UBC Theses and Dissertations

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UBC Theses and Dissertations

Bioenergetics of the rainbow trout (Salmo gairdneri) and the kokanee (Oncorhynchus nerka) populations… Sandercock, Frederick Keith 1969

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BIOENERGETICS OF THE RAINBOW TROUT ( S a l m o g a i r d n e r i ) AND THE KOKANEE ( O n c o r h y n c h u s n e r k a ) POPULATIONS OF MARION L A K E , B R I T I S H COLUMBIA b y FREDERICK K E I T H SANDERCOCK B . S c , U n i v e r s i t y o f T o r o n t o , 1962 M . S c , U n i v e r s i t y o f B r i t i s h C o l u m b i a , 1964 A THESIS SUBMITTED I N P A R T I A L FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF D o c t o r o f P h i l o s o p h y i n t h e D e p a r t m e n t o f Z o o l o g y We a c c e p t t h i s t h e s i s as c o n f o r m i n g t o t h e r e q u i r e d s t a n d a r d THE U N I V E R S I T Y OF B R I T I S H COLUMBIA D e c e m b e r , 1969 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 o f t h e r e q u i r e m e n t s f o r an advanced deg ree a t t he U n i v e r s i t y o f B r i t i s h C o l u m b i a , I a g r e e t h a t t h e L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e and S t u d y . I f u r t h e r ag ree 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 c o p y i n g o f t h i s t h e s i s f o r s c h o l a r l y p u r p o s e s may be g r a n t e d by the Head o f my Depar tment o r by h i s r e p r e s e n t a t i v e s . I t i s u n d e r s t o o d t h a t c o p y i n g o r p u b l i c a t i o n o f t h i s thes . is f o r f i n a n c i a l g a i n s h a l l n o t be a l l o w e d w i t h o u t my w r i t t e n p e r m i s s i o n . The U n i v e r s i t y o f B Vancouver 8 , Canada Depar tment i ABSTRACT The a n n u a l e n e r g y b u d g e t was d e t e r m i n e d f o r t h e r a i n b o w t r o u t a n d k o k a n e e p o p u l a t i o n s o f a s m a l l (13.3 h a ) B r i t i s h C o l u m b i a l a k e 0 P o p u l a t i o n s i z e o f t h e r a i n b o w t r o u t a g e d 1 + a n d o l d e r was 4000 b a s e d o n a 55 c a t c h - d a y S c h n a b e l e s t i m a t e . Number o f k o k a n e e p r e s e n t i n t h e l a k e (9000) was b a s e d o n t h e r a t i o o f o c c u r r e n c e o f k o k a n e e t o r a i n b o w t r o u t (2.25:1) i n c a t c h . M o n t h l y i n s t a n t a n e o u s g r o w t h a n d m o r t a l i t y r a t e s w e r e a p p l i e d t o e a c h age g r o u p 0 S e a s o n a l c h a n g e i n g o n a d w e i g h t as a p e r c e n t a g e o f b o d y w e i g h t was s i m i l a r i n b o t h s p e c i e s ( o v a r i e s 0.5$~13$» testes0»5#-4#)• S e x r a t i o s w e r e g e n e r a l l y e v e n i n k o k a n e e b u t t h e r e was a h i g h e r p e r c e n t a g e o f f e m a l e s (57.7$) i n t h e r a i n b o w t r o u t . The r e l a t i o n s h i p b e t w e e n egg number p e r f e m a l e ( F ) a n d f o r k l e n g t h ( L ) was c a l c u l a t e d ( r a i n b o w t r o u t , F = 18.7 L * ; k o k a n e e , , F * 2 .84 L l M ) . The p e r c e n t v o l u m e o f e a c h m a j o r f o o d i t e m was d e t e r m i n e d b y s t o m a c h a n a l y s i s . I n o n l y one month (May) was t h e f o o d i t e m o f h i g h e s t $> v o l u m e t h e same i n b o t h s p e c i e s . S e a s o n a l c h a n g e i n e n e r g y c o n t e n t o f t h e f o o d was e s t i m a t e d . C a l c u l a t i o n o f d a i l y r a t i o n ( a s a p e r c e n t a g e o f b o d y w e i g h t ) was b a s e d o n t u r n o v e r r a t e s ( y ) o f f o o d consumed a t d i f f e r e n t t e m p e r a t u r e s ( y = .5^ 5 e * ^ ^ * , w h e r e x i s i n ° C ) 0 F e e d i n g e x p e r i m e n t s u s i n g n a t u r a l f o o d s i n d i c a t e d a s s i m i l a t i o n e f f i c i e n c i e s o f 85.5$. C o n t i n u o u s (20 h r ) measurement o f r o u t i n e m e t a b o l i s m o f i n d i v i d u a l f i s h was made w i t h a r e s p i r o m e t e r s i t u a t e d i n M a r i o n L a k e , b e t w e e n , A u g u s t 1967 a n d O c t o b e r 1968. R a t e o f o x y g e n u p t a k e (Q) i n m l / h r , i i a t 15 C was p l o t t e d a g a i n s t w e i g h t (W) i n k g (Q » 62.6 W * ' ) . To d e t e r m i n e t h e amount o f e n e r g y t r a p p e d w i t h i n t h e t w o p o p u l a t i o n s , s a m p l e f i s h w e r e d r i e d ( g o n a d s s e p a r a t e l y ) a n d b u r n e d i n a c a l o r i m e t e r . The p e r c e n t m o i s t u r e o f t h e f i s h was 7*1—78$ d u r i n g t h e y e a r w i t h m a r k e d i n c r e a s e s a t s p a w n i n g t i m e ( r a i n b o w 8 2 $ , k o k a n e e 88$). The s e a s o n a l c h a n g e s i n e n e r g y c o n t e n t o f i m m a t u r e f i s h w e r e s m a l l ( r a i n b o w t r o u t 6050-6200 c a l / a s h - f r e e g ; k o k a n e e 6150-6300 c a l / a s h - f r e e g ) . However m a t u r e f i s h u n d e r w e n t s h a r p r e d u c t i o n s i n e n e r g y c o n t e n t p r i o r t o s p a w n i n g ( r a i n b o w t r o u t 6100 t o 5500 c a l / a s h - f r e e g ; k o k a n e e 6100 t o 5^00 c a l / a s h - f r e e g ) c The p a t t e r n o f s e a s o n a l c h a n g e i n e n e r g y c o n t e n t o f t h e gonads was s i m i l a r t o t h a t o f t h e b o d y . A v e r a g e b i o m a s s p r e s e n t i n t h e l a k e was 1.53 x 10^ K c a l / h a (12.3 k g / h a ) f o r t h e r a i n b o w t r o u t , a n d 2.23 x 10^ K c a l / h a (16.6 k g / h a ) f o r t h e k o k a n e e . A n n u a l p r o d u c t i o n f o r t h e t w o p o p u l a t i o n s was 2 . 8 1 x 10 K c a l / h a . The r a t i o o f p r o d u c t i o n t o b i o m a s s f o r r a i n b o w t r o u t was 0 . 8 0 , a n d f o r k o k a n e e was 0.71* i i i TABLE OF CONTENTS Page ABSTRACT i L I S T OF TABLES . v i L I S T OF ILLUSTRATIONS i x ACKNOWLEDGEMENTS x i INTRODUCTION I ' LIMNOLOGY OF MARION L A K E . . 2 P H Y S I C A L AND CHEMICAL CHARACTERISTICS 3 T e m p e r a t u r e . . • 3 O x y g e n 3 S P E C I E S I N LAKE 5 GENERAL MATERIALS AND METHODS 5 CAPTURE OF F I S H E S 5 MEASUREMENTS 6 AGE DETERMINATION 6 FECUNDITY 6 FOOD ANALYSIS 7 CALORIMETRY . „ 7 MEASUREMENT OF ROUTINE METABOLISM 9 RESULTS AND INTERPRETATION 13 V I T A L S T A T I S T I C S OF THE F I S H POPULATION 13 POPULATION ESTIMATES 13 AGE DISTRIBUTION 16 i v P a g e AGE STRUCTURE 16 MORTALITY 19 GROWTH 20 L e n g t h o f g r o w i n g s e a s o n 20 L e n g t h - w e i g h t r e l a t i o n s h i p 22 G r o w t h r a t e 22 REPRODUCTION 23 S e x r a t i o . „ 23 S e a s o n a l c h a n g e i n g o n a d w e i g h t f o r r a i n b o w t r o u t 26 S e a s o n a l c h a n g e i n g o n a d w e i g h t f o r k o k a n e e ...<>• 26 Age a t m a t u r i t y 31 F e c u n d i t y 31 FOOD 34 ANALYSIS OF F I S H STOMACHS 34 ENERGY CONTENT OF THE FOOD 39 FOOD TURNOVER 45 A S S I M I L A T I O N E F F I C I E N C I E S 57 ROUTINE METABOLISM O . o 61 A C T I V E METABOLISM 67 MOISTURE CONTENT 68 F I S H EXCLUDING GONADS o . 6 8 GONADS 69 ENERGY CONTENT ..<, 69 C A L O R I F I C CONTENT OF THE F I S H EXCLUDING GONADS 75 C A L O R I F I C CONTENT OF THE GONADS 78 V P a g e ASH CONTENT 8 1 ENERGY CHANGE I N MARION LAKE F I S H COMPARED WITH MIGRATORY SOCKEYE SALMON 83 DISCUSSION 92 U T I L I Z A T I O N OF ENERGY 92 EGG SURVIVAL . . . 8 . 93 ANNUAL ENERGY BUDGET 95 BIOMASS AND PRODUCTION 98 LITERATURE C I T E D 103 APPENDIX 115 v i L I S T OF TABLES P a g e T a b l e 1 . E s t i m a t e s o f r a i n b o w t r o u t p o p u l a t i o n s i z e ( n ) 15 T a b l e 2 . E s t i m a t e d number o f f i s h p r e s e n t i n M a r i o n L a k e a t t h e b e g i n n i n g o f e a c h age a n d a n n u a l i n s t a n t a n e o u s m o r t a l i t y -r a t e ( i ) 18 T a b l e 3« L e n g t h o f g r o w i n g s e a s o n c a l c u l a t e d f r o m d a i l y i n s t a n t -aneous g r o w t h r a t e s o f m a r k e d a n d r e c a p t u r e d r a i n b o w t r o u t . . . . 2 1 T a b l e 4 . A v e r a g e l e n g t h ( L ) a n d w e i g h t ( ¥ ) a t t h e b e g i n n i n g o f e a c h age ( J a n u a r y 1 s t ) a n d a n n u a l i n s t a n t a n e o u s r a t e s o f g r o w t h ( g L a n d g W ) . . 25 T a b l e 5« Gonad w e i g h t as a p e r c e n t a g e o f b o d y w e i g h t (mean ± s t a n d a r d d e v i a t i o n ) 29 T a b l e 6 . P e r c e n t a g e o f f i s h r e a c h i n g m a t u r i t y a t e a c h a g e . . . . . . 32 T a b l e ? . S e a s o n a l c h a n g e i n a v e r a g e egg d i a m e t e r (number i n b r a c k e t s i n d i c a t e s number o f f e m a l e s e x a m i n e d , egg d i a m e t e r i s b a s e d o n 60 e g g s / f e m a l e ) 35 T a b l e 8 . A v e r a g e number o f f o o d o r g a n i s m s p e r s t o m a c h (number o f f i s h e x a m i n e d i n b r a c k e t s ) . T a b l e 9. A v e r a g e e n e r g y c o n t e n t o f v a r i o u s f o o d i t e m s a n d t h e i r p e r c e n t a g e d r y w e i g h t 4 1 v i i P a g e T a b l e 10, A v e r a g e m o n t h l y v a l u e i n e n e r g y c o n t e n t o f f o o d ( K c a l / k g w e t ) 43 T a b l e 11. R e l a t i o n o f l o g t u r n o v e r r a t e o f f o o d t o .o t e m p e r a t u r e ( C) 50 T a b l e 12. C a l c u l a t e d d a i l y t u r n o v e r r a t e o f f o o d a t d i f f e r e n t t e m p e r a t u r e s . 52 T a b l e 13. D a i l y f o o d r a t i o n e x p r e s s e d as a p e r c e n t a g e o f t o t a l b o d y w e i g h t f o r M a r i o n L a k e r a i n b o w t r o u t (mean w e i g h t o f a l l t r o u t e x a m i n e d 91»1 g) a n d k o k a n e e (mean w e i g h t 45.1 g) (number o f f i s h i n b r a c k e t s ) 54 T a b l e 1 4 . A s s i m i l a t i o n e f f i c i e n c y i n r a i n b o w t r o u t b a s e d o n n a t u r a l f o o d s ( H y a l e l l a 3740 c a l / g d r y w e i g h t , C r a n g o n y x 3890 c a l / g d r y w e i g h t ) 58 T a b l e 15. S e a s o n a l c h a n g e i n r o u t i n e m e t a b o l i c r a t e b a s e d o n mean m o n t h l y t e m p e r a t u r e . 