  body  was  were a n a l y z e d  the  is  assay  together  were  precision  additional  a more  study  closer  on the  the  al.(1968)  ratio  This  groups  that  present  slopes  fed  distinguish  fed  each  error  improved i f  this  brought  found  slope  et  p a r a l l e l i s m between  coefficients  fed  and u n d o u b t e d l y  the  group.  In  diet  Hegsted  group was  this  primarily  animals  30%.  fed  reduce  were  al.(1968)  standard  approximately employed  lines  depended  that  values.  correlation  et  assay  to  protein-free  the  Hegsted  the  erroneous  the  diet  excluding  tended  response  for  ratio  one  1977).  concluded  protein-free  method  which  and K e i t h ,  method  the  24), the  sources.  assay  was  3.5  in  Summary of In  this  experiment  the  the  diet  of  salmon  growth,  food  chinook  conversion  investigated. measured  growth  conditions  and the  parameters. achieved  were 42  Rapid  of  by f r e e z e - d r y i n g  slightly  inferior  had a s e v e r e  fed  fish  growth  (75°C)  fry,  effect  adequate  in  as  measured  protein  High  on the  the  based  terms  of  protein  of  poor  by  on  various was  Cooking followed resulted  temperature  protein  (150"C)  quality diet,  quality,  drying  herring  although gave  results  because  between  fed  test  diets  and a c o m m e r c i a l d i e t (OMP)  protein  level  of  revealed  that  the  the  palability.  of  the  latter  by  in  inconsistent fish  of  to  that  utilization  h e r r i n g meal  In a d d i t i o n a c a s e i n - g e l a t i n  apparently  diets  showed  raw m a t e r i a l .  d r y i n g of  terms  had a p r o n o u n c e d e f f e c t  and b e s t the  in  source  was  experimental  protein  performance.  adverse  utilization  The r e s u l t s  meal  of  between p r o t e i n  and p e r f o r m a n c e  the  days.  utilization  temperature  meal.  for  relationship  and p r o t e i n  The f i s h  satiation  processing  low  Experiment 1  Comparisons  could  probably  be  lowered. An a t t e m p t techniques generally measured respect  for  at  more  slope  to  ratios  biologically  rather  compare protein  various  one  level  may have  the  of  when p r o t e i n dietary  than w e i g h t  Also, gain  The m a i n t e n a n c e  127  best  were  quality  protein.  the  was  assay  The r e s u l t s  p r o v i d e d the  proteins.  more v a l i d .  biological  quality.  be more r e l i a b l e  than  between  gain  made to  evaluating  found  comparison protein  was  In  means  to  requirements  this  of  measurement  considered  was  direct of  be for  protein  were e s t i m a t e d  protein  diet.  This  methods w h i c h the  This  for  was  the  the  a protein-free  estimation  endogenous  compared as  level  of  particular  the  in  of  loss  and a  protein of  assessed  adequately  level  intake  into  amount of  of  for  the  low  quality  nitrogen  of  by  from  various  intake  be  interpreted.  nutritive  value  of  the  their  conditions  by b i o a s s a y , predicted  available  of the  by the  the  protein  increased.  chemical  not  determination  a  Although  protein  have  Thus  was  content.  affected  could  with  proteins  lysine  meals  effect  test  for  excretion.  maxmimum e f f i c i e n c y  could  of  utilized  exogenous  how the  protein  amounts  source  terms  processing  and the  depiction  the  protein  protein  Lastly, evaluated  partitioned  and g r o w t h ,  provided a clear  sources  as  correct  intake  maintenance  the  enabled  fish  body. Protein  the  by f e e d i n g  quality  been of  available  lysine. In c o n c l u s i o n , preferable  to  experimental method  to  certain  others, protocol  measure  the  procedures  and s u b s t a n t i a l may p r o v i d e nutritive  128  judged  improvements  a more  value  f ish .  were  of  sensitive proteins  to  be  in and  precise  in diets  for  Chapter  4  EXPERIMENT 2 4.0  Protein  requirements  relation  4 .1  to  dietary  juvenile  energy  chinook  salmon  in  content  Introduction The  previous  pollock  muscle  experiment  with  a more p a l a t a b l e combination  of  The g r o s s  Long for  et  protein  have  fed  therefore  1979),  at  source lipid  of  of  light  in  the  juvenile  depress  to  This ratio  of  129  the  using  two  of  -  by De Long acids  of  (Yu and  salmonids It  protein FPE as  levels  protein  salmon  high  1% .  approach a l s o  salmon. -  excess  to  (De  protein  information rate  salmon  containing  optimal  chinook  in  diets  fatty  growth  investigate  chinook  diets  with  omega-6 t y p e  were  chinook  employed  more r e c e n t  concentrations  by m a r i n e o i l . of  of  known to  juvenile  protein  supplied  for  in  of  diets.  salmon  based  juvenile  diet  arginine  salmon  chinook  determined  basal  than a  with  casein-gelatin of  (FPE) provided  chinook  juvenile  freeze-dried  source  with a demonstrated  content  seemed w o r t h w h i l e  determination diets  is  dietary  requirements  of  M o r e o v e r , the  which,  euphausids  juvenile  been  products  of  supplemented  using  however  had a h i g h  corn o i l  Sinnhuber, when  not,  a blend  protein  The r e q u i r e m e n t s  fishery  al.,(1958)  in  requirements  value.  whole  and g e l a t i n ,  inclusion  1958).  from  biological  from  for  that  quality  (NRC 1 9 7 3 , 1 9 8 1 )  al . ,  derived  and h i g h e r  protein  established  showed  freeze-dried  casein  and m e t h i o n i n e ,  et  of  of  the  sole  dietary  enabled energy  in  In r e g a r d to one of  of  the  both  few  experiments  estimation  energy:total  Similar  Pacific  salmon  a i m , Lee and Putnam (1973) on s a l m o n i d s were  (PE:TE)  in  have  Their  ratio  practical not,  conducted  i n which d i e t a r y  varied.  an a c c e p t a b l e  estimates  of  methodology protein  diets  however,  levels  for  been  rainbow  obtained  for  diets.  M a t e r i a l s and  was  of  energy  trout.  4.2.1  latter  p r o t e i n and energy  provided  4.2  the  methods  Protocol  Whereas E x p e r i m e n t 1 was  conducted  continued  This  for  105  freshwater  residency  same  stock  of  used  i n E x p e r i m e n t 1.  time  and the  days.  for  43 d a y s ,  Experiment 2  period approximates  the  p e r i o d of  juvenile  chinook salmon.  c h i n o o k f r y were  employed  in Experiment 2 that  The f i s h  same manner as  were  d i s t r i b u t e d at  described  in  the  the  The was  same  previous  experiment . The e x p e r i m e n t factorial of  dietary  factors  each  replication  Each  assigned  row of  4.2.2  designed  p r o t e i n and two  tested.  randomly  dry  without  was  tanks  of  i n each  as  (Zar,  levels  the  the  constituted  1974).  of  eight  of  a 4 x 2 randomized block Four  dietary  dietary  concentrations  energy  were  treatments  two r o w s .  the  was  As i n E x p e r i m e n t 1,  a block.  Diets  The d i e t s  were  matter  c r u d e p r o t e i n from  as  f o r m u l a t e d to  -  contain FPE a t  130  -  17%, each  27%, of  two  37% or 47% of levels  of  available  dietary  energy,  level  29).  At each  kept  constant  at  form  of  proportions  equal  substituted  for  of  3150  in  the  of  the  is  shown  metabolizable  4.2.3  Data  The  previous  for  randomized  factorial  similar  that  random e f f e c t .  OMP was  included  in  described  criterion  energy  with of  the  basis  The p r o x i m a t e  concentration  the  analysis  calculated  the  diets.  consumption,  analysis  were  treated  included  the  tables of  was  also  on which  A mixed  for  in  polynominal  regression  growth  data  be  were  the  of  as  described  and i n t e r p r e t e d  were  and  and row a Duncan's  Data on  body w e i g h t s  the  fish  but  are  day  as  i n Experiment  1.  principal were  derived  according  131  protein  comparison.  requirements  equations  employed  to  analysis  g  as  was  subjected  log  selected  in  two-way  effects  (P = 0 . 0 5 ) .  for  a  described  model  fixed  then  as  feed  Dietary  statistical  conducted  protein  effects  to  and f i g u r e s  was  to  E x p e r i m e n t 1.  means  covariance  rate  were s u b j e c t e d  (1955)(DMR)  not  growth  rate  metabolizable  together  were assumed  Range T e s t  covariate  Specific  were  was  feed  ANOVA.  Treatment  An a n a l y s i s the  and g l u c o s e ,  weight,  The p a r a m e t e r s  concentration  New M u l t i p l e  body  carcass  1.  fed  energy  levels  Carbohydrate, in  experiment.  i n T a b l e 30,  fish  Experiment  energy  dextrin  lipid  (Table  analysis  and f i s h  to  of  on an e q u a l  and g r o s s  data  analysis  energy,  13% r e s p e c t i v e l y .  protein  described diets  metabolizable  6% or  as  and 3950 k c a l / k g d r y d i e t  to  based. (Zar,  with  response Linear 1974)  Cowey et  al.  and  from  Table 29.  Composition of diets containing various levels of protein and energy (Experiment 2).  % protein i n diet Metabolizable energy (kcal/kg)  Freeze-dried pollock and euphausid Herring o i l  27  17  47  37  3150  3950  3150  3950  3150  3950  315  3950  192.20  192.20  305.26  305.26  418.32  418.32  531.37  531.37  54.1  173.27  50.1  119.32  46.2  115.36  42.2  111.40  Dextrin  225.0  250.5  169.0  194.5  112.5  138.0  56.5  82.0  Glucose  225.0  250.5  169.0  194.5  112.5  138.0  56.5  82.0  Ground cellulose  224.5  85.5  235.0  127.52  248.15  142.15  259.95  116.73  Carboxy methyl cellulose  20.0  20.0  20.0  20.0  20.0  20.0  20.0  20.0  Mineral mix  71.1  71.1  62.7  62.7  54.3  54.3  45.7  45.7  Vitamin mix  7.3  7.3  7.3  7.3  7.3  7.3  7.3  7.3  Choline Chloride (50%)  3.5  3.5  3.5  3.5  3.5  3.5  3.5  3.5  1,2,3.  See Table (3).  T a b l e 30.  Proximate  composition and c a l c u l a t e d energy  Z protein i n diet M e t a b o l i z a b l e energy (kcal/kg)  17  Crude p r o t e i n (N x 6.25) T o t a l crude  lipid  Ash Digestible carbohydrate  Energy  27  37  47  3950  3150  3950  3150  3950  3150  3950  OMP  18.05  16.94  27.36  28.24  36.68  35.96  45.81  46.15  49.40  6.09  12.40  5.70  12.31  6.32  11.79  6.49  12.95  14.53  7.47  7.15  7.19  7.37  7.34  7.61  7.39  8.30  11.92  8.31  50.1  7.89  33.8  6.95  38.9  7.72  22.5  8.37  27.6  6.72  11.3  5.80  16.4  6.03  16.60  24.62  (kcal/kg)  Protein ME (4.5 k c a l / g )  812  762  1231  1126  1651  1618  2062  2077  2075  GE (5.7 k c a l / g )  1029  966  1560  1553  2091  2050  2611  2631  2816  Lipid ME & GE (9.5kcal/g)  579  1179  542  1169  600  1120  617  1230  1280  Carbohydrate ME & GE (4.0kcal/g)  1800  2004  1352  1556  800  1104  452  655  266  T o t a l ME  3191  3945  3125  3951  3151  3842  3130  3963  3721  T o t a l GE  3408  4145  3453  4278  3591  4274  3680  4517  4462  mg p r o t e i n / k c a l ME P r o t e i n energy (ME): t o t a l energy (ME) 1.-6.  2 .  3150  45.0  Moisture  contents of the d i e t s employed i n Experiment  See T a b l e ( 4 ) .  56.6 0.25  42.9 0.19  87.6 0.38  68.9 0.28  116.4 0.52  93.6 0.42  146.4 0.66  116.5 0.52  132.8 0.56  (1972)  and Z e i t o u n et  requirements  as  al.  discussed  were p e r f o r m e d by computer analysis  of  (GLM)(SAS, growth  variance 1982)  (1973,1976) in  the  text.  to  estimate  Statistical  through a general  program ( G E N L I N ) .  p r o c e d u r e was  used  rate.  134  to  protein  least  A general fit  analyses squares  linear  a regression  model model  4.3  Results  A.3.1  E s t i m a t i o n of salmon  from  the  protein  growth  data  The mean body w e i g h t s diets  and OMP i n c r e a s e d  period the  (Fig.  12).  different  apparent  in  that  0.001)  body w e i g h t  concentration weights 0.05)  were  with  significant were the  still  evident.  differences  containing  first the  requirement  half  of  their  in  contrast  weights  of  This  salmon  period in  fish  range  is  105, level  day  105  diets  longer  that  higher  water  P =  protein  receiving  suggests  fresh  test, At day  were no  fry  body  day A 2 , a t  between g r o u p s  31).  (P <  protein  level.  to  were  variance  dietary  respectively  chinook  growing  to  to  differences  new m u l t i p l e protein  due  and energy of  dietary  due  experimental  the during  than  the  during  second.  (day)  as  (P < 0 . 0 0 1 ) protein fed  for  with  (P < 0 . 0 0 1 )  (P > 0 . 0 5 ) ( T a b l e  A three-way time  final  protein  chinook  experimental  body w e i g h t  significant  in dietary  37% and 47% p r o t e i n  significant protein  were  (Duncan's  However,  in  the  The mean d i f f e r e n c e s  increment  differences  various  fish of  juvenile  An a n a l y s i s  associated 31).  of  during  i n wet  A2 t h e r e  significant  each  the  exponentially  experiment.  by day  (Table  fed  concentrations  the  indicated in  fish  Differences  dietary  early  of  requirement  in  analysis  the  covariate  slope  containing  covariance indicated  (specific  concentration.  diets  of  growth  However,  27% p r o t e i n  the were  135  of  log  wet  e  weights  a significant rate) slopes  due  to  difference dietary  obtained  similar  to  with  with  those  fish  with  fish  fed The  weight level fish et  gain for  was  been  et  Satia,  in  to  protein  the  the  specific  growth  highest  observed errors  at  considered  determined  from  1974;  al.,  for  fish  to  the  rate  to  means of  al.,  mean w i t h i n rate  was  intersection  point  two  lines  protein  these  level,  as  a percent  maximum growth  in  juvenile  method,  the  found  to  level  tested  Other fish  et  estimate  (Fig.  al.,  1979)  to  a second  et  (1972).  applied  of  line of  than  13).  the  two  as  this the  line  The  the minimum support  Employing in  of  a horizontal  taken  growth  et  By a v e r a g i n g  r e q u i r e d to  regardless  studying  1972; have  requirements.  fitted al.  al.,  for  was  line  by more  salmon.  Long  Ogata  procedure  response  drawn ( F i g . was  (De  the  study  above was  dietary  energy  13).  investigators et  chinook  35%,  method  level.  dry d i e t ,  requirement  be a p p r o x i m a t e l y  (Cowey  Murai  protein  of  1974;  treatments,  maximum growth  plots  for  a regression  differ  establishing  of  the  protein  which d i d not  the  type  1973,  fish,  of  maximum  requirements  F o l l o w i n g the  portion  dietary  attain  A similar  growing  31).  minimum r e q u i r e m e n t  "broken l i n e "  study.  ascending  be  protein  1981).  the  (Table  which  Z e i t o u n et  present  al.,(1973)  fitted  37 and 47% p r o t e i n  The q u a n t i t a t i v e  A n d e r s o n et  data  standard  of  commonly  growth.  1983;  Zeitoun  level  is  al.,1958;  the  containing  lowest  have  al . , to  diets  the  Z e i t o u n et employed  The f i t t e d  al.,  the  growth  p o l y n o m i n a l by the equation  136  for  requirements  1976;  non-linear  Therefore,  degree  nutrient  fish  Dabrowski,  growth rate  curves data  procedure fed  diets  of 1977; to  were of  Cowey  DIET Z protein - kcol/lOOg  47-315 47 -395 37-315 37-395 27-315 OMP 27-395  17-315 17-395  105  DAY  Fig. the  12. fry-  levels  of  Weight to  gain  (+  smolt-stage  protein  2 SE) of fed  diets  and e n e r g y ,  137  -  chinook  salmon  containing  and f e d OMP.  from  various  Table 31.  Wet f i s h body w e i g h t (mean + SE) a t day 43 and d a y 1 0 5 , and s p e c i f i c g r o w t h r a t e s o f d i e t s c o n t a i n i n g d i f f e r e n t l e v e l s o f p r o t e i n and e n e r g y a n d OMP.  Metabolizable  energy  content of d i e t (kcal/kg)  Percent P r o t e i n i n Diet 17  27  37  47  Mean  2.71  3.77  4.10  4.57  3.79  Mean body w e i g h t (+ S E ) Day  42  3150  + 0.068 3950  + 0.034  2.33  3.45 P  Mean  2.52 + 0.048  3.6i  q  3.94  4.48  4.02'  4.52*  OMP  3.55  3.59 + 0.025 Day  105  3150  5.66 + 0.23  3950  4.23  10.47  13.25  13.49  10.72 + 0.12  9.55  12.13  13.45  9.84  1 Mean  4.94 + 0.16  P  10.0i  q  12.69  r  13.47'  OMP + S p e c i f i c Growth 3150  3950  10.06 0.46  R a t e (Z wet w t / d a y ) 1.17 + 0.024 0.89  1.77  2.00  1.99  1.68  1.89  2.00  1.73'  1.9s'  2.00'  1.73 + 0.012 1.61  2 Mean  P  1.03 + 0.017  OMP  1.74 + 0.06  1. V a l u e s w i t h t h e same s u p e r s c r i p t f o r e a c h p a r a m e t e r w i t h d i f f e r s i g n i f i c a n t l y (DMR t e a t P - 0 . 0 5 ) . 2. S l o p e s w i t h t h e same s u p e r s c r i p t w i t h r e s p e c t s i g n i f i c a n t l y ( S c h e f f ^ ' s t e s t P - 0.05).  -  respect  to p r o t e i n l e v e l  138  -  to protein level  ( p - r ) do n o t  ( p - r ) e f f e c t s do n o t d i f f e r  Fig.  13.  weight fed  Specific  per  diets  energy. the  day)  growth  (+_ 2 SE) of  containing  at  straight  parallel  averaging  the  from  other  each  means.  line  p o r t i o n of  concentration line  both  highest  (percent  juvenile  different  A regression  ascending  rate  was  the  levels to  obtained  for  salmon  p r o t e i n and  fitted  to  the  response  dietary  abscissa means  by more t h a n two  The v a l u e  chinook of  of  observed  body  levels  growth  the  wet  comparison.  139  to  was  over  protein  energy.  A  drawn by  which d i d not  standard  OMP i s  data  errors  included  of  for  differ the  O O Z  DIET ME O 3150 kcal/kg A 3950 "  0-50  u LU  a. REQUIREMENT  0  10 20 30 40 PERCENT PROTEIN IN DIET  140  50  containing  3150  kcal/kg  was: 2  y = - 0 . 8 1 2 2 5 + 0.14212x (n = 4, and  r = 0.9991)  3950 k c a l / k g  was:  y = - 1 . 0 7 9 4 3 + 0.14671x (n = 4, where  growth  x = percent  P < 0.05  fit  protein kcals/kg al .  of  these  salmon  indicated  that  diets  not  was  tissue  values  for  the  diet  described  equations diets  of  the  variance  Fig.  (Table  30) .  and as  it  of  contributes,  to the  3150  curves  o c c u r r e d at 3150  c r i t e r i a of  protein  of  and 3950 k c a l / k g  containing  the  a  and 3950  Cowey  requirements  important 1.  a fish  et of  relative  to  in  weights  growth  of  was  Since  the  (1979)  total  141  of  of  the  chinook  p r o v i d e d by p r o t e i n  and  protein  both  source,  nitrogen  terms  juvenile  is it  utilized is  and n o n - p r o t e i n  The o b s e r v a t i o n s  diet  body  concentration  energy  in  on f i s h  energy  an energy  c a s e made by Cowey and S a r g e n t requirement  diets  energy  between p r o t e i n  in  response  conducted  affecting  Dietary  synthesis  is  be  metabolizable  31).  