66 T a b l e 16. S e a s o n a l c h a n g e i n p e r c e n t a g e m o i s t u r e c o n t e n t o f r a i n b o w t r o u t a n d k o k a n e e ( e x c l u d i n g g o n a d s ) (number o f f i s h i n b r a c k e t s ) 71 T a b l e 17. S e a s o n a l c h a n g e i n p e r c e n t a g e m o i s t u r e c o n t e n t o f r a i n b o w t r o u t a n d k o k a n e e gonads (number o f f i s h i n b r a c k e t s ) 73 v i i i P a g e T a b l e 1 8 . E n e r g y l o s s ( c a l / g d r y w e i g h t ) d u r i n g m a t u r a t i o n i n M a r i o n L a k e r a i n b o w t r o u t a n d k o k a n e e , a n d m i g r a t i n g F r a s e r R i v e r s o c k e y e s a l m o n 86 T a b l e 19. C a l c u l a t e d m e t a b o l i c c o s t o f m i g r a t i o n i n F r a s e r R i v e r s o c k e y e s a l m o n ( S t u a r t r u n ) . .<> 91 T a b l e 20. A n n u a l e n e r g y b u d g e t f o r M a r i o n L a k e r a i n b o w t r o u t 2 4 a n d k o k a n e e , K c a l / m ( x 10 = K c a l / h a ) . 96 A p p e n d i x T a b l e 1 . E n e r g y b u d g e t o f m a t u r e a n d i m m a t u r e r a i n b o w t r o u t o f b o t h s e x e s b y m o n t h . 115 A p p e n d i x T a b l e 2. S u m s , b y m o n t h , a g e , s e x , m a t u r i t y a n d t o t a l , o f e a c h component o f t h e e n e r g y b u d g e t f o r r a i n b o w t r o u t . 128 A p p e n d i x T a b l e 3. E n e r g y b u d g e t o f m a t u r e a n d i m m a t u r e k o k a n e e o f b o t h s e x e s b y month . , 139 A p p e n d i x T a b l e 4. S u m s , b y m o n t h , a g e , s e x , m a t u r i t y a n d t o t a l , o f e a c h component o f t h e e n e r g y b u d g e t f o r k o k a n e e 152 i x L I S T OF ILLUSTRATIONS F i g u r e 1 . S e a s o n a l c h a n g e i n t e m p e r a t u r e a t 3 n d e p t h (1967-68 d a t a c o m b i n e d ) . 4 F i g u r e 2 . D e t a i l s o f c o n s t r u c t i o n o f r e s p i r o m e t e r 10 F i g u r e 3* R e l a t i v e numbers o f f i s h i n e a c h age g r o u p b a s e d o n " c a t c h c u r v e ' * . . . . . < > . . . . • 17 F i g u r e 4 . Change i n l e n g t h (mm) b e t w e e n e a c h a g e g r o u p . . . . . . 2 4 F i g u r e 5« Change i n w e i g h t ( l o g ) b e t w e e n e a c h age g r o u p . . . . 2 4 F i g u r e 6. S e a s o n a l c h a n g e i n g o n a d w e i g h t as a p e r c e n t a g e o f b o d y w e i g h t i n r a i n b o w t r o u t 0 27 F i g u r e 7. S e a s o n a l c h a n g e i n g o n a d w e i g h t as a p e r c e n t a g e o f b o d y w e i g h t i n k o k a n e e . o . 2 8 F i g u r e 8 . T o t a l number o f e g g s / f e m a l e ( l o g e ) a g a i n s t f o r k l e n g t h ( l o g e , cm) 33 F i g u r e 9. C o m p a r i s o n o f t h e p e r c e n t a g e v o l u m e o f v a r i o u s f o o d i t e m s consumed b y r a i n b o w t r o u t a n d k o k a n e e , w h e r e ( L ) = l a r v a e , ( P ) = p u p a e , (N) = n a i a d s , M I S C . = m i s c e l l a n e o u s a n d TERR. = t e r r e s t r i a l 37 F i g u r e 10. S e a s o n a l change i n e n e r g y c o n t e n t o f f o o d ( c a l / g X F i g u r e 1 1 . T u r n o v e r r a t e s i n f o o d ( l o g ) a g a i n s t t e m p e r a t u r e ( ° C ) . o 4 8 F i g u r e 1 2 . S e a s o n a l c h a n g e i n d a i l y r a t i o n as a p e r c e n t a g e of b o d y w e i g h t o e o . . . . . © o o . o o . . . » o o o o o . . 0 0 . 0 0 0 . 0 0 0 . 0 . 0 . 0 . 5^ F i g u r e 13* S e a s o n a l c h a n g e i n p e r c e n t a g e m o i s t u r e c o n t e n t o f t h e b o d y ( l e s s g o n a d ) o 0 0 0 0 . o . . . . . . . . . . . . . o o o . . . . . o . o o o . o 70 F i g u r e 14 0 S e a s o n a l c h a n g e i n p e r c e n t a g e m o i s t u r e c o n t e n t o f t h e gonads o o . . e « « o . o o « » o . . o a o « o . « . . o o o o o o o o o o o o o e » o a . . o o 72 F i g u r e 15. S e a s o n a l change i n e n e r g y c o n t e n t ( c a l / a s h - f r e e g , mean t s t a n d a r d d e v i a t i o n ) o f t h e f i s h e x c l u d i n g gonad i n r a i n b o w t r o u t o o o o o o a o o o . o . e © o a . . » o e o o a o o e o o a o o . « o . . o o © o o 76 F i g u r e 16. S e a s o n a l c h a n g e i n e n e r g y c o n t e n t ( c a l / a s h - f r e e g , mean i s t a n d a r d d e v i a t i o n ) o f t h e f i s h e x c l u d i n g g o n a d i n kolcanOe o • . • o . e a o . o . o a « o o . . o o o . « » « e . . . . . . . o o o . o o . 00000.o« 77 F i g u r e 17o S e a s o n a l c h a n g e i n e n e r g y c o n t e n t ( c a l / a s h - f r e e g , mean £ s t a n d a r d d e v i a t i o n ) o f t h e gonads i n r a i n b o w t r o u t . 79 F i g u r e 1 8 . Seasonal c h a n g e i n e n e r g y content ( c a l / a s h - f r e e g , mean « s t a n d a r d d e v i a t i o n ) o f t h e gonads i n kokanee. . o o o . 80 x i ACKNOWLEDGEMENTS T h i s s t u d y was c o m p l e t e d u n d e r t h e d i r e c t i o n o f Dr„ J . D. M c P h a i l , D e p a r t m e n t o f Z o o l o g y , U n i v e r s i t y o f B r i t i s h C o l u m b i a , t o whom I w i s h t o e x p r e s s my t h a n k s * I am a l s o i n d e b t e d t o D r B Io E B E f f o r d , o f t h e same D e p a r t m e n t , f o r h i s c o n t i n u a l e n t h u s i a s m a n d s u p p o r t f o r t h i s p r o j e c t o I w o u l d l i k e t o a c k n o w l e d g e t h e h e l p g i v e n me i n t h e f i e l d b y M e s s r s , K» T s u m u r a , G, Io McTo Cowan a n d Do M . W a r e . I am p a r t i c u l a r l y g r a t e f u l t o M r 0 Go H a n e y f o r h i s s k i l l i n m o d i f y i n g t h e e l e c t r o n i c e q u i p m e n t u s e d i n t h i s s t u d y e A n a l y s i s o f t h e d a t a b y c o m p u t e r was made p o s s i b l e b y t h e a s s i s t a n c e o f Mr» So B o r d e n . D r . Io McEwan a n d Mr. Po W h i t e h e a d o f t h e C a n a d i a n W i l d l i f e S e r v i c e made a v a i l a b l e e q u i p m e n t u s e d f o r c a l o r i m e t r y 0 The c r i t i c a l r e v i e w o f t h e m a n u s c r i p t b y D r s 0 P. A. L a r k i n , J . Do M c P h a i l , I. Eo E f f o r d , Do Jo R a n d a l l a n d Co Jo W a l t e r s was m o s t • a p p r e c i a t e d o T h i s i n v e s t i g a t i o n was s u p p o r t e d b y g r a n t s t o D r s . Jo D 0 M c P h a i l , Io Eo E f f o r d a n d P« Ao L a r k i n f r o m t h e N a t i o n a l R e s e a r c h C o u n c i l o f C a n a d a a n d t h e F i s h e r i e s R e s e a r c h B o a r d o f C a n a d a , P e r s o n a l s u p p o r t was p r o v i d e d b y N a t i o n a l R e s e a r c h C o u n c i l o f C a n a d a s c h o l a r s h i p s a n d a n Ho Ro M a c M i l l a n F a m i l y S c h o l a r s h i p o 1 INTRODUCTION The p o s s i b i l i t i e s o f p r e d i c t i o n a n d u l t i m a t e m a n i p u l a t i o n o f b i o l o g i c a l p r o d u c t i o n p r o v i d e d a f o c a l p o i n t f o r t h e e s t a b l i s h m e n t o f t h e I n t e r n a t i o n a l B i o l o g i c a l P r o g r a m ( I B P ) . As a p a r t o f C a n a d a ' s c o n t r i b u t i o n t o I B P , t h e M a r i o n L a k e P r o . j e c t was s e t up u n d e r t h e d i r e c t i o n o f D r . I a n E . E f f o r d , U n i v e r s i t y o f B r i t i s h C o l u m b i a , t o u n d e r t a k e a m i n u t e d i s s e c t i o n o f a f r e s h w a t e r e c o s y s t e m w i t h t h e o b j e c t o f d e v e l o p i n g a p r e d i c t i v e m o d e l t h a t w o u l d be a p p l i c a b l e t o o t h e r f r e s h w a t e r c o m m u n i t i e s . T h i s s t u d y was u n d e r t a k e n t o d e t e r m i n e t h e e n e r g y p a t h w a y s a n d r a t e s o f e n e r g y f l o w t h a t e x i s t w i t h i n t h e p o p u l a t i o n s o f r a i n b o w t r o u t , S a l m o g a i r d n e r i R i c h a r d s o n a n d o f k o k a n e e , O n c o r h y n c h u s n e r k a (Walbaum) i n M a r i o n L a k e , B r i t i s h C o l u m b i a , , The b a s i c a p p r o a c h was t o e x a m i n e t h e amount o f e n e r g y " t i e d u p " i n t h e f i s h p o p u l a t i o n s , as w e l l as t h e d y n a m i c s o f t h e e n e r g y f l o w t h r o u g h t h e s y s t e m i n much t h e same way a s d i d W i n b e r g (1956) a n d Mann (1965). s i n c e t h e p o t e n t i a l p r o d u c t i o n o f a p o p u l a t i o n c a n n o t be d e t e r m i n e d u n l e s s t h e e n e r g y r e q u i r e m e n t s a n d i t s a v a i l a b i l i t y t o t h e p o p u l a t i o n a r e known ( M a n n , 1965). ; W i n b e r g ' s m e t h o d o f s t u d y i n g t h e b i o e n e r g e t i c s o f a f i s h p o p u l a t i o n i s b a s e d o n t h e f a c t t h a t t h e e n e r g y c o n t e n t o f t h e f o o d i s e q u a l t o t h e sum o f t h e e n e r g y r e t a i n e d i n g r o w t h , p l u s t h e amount l o s t t o m e t a b o l i s m a n d w a s t e p r o d u c t s . Mann r e - c a l c u l a t e d t h e d a t a p r e s e n t e d i n a s e r i e s o f p a p e r s t o d e m o n s t r a t e c l o s e a g r e e m e n t w i t h W i n b e r g ' s p r e d i c t e d v a l u e s . I n o r d e r t o i m p r o v e r e l i a b i l i t y o f t h e measurements 2 o f e n e r g y t r a n s f o r m a t i o n o f R i v e r Thames f i s h , M a n n d e t e r m i n e d e x p e r i m e n t a l l y t h e s t a n d a r d m e t a b o l i c r a t e o f t h e s e f i s h a n d t h e e n e r g y c o n t e n t o f t h e g o n a d m a t e r i a l . O t h e r v a l u e s , s u c h a s e n e r g y r e q u i r e m e n t s f o r a c t i v e m e t a b o l i s m , b o d y e n e r g y , l o s s t h r o u g h e g e s t i o n a n d e x c r e t i o n f o l l o w e d t h e same a s s u m p t i o n s made b y W i n b e r g . M a r i o n L a k e was c h o s e n f o r t h e p r e s e n t s t u d y b e c a u s e o f