that  probability levels  containing  the  would  with  maximum g r o w t h .  sources  relationship  diet.  According  a factor  (Table  non-protein for  for  the  40% and 42% f o r  The a n a l y s e s  salmon  above  respectively.  ( 1972)  chinook  the  in  The maximum of  level  0.00174x ,  rate,  protein  and P < 0.10  respectively.  2  -  r = 0.9956)  y = specific  The d a t a  0.00177x ,  balance  made above for the  dietary  stating  the energy scheme  support the  the  protein  p r o p o r t i o n of energy.  in  When  energy the  Fig.  14.  weight fed  per day)  diets  energy  growth  by a s i m i l a r  (percent juvenile  different  (PE:TE).  The v a l u e  rate  (+_ 2 SE) of  containing  ratios  obtained (13).  Specific  method  obtained  for  to  that  OMP i s  comparison.  142  body  chinook  protein  The p r o t e i n  wet  salmon  energy:total  requirement described included  was in F  for  CO  200  y=i 97  CN +1  4 — i  < OMP"  1-50  < £  100  O O U  DIET ME  LU  a.  CO  0-50  O 3150 kcal/kg A 3950 » REQUIREMENT  010  0-20  0-30 0-40 PE:TE  143  0-50  0-60  0-70  protein  requirement  was  dietary  energy,-the  minimum P E : T E  salmon was The  found  protein  quantified quadratic  to  equation  both  0.05).  Cowey et  curve. achieved  at  the  salmon  (Table  consider  gain  the  within  the  present  of  the  means  chinook  method  PE:TE  was  (Fig.  14).  also  polynomial equation.  2  The  ,  study,  (x  the  which  Fig.  on  fish  15).  al.  the  was  This  range  adapt  growth  value  to  of  dose  confidence  it  of  approach  a response  drawn  of  to  that  response.  limits  was  in  nor does  maximum  line  to  a range  provide the  pointed  differences  a statistical  and a s t r a i g h t  144  (1976)  maximum p o i n t ,  could  upper and lower  plotted,  point  the  insignificant  They a p p l i e d  confidence  of  relating  a n i m a l to  level  (P <  protein  highest  max.,  curve  and beyond the  significant  dietary  Z e i t o u n et  practically  of  the  (P <  i n OMP, a p o p u l a r c o m m e r c i a l  quadratic  levels."  were  0.55  However,  below  being  maximum growth  30).  the  adequately  p r o d u c e d the  study, of  data  terms  defined  ratio  a certain  the  of  8.37872x  the  PE:TE  nutrient  In  as  order  fit  that  ratio  ability  protein  determine lay  the  reflect  percentage  growth  rate,  (1972)  x max on the  "does not  total  was:  15)  level  a PE:TE  feed  of  broken l i n e  expressed  and q u a d r a t i c  present  to  dietary  (Fig.  al.  corresponds  that  growth  linear  In the  for  that  ratio •  0.005),  as  to  r = 0.9647),  obtained  requirement  ratio  0 . 5 0 6 9 9 + 9.25398x -  x = PE:TE  out  derived  y = specific  curve  relative  by the  requirement  (n = 8,  The  be 0.41  by a p p l y i n g a second  y = -  where  estimated  95%  Fig.  15.  curved  The second  line)  confidence chinook Dashed  fitted  limits  salmon lines  relationship.  for  maximum g r o w t h . PE:TE  relatively (y  max).  PE:TE  diets  with  the  would  of  juvenile  the  result  OMP i s  in  large.  included  145  the The for  of  ratio range  of  in  from maximum growth reduction  ratios.  limits  PE:TE  represent  predictably  relatively  fed  the  (+_ 95%  PE:TE  95% c o n f i d e n c e  xO and x l ,  would be fish  different  (solid  rate  means)  x max r e p r e s e n t s  The c o n c o m i t a n t  with  growth  estimated  minor d e c r e a s e s  ratio  obtained  that  polynomial curve  specific  the  represent  the  dietary  to  of  fed  order  rate  dietary value comparison.  010  0-20  -  0-30 0-40 PE :TE  146  -  0-50  0-60 0-70  parallel of  the  to  lower  polynomial at  the  xO.  abssissa  confidence  curve  at  Statistically,  expected include  with the  a diet  of  0.41  (xl)  ratio  of  0.35  (xO).  (1982)  the  PE:TE  investigators differently the  PE:TE  requirement  4.3.2  et the the  0.55  (x  with  the  criterion  1979)  salmon  would  a  PE:TE  0.55.  interpret that  al.  Other  the  data  corresponds  (xl,  method  found  to  be 0.41  Other i n t e r p r e t a t i o n s  of  the  was  a PE:TE  Cowey et  was  ratio  limit  max)  response  of  the  response  maximum r e s p o n s e  conversion  and g r o s s  food  the  dietary  conversion  protein  (Table  32).  protein energy  Fig.  15). in  curve  The the  will  be  level  and the  Differences  of  levels  concentration the  diet  was  dietary  food  (GFC) was  energy  level  i n GFC between  containing  low  and e n e r g y  (P > 0 . 0 5 ) protein.  conversion  increased.  147  level  on  food  utilization.  efficiency  significant  of  the  limit  chinook  3950 k c a l / k g were not  level  of  of  having  minimum P E : T E  of  protein  crossed  a diet  to  of  al . ,  The e f f e c t  both  raised  upper  line  level  upper c o n f i d e n c e  ratio with  maximum  later.  Gross as  a PE:TE  obtained  by t h i s  experiment.  discussed  limits  confidence  requirement  of  This  the  According  and s e l e c t  15). the  and the  limit  (Fig..  t h r o u g h the  and i n t e r s e c t s  having  (Robbins  lower  present  limit  mean r e s p o n s e  ratio  to  xl  and p a s s i n g  in  noted of  increased  the  fish  as  increase  diets  fed  fish  At h i g h  to  were  3150  fed  diets  dietary the  and  energy  Gross level the  of  the of  dietary  diet  protein  energy  (Table level  protein energy  laws of  utilization protein 32).  and the  GEU was  increased  diminishing  of  in  fish  content  a reduction  of  GEU i n  the  observation  suggests  employed  in  this  case  the  for  4.3.3.  diets  The e f f e c t protein  used  for  followed  Because  33).  not  point  of  protein  of  dietary at  The  which  efficiency the  protein,  by 800  energy  of  dietary  the  kcal/kg  magnitude.  limiting.  protein  This  of  was  (Table  and energy  caused  This  content  the  diet  of  the  and the  pattern  dietary  diets  certainly  the  32).  level  on  the  that  protein  protein  lower  in  to  which  described  in  a maximum at  fish  148  fed  -  dietary  was  diets  was  protein  previous increased,  a lower for  PER and PPV is  protein  the  level  maintenance  by PER  protein of  concentration  levels  for  indicated  between the  minimum r e q u i r e m e n t  utilization not  to  extent  as  utilization  and r e a c h e d  At h i g h e r  are  utilization,  The . r e l a t i o n s h i p  than  protein  NPU-2 v a l u e s  diet  3950 k c a l / k g  increased,  concentration (Table  of  As the  efficiency  was  an i n d i c a t i o n  a similar  chapter.  of  containing  of  growth.  concentration  levels  of  content  by GEU f o l l o w s  At a l l  total  the  the  utilization  The e f f i c i e n c y and PPV, g i v e s  the  as  reached.  indicated  the  by b o t h  energy  increase  was  same o r d e r  that  study  fish as  returns.  energy  to  approximately  the  the  influenced  metabolizable  found  until  requirement  utilization  increasing  (GEU) was  dietary  growth  decreased.  accounted  containing  for, protein  T a b l e 32.  Gross food c o n v e r s i o n e f f i c i e n c y (GFC)and gross d i f f e r e n t l e v e l s o f p r o t e i n and e n e r g y .  M e t a b o l i z a b l e energy content of d i e t (kcal/kg)  17  e n e r g y u t i l i z a t i o n (GEU) o f d i e t s c o n t a i n i n g  Z Crude P r o t e i n i n D i e t 37  27  47  Mean  GFC 3150  11.56 +  3950  ,1  +  23.48  24.70  19.76 +  11.26°  Mean  19.31  0.89  11.41 0.63  P  21.58 20.44  b  q  27.53 25.5l  d e  30.06*  r  0.45 22.61*  27.38'  OMP +  24.40 0.20  GEU 3150 + 3950 Mean +  D C  20.49° 1.59  36.71  o 16.41  b 31.25  18.45° 1.12  33.98  44.01  d  45.77  d  +  q  39.86  cd  cd 41.20  41.94'  43.48'  OMP +  36.74* 0.79 v 32.18  32.03 1.54  1. V a l u e s w i t h t h e same s u p e r s c r i p t f o r e a c h p a r a m e t e r w i t h r e s p e c t t o p r o t e i n l e v e l x e n e r g y l e v e l c ) , p r o t e i n l e v e l ( p - s ) and e n e r g y l e v e l ( v , w ) e f f e c t s do n o t d i f f e r s i g n i f i c a n t l y (DMR t e s t , P " 0.05).  149  (a -  levels three was  below  minimum r e q u i r e m e n t s .  indicators  increased  content  of  from  the  more d i e t a r y  was  i m p r o v e d when  3150  diet  Protein u t i l i z a t i o n energy  content  3950 k c a l / k g .  The h i g h e r  had a s p a r i n g e f f e c t  on p r o t e i n ,  protein  to  the  to  be u t i l i z e d  for  synthesis  by  of  all  diets  energy allowing  of  tissue  protein.  4.3.4  The e f f e c t  of  proximate  body  A pooled  sample  79.88% m o i s t u r e ; lipid  dietary  composition  of  sampled  dietary  at  energy  day  day 0,  decrease  as  decreasing also  in  As one resulted  the  as  (Table  fish  fish  dietary  day 42  energy  9.99%  ash,  the 34  of  and 3 5 ) .  was  protein  16.6%  chinook  experiment  varied  Overall,  slightly The  and 105  from  with the  samples  moisture  was  found  raised.  due  to  increasing  the  dietary  to  A trend  concentration  been  expect,  in  body  level  lipid.  between  dietary  energy  Dietary  protein  concentration  was  or  day 42.  