i t s p r o x i m i t y , c o n t r o l l e d a c c e s s , a n d b a c k l o g o f p r e l i m i n a r y i n f o r m a t i o n . S t u d i e s o f o t h e r a s p e c t s o f t h i s l a k e e i t h e r c o m p l e t e d o r i n p r o g r e s s p r o v i d e d c o n s i d e r a b l e b a c k g r o u n d d a t a ( H a m i l t o n , M S , 1965; M a t h i a s , M S , 1964 a n d M S , 1967; E f f o r d , 1967; H a r g r a v e , M S , 1969; a n d o t h e r s ) . P o p u l a t i o n s t r u c t u r e , g r o w t h r a t e , m o r t a l i t y r a t e a n d f e c u n d i t y w e r e e x a m i n e d i n a n a t t e m p t t o p r o v i d e a c o m p l e t e s e t o f v i t a l s t a t i s t i c s o f t h e t w o p o p u l a t i o n s . F o o d h a b i t s w e r e d e t e r m i n e d b y : s t o m a c h c o n t e n t a n a l y s i s a n d s u b s e q u e n t l y e x p r e s s e d i n t e r m s o f e n e r g y v a l u e s b y a s s i g n i n g c a l o r i f i c e q u i v a l e n t s t o e a c h f o o d i t e m * F o o d -t u r n o v e r r a t e s a n d a s s i m i l a t i o n e f f i c i e n c i e s w e r e d e t e r m i n e d e x p e r i m e n t a l l y e S e a s o n a l c h a n g e s i n r o u t i n e m e t a b o l i s m a n d e n e r g y c o n t e n t o f t h e f i s h w e r e d e t e r m i n e d . A n i n t e g r a t e d m o n t h l y a n a l y s i s o f t h e e n e r g y f l o w t h r o u g h t h e f i s h p o p u l a t i o n p r o v i d e d a m e a s u r e o f t h e f i s h p r o d u c t i o n a s r e l a t e d t o t h e e n e r g y a v a i l a b l e f r o m l o w e r t r o p h i c l e v e l s 0 LIMNOLOGY OF MARION LAKE M a r i o n L a k e (300 m A S L ) i s s i t u a t e d i n s o u t h w e s t e r n B r i t i s h C o l u m b i a (49°19 f N , 122°33* W) o n t h e n o r t h s i d e o f t h e F r a s e r v a l l e y a b o u t 50 km e a s t o f V a n c o u v e r . E f f o r d (1967) h a s d e s c r i b e d t h e 3 g e o l o g i c h i s t o r y , p r e s e n t l a n d f o r m s , c l i m a t e a n d v e g e t a t i o n o f t h e s u r r o u n d i n g a r e a . The l a k e i s s m a l l , a p p r o x i m a t e l y 800 m i n l e n g t h a n d 200 m i n maximum w i d t h . S u r f a c e a r e a i s a b o u t 13*3 b a (32.9 a c r e s ) , maximum d e p t h a b o u t 6 m a n d mean d e p t h 2.4 m. The v o l u m e o f w a t e r i n t h e l a k e i s a b o u t 330,000 v?. D u r i n g h e a v y r a i n s t h e i n f l o w i n t o t h e l a k e o v e r a 2\ d a y p e r i o d may e q u a l t h e t o t a l v o l u m e o f t h e l a k e ( E f f o r d , 1967). I n t h e v i c i n i t y o f t h e i n l e t a n d o u t l e t t h e r e i s a r e s t r i c t e d z o n e ( c o m b i n e d t o t a l a b o u t 25.5 m^) w h e r e t h e b o t t o m t y p e c o n s i s t s o f g r a v e l a n d w h i c h r e p r e s e n t s t h e o n l y a v a i l a b l e s p a w n i n g a r e a f o r t h e f i s h . E x c e p t f o r t h e a r e a n e a r a l a r g e s p r i n g i n t h e s o u t h h a l f o f t h e l a k e , t h e r e m a i n d e r o f t h e l a k e b o t t o m c o n s i s t s o f 1-2 m d e p t h o f f l o c c u l e n t o o z e . M a c r o p h y t e s ( N u p h a r v a r i a g a t u m , P o t a m o g e t o n s p p . ) a r e s p a r s e l y d i s t r i b u t e d a b o u t t h e l i t t o r a l z o n e . C h a r a i s f o u n d ; i n t h e v i c i n i t y o f t h e s p r i n g s a n d I s o e t e s o c c i d e n t a l i s m o s t l y a t d e p t h s o v e r 2m. P H Y S I C A L AND CHEMICAL CHARACTERISTICS T e m p e r a t u r e — S e a s o n a l c h a n g e s i n t e m p e r a t u r e o v e r t h e e n t i r e w a t e r c o l u m n (1963-1964) a r e g i v e n b y E f f o r d (1967). I n t h e p r e s e n t s t u d y t h e t e m p e r a t u r e c h a n g e s a t t h e 3 m d e p t h z o n e w e r e r e c o r d e d a t a b o u t w e e k l y i n t e r v a l s t h r o u g h m o s t o f 1967-1968 ( F i g . 1). S u r f a c e t e m p e r a t u r e s r e a c h e d a maximum o f 27°C i n A u g u s t 1967 a n d 23°C i n A u g u s t 1968. O x y g e n — The l e v e l o f d i s s o l v e d o x y g e n was g e n e r a l l y c l o s e F i g u r e 1 . S e a s o n a l c h a n g e i n t e m p e r a t u r e a t 3 m d e p t h (1967-68 d a t a c o m b i n e d ) . 4 5 t o s a t u r a t i o n a t a l l t e m p e r a t u r e s . A l t h o u g h t h e l a k e may s t r a t i f y b r i e f l y i n m i d s u m m e r , o x y g e n d e p l e t i o n n o r m a l l y o c c u r r e d o n l y i n d e e p w a t e r c l o s e t o t h e mud b o t t o m . The h i g h f l u s h i n g r a t e i n t h e l a k e a n d t h e p r e s e n c e o f numerous s p r i n g s p r e c l u d e a n i n t e n s i v e o x y g e n d e p l e t i o n i n t h e e p i l i m n i o n 0 T o t a l d i s s o l v e d s o l i d s r a n g e d f r o m 12-15 p p m , t h e amount o f f r e e C 0 2 r a n g e d f r o m 5-15 m g / l ( E f f o r d , 1967) a n d t h e p H , w h i l e m o s t l y a t 7.0 i n summer a n d 6.5 i n w i n t e r v a r i e d b e t w e e n 5*9 a n d 7.4 ( H a m i l t o n , M S , 1965). S P E C I E S I N LAKE I n a d d i t i o n t o t h e r a i n b o w t r o u t a n d k o k a n e e o t h e r v e r t e b r a t e s f o u n d i n t h e l a k e a r e as f o l l o w s : R o u g h - s k i n newt T a r i c h a g r a n u l o s a ( S k i l t o n ) , N o r t h w e s t e r n s a l a m a n d e r Ambystoma g r a c i l e ( B a i r d ) , a s w e l l a s t h e t a d p o l e s o f B u f o b o r e a s B a i r d & G i r a r d , H y l a r e g i l l a B a i r d & G i r a r d a n d R a n a a u r o r a B a i r d & G i r a r d . GENERAL MATERIALS AND METHODS CAPTURE OF F I S H E S B e t w e e n 1963 a n d 1968 f i s h w e r e c o l l e c t e d b y t h r e e g e n e r a l m e t h o d s . The m a j o r i t y o f t h e f i s h t a k e n d u r i n g t h i s p e r i o d w e r e c a u g h t u s i n g m o n o f i l a m e n t g i l l n e t s r a n g i n g i n s i z e f r o m 2 cm s t r e t c h e d m e s h , b y 1 cm i n t e r v a l s , u p t o a maximum s i z e o f 9 c m . I n t h e e a r l y p a r t o f t h e s t u d y , f i s h o f a l l s i z e s w e r e a l s o 6 c o l l e c t e d i n l a r g e t r a p s p l a c e d n e a r t h e i n l e t a n d o u t l e t s t r e a m s 0 A b o u t 50 s m a l l , w i r e - m e s h t r a p s d i s t r i b u t e d a b o u t t h e l a k e a s a p a r t o f t h e a m p h i b i a n s t u d y w e r e u s e f u l i n p r o v i d i n g s m a l l s a m p l e s o f t h e 0 + age g r o u p . D u r i n g 1 9 6 8 , p a r t o f e a c h c a t c h was f r o z e n s h o r t l y a f t e r c a p t u r e a n d s t o r e d f o r l a t e r a n a l y s i s . MEASUREMENTS The l e n g t h s o f a l l f i s h a r e t a k e n t o be t h e d i s t a n c e b e t w e e n t h e s n o u t a n d t h e p o s t e r i o r e n d o f t h e m i d d l e c a u d a l r a y ( f o r k l e n g t h ) m e a s u r e d t o t h e n e a r e s t m i l l i m e t r e . A l l f i s h w e i g h t s a r e t o t h e n e a r e s t 0 . 1 g . AGE DETERMINATION The a g e s o f t h e r a i n b o w t r o u t s a m p l e d w e r e d e t e r m i n e d b y s c a l e r e a d i n g . H o w e v e r t h e s c a l e s o f t h e k o k a n e e , p a r t i c u l a r l y t h o s e o f t h e o l d e r f i s h p r o v e d t o be much more d i f f i c u l t t o r e a d d u e t o r e s o r p t i o n F o r t h e s e f i s h a g i n g was c o m p l e t e d b y e x a m i n a t i o n o f t h e o t o l i t h s -( E f f o r d - p e r s . c o m m . ) . FECUNDITY The o v a r i e s o f f e m a l e r a i n b o w t r o u t a n d k o k a n e e c o l l e c t e d d u r i n g r o u t i n e s a m p l i n g w e r e e x a m i n e d f o r egg s i z e a n d number i n t h e l a t e r s t a g e s o f f o l l i c l e m a t u r a t i o n . F o r r a i n b o w t r o u t t h e p e r i o d o v e r w h i c h t h e eggs c o u l d be c o u n t e d r e a d i l y was e a r l y A u g u s t u n t i l s p a w n i n g t i m e i n m i d - A p r i l o f t h e f o l l o w i n g y e a r , f o r k o k a n e e t h e 7 t h e p e r i o d e x t e n d e d f r o m m i d - M a y u n t i l m i d - O c t o b e r . E a c h o v a r y was w e i g h e d s e p a r a t e l y , t h e n t h e o u t e r membrane was s e p a r a t e d f r o m t h e e g g s . Once t h e eggs w e r e f r e e d f r o m t h e r e m a i n i n g t i s s u e t h e y w e r e d i v i d e d i n t o g r o u p s o f t e n a n d c o u n t e d . T h r e e o f t h e s e , s u b - g r o u p s p e r o v a r y w e r e c h o s e n a t random a n d t h e t e n eggs a r r a n g e d i n a s t r a i g h t l i n e . T h i s l e n g t h was m e a s u r e d w i t h d i a l c a l i p e r s i n c m . A n a v e r a g e o f 60 eggs p e r f e m a l e was m e a s u r e d a n d e x p r e s s e d as t h e a v e r a g e d i a m e t e r p e r egg i n mm. S e a s o n a l c h a n g e s i n t e s t e s w e i g h t w e r e r e c o r d e d f o r b o t h r a i n b o w t r o u t a n d k o k a n e e d u r i n g 1968. FOOD A N A L Y S I S S t o m a c h s w e r e r e m o v e d b y s e v e r i n g t h e oesophagus a n d t h e p o s t e r i o r s e c t i o n o f t h e p y l o r i c s p h i n c t e r . The w a l l o f t h e s t o m a c h was s p l i t a n d t h e c o n t e n t s t r a n s f e r r e d t o a v i a l c o n t a i n i n g 4 0 $ i s o p r o p y l a l c o h o l . The v o l u m e o f e a c h o f t h e m a j o r f o o d i t e m s a n d t h e t o t a l v o l u m e o f t h e f o o d f r o m e a c h s t o m a c h w e r e d e t e r m i n e d b y d i s p l a c e m e n t i n a l c o h o l u s i n g g r a d u a t e d c e n t r i f u g e t u b e s . N o n - e d i b l e m a t e r i a l s w e r e e x c l u d e d f r o m t h e v o l u m e t r i c a n a l y s i s . The c o n t e n t s o f 98 r a i n b o w t r o u t a n d 107 k o k a n e e s t o m a c h s w e r e , a n a l y s e d . CALORIMETRY F i s h f r o z e n a t c a p t u r e w e r e t h a w e d i n c o o l w a t e r . The f i s h w e r e s l i t v e n t r a l l y a n d s e x a n d s t a t e o f m a t u r i t y r e c o r d e d . The gonads w e r e 8 t h e n r e m o v e d f o r s e p a r a t e w e i g h i n g a n d d r y i n g . The c o n t e n t s o f t h e s t o m a c h a n d i n t e s t i n e w e r e removedo The w h o l e f i s h , e x c l u d i n g t h e g o n a d s , was p l a c e d i n a b l e n d e r w i t h a p p r o x i m a t e l y 150 m l d i s t i l l e d w a t e r a n d r e d u c e d t o a homogeneous s l u r r y . The r e s u l t a n t m i x t u r e was p o u r e d i n t o P e t r i d i s h e s ( 1 4 cm d i a m e t e r ) f o r d r y i n g . A l l s a m p l e s w e r e d r i e d i n a c o n s t a n t t e m p e r a t u r e o v e n a t 70°C f o r a minimum o f 60 h r ( r a n g e 60-75 h r ) . Once d r i e d , t h e s a m p l e s w e r e r e m o v e d f r o m t h e o v e n a n d c o o l e d i n a d e s i c c a t o r t o room t e m p e r a t u r e . The d r i e d m a t e r i a l was r e m o v e d f r o m t h e d i s h e s a n d w e i g h e d t o t h e n e a r e s t 100 mg. The d r y w e i g h t o f t h e g o n a d m a t e r i a l was d e t e r m i n e d s e p a r a t e l y t o t h e n e a r e s t 10 m g . The d r i e d , w h o l e - f i s h m a t e r i a l , e x c l u d i n g g o n a d s , was p o w d e r e d w i t h a m o r t a r a n d p e s t l e . Two 1 g s a m p l e s o f p o w d e r e d m a t e r i a l f r o m -e a c h f i s h w e r e c o m p a c t e d i n t o p e l l e t s w i t h a Parr^" p e l l e t - p r e s s i n p r e p a r a t i o n f o r c o m b u s t i o n . P e l l e t s w e i g h i n g 30-400 mg w e r e p r e p a r e d f r o m d r i e d gonad m a t e r i a l . A G a l l e n k a m p ^ ( M o d , CB-110 a u t o a d i a b a t i c ) bomb c a l o r i m e t e r was u s e d t o d e t e r m i n e t h e e n e r g y c o n t e n t o f t h e w h o l e - f i s h s a m p l e s . A t o t a l o f 2 0 4 r a i n b o w t r o u t a n d 222 k o k a n e e s a m p l e s w e r e c o m b u s t e d . A l l s a m p l e s s m a l l e r t h a n 500 mg w e r e b u r n e d i n a Parr"* - c a l o r i m e t e r (Modo 1411 C o m b u s t i o n C a l o r i m e t e r ) , A t o t a l o f 133 d e t e r m i n a t i o n s w e r e made c o m p r i s i n g s a m p l e s o f gonad m a t e r i a l , f o o d i t e m s , f e c a l m a t e r i a l p l u s a 1 , P a r r I n s t r u m e n t C o . , M o l i n e , I l l i n o i s . 2 , A . G a l l e m k a m p & C o . L t d . , L o n d o n , E n g l a n d , 9 few w h o l e f i s h s a m p l e s o f u n d e r - y e a r l i n g f i s h , w h i c h when d r i e d d i d n o t y i e l d s u f f i c i e n t m a t e r i a l t o be b u r n e d i n t h e l a r g e r c a p a c i t y c a l o r i m e t e r . A f e w s a m p l e s o f i d e n t i c a l m a t e r i a l w e r e b u r n e d i n b o t h c a l o r i m e t e r s as a means o f c h e c k i n g t h e p e r f o r m a n c e o f one c a l o r i m e t e r a g a i n s t t h e o t h e r . I n e a c h c a l o r i m e t e r , c o r r e c t i o n f a c t o r s w e r e a p p l i e d t o c o m p e n s a t e f o r t h e h e a t p r o d u c e d b y t h e f u s e w i r e , h o w e v e r no c o r r e c t i o n was made f o r t h e n i t r i c a n d s u l p h u r i c a c i d b y - p r o d u c t s o f c o m b u s t i o n . P e r c e n t a s h v a l u e s a r e b a s e d o n t h e amount o f m a t e r i a l r e m a i n i n g i n t h e bomb c r u c i b l e f o l l o w i n g c o m b u s t i o n . MEASUREMENT OF ROUTINE METABOLISM A f l o a t i n g p l a t f o r m (16 s q m) was a n c h o r e d i n t h e m i d d l e o f M a r i o n L a k e . W a t e r d e p t h was a p p r o x i m a t e l y 4 m. Power was s u p p l i e d t o t h e p l a t f o r m b y c a b l e f r o m a s h o r e - b a s e d d i e s e l g e n e r a t o r . P o s i t i o n e d i n t h e c e n t r e o f t h e p l a t f o r m was a s t y r f o a m i n s u l a t e d w e l l (125 cm x 75 cm x 40 cm) t h e t o p o f w h i c h was a b o u t 10 cm a b o v e t h e l a k e s u r f a c e (Figo 2 ) . The w e l l was c o v e r e d w h i l e i n u s e w i t h a l a r g e p l y w o o d c a n o p y w h i c h h a d l o u v r e d v e n t s o n f o u r s i d e s , a l l o w i n g f o r a m p l e v e n t i l a t i o n b u t p r e v e n t i n g d i r e c t s u n l i g h t f r o m r e a c h i n g t h e w e l l 0 W a t e r was pumped f r o m a d e p t h o f 3 m i n t o a n a c r y l i c p l a s t i c c y l i n d e r f u n c t i o n i n g as a c o n s t a n t h e a d w h i c h c o u l d be a d j u s t e d v e r t i c a l l y t o r e g u l a t e t h e f l o w r a t e t h r o u g h t h e r e s p i r a t i o n chamber,, F l o w r a t e t h r o u g h t h e chamber was v a r i e d o v e r t h e r a n g e o f 6 l / h r t o 40,5 l / h r d e p e n d i n g o n w a t e r t e m p e r a t u r e a n d t h e s i z e o f t h e e x p e r i m e n t a l f i s h . F i g u r e 2. D e t a i l s o f c o n s t r u c t i o n o f r e s p i r o m e t e r . X INTAKE AT 3 m DEPTH 11 The e x c e s s w a t e r f r o m t h e h e a d k e p t t h e i n s u l a t e d w e l l f i l l e d . The w a r m e r w a t e r i n t h e s u r f a c e o f t h e w e l l s p i l l e d o v e r i n t o t h e l a k e D The w a t e r f r o m t h e h e a d f l o w e d t h r o u g h v i n y l t u b i n g i n t o t h e r e s p i r a t i o n c h a m b e r . T h i s chamber c o n s i s t e d o f a n a c r y l i c p l a s t i c c y l i n d e r 75 cm i n l e n g t h a n d 12 .5 cm i n s i d e d i a m e t e r (9 .2 1 c a p a c i t y ) 0 The ends o f t h e c y l i n d e r w e r e c l o s e d w i t h t i g h t l y f i t t i n g c o n e s c o n s t r u c t e d o f m o u l d e d f i b r e g l a s s . D o u b l e " 0 " r i n g s s e a l e d e a c h e n d p r e v e n t i n g a n y e x c h a n g e o f w a t e r b e t w e e n t h e i n t e r i o r o f t h e c y l i n d e r a n d t h e w e l l . The c o n i c a l ends w e r e d e s i g n e d t o e l i m i n a t e a n y " d e a d " w a t e r a r e a s w i t h i n t h e c y l i n d e r . W a t e r f l o w i n g t h r o u g h t h e r e s p i r o m e t e r was e x h a u s t e d i n t o t h e w e l l t h r o u g h a v i n y l t u b e . The o x y g e n c o n c e n t r a t i o n w i t h i n t h e r e s p i r a t i o n chamber was m e a s u r e d c o n t i n u o u s l y o v e r a 2 0 - 2 4 h r p e r i o d u s i n g a m o d i f i e d YSI^ o x y g e n m e t e r ( M o d . 54RC) a n d t w o YST t e m p e r a t u r e - c o m p e n s a t e d o x y g e n e l e c t r o d e s ( M o d . 5 4 1 9 ) . The o x y g e n a n a l y s e r was m o d i f i e d b y t h e i n s t a l l a t i o n o f a d i f f e r e n t i a l a m p l i f i e r a n d a n a d d i t i o n a l o u t l e t ; s o t h a t b o t h p r o b e s c o u l d b e o p e r a t e d o n t h e same u n i t . The a n a l y s e r was c a l i b r a t e d b y i n s e r t i n g b o t h p r o b e s i n t o t h e same w a t e r s u p p l y a n d s e t t i n g t h e o x y g e n s a t u r a t i o n f o r a g i v e n t e m p e r a t u r e . The p r o b e s w e r e t h e n i n s e r t e d i n t o p l a s t i c h o l d e r s l o c a t e d i n t h e i n l e t a n d o u t l e t t u b e s o f t h e r e s p i r a t i o n c h a m b e r . I t was p o s s i b l e t o m e a s u r e t h e d i f f e r e n c e b e t w e e n t h e o x y g e n c o n c e n t r a t i o n o f t h e w a t e r e n t e r i n g a n d t h a t l e a v i n g t h e chamber c o n t i n u o u s l y . The r a t e o f f l o w o f e l e c t r i c a l 3. Y e l l o w S p r i n g s I n s t r u m e n t C o „ , Y e l l o w S p r i n g s , O h i o . • 12 c u r r e n t t h r o u g h a n o x y g e n e l e c t r o d e i s p r o p o r t i o n a l t o t h e c o n c e n t r a t i o n o f d i s s o l v e d o x y g e n . S t a b i l i t y i n t h e p r o b e s was a c h i e v e d b y t h o r o u g h l y c l e a n i n g t h e o x i d a t i v e m a t e r i a l o f f t h e s i l v e r c o i l w i t h i n t h e p r o b e a n d t h e n r e p l a c i n g t h e e l e c t r o l y t e a n d membrane ( s u p p l i e d b y t h e m a n u f a c t u r e r ) . The p r o b e s w e r e p l a c e d i n r u n n i n g w a t e r a n d l e f t i n a n " o n " s t a t e f o r 2h h r . Use o f t h e p r o b e s f o l l o w i n g t h i s t r e a t m e n t p r o v i d e d g o o d s t a b i l i t y a n d d r i f t - f r e e p e r f o r m a n c e . Once c l e a n e d i n t h i s manner t h e p r o b e s c o u l d b e u s e d r e l i a b l y f o r more t h a n 10 d a y s ( a b o u t 250 o p e r a t i n g h o u r s ) b e f o r e r e c l e a n i n g o The s i g n a l f r o m t h e a n a l y s e r was r e c o r d e d o n a H e w l e t t P a c k a r d 5 i n (12.7 cm) s t r i p c h a r t r e c o r d e r ( M o d 0 H01-680). T e m p e r a t u r e was r e c o r d e d c o n t i n u o u s l y u s i n g a R u s t r a k ^ t h e r m i s t o r p r o b e r e c o r d e r ( M o d . 133). F l o w r a t e o f t h e w a t e r t h r o u g h t h e chamber was m e a s u r e d w i t h a s t o p - w a t c h a n d g r a d u a t e d c y l i n d e r . F o r e a c h h e i g h t a d j u s t m e n t o f t h e h e a d , t h e f l o w r a t e was u n i f o r m . The p r o c e d u r e f o r m e a s u r i n g r o u t i n e m e t a b o l i s m was t o s e t a ; s e r i e s o f m o n o f i l a m e n t g i l l n e t s (2.5 t o 8 cm s t r e t c h e d mesh) i n w a t e r 3-^ m deepo The n e t was c h e e k e d e v e r y 15-20 m i n u n t i l one o r m o r e f i s h w e r e c a u g h t . The f i s h was t h e n q u i c k l y t r a n s f e r r e d t o t h e w e l l o n t h e p l a t f o r m . The f i s h was a l l o w e d t o swim f r e e l y a b o u t t h e w e l l f o r 30-60 m i n . The t e m p e r a t u r e o f t h e w a t e r i n t h e w e l l was w i t h i n t 1 C ° o f t h e w a t e r a t a d e p t h o f 3 m . 4 . H e w l e t t - P a c k a r d , M o s e l e y D i v . , P a s a d e n a , C a l i f . . 