lipid  basis  of  increased  concomittant  size.  i n an i n c r e a s e  body a s h ,  contained  composition  level  dietary  the  day 0  respectively.  at  on  at  decreased  which may have  fish  would  of  and 105  moisture  noted,  increases  42,  of  the  taken  of  level  fish  The g r o s s  concentration  taken  concentration  of  fish  42 and 105  concentration  at  and energy  and on a dry m a t t e r  moisture  was  whole  and 76.54% p r o t e i n .  salmon  protein  protein  at  and body  150  not  energy  An i n v e r s e protein found  to  At day  was  relationship observed.  have 105  level  an e f f e c t  on  differences  in  T a b l e 33.  P r o t e i n e f f i c i e n c y r a t i o n (PER), p r o t e i n productive value (PPV), and net p r o t e i n (NPU-2) of d i e t s c o n t a i n i n g d i f f e r e n t l e v e l s of p r o t e i n and energy.  M e t a b o l i z a b l e energy content of d i e t ( k c a l / k g )  17  1 PER 3150  ab 0.64 + 0.025  3950  0.66  2  be  Mean  X Crude P r o t e i n l a D i e t 37  27  0.65 + 0.18  Pq  _ bed 0.71  , ab 0.64  d 0.79  0.77  0.75  f  cd  0.72  q  OMP  47  utilization  Mean  _a 0.54 ob 0.65 0.60  0.63 + 0.013 v  0.72  P  0.49 + 0.02 1  PPV 3150  abc 42.79 + 1.69 abc  3950  42.77  Mean  42.78 + 1.20  OMP  Pq  abc 44.73  abc 43.27  c 48.87  46.48  46.80  qf  be  44.87  q  a 38.37 ab 41.20 39.78  31.42 + 1.05  P  w 42.29 + 0.86 v 44.83  Table 33.  cont'd...(2)  Metabolizable  X Crude P r o t e i n  energy  content of d i e t  (kcal/kg)  17  27  37  i n Diet  47  Mean  a 49.24 ab 54.10  v> 56.38 + 1.10 v 61.38  NPU-2 3150 3950 i—  1  Mean  K>  be 60.60 + 2.20be 63.63 q  62.1i + 1.55  be 59.60 c 65.91 62.75  q  abc 56.08 be 61.89 58.99  q  51.67  P  —  OMP  54.80 + 1.35  1. A two-way randomized ANOVA i n d i c a t e d s i g n i f i c a n t d i f f e r e n c e s i n PER, PPV and NPU-2 due to p r o t e i n l e v e l (P > 0.005) and energy l e v e l (P > 0.05). D i f f e r e n c e s due to p r o t e i n l e v e l x energy l e v e l and block e f f e c t s were not s i g n i f i c a n t (P < 0.05). 2. Values with the same s u p e r s c r i p t f o r each parameter with respect t o p r o t e i n l e v e l x energy l e v e l d ) , p r o t e i n l e v e l (p - r ) and energy l e v e l (v,w) e f f e c t s do not d i f f e r s i g n i f i c a n t l y (DMR t e s t , P 0.05).  (a -  T a b l e 34.  Whole body p r o x i m a t e c o m p o s i t i o n and e n e r g y , and OMP.  M e t a b o l i z a b l e energy content of d i e t (kcal/kg)  Z  a t day 42 o f t h e d i e t s c o n t a i n i n g d i f f e r e n t l e v e l s o f p r o t e i n  Percent 17  27  Protein i n Diet 37  47  Mean  Moisture (+ SE)  3150  1  78.09  ab  a  abc  77.54  72.09  77.68  + 0.27 3950  +  77.80°  Mean +  w  77.80  P  77.94 0.19  7  6  .  77.04  q  7  6  .  1  1  C  76.60°  76.99  OMP +  0.14  c  76.18  76.65  V  q  77.38 0.27  Z Ash (Dry B a s i s )  ab  a 10.41 + 0.26  3150  9.78  9.59  Mean +  b  10.00 0.18  P  b  9.04  9.38  9.41°°-  9.56  b  P q  9.23  +  9.87 0.13  v  8.91  OMP  Z Lipid (+ SE)  w  ab 9.55 +  ab 3950  ab 9.74  9.23 q  9.86 0.07  (Dry B a s i s )  c 3150  17.89  c 17.18  c  be 19.75  + 1.12 3950 Mean +  w  17.15  17.99 +  22.05°°  24.52°  23.94°  24.81°  19.97 0.79  20.85  21.85  20.98  OMP +  0.56 23.83  V  20.96 0.78  Z P r o t e i n (Dry B a s i s ) (+ SE) 3150  72.98  73.02  67.47  71.02  + 2.27 3950 Mean +  71.12*' +  65.57  67.71  67.06  68.69  69.27 1.61  70.36  67.26  69.85  OMP +  1.14 67.25*  68.67 1.94  1. V a l u e s w i t h t h e Bane s u p e r s c r i p t f o r e a c h p r o x i m a t e component w i t h r e s p e c t t o p r o t e i n l e v e l x e n e r g y l e v e l ( a - c ) , p r o t e i n l e v e l ( p , q ) and e n e r g y l e v e l (v,w) e f f e c t s do n o t d i f f e r s i g n i f i c a t n l y (DMR t e s t , P - 0.05).  Table 35. Whole body proximate composition at day 105 of the diets containing different levels of protein and energy, and OMP. Metabolizable energy content of diet (kcal/kg)  17  Percent Protein In Diet 37 47  27  Mean  Z Moisture (+ SE) 1 78.60 + 0.27 b 77.38  3150 3950  P  77.99 + 0.19  Mean  76.12  „„ , be 76.73  b 77.52  d 75.43  d 75.23  d 75.40  r  75.98  q  75.77  OMP  77.24 + 0.13 v 75.86  0,  76.46 76.05 + 0.15  Z Ash (Dry Basis) 3150 3950 Mean  a 11.70 + 0.72 ab 10.18  ob 10.53  ab 9.52  ab 9.97  ab 9.65  ab 10.06  b 9.57  10.94 + 0.51  10.09  9.79  9.19  OMP  10.43 + 0.36 9.57  8.89 + 0.43 Z Lipid (Dry Basis) (+ SE)  3150  de  DC  18.70  23.42  cd  21.08  18.17*  + 0.85  20.34* + 0.42  3950  22.38  C  26.59°  27.48°  25.21°°  Mean  20.54°+ 0.60  25.01°  24.28°  21.69°"  OMP  V  25.41  23.23 + 0.68 Z Protein (Dry Basis) (+ SE)  3150  C  6  66.61  62.43°  68.20°  74.91°  + 1.39 3950 Mean  68.04* + 0.70  60.12  58.58 q  63.37°" + 0.98  60.50  OMP  58.32  64.23  63.26°  69.57°  60.31  63.58 + 0.41 1. Values with the same superscript for each proximate component with respect to protein level x energy level (a - c), protein level (p.q) and energy level (v,w) effects do not differ slgnificatnly (DMR teat, P - 0.05).  -  154  body  lipid  found  A.A  but  and p r o t e i n no c l e a r  due  trends  to  dietary  were n o t e d  protein (Table  concentration  were  35).  Discussion  A.A.I  The d i e t a r y  protein  requirement  of  juvenile  chinook  salmon The  results  of  the  growth  salmon may r e q u i r e  diets  protein  fry  d u r i n g the  freshwater (197A)  rearing.  who found  dropped  fingerling  stage.  requirements, several Page  species  hand,  rainbow  water  trout  (Ogino  higher the  A0% as  of  fry  may be recommended  in  fish  diet,  in  size  only feed  support  maximum growth  contain  a minimum P E : T E  latter  with  grew  trend  1970;  (x  the  a report for  of  almost  the  chinook of  diets max,  ratio  salmon.  with  Fig. of  entire  1980).  15.5  fish  this  It is  is  on  Diets  the of  study,  likely  this  the  fresh  indicated  During  of  1972;  p e r i o d of  a protein  15).  0.55.  (1977)  In  to  protein  and A r a i ,  70g.  Satia trout  fry  requirements  salmonids  (Fowler,  rainbow  lower  of of  by  and l a r g e r  Nose  protein  chinook  stage  from the  in older  from 5 to  requirement  to  the  the  concentration  G u l b r a n d s e n and Utne  juvenile  stage  that  requirement  and S a i t o ,  period covered  protein  agreement  a general  no change  for  in  a higher  d u r i n g the  protein  1971).  ranging  residency  during  the  a percent  found  experimental  the  as  than  is  There i s  and A n d r e w s ,  other  stage  down from 50 to  suggested  containing  This  that  trial  that  for  period  it  concentration  to  for  fry  should  The e m p i r i c a l r e q u i r e m e n t on the  physiological  experiment,  but  requirement  (Robbins  percentage for  of  line"  chinook  salmon,  water the  (40%)  et  to  al . ,  a r r i v e at  (Dab r o w s k i , that  the  abrupt method.  growth  to  high  was  al.  dietary  rainbow  In tested  the  the  reported  a decline  containing  more  et  to  et  levels  These  study,dietary  in  than  47%. growth  reared  in 7°C  156  In  growth  for  (Cowey  plaice  1976),  grass  They  the  growth  line  maximum  level found in  found  the  line"  response of  is  reached.  that  very  and  that  their  polynominal.  Delong  et  salmon  The d i e t a r y  al. with  protein (1958) diets  protein  was  average  plaice  of  carp  stated  exhibit  "broken  (1976)  chinook  60% p r o t e i n .  1958).  for  continuous  concentrations  However, of  stated  d i d not  authors  degree  the  that  1979).  requirement  depressed  by  al . ,  al.,  by the  al.  a  c  a1.,  of  as  10.5 C.  used  dose  implied  by a second  exceed  et  and r e p r e s e n t a t i v e  respectively.  d i d not  have  and Z e i t o u n et  protein  present  method,  (Zeitoun  growth  the  concentration  diet  than  requirements  minimum p r o t e i n  described  lower  right-hand section  (1972)  trout  best  the  be h o r i z o n t a l  after  Cowey et  of  the  r e a r e d at  (Murai  from l i n e a r i t y  Also,  assumed  were  trout  protein  only  the  estimate  (50%)(Delong  protein  and shad  relationship  change  is  investigators the  35% of  not  of  When e x p r e s s e d  by a s i m i l a r  fish  rainbow  1977)  be  value  1 5 ° C water  other  1972),  This  the  to  minimum d i e t a r y to  depends,  conditions  used  1979).  found  obtained  study  method  al . , the  method.  Alternatively, curves  et  was  and i n  present  on the  dry d i e t  "broken  a nutrient  and e n v i r o n m e n t a l  also  maximum growth  for  data  concentration degree  polynomial equation  containing  3150  requirements arrived  at  fish,  are  approximately  as  (Lee  and g e l a t i n  in  implies  be  fish  that  protein in  level  the  was  not  fish  dietary due  to  depressed  than  a protein  not  the  source  diets  requirement using  diets  (Zeitoun  et  diets  energy  fed  energy  note  above  dietary  level  had the in the  Protein  by the that  dietary  energy growth  the  though  of  intake. energy  intake content rates  of  the  Growth level  their  fish  composition  157  and  energy of  of  the  fish. increasing  depressing  same p r o p o r t i o n of the  the  dietary  levels,  by t h e s e of  for to  The  This  content  of  protein  effect  protein  level  energy  growth  low  due  growth  rate in  was  groups diet.  total  not  diets of It  were s i m i l a r  the  of  was  energy.  