5. R u s t r a k I n s t r u m e n t C o . I n c . , M a n c h e s t e r , New H a m p s h i r e . 13 The f i s h was g u i d e d i n t o t h e o p e n e n d o f t h e r e s p i r a t i o n chamber w i t h o u t f u r t h e r h a n d l i n g a n d t h e c o n i c a l e n d r e p l a c e d . W i t h i n a f e w m i n u t e s o f t h e f i s h e n t e r i n g t h e chamber a d i f f e r e n c e c o u l d b e r e c o r d e d b e t w e e n t h e i n l e t a n d o u t l e t e l e c t r o d e s r e l a t e d t o t h e o x y g e n u p t a k e o f t h e f i s h . The c a n o p y o v e r t h e w e l l was c l o s e d a n d t h e e x p e r i m e n t a l l o w e d t o c o n t i n u e u n d i s t u r b e d f o r 20-24 n r . A t t h e e n d o f t h i s t i m e t h e f i s h was r e m o v e d f r o m t h e chamber a n d w e i g h e d . M o s t o f t h e f i s h w e r e t a g g e d a n d r e l e a s e d . RESULTS AND INTERPRETATION . V I T A L S T A T I S T I C S OF THE F I S H POPULATION POPULATION ESTIMATES B e c a u s e o f t h e d i f f i c u l t i e s a s s o c i a t e d w i t h a n a c c u r a t e m e a s u r e o f p o p u l a t i o n s i z e , t h e e s t i m a t e d number o f f i s h p r e s e n t i n M a r i o n L a k e h a s b e e n d e t e r m i n e d o n l y o n c e . I n 1963* a S c h n a b e l o r M u l t i p l e C e n s u s e s t i m a t e o f t h e r a i n b o w t r o u t p o p u l a t i o n was u n d e r t a k e n . B e t w e e n A p r i l 30th a n d November 27th ( i n c l u s i v e ) t h e r e w e r e 55 c a t c h d a y s d u r i n g w h i c h t i m e 536 f i s h w e r e f i n c l i p p e d o r t a g g e d a n d r e l e a s e d , . S i n c e f i s h w e r e c a u g h t b o t h b y t r a p a n d g i l l n e t i t was f e l t t h a t a l l a g e g r o u p s e x c e p t t h e y o u n g e s t (0 + ) w e r e s a m p l e d a d e q u a t e l y . A b a s i c a s s u m p t i o n o f t h e S c h n a b e l m e t h o d i s t h a t t h e p o p u l a t i o n be c o n s t a n t , w i t h no r e c r u i t m e n t a n d no m o r t a l i t y d u r i n g t h e e x p e r i m e n t a l p e r i o d . However R i c k e r (1958) i n h i s d i s c u s s i o n o f t h e m e t h o d s u g g e s t s 14 t h a t " i t i s o f t e n u s e f u l e v e n i f t h e s e c o n d i t i o n s a r e o n l y a p p r o x i m a t e l y s a t i s f i e d " . I t w o u l d a p p e a r t h a t m o s t o f t h e 1 + a n d o l d e r f i s h w e r e f u l l y v u l n e r a b l e t o t h e g e a r , s o i t may b e assumed t h a t r e c r u i t m e n t t o t h e p o r t i o n o f t h e p o p u l a t i o n b e i n g e s t i m a t e d was s m a l l . L o s s e s due t o n a t u r a l m o r t a l i t y o f b o t h m a r k e d a n d u n m a r k e d f i s h may h a v e h a d some i n f l u e n c e o n t h e f i n a l e s t i m a t e . P o p u l a t i o n s i z e was c a l c u l a t e d u s i n g t h e o r i g i n a l S c h n a b e l , e q u a t i o n ( R i c k e r , 1958 - e q u a t i o n 3*12) p l u s t h r e e m o d i f i c a t i o n s o f t h i s e q u a t i o n ( R i c k e r ' s e q u a t i o n s 3.16, 3 . 1 8 , 3.21) ( T a b l e 1 ) 8 I n a l l s u b s e q u e n t c a l c u l a t i o n s , t h e mid-summer p o p u l a t i o n s i z e o f 1 + a n d o l d e r r a i n b o w t r o u t was a s s u m e d t o b e 4 0 0 0 . U n d o u b t e d l y t h i s number v a r i e d f r o m y e a r t o y e a r a s a r e s u l t o f t h e d i f f e r e n t i a l s u c c e s s o f v a r i o u s y e a r c l a s s e s , h o w e v e r t h e f l u c t u a t i o n may h a v e b e e n s m a l l w i t h no f i s h i n g m o r t a l i t y o c c u r r i n g . M a r k a n d r e c a p t u r e e s t i m a t e s o f t h e k o k a n e e p o p u l a t i o n w e r e • c o m p l e t e l y u n s u c c e s s f u l d u e t o t h e v e r y h i g h m o r t a l i t y o f f i s h t h a t w e r e h a n d l e d , r e g a r d l e s s o f w h e t h e r o r n o t t h e y w e r e m a r k e d . A s s u m i n g e q u a l v u l n e r a b i l i t y t o t h e g e a r , t h e e s t i m a t e d number o f k o k a n e e p r e s e n t was b a s e d o n t h e r a t i o o f k o k a n e e t o r a i n b o w t r o u t i n t h e c a t c h ( e x c l u s i v e o f t h e 0 + a g e g r o u p ) . The c o m b i n e d c a t c h d a t a f o r 1963-64 y i e l d e d a r a t i o 2.12 k o k a n e e s t o e a c h r a i n b o w t r o u t c a u g h t . The r a t i o f r o m t h e 196?-68 c a t c h d a t a was 2.25 k o k a n e e s t o r a i n b o w s . I t was •; d e c i d e d a r b i t r a r i l y t h a t t h e m o r e r e c e n t d a t a w o u l d be u s e d ( b a s e d o n 15 T a b l e 1 . E s t i m a t e s o f r a i n b o w t r o u t p o p u l a t i o n s i z e ( n ) . K i n d o f e s t i m a t e ^ n 95$ c o n f i d e n c e l i m i t s O r i g i n a l S c h n a b e l (3 .12) 4084 3072-6093 M o d i f i e d S c h n a b e l (3 .21) 3977 S c h u m a c h e r ( 3 . 1 8 ) 3874 2821-6180 W e i g h t i n g f o r m u l a ( 3 . 1 6 ) 4065 1 . e q u a t i o n s g i v e b y R i c k e r ( 1 9 5 8 ) 16 700 f i s h . T h e r e f o r e , t h e mid-summer p o p u l a t i o n s i z e o f 1 a n d o l d e r k o k a n e e was assumed t o b e 4000 x 2.25 = 9000. AGE D I S T R I B U T I O N F o r c o n v e n i e n c e , t h e c h a n g e - o v e r i n a g e o f f i s h e s f r o m M a r i o n L a k e i s t a k e n t o b e t h e e n d o f t h e c a l e n d a r y e a r . Thus a f i s h becomes a g e 1 o n J a n u a r y 1st o f t h e y e a r f o l l o w i n g h a t c h i n g ( R i c k e r , 1958). I n t h e r a i n b o w t r o u t p o p u l a t i o n , r e c r u i t m e n t o c c u r s a b o u t J u n e 15th. The l i f e s p a n o f t h e s e f i s h i n M a r i o n L a k e u s u a l l y t e r m i n a t e s m i d - w a y t h r o u g h t h e i r f i f t h y e a r o f l i f e . K o k a n e e r e c r u i t m e n t p r o b a b l y t a k e s p l a c e a b o u t M a y 1st w i t h a maximum l i f e s p a n o f f o u r y e a r s . AGE STRUCTURE R o u t i n e c a t c h c u r v e s w h e r e l o g o f t h e number o f f i s h i s p l o t t e d a g a i n s t l e n g t h o r a g e i n t e r v a l s ( R i c k e r , 19^8) a r e d i f f i c u l t t o i n t e r p r e t when t h e l i f e s p a n o f t h e f i s h i s f e w e r t h a n 5 y e a r s . I n o r d e r t o e s t i m a t e t h e number o f 0 + a n d 1 + r a i n b o w t r o u t p r e s e n t i n t h e l a k e , t h e s e a g e g r o u p s w e r e assumed t o h a v e t h e same m o r t a l i t y r a t e as t h e 2 + g r o u p * t h e y o u n g e s t f o r w h i c h t h e d a t a i s r e l i a b l e ( F i g . 3). I t was f u r t h e r assumed t h a t t h e m o r t a l i t y r a t e s i n y o u n g k o k a n e e w e r e t h e same as t h o s e o f t h e t r o u t f r o m r e c r u i t m e n t t o a g e 2 +. + I n c o r p o r a t i o n o f t h e p o p u l a t i o n e s t i m a t e s o f t h e 1 a n d o l d e r a g e g r o u p s w i t h t h e p r o p o s e d a g e s t r u c t u r e p r o v i d e d a n e s t i m a t e o f ;. t h e numbers o f f i s h o f e a c h a g e p r e s e n t i n t h e l a k e o n J a n u a r y 1st ( T a b l e 2). F i g u r e 3. R e l a t i v e numbers o f f i s h i n e a c h age g r o u p b a s e d o n " c a t c h c u r v e " . T a b l e 2 . E s t i m a t e d number o f f i s h p r e s e n t i n M a r i o n L a k e a t t h e b e g i n n i n g o f e a c h a g e a n d a n n u a l i n s t a n t a n e o u s m o r t a l i t y r a t e ( i ) e R a i n b o w t r o u t K o k a n e e Age Number o n i J a n u a r y 1st a 0+ 0 ( r e c r u i t s 2350 .43 o n J u n 15th) l + a 1848 .43 2+ 1202 .43 3+ 782 1.53 4+ 169 5.13 A g e a 0+ 1 + 3 2+ 3+ Number o n i J a n u a r y 1st 0 ( r e c r u i t s 6856 .43 o n M a y 1st) 5130 .43 3337 lo84 530 6.27 Age g r o u p n o t f u l l y v u l n e r a b l e t o gear. 19 MORTALITY The t o t a l i n s t a n t a n e o u s m o r t a l i t y r a t e ( i ) i s e q u a l t o t h e sum o f t h e m o r t a l i t y due t o f i s h i n g (p_) p l u s t h e m o r t a l i t y f r o m n a t u r a l c a u s e s (c|) ( R i c k e r , 1958) o F i s h i n g m o r t a l i t y , a p a r t f r o m t h o s e f i s h e s c o l l e c t e d d u r i n g t h i s s t u d y , h a s b e e n i n s i g n i f i c a n t i n M a r i o n L a k e p r i m a r i l y d u e t o t h e s m a l l s i z e o f t h e f i s h . T h e r e f o r e a l l d e a t h s h a v e b e e n a t t r i b u t e d t o n a t u r a l m o r t a l i t y . The a b s e n c e o f m a j o r p r e d a t o r s , w i t h t h e e x c e p t i o n o f l a r g e r a i n b o w t r o u t o c c a s i o n a l l y c o n s u m i n g s m a l l k o k a n e e , s u g g e s t s t h a t n a t u r a l m o r t a l i t y i n M a r i o n L a k e f i s h m i g h t r e a s o n a b l y be d i v i d e d i n t o t w o c a t e g o r i e s , ( 1 ) d e a t h s a s s o c i a t e d w i t h s p a w n i n g a n d p o s t - s p a w n i n g c o n d i t i o n ( q ] ) , a n d ( 2 ) d e a t h s a t t r i b u t a b l e t o a l l o t h e r c a u s e s ; d i s e a s e , i n j u r y , t h e r m a l s t r e s s , p l u s s m a l l l o s s e s t o p r e d a t o r s s u c h as f i s h , m e r g a n s e r s a n d p o s s i b l y m i n k ( q 2 ) » I n i m m a t u r e f i s h i e q u a l s q 2 ( e q u a l s 0 o 4 , s e e T a b l e 2 ) . The o n s e t o f m a t u r i t y i s a s s o c i a t e d w i t h a n i n c r e a s e i n m o r t a l i t y r a t e a n d q-^ becomes g r e a t e r t h a n q 2 . F o r s i m p l i f i c a t i o n i t h a s b e e n , a s s u m e d t h a t t h e v a l u e f o r f o r m a t u r e r a i n b o w t r o u t a n d k o k a n e e i s s i m i l a r t o t h a t o f t h e i m m a t u r e f i s h a n d t h a t i _ OA = q i . I n t h e c a l c u l a t i o n o f m o n t h l y p r o d u c t i o n v a l u e s s p a w n i n g a n d p o s t - s p a w n i n g m o r t a l i t y r a t e s (q^) h a v e b e e n a p p l i e d t o t h e m o n t h i n w h i c h s p a w n i n g o c c u r s a n d t h e t w o months f o l l o w i n g ; f o r r a i n b o w t r o u t t h e months o f A p r i l , May a n d J u n e , a n d f o r t h e k o k a n e e , O c t o b e r , 20 November and December. Natural mortality from other causes (q 2) has been applied evenly over the f u l l 12 months for a l l age groups. In calculating monthly rates, adjustments were made to compensate for differing number of days. GROWTH Length of growing season — During 1963 and 1964 a total of 66 tagged rainbow trout were recaptured in Marion Lake. These fish had been free in the lake for intervals of 7 to 180 days (Age 1 + % A, mid-point of time at large June 11th, mean number of days 30•8 3.I; Age 2 + : mid-point June 15th, mean number of days 46.7 - 9.9). Five fish tagged in I963 and recaptured in 1964 were excluded from the following calculations. On the basis of size at tagging, 44 trout were considered to be 1+ while the remaining were probably 2 + . If the population^ annual instantaneous growth (in length) rate, (gy)» for the 1 and 2 age groups is divided by the daily instantaneous growth, (g^)* of the recaptured fish, the result should be an estimate of the number of days in the growing season (Table 3 ) . This rather crude estimate would be affected by any temporary decline in growth rate as a result of the tagging (Ricker, 1956) and would result in an overestimate of the length of the growing season. Kohler (I963) demonstrated however that the presence or absence of a tag in cod bore l i t t l e relation to growth rate. The number of growing days in Marion Lake fish as calculated 21 T a b l e 3» L e n g t h o f g r o w i n g s e a s o n c a l c u l a t e d f r o m d a i l y i n s t a n t a n e o u s g r o w t h r a t e s o f m a r k e d a n d r e c a p t u r e d r a i n b o w t r o u t . A g e A n n u a l i n s t . D a i l y i n s t . Number o f g " y «= d a y s g r o w t h r a t e g r o w t h r a t e r e c a p t u r e s g^ f y § d 1 + 0.3249 0.002?29 44 119 2 + 0.1769 0.001678 22 105 22 ( T a b l e 3) w o u l d a p p e a r t o b e a n u n d e r e s t i m a t e . I f t h e g r o w t h c h a r a c t e r i s t i c s o f t h e s e f i s h w e r e h i g h e s t i n J u n e a n d g r a d u a l l y -d e c l i n e d t o w a r d t h e e n d o f t h e s e a s o n , t h e n t h e f a c t t h a t t h e t a g g e d f i s h w e r e f r e e i n t h e l a k e o n l y 4-6 weeks a n d t h a t t h i s i n t e r v a l was c e n t r e d a b o u t t h e m i d p a r t o f J u n e , w o u l d c o m b i n e t o p r o d u c e a n u n d e r e s t i m a t e o f t h e a c t u a l g r o w i n g p e r i o d . The l a c k o f d a t a o n t h e r a t e o f g r o w t h w i t h i n a g i v e n g r o w i n g s e a s o n p r e c l u d e s t h e a p p l i c a t i o n o f d i f f e r e n t s e a s o n a l i n s t a n t a n e o u s g r o w t h r a t e s . A s a r e s u l t , the f o l l o w i n g s i m p l i f y i n g a s s u m p t i o n s h a v e b e e n made: t h e g r o w i n g s e a s o n i n M a r i o n L a k e f i s h e s e x t e n d s f r o m M a y 1st t o A u g u s t 31st (123 d a y s ) a n d t h e g r o w t h i s u n i f o r m d u r i n g t h i s p e r i o d . L e n g t h - w e i g h t r e l a t i o n s h i p — A n a l y s i s o f t h e l e n g t h - w e i g h t r e l a t i o n s h i p b y s p e c i e s , b y s e x , b y s e a s o n a n d b y y e a r d i d n o t y i e l d a n y s i g n i f i c a n t d i f f e r e n c e s w h e r e s a m p l e s i z e was a d e q u a t e , e x c e p t f o r b e t w e e n s p e c i e s . T h u s , a l l l e n g t h - w e i g h t d a t a w e r e c o m b i n e d i n t h e c a l c u l a t i o n o f t h e e q u a t i o n s f o r e a c h s p e c i e s : R a i n b o w t r o u t W = 0.0109 L 3.016 K o k a n e e W = 0.0058 L' 3.232 f o r w e i g h t (W) i n g a n d l e n g t h ( L ) i n cm. G r o w t h r a t e — R a t e s o f g r o w t h i n b o t h l e n g t h a n d w e i g h t w e r e d e t e r m i n e d b y p l o t t i n g t h e s e d e p e n d e n t v a r i a b l e s a g a i n s t a g e a t t i m e 23 o f c a p t u r e . F o r e a c h a g e g r o u p t h e mean l e n g t h o r w e i g h t was p l o t t e d a g a i n s t t h e mean d a t e o f c a p t u r e . The g r a p h i c r e p r e s e n t a t i o n ( F i g s . 4 & 5) o f g r o w t h i n t h e s e f i s h e s was a c h i e v e d b y d r a w i n g a s t r a i g h t l i n e t h r o u g h t h e p o i n t s ( l e n g t h o r w e i g h t a t a g i v e n a g e ) a n d w h i c h p a s s e d t h r o u g h t h e mean o f a l l t h e v a l u e s f o r t h a t a g e . A n n u a l i n s t a n t a n e o u s r a t e s o f g r o w t h w e r e c a l c u l a t e d b y s u b t r a c t i n g t h e l o g ( s i z e ) a t t h e e n d o f e a c h a g e f r o m t h e l o g ( s i z e ) a t t h e b e g i n n i n g . E s t i m a t e s o f t h e g r o w t h r a t e s i n t h e 0 + age g r o u p w e r e made b y a s s u m i n g t h e s i z e a t r e c r u i t m e n t , f o r b o t h r a i n b o w t r o u t a n d k o k a n e e , t o b e 30 ram f o r k l e n g t h a n d 0.3 g ( b a s e d o n d a t a f r o m F r a s e r , 1969). As d i s c u s s e d e a r l i e r , t h e a n n u a l i n s t a n t a n e o u s r a t e s o f g r o w t h ( T a b l e 4) w e r e a p p l i e d u n i f o r m l y o v e r a 123 d a y p e r i o d commencing M a y 1st ( J u n e 15th f o r 0 + r a i n b o w t r o u t ) i n c a l c u l a t i n g p r o d u c t i o n v a l u e s . REPRODUCTION S e x r a t i o — The s e x r a t i o o f r a i n b o w t r o u t i s s i g n i f i c a n t l y d i f f e r e n t f r o m 1:1 a t t h e 95$ l e v e l when d a t a f o r a l l y e a r s (1963-1964, 1966-1968) a r e c o m b i n e d , w i t h a p r e p o n d e r a n c e o f f e m a l e s (57*7$). A f u r t h e r t e s t o f h o m o g e n e i t y u s i n g a d j u s t e d c h i s q u a r e v a l u e s ( S n e d e c o r , 1956) i n d i c a t e d t h a t t h e s e x r a t i o i s s i m i l a r l y l o p s i d e d f r o m y e a r t o y e a r . The s e x r a t i o i n t h e k o k a n e e , c o m b i n i n g d a t a f o r a l l y e a r s i s n o t s i g n i f i c a n t l y d i f f e r e n t f r o m 1:1 a t t h e 95$ l e v e l , h o w e v e r a t e s t F i g u r e 4 . Change i n l e n g t h (mm) b e t w e e n e a c h a g e g r o u p . F i g u r e 5» Change i n w e i g h t ( l o g ) b e t w e e n e a c h age g r o u p . T a b l e 4 . A v e r a g e l e n g t h ( L ) a n d w e i g h t (W) a t t h e b e g i n n i n g o f e a c h a g e ( J a n u a r y 1 s t ) a n d a n n u a l i n s t a n t a n e o u s r a t e s o f g r o w t h ( g ^ a n d g ^ ) . R a i n b o w t r o u t Age L (mm) 0 + ( J u n e 30 15th) 1.317 W ( g ) 0.3 w 112 17.8 0.325 0o890 155 43.4 0.177 0.600 185 79.0 0.301 0.910 250 196.4 Age L (mm) K o k a n e e L g 0 + (May 30 1 s t ) 4.08 1.308 W ( g ) 0.3 w 3.95 111 15 .6 0 .218 0.570 138 27.7 0.173 0.570 164 4 8 . 9 0 . 147 0.580 190 87.4 to 26 o f h o m o g e n e i t y i n d i c a t e s t h a t t h e r e may b e y e a r t o y e a r d i f f e r e n c e s i n t h e s e x r a t i o w i t h one o r t h e o t h e r o f t h e s e x e s p r e d o m i n a t i n g . S e a s o n a l c h a n g e i n g o n a d w e i g h t f o r r a i n b o w t r o u t «•« The g o n a d w e i g h t r e a c h e s i t s l o w e s t l e v e l i n m a t u r e r a i n b o w t r o u t i m m e d i a t e l y a f t e r s p a w n i n g ( a b o u t A p r i l 15th) when i t s w e i g h t i s l e s s t h a n 0.5$ o f t h e b o d y w e i g h t ( F i g s . 6 & 7, T a b l e 5 ) . The m a t u r e f i s h ( t h o s e h a v i n g spawned p r e v i o u s l y ) a n d m a t u r i n g i n d i v i d u a l s b e g i n i n c r e a s i n g t h e i r g o n a d w e i g h t , a n d b y l a t e December t h e a v e r a g e w e i g h t o f t h e , o v a r i e s i n t h e f e m a l e i s a b o u t 10$ o f t h e b o d y w e i g h t . Maximum o v a r y w e i g h t p r i o r t o s p a w n i n g i s e s t i m a t e d t o be a b o u t 13$ o f t h e b o d y w e i g h t . The t e s t e s w e i g h t i n t h e m a l e t r o u t i n c r e a s e s t o a maximum v a l u e o f a l m o s t 3$ b y O c t o b e r . T h e r e i s t h e n a d e c l i n e i n w e i g h t f r o m t h i s p e a k l e v e l t o j u s t u n d e r 2$ o f t h e b o d y w e i g h t , a n d i t a p p e a r s t o r e m a i n a t t h i s l e v e l u n t i l s p a w n i n g b e g i n s . Gonad w e i g h t i n i m m a t u r e r a i n b o w t r o u t i s l e s s t h a n 0.5$ o f b o d y w e i g h t i n a l l months ( T a b l e 5 ) . S e a s o n a l c h a n g e i n g o n a d w e i g h t f o r k o k a n e e — The o v a r y w e i g h t i n f e m a l e k o k a n e e i n c r e a s e s f r o m a b o u t 1$ o f b o d y w e i g h t i n J u n e t o a maximum o f more t h a n 13$ i m m e d i a t e l y p r i o r t o s p a w n i n g ( F i g . 7» T a b l e 5 ) . A f t e r s p a w n i n g t h e g o n a d ( o v a r y w i t h o u t e g g s ) c o n s t i t u t e s l e s s t h a n 0.5$ o f t h e b o d y w e i g h t a n d r e m a i n s a t a b o u t t h a t l e v e l u n t i l d e a t h . K o k a n e e t e s t e s a r e s t i l l l e s s t h a n 1$ o f b o d y w e i g h t i n J u n e F i g u r e 6 . S e a s o n a l c h a n g e i n gonad w e i g h t as a p e r c e n t a g e o f body-w e i g h t i n r a i n b o w t r o u t . </) > m PERCENT OF BODY WEIGHT ro O) 00 ro Q.IO 0)|<3) m CP 2 o | o > > >|ro u z ol3 $ Wioi S= ^Icji c ro|<o c col*, c o m oo|w o o>l5> O < N>|W m \ MAI m m Z > m Lz F i g u r e 7* S e a s o n a l c h a n g e i n g o n a d w e i g h t a s a p e r c e n t a g e o f body-w e i g h t i n k o k a n e e . (/) > m TJ i— m PERCENT OF BODY WEIGHT > "n u>|o m 03 70 N)|N> J= oo ro c oo|ui m ~0 . 