responsible  protein  the  containing  contained even  rate,  stated  by  content  1973).  was  of  energy  and A n d r e w s ,  of  that  Feed i n t a k e  The e f f e c t s  limiting  a reduction  to  levels  rate.  be e m p h a s i z e d  by the  The m e t a b o l i z a b l e  47% p r o t e i n .  interesting  protein  fed  Page  alone  must  satiation.  governed  interaction  a factor  restricted  diets  growth  it  on growth  both  by i n c r e a s i n g  containing  the  second  These  rainbow t r o u t  to  1973;  intake  and the  diets  is  based  same a t  were m i n i m a l .  was  respectively.  for as  from a  be 40 and 42% f o r  10% lower  were f e d  animals,  level  rate  to  discussion,  and Putnam,  the  differences  In  the  requirement,  to  energy  the  by o t h e r  diet  found  point  study  protein  the  found  and 3950 k c a l s / k g  casein  this  this  the  was  rate  1976). At  in  maximum growth  by a s i m i l a r method  containing al.,  providing for  energy  diets  fish is when as  differed  (i.e.,  diets  with  37% p r o t e i n  and 3150  and 3950 k c a l / k g r e s p e c t i v e l y ) . the  dietary  trout  was  by l i p i d  protein  30  to  (1:1,  shifted  to  In  non-protein  contrast,  energy  requirements detail  for  was  the  not  growth.  dietary  liver  40% w i t h in  oil)  energy  but  a major  study  factor  aspect  found  the  was  (starch  the  in  provided  requirement and  source  affecting  will  that  maximum growth  carbohydrate  present  This  (1976)  of  protein  be d i s c u s s e d  i n more  later.  Furthermore, content (1961) diets  and cod  al.  required for  35% when n o n - p r o t e i n  approximately  dextrin).  Ogino et  concentration  soybean  k c a l / k g and 47% p r o t e i n  of  depressed  increasing  concluded  that  less  10% of  than  little  also  the  growth of  fiber  diet  Higgs  et  markedly i n  29).  rates  B u h l e r and of  chinook  cellulose.  levels  because  al.  mineral uptake,  differed  (Table  levels  dietary  fiber.  depress  diets  ground c e l l u l o s e  reported with  the  for  adversely  Halver"  salmon  Hilton  et  affect  that  transit  (1983)  should  natural diet  suggested  fed  al.  rainbow t r o u t  their  (1983)  their  be  contains  fiber time  may  of  digesta  *  and  decrease  study, with  the  however,  other  were a b l e  to  above,  protein  Putnam  in  nutrients.  fish  fed  growth  feed  al.,  that  of  In the  low  to  catfish  is  -  as  in  (1966)  and r a i n b o w t r o u t  158  fed  diets high noted  evacuation  This  made by Dupree and Sneed  channel  those  gastric  interacted  energy  and h e n c e ,  1983).  present  cellulose  the  rate  intake,  by a l t e r a t i o n  ( H i l t o n et  conclusions  (1973)  Rather,  a similar  intake  of  seem u n l i k e l y  by i n c r e a s i n g  distention  the  would  attain  diets  with  it  nutrients.  energy  stomach  digestibility  and  agreement and Lee and  respectively.  Discrepancies temperature al. ,  1973) The  with  (Brett  the  p r o p o r t i o n of 14)  (1979)  the  be 0 . 4 1 . chinook  for  is  PE:TE  for  1964).  Discrepancies  from  several  sources  fish  stocks,  water  nutrients.  ratio  in  0.55.  the  However,  some of fish.  reported 1977)  for  the  newly  for  on the  0.41  char,  finding  proportion  metabolizable  data  a single  (Fig.  described a PE:TE be  15). above,  diets  second With one  requirement.  physiologically  -  -  the  was  not  While  correct,  the  Fowler  this  et  assigned  to  the  PE:TE  best  0.51  to  ingredients,  very  small  and W a n d s v i k ,  response  supplied  that  to  the  value  the  by changes  1983). protein in  the  by c a r b o h y d r a t e and  such  estimate  159  for  resulted  agrees with  influenced  PE:TE  to  ( G u l b r a n d s e n and U t n e ,  (Jobling  with  of  different  f r y was  p o l y n o m i a l c u r v e was  confronted  found  likely  that  study  0.41  et  method  reported  1962;  by the  a polynomial curve is  was  values  found  terms  broken l i n e  ingredients,  growth  energy  order  values  chinook  this  0.45  that  of  lipid,  the  0.37  0.35  in  p r o v i d e d by s e v e r a l  in  (Adron  c a s e made by Cowey  al.,  digested  of  1979).  previously  et  (1980)  been  found  concentration of  that  hatched  rainbow t r o u t ,  and a r c t i c  Based  of  the  and c a l o r i c  p r o t e i n was have  (Brett,  requirement  between the  Fowler  effects  frequency  By the  (Combes  temperature  which may not The P E : T E  protein  including dietary  Recently  diet  support  than  0.50  al . ,  the  factors  to  maximum growth r a t e  lower  of  due  feeding  contributed.  ratio  value  salmon,  stating  energy  may be  1970),  made above  PE:TE  This  studies  environmental  observations  Sargent  other  and H i g g s ,  and o t h e r  and  (Fig.  the  as  task of  is  fitted the of  0.55  to  the  one recommending  (x  max) may  unsatisfactory  for  practical  use  identical  dilemma e x i s t s  requirements example, to  because  from  models  requirements  of  Despite  the  authors  concluded  although  that  response  determined  response  at  must  in  bear  mind  regression  Zeitoun  al.  et  "statistical  PE:TE most  that  the  chose  total  were  at  xl  is  namely  these  preferable,  by b o t h methods  ratio  the  response.  "requirement"  models  models  and v i t a m i n  were  may be drawn from  PE:TE  is  the  not  expensive  view  the  nearly results  identical PE:TE =  on the  one  cannot  basis  that  statistically  to 0.41  increasing  number of  may have  very  the  had the  shift  succinctly is  somewhat What i s  importance." could  be  components  to  the  growth  out  relevant  Also,  in  of  the  narrowing right.  that  is  for a  regressions difference  E c o n o m i c a l l y , the since diet.  one  observations  the  irrelevant  a fish  160  1976).  effect  point  important of  al,  xO and x l  establish  significantly  et  parameters.  diets  of  x max ( Z e i t o u n  significance  a practical of  point  xO or x l  and hence  (1976)  significant  which has  at  analysis  limits  at  levels  that  confidence  the  the  non-linear  statistical  these  f rom  in  For  arbitrarily  b r o k e n - l i n e model,  requirement  different  with  livestsock.  95% of  defining  the  nutrient  amino a c i d they  An  14,15).  protein  the  as  of  compared n o n - l i n e a r  the  conclusion  by the  From a s t r i c t the  (1979)  requirements  An i d e n t i c a l study  other  chick,  in  considerations.  estimation  reached  the  estimated  present  (Figs.  al.  growing  that  economic  data with  subjectivity  the  same.  the  the  re-evaluate  the  which  the  in  growth  to  dose a t  in  ignores  when R o b b i n s et  linear  the  it  protein  difference is  one  The i m p l i c a t i o n  of  is  that  each  the  dietary  monetary the  curve  cost  of  is  Feed  this  relative  growth  for  was  conversion  approximately 32).  utilization  when  rate  of  the  when  cost  of  corresponding  to  fish.  cost  Thus  benefit  i n f o r m a t i o n on price  increased  increased  preferred  source  GEU by f i s h due  to  the  other  not  affected  fed  increased hand,  for  requirement  of  as  was  of  the  the  at of  the  the  of  lipid  fish,  Takeuchi  to  previously.  161  the  dietary The t r e n d was  protein  is  The  density  digestibility  support  energy  diets  et  The  level  energy  1982).  energy  Hence,  protein  diet.  the  concentration.  GFC and GEU s e r v e d  trout.  protein  GFC as  the of  for  a lower  lipid  of  dietary  ( W a l t o n and Cowey,  that  rainbow  dietary  to  ratio  principal  considered.  total  37% of  digestibility.  made  of  a dietary  manner  that,  containing  by d i e t a r y  to  PE:TE  point  energy  are  efficiency  raised  the  reported  estimate  diets  levels  the  an i n d i c a t o r  obtained  both  was  lipid  as  be used  in a similar  diets  (1974)  requirements  GEU as  may s u p p o r t  by S a t i a  protein  Predictably,  increased  obtained  on the  the  and s e l l i n g  isocaloric  3 7% at  concentration  of  to  be a d a p t e d  fish  used  GFC c o u l d  maximum f o o d  noted  the  determining  study,  protein  growth  based  available.  only  (Table  the  15 c o u l d  of  requirements  of  to  be  production enterprise  conversion  criteria  increment  in F i g .  a fish  feed,  product  could  assigned  shown in  requirement  protein  value  analysis  In  protein  al. in  higher  may be (1978)  trout  results  protein  the  was  on  A.4.2  The e f f e c t  of  dietary  energy  on the  efficiency  of  protein  utilization The  replacement  furnish  the  means f o r sparing and  effect  Putnam, et  Utne,  1977;  levels  1976;  dietary  lipid,  the  from  to fed  up to  43% of  ratios  the  the  growth  present  carbohydrate highest PER  from 71 diet.  were i n c r e a s e d  although In  no l o s s  rate  of  PE:TE  utilization  increased  increasing  the  was  similar in  the  ratio  the  162  diet  were  of  be  reduced  the of  efficiency protein treatments.  