2 Ui|cD O < o w m ' o 82 29 Table 5» Gonad weight as a percentage of body weight (mean - standard deviation)• Rainbow trout (mature) Month Female Male Jan. 10.9 + 2.1 a (4) 1.90 .14 (2) Feb. 12.3 + 1.6 (6) 1.82 + .3^ (6) Mar. (0) (0) Apr. 0.36 + .07 (2) 0.15 (1) May 0.54 .38 (17) 0.44 + .38 (10) June 0.40 + .14 (55) 0.25 + .15 (34) July 0.38 + .17 (9) 0.14 + .05 (2) Aug. I.05 + .42 (1*0 0.67 + .58 (9) Sep. 1.13 (1) 3.05 + .35 (3) Oct. 4.72 + mm 3.14 (12) 2.82 + 1.59 (8) Nov. 7.60 + mm 1.48 (16) 1.83 + .65 (6) Dec. 8.26 + 1.53 (3) 1.69 .04 (2) Immature A l l months 0.25 - 0.14 (56) 0.10 ± .10 (55) Number in brackets indicates sample size. 30 T a b l e 5« ( c o n t i n u e d . ) Gonad w e i g h t a s a p e r c e n t a g e o f b o d y w e i g h t (mean - s t a n d a r d d e v i a t i o n ) . Kokanee (mature) Month Female Male Jan. 0.77 + .21 ( 3 ) a 0.05 - o02 (3) Feb. (o) 0.09 i .02 (5) Mar. 0.61 + .12 (4) (o) Apr. 0.82 + .11 (16) 0.12 t .10 (24) May 0.69 .16 (11) o . i i i .01 (4) June 0.93 + .19 (22) 0.35 ± .11 (20) July 3.^3 + 1.60 (2) 1.64 i .83 (8) Aug. 5.73 + 1.51 (46) 3.07 - 1.02 (32) Sep. 11.2 + 2.4 (5) 3.90 ± .52 (8) Oct. 13.2 + 2.1 (27) 1.99 - .90 (20) Nov. 0.46 i .11 (9) 0.06 - .03 (5) Dec. 0.53 ± .14 (3) (o) Immature months 0.47 + .14 (44) 0.14 t .10 (30) aNumber in brackets indicates sample size. 31 b u t c o n t i n u e i n c r e a s i n g i n w e i g h t up t o a maximum o f a b o u t 4 $ by-S e p t e m b e r . F o l l o w i n g s p a w n i n g t h e t e s t e s w e i g h t i s l e s s t h a n 0.1$ u n t i l d e a t h . I m m a t u r e k o k a n e e i n a l l y e a r s h a v e a g o n a d w e i g h t o f l e s s t h a n 0.5$ i n a l l m o n t h s . A g e a t m a t u r i t y —- M a t u r i t y i n M a r i o n L a k e f i s h i s n o t r e a c h e d u n t i l t h e i r t h i r d y e a r o f l i f e ( T a b l e 6 ) . I n t h e r a i n b o w t r o u t o n l y t h e m a l e s a r e m a t u r e a t age Z+ w i t h none o f t h e f e m a l e s r e a c h i n g m a t u r i t y + + u n t i l 3 • I n t h e k o k a n e e s , some f i s h o f b o t h s e x e s m a t u r e b y age 2 w i t h a h i g h e r p e r c e n t a g e o f m a t u r e f e m a l e s t h a n m a l e s . T h i s i n e q u a l i t y i s p r o b a b l y a r e s u l t o f t h e d i f f i c u l t y i n j u d g i n g c o n c l u s i v e l y w h e t h e r o r n o t a m a l e i s i n t h e p r o c e s s o f m a t u r i n g , w h e r e a s a m a t u r i n g f e m a l e i s much more r e a d i l y a p p a r e n t . I n t h e o l d e s t age g r o u p s ( r a i n b o w 4 a n d k o k a n e e 3 ) a l l t h e f i s h e x a m i n e d w e r e m a t u r e . >. F e c u n d i t y ««-A l i n e a r r e g r e s s i o n , b a s e d o n l e a s t s q u a r e s m e t h o d , o f t o t a l egg number p l o t t e d a g a i n s t f o r k l e n g t h o n a l o g - l o g p l o t ( F i g . 8) y i e l d s t h e f o l l o w i n g e q u a t i o n s : R a i n b o w t r o u t F = 18<>7 L * 9 5 9 1 .414 K o k a n e e F = 2 . 8 4 L w h e r e F = t o t a l number o f eggs p e r f e m a l e , a n d L = f o r k l e n g t h i n c m . S i m i l a r r e l a t i o n s h i p s h a v e b e e n d e m o n s t r a t e d f o r a number o f s a l m o n i d s p e c i e s ( R i c k e r , 1932; R o u n s e f e l l , 1957). The maximum s i z e r a n g e o f f e m a l e f i s h , f o r w h i c h egg c o u n t s 32 T a b l e 6. P e r c e n t a g e o f f i s h r e a c h i n g m a t u r i t y a t e a c h a g e . Age a t s p a w n i n g t i m e R a i n b o w t r o u t N o . e x a m i n e d % m a t u r e 39 30 f e m a l e s 30 m a l e s 107 f e m a l e s 76 m a l e s 63 f e m a l e s 29 m a l e s 0 0 36.7 72.9 44.7 100 100 K o k a n e e 1 + 22 0 2 + 144 f e m a l e s 72 .2 129 m a l e s 62.0 3 + 21 f e m a l e s 100 19 m a l e s 100 F i g u r e 8 . T o t a l number o f e g g s / f e m a l e ( l o g Q ) a g a i n s t f o r k l e n g t h ( l o g , c m ) . 500 FORK LENGTH (cm) 34 w e r e a v a i l a b l e , was i n t h e r a i n b o w t r o u t 16.6-26.7 cm ( c a l c u l a t e d egg number 277-436) a n d i n t h e k o k a n e e 12.2-19.1 cm ( c a l c u l a t e d egg number 98 -184) . One p o s s i b l e s o u r c e o f e r r o r i n t h e egg c o u n t s i s t h e g r a d u a l l o s s o f m a t u r i n g o v a d u r i n g d e v e l o p m e n t , w h i c h w o u l d t e n d t o c a u s e a n o v e r e s t i m a t e o f t h e number o f eggs a v a i l a b l e a t s p a w n i n g t i m e ( S c o t t , 1962; H e n d e r s o n , I963). T h i s e r r o r , i f r e a l , w o u l d h a v e a g r e a t e r e f f e c t o n t h e r a i n b o w d a t a as egg c o u n t s w e r e made o v e r a n 8 month p e r i o d p r i o r t o s p a w n i n g , i n c o n t r a s t t o t h e k o k a n e e f o r w h i c h c o u n t s w e r e p o s s i b l e f o r o n l y 5 m o n t h s . However i n b o t h s p e c i e s t h e egg s i z e i n a n y g i v e n o v a r y was u n i f o r m , s u g g e s t i n g t h a t t h e amount o f egg r e s o r p t i o n b e f o r e s p a w n i n g was p r o b a b l y s m a l l . S e a s o n a l c h a n g e s i n egg d i a m e t e r s a r e g i v e n i n T a b l e 7» The number o f eggs r e t a i n e d b y t h e f e m a l e i n t h e b o d y c a v i t y a f t e r s p a w n i n g was p r o b a b l y l e s s t h a n 1$ o f t h e o r i g i n a l number b a s e d o n f o u r s p e n t f e m a l e s (2 o f e a c h s p e c i e s ) . FOOD ANALYSIS OF F I S H STOMACHS The s e a s o n a l c h a n g e i n a v e r a g e numbers o f f o o d o r g a n i s m s p e r s t o m a c h i s g i v e n i n T a b l e 8. The p e r c e n t v o l u m e o f e a c h m a j o r f o o d i t e m h a s b e e n r a n k e d i n o r d e r o f d e c r e a s i n g v o l u m e (#) f o r t h e r a i n b o w t r o u t a n d b y i n c r e a s i n g v o l u m e f o r k o k a n e e f o r a l l f o o d i t e m s n o t s h a r e d ( F i g . 9 ) . The p e r c e n t v o l u m e o f t h o s e f o o d i t e m s e a t e n b y b o t h s p e c i e s h a v e b e e n p l o t t e d w i t h t h e v a l u e s f o r t h e k o k a n e e b e i n g 35 T a b l e 7* S e a s o n a l c h a n g e i n a v e r a g e egg d i a m e t e r (number i n b r a c k e t s i n d i c a t e s number o f f e m a l e s e x a m i n e d , egg d i a m e t e r i s b a s e d o n 60 e g g s / f e m a l e ) • M o n t h R a i n b o w egg K o k a n e e egg d i a m e t e r d i a m e t e r (mm) (mm) May - 1.44 ( 2 ) J u n e - 1.67 (3) J u l y - 2 . 1 8 (3). A u g . 1.53 (2) 2.61 (4) S e p . - 3.51 (5) O c t . 2.57 (6) 4 . 18 (32) N o v . 3.09 (4) D e c . 3.07 (4) J a n . F e b . M a r . 4.26 (1) A p r . 4.95 (2) 36 T a b l e 8. A v e r a g e number o f f o o d o r g a n i s m s p e r s t o m a c h (number o f f i s h e x a m i n e d i n b r a c k e t s ) . M o n t h R a i n b o w K o k a n e e t r o u t J a n . mm Oft F e b . 18 (11) 40 ( 1 7 ) M a r . - -A p r . 15 ( 14) 39 (7) May 97 (13) 239 ( 1 4 ) J u n e 2 4 (12) 90 (9) J u l y 22 (12) 117 (12) A u g . 23 (10) 19 (12) S e p . 86 ( 6 ) 22 (13) O c t . 14 (9) 35 ( 8 ) N o v . 4 ( 6 ) 11 (9) D e c . 9 (5) 6 ( 6 ) M e a n 32 (98) 70 (107) F i g u r e 9» C o m p a r i s o n o f t h e p e r c e n t a g e v o l u m e o f v a r i o u s f o o d i t e m s consumed b y r a i n b o w t r o u t a n d k o k a n e e , w h e r e ( L ) «= l a r v a e , ( P ) = p u p a e , (N) = n a i a d s , M I S C . *= m i s c e l l a n e o u s a n d TERR. = t e r r e s t r i a l . 37 F E B JUL OCT TRICHOPTERA MOLLUSCA MISC. AQUATIC INSECTS EPHEMEROPTERA SIMULIUM AMPHIPODA PLECOPTERA CHIRONOMID (L) AMPHIPODA MISC. AQUATIC INSECTS PLANORBIDS TRICHOPTERA EPHEMEROPTERA ANISOPTERA (N) ZYGOPTERA (N) CHIRONOMID (L) CHIRONOMID (P) CHIRONOMID (P) ANTS LEECHES AMBYSTOMA TRICHOPTERA MISC. AQUATIC INSECTS CRANGONYX ANTS TRICHOPTERA ZYGOPTERA (N) PLANORBIDS MISC. AQUATIC INSECTS ANISOPTERA (N) CRANGONYX APR CLADOCERA PISIDIUM CHIRONOMID (P) HYALELLA TRICHOPTERA PLANORBIDS CRANGONYX H ANISOPTERA (N) MISC. AQUATIC INSECTS ZYGOPTERA (N) MISC. TERR. INSECTS CHIRONOMID (P) NOV J 1 1 PLANORBIDS J ANISOPTERA (N) MISC. AQUATIC INSECTS TRICHOPTERA HYALELLA AUG CHIRONOMID (L) CRANGONYX MAY PISIDIUM SEP TRICHOPTERA AMBYSTOMA NOTONECTIDS ZYGOPTERA (N) MISC-AQUATIC INSECTS CRANGONYX  HYALELLA CHIRONOMID (P) MISC. TERR. INSECTS TRICHOPTERA PSEUDOSUCCINEA DEC J CRANGONYX TIPULID (L) ANISOPTERA (N) ~l MISC. AQUATIC INSECTS ZYGOPTERA (N) M I S C TERR.INSECTS JUN TRICHOPTERA MISC. TERR- INSECTS LEECHES ZYGOPTERA (N) CHIRONOMID (P) CRANGONYX MISC. AQUATIC INSECTS CERATOPOGONID (L) CHIRONOMID (P) AMBYSTOMA ZYGOPTERA (N) MISC. AQUATIC INSECTS SAMPLE SIZE J F M A M J J A S O N D RAINBOW TROUT Q - 11 - 14 13 12 12 10 6 9 6 5 KOKANEE K | - 17 - 7 14 9 12 12 13 8 9 6 38 s u p e r i m p o s e d o n t h o s e o f t h e r a i n b o w t r o u t ( F i g . 