growth  as  33).  beyond  protein  dextrin  replaced  by  lower  The  to  minimum  by  0.42.  when  Therefore the  both  measured  from 0 . 2 8  (P < 0 . 0 5 )  (Table  For  lipid  gains  basis.  ranged  ratio  of  energy  PE:TE  of  for  was  for  0.66  levels  among d i e t a r y  utilization  to  could  protein  diets  1975;  1980).  level  i n which  dextrin  significantly  from 0.42 PE:TE  that  al.,  B u h l e r and H a l v e r  diets  on a m e t a b o l i z a b l e  protein  diet  40% by i n c r e a s i n g  protein  and PPV o c c u r r e d when  Protein  of  (Lee  containing  weight  gain.  by s u b s t i t u t i n g  study,  protein  al.,  At t h i s  the  i n weight  remained  potential  48% and l e v e l s  of  They showed  and l i p i d  values  to  et  best  to  G u l b r a n d s e n and  diets  18% l i p i d .  a series  This  1976;  that  content  a  T a k e d a et  trout  to  is  energy  investigators  Shimeno  They found  salmon  fish.  1973;  fed  from 16  protein  chinook  decreased  (1978)  non-protein  by s e v e r a l  1978;  contained  35% w i t h  level  rearing  and W i l s o n ,  al.,  al.  25%.  diets  (1961)  Garling  ranging  when  48  of  with  requirements,  and A n d r e w s ,  T a k e u c h i et  from 5 to  obtained  energy  cost  Page  protein  energy  been o b s e r v e d  T a k e u c h i et  of  ranging  the  has  1973;  al.,  example,  protein  metabolizable  reducing  Adron  of  the  protein  requirement  reducing any  the  fed  the  in  not  diets  sparing  of  use  limited  by the  high  growth  with  a PE:TE and  lipid  spared  27.6%  fish  the  fed  other  diet  of  lower  rate to  This  than  did  that  of  33.8%  and  protein,  (OMP, T a b l e  diets  the  and  33).  In  the fish  digestible which  both  and  promoted with  disadvantageous to  oxidative  carbohydrates  fed  amount study,  either  of  30).  163  identical  of  two  0.42  fish  the  growth a PE:TE  at  further that  33).  with  only  a  of  and .only  was  contained further a  PE:TE  slightly 0.42  However,  markedly with  and  spared  Still  diet  ratio  diets  22.5%  protein  11.79% l i p i d .  carbohydrate, contrasts  was  glucose  carbohydrate  dietary  protein  is  of  One c o n t a i n e d  ratio  fish  carbohydrate.  present  diets  OMP ( T a b l e  of  demonstrated  12.95% d i g e s t i b l e  a PE:TE  fed  the  of  lower,  that  have  and the  31).  Dietary  diet  of  by f i s h  by f e e d i n g  diet  use  rate  31).  can be  starch  (Table  With  achieved  0.38.'  and  of  carbohydrate  contained  diet  0.52  with  (Table  and s u s c e p t i b i l i t y  16.4%  (Table  the  may be  27.4%  of  slightly  by l i p i d  were a c h i e v e d  ratio  was  than  of  promoting  0.42  2)  hand,  without growth  (P > 0 . 0 5 ) ,  tolerate.  respectively.  ratio  similar  other  effect  the  of  0.52  the  fact,  i n Chapter  fish  cost  can  rates  digestible  savings  high  in  digestibility  equally  by f i s h  of  protein  lipids  In  lower  has  utilization  ratio  (reviewed  On the  carnivourous  6.32%,  of  their  rancidity.  rate.  ratio  dietary  PE:TE = 0.41)  protein  a PE:TE  a PE:TE  studies  of  of  growth  with  Several  because  (i.e:  significantly  with  Increased  and  the  a diet  although fed  efficiency  increase  fish  level  and it  5.70% the  lipid  commercial  The above Halver  observations  (1961)  carbohydrate  who found was  tolerate  diets  variance  with  (1982)  containing  who c l a i m t h a t not  major  differences  exist  that  rainbow t r o u t  are  were  the  chinook  variable  inclusion depending  was  authors  inclusion  to  a lesser  level  in  1967). that  diet  Therefore,  juvenile  carbohydrate  this  levels amply  as  to  be  the  to  demonstrated  that  energy the  not  salmon  it  This of  results  present  in  inclusion the  fish  1976;  utilize  that  Pieper  164  of  and diet.  glucose  by the  as  and  level and the Nose,  indicate  absorbed thought  starch-containing in  reliance  diets.  fat  the  study  previously  to  of  be  (Singh  than  recommended  than  optimum  decreases  increased of  that  raw s t a r c h  extent  may l e s s e n  (Smith,  of  to  energy  the  affected  able  prior  trout  considering  protein,  are  may be  in  study.  that  and d e x t r i n is  at  salmonid  digestibility  and was  rainbow t r o u t the  of  content  starch,  processed  a source  suggested  cooked  since  is  and A t k i n s o n  and  present  of  could  seem d o u b t f u l  species  The d i g e s t i b i l i t y  of  this  levels  species  balance  total  99%)  salmon  c a r b o h y d r a t e may w e l l  overall  a greater  species.  (1982)  dietary  and H i l t o n  would  the  B u h l e r and  chinook  diet.  chinook  1969),  ingredients  as  (96  the  It  in  digestible  reported  extent,  the  (Phillips,  of  well  also  in  employed al.,  of  carbohydrate  14%.  of  However,  (1969)  a more d o m e s t i c  et  level  levels.  between r e l a t e d  on the  uniformly high  of  for  as  to  findings  20% a l t h o u g h  higher  12  salmon  Hilton  carbohydrate These  exceed  the  optimal  digestible  should  of  the  by P h i l l i p s  diets  levels  that  approximately  reports  Subsequently,  support  practical on h i g h  This  point  and P f e f f e r ,  diets lipid has  1980;  been  Hilton  et  adopted  al.,  by f i s h  Although applied livers  to  high  size  tolerance function  4.4.3  farming, fish  for  these  (Hilton,  (1970)  by the was  time  not  the  noted.  and G r o v e s , with  of  to  However,  to  the  it  has  content  been  widely  develop  to  the  shown  trout  1969;  health that  reduce  toxicants  when  enlarged  (Phillips,  been in  waterborne  source  study  salmon al.,  that  salmon  in  the is  Lee  of  abnormal the  and i m p a i r  as  liver  rate  of  also  1984)  above  Increasing  where  study  of  of  of  early  size  life  observed  energy  165  on  in  the  with  content  growth  further  fish  but  change gross in  body  (Buckley  present  steelhead  study trout  and c o m m e r c i a l  effect of  of  body  variations  the  the  in  salmonids  both e x p e r i m e n t a l  is  those  increase  smolt  a result  observations the  with  during  the  energy  fish  non-lipid fraction was  dietary  agreement  reached  throughout  of  of  rapid  and i n a r e c e n t  employed. on the  and l e v e l  are  do o c c u r  This  diets  composition.  levels  body c o m p o s i t i o n  sockeye  et  Imposed  to  diets  be h a z a r d o u s  since  body water  (Fagerlund were  the  this  1979).  chinook  have  1982).  changes  relative  to  may be a d v a n t a g e o u s  these  glycogen  could  who o b s e r v e d  compositional lipid  fish  of  relative  fed  glycogen  proximate  The r e s u l t s  protein  This release  The e f f e c t  Groves  liver  and l i v e r  the  appear  carbohydrate diets  1973).  of  does not  manufacturers.  and i n c r e a s e d  destined  liver  yet  feed  fish  and Putnam, fish  1981)  the  of diet  diet with  herring also  oil  been  noted  Watanabe, protein level et  had the  effect  by o t h e r  1982).  and ash  (1983)  investigators  is  essentially  cite  are  several  of  and body  the  diet,  other  hand o b s e r v e d  were p o s i t i v e l y range  of  content of  the  tended fish  to  or  study  between diet  composition.  in  species,  and p r o t e i n  source  of  composition  characteristics suggest other  that  hand,  production environment  of  lean high  of  have the flesh lipid  or  protein  protein  the  lipid.  caloric  body  of  fall the  fat.  identical  body  regardless  diets is  Over  study  in  PE:TE  the  is  a function  flesh.  bearing Current  amount of  of the  in  quality trends  consumers.  may be a d v a n t a g e o u s  fish  on l i p i d  to  but  body  human d i e t a r y  more d e s i r a b l e  retention  in  changes  on the  35).  because  ratios  and the  These  32,  present  content  energy.  age  environmental  moisture  also  the  of  (Table  and  on  lipid  complicated  ingredients  fish  ratio  ratios  differences  in  Higgs  inverse  revealed  a direct  is  lipid  calorie  constituents  with  moisture,  an to  to  present  studies  proximate diets  report  1979;  Lee and Putnam (1973)  the  dietary  hatchery-reared  may depend  in  content  non-protein  would  that  percentage  between  deposited  dietary  body  with  composition Clearly,  in  and s u b s e q u e n t l y  results  fed  dietary  higher  tested  of  fish  of  changes  has  and S a r g e n t ,  independent  content.  energy  C o m p a r i s o n of  differing  lipid  rise  total  differences  factors.  levels  (Cowey  This  body  level,  that  correlated  protein  Comparison of  lipid  lipid.  in  studies  between p r o t e i n  body  decrease  made f o r  relationship the  increasing  The c o n c o m i t a n t  when a l l o w a n c e s  al.,  of  to  the  where  survival  in  reserves  (Buckley  and G r o v e s ,  166  the  On the  wild  1979). energy  The r e s u l t s sources  as  of  well  the as  energy  diet  may be m a n i p u l a t e d to  with  respect  previously, release  to  protein,  high l i v e r  present  alter  lipid  glycogen  c o u l d be a f a c t o r  that  -  study  demonstrate  c o n c e n t r a t i o n of the  that  the  salmonid  body c o m p o s i t i o n of  and g l y c o g e n .  in fish  destined  adversely  affects  ocean  -  fish  As m e n t i o n e d  reserves  167  the  for  survival.  4.5  Summary of  Experiment  Determining chinook  salmon  rate  dilemma to  "economic  protein  modified study.  order  The m o d i f i c a t i o n  percentage  in  of  terms  the  fish  culturist,  suit  a particular release  may r e q u i r e protein. a PE:TE  of  improvements manufacture  of  table  fish  input  costs.  safe  to  just  within The  an o p t i m a l range  above P E : T E  of  that  the  of  results  the  fish  salmon  a wild  is of  retention that  farmer  the  available This  found  to  and producing  returns  was  -  (PE:TE  and  ingredients  0.41.  for  major  maximizes not  to  contained  maximum growth  of  the  smolts  (OMP)  utilization  168  of  a  environment  protein not  as  requirement  diet  The f i s h  that  ratio  relationship  to  than  may be a d j u s t e d  chinook  energy  information  optimal  protein  diets  indicate  ratio  present  objectives  for  feeds.  was  on the  selection  PE:TE  of  an the  It  commercial  dietary  of  involved  physiological  required  the  concept  equation.  adaptability  meets  with  (1976)  rather  fish  of  al.,  energy  Thus p r o d u c i n g  a PE:TE  Where t h i s  the  ratio  be made i n  may s e l e c t  select  Depending  objectives A  expressing  utilization  commercial  the  juvenile  efficiency.  method  metabolizable  on g r o w t h ,  protein can  to  to  the  the  polynomial  a successful  similar  results  efficiency  which  Their  involved  an enhanced  Although,  = 0.55);  PE:TE  need.  a diet  ratio  of  et  suggested  applied  dry d i e t .  the  with  by Z e i t o u n  who  for  between  utilization  requirement."  and s u c c e s s f u l l y  requirements  for  trout  a second  requirement  a compromise  discussed  rainbow  of  protein  and p r o t e i n  was  reference  application  gross  presents  maximum growth similar  the  2  it  over  may  value  be  lies  efficiency.  be a f f e c t e d  by  the  source  of  proportions dextrin) PE:TE  non-protein  of  and  carbohydrate  lipid  ratios,  energy  (herring  growth  and  in  the  (supplied oil)  the  diet.  When  by g l u c o s e  were v a r i e d  efficiency  of  in  the  and  hydrolyzed  diets  dietary  of  energy  similar and  I protein the  utilization  fish  having  reflected  higher  fish  can  be  their  content  by m a n i p u l a t i n g carbohydrate  were s i m i l a r .  the  PE:TE  and l i p i d reared  of  to  in  However,  diet,  the  body  lipid.  ratio the  and  diet  a prescribed  composition.  169  higher  body  lipid  diet  Experiment the  of  composition  2 showed  proportions  juvenile  performance  fed  fish that  of  chinook and  of  body  salmon,  CHAPTER 5  5 .0  Conclusions The  subject  involves  several  draw a t t e n t i o n protein of  of  quality  the  of  of  investigators,  serve  to  aid  juvenile  successful  protein  other  in  the  chinook  salmonids  have  deficient  diets  amino a c i d s  of  the  have  fish  dietary  aspects  s c o p e and  this  concerning  protein  on growth and with  the  drawn from  diets  for  the  may  requirements  supplemented  with  pure  nutrient  of  r e a r i n g of  experiments  of  effects  study  from  level  to  the  findings  this  the  protein  are  largely  the  thesis  These  The p r e s e n t  been  in  made i n  Along  f o r m u l a t i o n of  determine  was  wide  diets.  conclusions  calculated  very  relevant  utilization.  salmon.  been  to  An a t t e m p t  two most  and q u a n t i t y  efficiency  nutrition is  factors.  to  content  protein  for  in  proteins  which and  g i v i n g maximum  growth. Experiment conversion response The  and p r o t e i n  to  both  proteins  mixture  of  on the  to  nature  satisfy  of  to  casein the  of  and g e l a t i n  sources  protein  170  source  the  latter  -  feed in  protein.  (i.e.  requirements  and p r o c e s s i n g  re-evaluated  -  dietary  d e r i v e d from  with  raw m a t e r i a l  may d i f f e r  supplemented  known amino a c i d  obtained  which g r o w t h ,  efficiency  and c o n c e n t r a t i o n  In E x p e r i m e n t 2 the  salmon were  extent  utilization  and p r o t e i n  the  the  i n c l u d e d a pure p r o t e i n  The r e s p o n s e s  employed. chinook  tested  salmon)  fishery.  source  vitamin-free  amino a c i d s chinook  1 demonstrated  the  a with for  commercial  were  dependent  conditions  requirements  for  juvenile  employing protein  a freeze-dried  source  1.  In  both  the  test  The  results  found  experiments  diets  commercial  The  showed t h a t fish  utilization  concluded  that  present  the  (FPE) i s  valid  if  enhancing  the  pollock  that  mix  in the  acids  drying found that  of  jacketed  the the  fish  product  of  meal  growth fed  diet.  of  for  Mild  made  a freeze-dried  that  in  the  satisfactory  (9:1)  as  a  that  it  of may  acids  be  present  in  and p r e f e r r e d  conclusion  would  no o t h e r  growth  protein  quality  over only  or  protein  from the  considered plants  would  meal  171  loss  showed of  quality  same l o t to  be  high C  (75 C)  was  compared  of  to  raw  similar  equipped  with  seem d o u b t f u l  that  would  that  some  i m p a i r e d by  drying temperature in  by l o c a l It  salmon  drastically  reduction  dryers.  improvement  protein  amino  amino a c i d s  was  may be  (OMP).  in FPE.  bioassays  meal  and  a mix  This  were made  fish  be  Therefore  chinook  present  be m a n u f a c t u r e d  production  for  for  fed  evaluation.  available  (150°C).  a freeze-dried  can  of  the  Experiment  diet  indicated  may not  by g r o u p s  purified  a slight  This  of  (9:1),  in  salmonid scope  quality  terms  from h e r r i n g meal  cause  material. which  to  temperatures to  in  quality  also  and e u p h a u s i d .  were  of  ample  source  protein  assumption  results  is  mix  were made between g r o u p s  The r e s u l t s  optimal  factors  highest  commercial  there  pattern  the  availability  amino  the  protein  for  and e u p h a u s i d  comparisons  were o b t a i n e d  this  The  based  responses  freeze-dried  be  feeds.  standard  best  have  and a p o p u l a r  casein-gelatin reference  to  pollock  be c o m m e r c i a l l y  to  that  steam  viable.  Definitive protein this  quality  study.  protein  intake.  bioassays  on p r o t e i n  compared.  through  the  It  various  is  used  also  for  exogenous  protein.  The d e t e r m i n a t i o n of  of  the  bioassay fish  to  diet  described quality  for  that  the  the  in  this all  of  heat  of  fishmeal.  damage  of  improvements thesis  for  quantities  of  all  gain  or body  adequately  amount  intakes  lysine  In c o n c l u s i o n ,  means  of  the  test  not  found  that  essential  the  to  predictor the  use  a l i m i t i n g amino a c i d  testing  intake  lost  of  was  i n methodology.  in  protein  nor a r e l i a b l e  biological  non-specific  of  an i n t e r e s t i n g  different  of  of  partitioning protein  available  presence  results  be more  level  may be  for  of  a  in a  The p r o c e d u r e s of  protein  p r o p o r t i o n s and  amino a c i d s  were  tested  simultaneously. The  of  define  requires  were  available  value  to  growth and the  u t i l i z a t i o n at  measure  nutritive  sources  provides  protein  the  body w e i g h t  that  maintenance, excretions  a single  with  concluded  depicting  be an a c c u r a t e  considered  made a t  protein  methodology  involving regression  are  along  best  be drawn from  methods  than e s t i m a t i o n s  gain  that  can not  intake  When c o n s i d e r e d  protein  r e g a r d i n g the  Nevertheless,  gain  meaningful  into  conclusions  results  the  the  total  protein  Experiment 2 provided c o n v i n c i n g  requirements  p r o p o r t i o n of  fish  needs,  of  in  this  study  protein intake energy  proportions  ratio of  dietary  of  were  for  fish  energy fed  to  was  governed  the  diet.  should  stated  in  s u p p l i e d by p r o t e i n . satisfy by the  Although  c a r b o h y d r a t e and l i p i d  172  be  evidence  their  total  energy  Since  energy  p r o t e i n energy the  terms  to  varying in  the  diets  could  have  confounded  effect  was  study.  are  source  of  of  above  observations,  detected  under  the  Therefore,  salmon  diet  the  able  to  It  is  adapt  This  salmon also  requirements  that  chinook  salmon  requirement  This  described  response  establishment fish range  chinook other  to  of  nutritionist for  the  salmon  energy,  a species  and economic  growth  either  of  use  the  dietary  suitable  of  quantitative found  investigators  in for  depend, fish,  limits  for  protein study  carnivorous  173  this  of  this  chinook  lipid of  the  as  a  practical  suitability  protein  not  only  on  the  but  also  on  the  of  the  a polynomial energy  confidence  this  the  or  e m p i r i c a l gross  protein  with a basis  juvenile  of  domestication.  considerations  the  that  would advance  for  the  conditions  carbohydrate  within  conclusion  objectives study  to  concluded  of  physiological  as  experimental  may be c o n c l u d e d  metabolizable  formation.  chinook  it  no i n d i c a t i o n  fish curve  level,  limits,  requirements agrees fish  with  relating  which,  would  formulating  culturist.  with  provide  fish  a  the  feeds.  for  juvenile  those  stated  species.  the  The  by  6.0  Bibliography  A l b a n e s e , A . 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ANOVA of  Source  Protein  df  (PS)  day  42  Mean  body  weights  squares  in  Experiment  Prob.>F  Test  1.  term  4  106. .33  0. .0002  PSxRow  2  301 ,.82  0, .005  PLxRow  PSxPL  8  9, .16  0, .006  PSxPLxRow  Row  1  0. .0914  0. .62  Residual  PSxRow  4  0, .774  0. .078  Residual  PLxRow  2  1. .730  0. .009  Residual  PSxPLxRow  8  1, .309  0, .0004  Residual  Level  (PL)  Residual  1770  Total  1799  0. .3688  -  189  -  Table  2.  A n a l y s i s of v a r i a n c e and c o v a r i a n c e f o r s l o p e (GR%) t e s t of l o g body w e i g h t s i n E x p e r i m e n t 1.  Source  df  Mean  squares  Prob.>F  Test  term  4  12.5495  0.001  PSxRow  L e v e l (PL)  2  29.0717  0.01  PLxRow  PSxPL  8  0.4654  0.1  PSxPLxRow  Row  1  0.02686  0.5  Residual  PSxRow  4  0.2592  0.0000  Residual  PLxRow  2  0.4192  0.0000  Residual  PSxPLxRow  8  0.2122  0.0000  Residual  Protein  (PS)  Residual  5369  0.00517  DayxPS  4  8.3211  0.05  DayxPSxPLxRow  DayxPL  2  18.2168  0.01  DayxPSxPLxRow  14  5.1357  0.01  DayxPSxPLxRow  0.00140  0.8  Residual  2.5032  0.00.00  Residual  DayxPSxPL DayxRow DayxPSxPLxRo w  1 29  Residual  5349  Total  5399  0.03840  -  190  -  Table  3.  Analysis protein  Source  o f c o v a r i a n c e o f body w e i g h t intake (Experiment 1).  gain  d_f  P r o b . >F  Mean s q u a r e s  against  4  1.3426  0.0001  Row  1  0.0053  0.63  PSxRow  4  0.006  0.89  1  31 .5088  0.0001  PIxPS  4  0.2593  0.0001  PIxRow  1  0.0006  0.88  Error  24  0.0219  Total  39  Model  15  2.5297  Error  24  0.0219  Total  39  Protein  Prot.  Table  (PS)  i n t a k e (PI)  4.  ANOVA of  GFC and  GEU of  Parameter Source Protein  fed in  Experiment  Mean s q u a r e s  1.  GEU  GFC df  Prob.>F  Mean s q u a r e s 478.34  Prob.>F 0.0016  4  213. 62  0.0001  L e v e l (PL)  2  579. 18  0.014  PSxPL  8  9 . 26  0.0002  Row  1  0 . 584  0.25  0.66  0.63  PSxRow  4  2. 26  0.017  6.90  0.11  PLxRow  2  2. 73  0.017  6.25  0.15  PSxPLxRow  8  0. 388  Total  (PS)  diets  0.0001  17.70  2 .63  29  -  1120.04  191  -  0.015 0.0076  Table  5.  ANOVA of PER and NPR o f  Parameter  fed i n Experiment  PER df  Source Protein  diets  (PS)  Mean  squares  1.  NPR Prob.>F  Mean  squares  Prob.>F  4  0.3097  0.0013  0.2267  0.0019  L e v e l (PL)  2  0.0679  0.077  0.0144  0.064  PSxPL  8  0.0129  0.0004  0.0162  0.0001  Row  1  0.00348  0.057  0.00864  0.0049  PSxRow  4  0.00364  0.025  0.00373  0.0133  PLxRow  2  0.00508  0.017  0.000836  0.2921  PSxPLxRow  8  0.000715  Total  Table  0.000579  29  6.  ANOVA of  PPV o f  d i e t s fed  in  Experiment 1 •  Parameter  PPV Mean s q u a r e s  P:r o b . > F  4  1391 .96 ,  0 .002  L e v e l (PL)  2  468 ,.87  0 .097  PSxPL  8  49. .61  0 .04  Row  1  97. .64  0 .03  PSxRow  4  22 .91 ,  0 .24  PLxRow  2  46. .76  0 .08  PSxPLxRow  8  13, .45  Source Protein  Total  df (PS)  29  192  Table  7.  ANOVA of NPU-1 and NPU-2 o f  Parameter  fed i n Experiment  NPU- 1  Source  Mean  df  Protein  diets  (PS)  squares  NPU- 2  Prob.>F  1372 .41  Prob .>F  1011. .84  L e v e l (PL)  2  69. .70  0 . 17  228 .83  PSxPL  8  58, .52  0 . 02  98 .00  0 .003  Row  1  1,.03  0. 77  164 .39  0 .004  PSxRow  4  24, .54  0 . 20  24 .04  0 .15  PLxRow  2  13, .81  0 . 39  77 .75  0 .016  PSxPLxRow  8  12 .95  Table  0. 003  Mean s q u a r e s  4  Total  1.  0 .002 0 .25  10 .56  29 '  8.  Summary of s t a t i s t i c a l a n a l y s i s f o r the s l o p e s o f d r y body w e i g h t and body p r o t e i n g a i n . S l o p e s were a n a l y z e d b o t h i n c l u d i n g and e x c l u d i n g d a t a f o r t h e p r o t e i n - f r e e diet (PF)(Experiment 1).  Treatment df  Error df  Treatment MS  Error MS  Prob .>F  Dry w e i g h t Rain Slope ( e x c . PF)  15  0.00967  0.00109  0.0007  Slope ( i n c . PF)  25  0.01901  0.00149  0.0000  Protein £a i n Slope ( e x c . PF)  4  15  0.00362  0.00050  0.0019  Slope ( i n c . PF)  4  25  0.00786  0.00082  0.0000  193  Table  9.  ANOVA of  Source  df.  day 42 body  weights  Mean :s q u a r e s  i n Experiment  Prob.>F  Test  term  0 .0002  PLxRow  0 .32  ELxRow  3  174 .32  Energy ( E L )  1  13 .52  PLxEL  3  1 . 142  0 .80  PLxELxRow  Row  1  0 .527  0 .33  Residual  PLxRow  3  0 .334  0 .62  Residual  ELxRow  1  4 .021  0 .0076  Residual  PLxELxRow  3  3 .317  0 .0005  Residual  Residual  944  0 .561  Total  959  Protein  Table  (PL)  10.  ANOVA of  Source Protein  df (PL)  3  day 105 body  weights  Mean :s q u a r e s 3561 .3  i n Experiment  P:r o b . > F  Test  term  0 .0004  PLxRow  Energy ( E L )  1  184 .35  0 .12  ELxRow  PLxEL  3  21 .59  0 .35  PLxELxRow  Row  1  58 .44  0 .003  Residual  PLxRow  3  13 .83  0 .09  Residual  ELxRow  1  0 .32  Residual  PLxELxRow  3  13 .07  0 .11  Residual  Residual  944  6 .46  Total  959  6 .286  -  194  -  2.  2.  Table  11.  Analysis  Source  df  of c o v a r i a n c e  f o r body  w e i g h t s i n E x p . 2.  Mean s q u a r e s  Prob.>F  Test  0.0005  PLxRow  3252. 7  term  PL  3  EL  1  154. 17  0.14  ELxRow  PLxEL  3  23. 74  0.37  PLxELxRow  Row  1  5 7 . 14  0.0000  Residual  PLxRow  3  8. 534  0.014  Residual  ELxRow  1  7. 466  0.079  Residual  PLxELxRow  3  0.0002  Residual  Day  1  0.021  DayxRow  DayxPL  3  0.0005  DayxPLxRow  DayxEL  1  107 . 29  0.106  DayxELxRow  DayxPLxEL  3  17 . 1  0.33  DayxPLxELxRow  DayxRow  1  50. 49  0.0000  Residual  Da yxPLxRow  3  10. 40  0.0048  Residual  DayxELxRow  1  3 . 002  0.26  Residual  DayxPLxELxRow  3  9. 976  0.006  Residual  Residual  5727  2 . 412  Total  5758  15. 59 47510. 2336. 1  -  195  -  Table  12.  Dietary  S t a t i s t i c a l a n a l y s i s f o r t h e second o r d e r p o l y n o m i a l model to e s t i m a t e p r o t e i n r e q u i r e m e n t s i n E x p . 2.  ener gy  Source  df  3950  3150 k c a l / k g Mean  square  Model  2  0.22743  Error  1  0.00081  Total  3  kcal/kg  Prob.>F  Mean s q u a r e  Prob.>F  0.042  0.37356  0.093  0.00658  1  0.36332  0.030  0.63271  0.065  Quadratic term  1  0.09155  0.059  0.11441  0.150  Error  1  0.00081  Total  3  Linear  Table  term  13.  S t a t i s t i c a l a n a l y s i s f o r the second o r d e r p o l y n o m i a l model to e s t i m a t e the P E : T E r e q u i r e m e n t s i n E x p e r i m e n t 2.  Source  df  Mean  square  Model  2  0.57562  Error  5  0.01715  Total  7  Prob.>F 0.0013  1  0.90859  0.0008  1  0.24265  0.013  Error  5  0.01715  Total  7  Linear  term  Quadratic  term  196  Table  1A .  ANOVA of GFC and GEU i n E x p e r i m e n t 2.  Parameter  GFC df  Source  Mean  P r o t e i n (PL)  3  Energy ( E L )  1  PLxEL  3  Row  GEU squares  df  20A.23**  Mean  3  squares  525 .22**  1  83.25  6.003  3  0.A0  1  0.360  1  1 .438  PLxRow  3  0.708  3  2.A51  ELxRow  1  1 .183  1  2.A95  PLxELxRow  3  1 .596  3  5.033  32.29  15  Total Level  of  Table  15.  significance  15 ** =  0.01.  ANOVA of PER and  Parameter  PPV i n E x p e r i m e n t 2.  PPV  PER  Source  df  Mean  square  df  Mean  square  P r o t e i n (PL)  3  0.017A*  3  36.136*  Energy ( E L )  1  0.0302  1  25.702  PLxEL  3  0.00202  3  3.223  Row  1  0.000A05  1  A.852  PLxRow  3  0.000769  3  1 .925  ELxRow  1  0.00132  1  7.930  PLxELxRow  3  0.00127  3  5. 738  Level  15  15  Total of  significance  * = 0.05.  -  197  -  Table  16.  ANOVA o f NPU-2 i n E x p e r i m e n t  NPU-2  Parameter df  Source  Mean  square  3  103.36 * *  Energy ( E L )  1  100.11  PLxEL  3  2 .089  Row  1  10.652  PLxRow  3  3.269  ELxRow  1  10.115  PLxELxRow  3  9.656  Protein  (PL)  15  Total  Level  of  significance  ** = 0.01.  198  2.  Table  17.  T a b l e of mean s q u a r e s f o r f i s h body m o i s t u r e , l i p i d and p r o t e i n a t day 42 ( E x p e r i m e n t 2) •  Mean  df  Source  Ash  Moisture  ash,  squares Lipid  Protein  P r o t e i n (PL)  3  1.2915**  0.4367  2.355  7 .368  Energy ( E L )  1  3.8025**  1.6448  136.422**  59.714*  PLxEL  3  0.2934  0.0403  3.713  9.661  Row  1  0.4356  0.1278  0.483  50.730  PLxELxRow .  7  0.1473  0.1325  2.527  10.331  15  Total  Level  of  Table  18.  signi ficance  **  * = 0.05,  = 0 .01.  T a b l e o f mean s q u a r e s f o r f i s h body m o i s t u r e , a s h , l i p i d and p r o t e i n a t day 105 ( E x p e r i m e n t 2 ) .  Mean  df  Source  Ash  Moisture  squares Lipid  Protein  "3  4.0058**  2.1109  17 .8373**  58.782**  Energy ( E L )  1  7 .6591**  2.9241  102 .8703**  238 . 7 8 0 * *  PLxEL  3  0.3493  0.9528  3.7150  9.959  Row  1  0.6931  0.3080  0.1871  7 .798  PLxELxRow  7  0.1436  1.0499  1.4302  3 .868  P r o t e i n (PL)  Total  Level  15  of  significance  ** = 0.01.  -  199  -  

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