9)« A l l f o o d i t e m s w i t h a p e r c e n t v o l u m e l e s s t h a n 5$ h a v e b e e n o m i t t e d . I n a l l months f o r w h i c h t h e r e a r e d a t a , some p o r t i o n o f t h e f o o d r e s o u r c e i s s h a r e d b y b o t h f i s h s p e c i e s e x c e p t i n D e c e m b e r . However o n l y i n May i s t h e f o o d i t e m o f h i g h e s t # v o l u m e t h e same i n b o t h s p e c i e s ( c h i r o n o m i d p u p a e ) . The d i f f e r e n t u t i l i z a t i o n o f t h e f o o d r e s o u r c e b y t h e r a i n b o w t r o u t a n d k o k a n e e i s v e r y s u g g e s t i v e o f i n t e r a c t i v e s e g r e g a t i o n as d e s c r i b e d b y N i l s s o n (I965). The c o n s u m p t i o n o f t h e s a l a m a n d e r , Ambystoma g r a c i l e , b y t h e r a i n b o w t r o u t i s o f p a r t i c u l a r i n t e r e s t . I n t h e months o f M a y , November a n d December t h i s f o o d i t e m e x c e e d s 5$ o f t h e v o l u m e e a t e n . Of a l l t h e s t o m a c h s e x a m i n e d f r o m M a r i o n L a k e o n l y 10 ( a l l r a i n b o w t r o u t ) c o n t a i n e d v e r t e b r a t e r e m a i n s . N i n e o f t h e s e c o n t a i n e d A m b y s t o m a , one c o n t a i n e d f i s h r e m a i n s ( p r o b a b l y k o k a n e e ) , a n d one c o n t a i n e d b o t h Ambystoma a n d f i s h . T h o s e Ambystoma w h i c h w e r e u n d i g e s t e d r a n g e d i n s i z e f r o m 7-7 t o 10 cm ( t o t a l l e n g t h ) a n d t h e f i s h w h i c h w e r e consumed w e r e a p p r o x i m a t e l y 9 cm ( f o r k l e n g t h ) . The s m a l l e s t f i s h known t o h a v e consumed v e r t e b r a t e m a t e r i a l m e a s u r e d 19«3 cm. T h e r e a p p e a r s t o b e a " t h r e s h o l d s i z e " a b o v e w h i c h f i s h commence f e e d i n g o n o t h e r v e r t e b r a t e s . F o r e x a m p l e , Clemens (1951) f o u n d t h a t K a m l o o p s t r o u t b e g a n f e e d i n g o n s m a l l f i s h o n c e t h e y r e a c h e d a l e n g t h o f 8 i n c h e s (20.3 c m ) , N i l s s o n (19&5) f o u n d t h a t c h a r o v e r 20.1 cm consumed f i s h i n S w e d i s h l a k e s , a n d L a r k i n e t a l (1957) o b s e r v e d t h a t r a i n b o w t r o u t 20-30 cm w e r e e a t i n g f i s h when a v a i l a b l e . A l t h o u g h t h i s t h r e s h o l d v a l u e i s p r o b a b l y c e n t r e d a b o u t 20 c m , i t may c o v e r a 39 b r o a d r a n g e o f s i z e s i n v a r i o u s p o p u l a t i o n s d e p e n d i n g o n b e h a v i o u r a l d i f f e r e n c e s ( L a r k i n e t a l , 1957). I t i s s i g n i f i c a n t h o w e v e r t h a t t h e r e i s a m a r k e d i n c r e a s e i n t h e i n s t a n t a n e o u s g r o w t h r a t e i n t h e r a i n b o w t r o u t o f t h e 3 + age g r o u p i n M a r i o n L a k e , w h e r e t h e l e n g t h r e a c h e d a t t h e b e g i n n i n g o f t h i s a g e , i s a n a v e r a g e 18.5 cm ( F i g . 4 ) . L a r k i n e t a l (1957) o b s e r v e d a s i m i l a r i n c r e a s e i n g r o w t h r a t e s f o r b o t h P a u l L a k e a n d K o o t e n a y L a k e r a i n b o w s ( o v e r 20 c m ) , b u t i t was n o t e d t h a t t h e r e i s no s h a r p g r o w t h i n f l e c t i o n a t t h e t h r e s h o l d s i z e s i n c e t h e g r o w t h r a t e s a r e c a l c u l a t e d o n a n ,: a n n u a l b a s i s a n d t h e c h a n g e o v e r i n d i e t a n d g r o w t h may o c c u r p a r t way t h r o u g h a s e a s o n . ENERGY CONTENT OF THE FOOD I n o r d e r t o d e t e r m i n e t h e s e a s o n a l c h a n g e i n t h e e n e r g y c o n t e n t o f t h e f o o d , t h e % v o l u m e o f e a c h i t e m o c c u r r i n g i n t h e s t o m a c h s was e q u a t e d t o t h e same number o f grams (wet w e i g h t ) . F o r e x a m p l e , c h i r o n o m i d p u p a e c o m p r i s e d 25$ o f t h e v o l u m e o f f o o d consumed b y k o k a n e e d u r i n g t h e month o f J u n e . T h u s , f o r e v e r y 100 g ( w e t ) o f f o o d e a t e n , 25 g c o n s i s t e d o f c h i r o n o m i d p u p a e . The r e l a t i o n s h i p b e t w e e n v o l u m e a n d w e i g h t w i l l b e d i s c u s s e d l a t e r . The grams w e t w e i g h t o f f o o d w e r e c o n v e r t e d t o grams d r y w e i g h t a n d m u l t i p l i e d b y t h e e n e r g y c o n t e n t ( K c a l o r i e s / g d r y w e i g h t ) o f t h a t f o o d i t e m . The c r u d e e s t i m a t e s o f t h e e n e r g y c o n t e n t o f t h e v a r i o u s f o o d i t e m s consumed b y M a r i o n L a k e f i s h a r e b a s e d o n a v e r a g e s o f c a l o r i f i c v a l u e s p u b l i s h e d b y Geng ( i n W u n d e r , 1936), R i c h m a n (1958), Cummins ( M S , 1967), M a t h i a s ( M S , 1967), S w i f t ( M S , 1968), W i s s i n g a n d H a s l e r (1968), a n d H a r g r a v e ( p e r s . c o m m . ) , p l u s bomb c a l o r i f i c d e t e r m i n a t i o n s made i n t h e p r e s e n t s t u d y ( f o r C l a d o c e r a , T r i c h o p t e r a ) ( T a b l e 9) 0 F o r a n y g i v e n m o n t h , t h e sum o f a l l t h e e n e r g y v a l u e s r e p r e s e n t s t h e number o f K c a l o r i e s p e r 100 g w e t ( T a b l e 10). The m o n t h l y c h a n g e i n a v e r a g e e n e r g y c o n t e n t o f t h e f o o d ( F i g . 10) c o n t a i n s t h r e e d i s t i n c t f e a t u r e s . F i r s t , t h e r e a r e t w o p e r i o d s when t h e e n e r g y c o n t e n t o f t h e f o o d i s g r e a t e s t , i n J u n e a n d d u r i n g t h e w i n t e r m o n t h s . S e c o n d , a t no t i m e d u r i n g t h e y e a r i s t h e e n e r g y c o n t e n t o f t h e k o k a n e e f o o d as g r e a t a s t h a t o f t h e r a i n b o w t r o u t . T h i r d , t h e r a i n b o w t r o u t a n d k o k a n e e b o t h e x h i b i t t h e same s e a s o n a l p a t t e r n w i t h t h e e x c e p t i o n o f some s m a l l v a r i a t i o n i n J u l y a n d A u g u s t . On t h e a v e r a g e , i n s e c t s , w h i c h make up t h e b u l k o f t h e t r o u t f o o d , a r e h i g h e r i n e n e r g y c o n t e n t t h a n t h e o t h e r m a j o r f o o d i t e m s , t h e m o l l u s c s a n d t h e c r u s t a c e a n s . B a s i c a l l y i t i s t h i s r e l a t i o n s h i p w h i c h a c c o u n t s f o r t h e o b s e r v e d s e a s o n a l d i f f e r e n c e s i n e n e r g y c o n t e n t o f t h e f o o d . The p e a k v a l u e s a r e r e a c h e d a t a t i m e when t h e f i s h a r e c o n s u m i n g t h e m o s t i n s e c t m a t e r i a l . L i k e w i s e t h e d i f f e r e n c e s b e t w e e n t h e r a i n b o w t r o u t a n d k o k a n e e c a n b e a c c o u n t e d f o r i n t h e f a c t t h a t i n s e c t s make up a l a r g e r p o r t i o n o f t h e t o t a l f o o d o f t h e r a i n b o w t r o u t t h a n t h e k o k a n e e . B e c a u s e t h e e s t i m a t e s a r e c r u d e no d e f i n i t i v e s t a t e m e n t c a n b e made r e g a r d i n g t h e p a r a l l e l c h a n g e s w h i c h o c c u r s e a s o n a l l y , h o w e v e r i t i s t e n t a t i v e l y s u g g e s t e d t h a t t h i s c h a n g e r e f l e c t s t h e ^ a v a i l a b i l i t y o f " h i g h e n e r g y " f o o d s t o e a c h o f t h e f i s h s p e c i e s . 4 1 T a b l e 9 . A v e r a g e e n e r g y c o n t e n t o f v a r i o u s f o o d i t e m s a n d t h e p e r c e n t a g e d r y w e i g h t . F o o d i t e m A p p r o x . e n e r g y A p p r o x . amount K c a l / g d r y w t . d r y m a t e r i a l as $ o f w e t w t . H i r u d i n e a 5 .440 2 0 . 0 C l a d o c e r a 4.450 6 . 2 A m p h i p o d a 3 .820 2 1 . 7 C r a n g o n y x 3 .890 2 1 . 7 H y a l e l l a 3 .740 2 1 . 7 E p h e m e r o p t e r a (nymphs) 5 .910 1 7 . 9 O d o n a t a - A n i s o p t e r a ( n a i a d s ) 5 . 0 8 0 19.5 - Z y g o p t e r a ( n a i a d s ) 5 .280 19.5 P l e c o p t e r a - (nymphs) 5 . 160 1 6 . 6 H e m i p t e r a - N o t o n e c t i d a e ( a d u l t s ) 5.350 25.0 C o l e o p t e r a - ( a q u a t i c l a r v a e ) 5 . 4 2 0 25.0 T r i c h o p t e r a - ( e x c l u d i n g c a s e ) ( l a r v a e ) 5 .460 2 2 . 5 D i p t e r a - C h i r o n o m i d a e ( l a r v a e ) 5.300 1 2 . 8 ( p u p a e ) 5 . 0 2 0 12 .,8 - C e r a t o p o g o n i d a e ( l a r v a e ) 5 . 4 2 0 12 .,8 - S i m u l i i d a e ( l a r v a e ) 5.250 15.0 - T i p u l i d a e ( l a r v a e ) 5.090 15.0 42 T a b l e 9. ( c o n t i n u e d ) . A v e r a g e e n e r g y c o n t e n t o f v a r i o u s f o o d i t e m s a n d t h e p e r c e n t a g e d r y w e i g h t . F o o d i t e m A p p r o x . e n e r g y K c a l / g d r y w t . H y m e n o p t e r a ( a d u l t a n t s ) 5-490 M i s c e l l a n e o u s a q u a t i c i n s e c t s ( m o s t l y n o n - a d u l t s ) 5.330 M i s c e l l a n e o u s t e r r e s t r i a l i n s e c t s ( m o s t l y a d u l t s ) 5.130 M o l l u s c a ( i n c l u d i n g s h e l l s ) 3.560 M o l l u s c a - P l a n o r b i d s 3.5&0 - P s e u d o s u c c i n i a s p . 3.570 - P i s i d i u m s p p . 3 .800 A m p h i b i a - Ambystoma g r a c i l e 5 .340 A p p r o x , amount d r y m a t e r i a l a s # o f w e t w t . 25.5 17.1 25.0 25.0 25.0 25.0 2 4 . 2 23,0 43 T a b l e 10. A v e r a g e m o n t h l y v a l u e i n e n e r g y c o n t e n t o f f o o d ( K c a l / k g w e t ) . M o n t h R a i n b o w K o k a n e e t r o u t J a n . 1130a 915a F e b . 1091 812 M a r . 1015a 760a A p r . 932 704 May 968 655 J u n e 1140 9 3 ^ J u l y 1057 785 A u g . 932 893 S e p . 898 8 7 1 O c t . 1022 868 N o v . 1192 947 D e c . 1172 1014 E s t i m a t e d v a l u e s . F i g u r e 1 0 . S e a s o n a l c h a n g e i n e n e r g y c o n t e n t o f f o o d ( c a l / g w e t ) . 6 0 0 5 0 0 1 1 1 1 1 1 i 1 1 1 i 1 — J A N F E B M A R A P R M A Y J U N J U L A U G S E P O C T N O V D E C 45 FOOD TURNOVER The l i t e r a t u r e o n t h e e c o l o g y o f f i s h e s i n c l u d e s many s t u d i e s d e t a i l i n g t h e q u a n t i t a t i v e a n d q u a l i t a t i v e f e a t u r e s o f f o o d c o n s u m p t i o n ( C l e m e n s e t a l , 1923; H a r t l e y , 1 9 4 8 ; L e o n a r d & L e o n a r d , 1949). However t h e r e a r e f e w s t u d i e s w h i c h r e l a t e t h e q u a n t i t y o f f o o d o b s e r v e d i n t h e s t o m a c h a t a g i v e n t i m e t o t h e d a i l y c o n s u m p t i o n o r b e t t e r , t h e d a i l y r a t i o n ( R i c k e r , 1946). Much o f t h e d i f f i c u l t y i n d e t e r m i n i n g f o o d " t u r n o v e r " r a t e s , e s p e c i a l l y u n d e r f i e l d c o n d i t i o n s , i s t h e l a r g e number o f v a r i a b l e s i n v o l v e d . J u s t a s t e m p e r a t u r e r e g u l a t e s t h e m e t a b o l i c r a t e i n f i s h e s , i t a l s o a f f e c t s t h e r a t e o f f o o d i n t a k e ( H a t h a w a y , 1927). The d i s t r i b u t i o n o f t h e f o o d , b o t h i n t e r m s o f a v e r a g e c o n c e n t r a t i o n s a n d a v a i l a b i l i t y , a l s o a f f e c t s t h e f o o d i n t a k e ( I v l e v , 1961). The p o p u l a t i o n d e n s i t y o f t h e f i s h a n d t h e r e s u l t a n t i n f l u e n c e o f i n t e r - a n d i n t r a - s p e c i f i c c o m p e t i t i o n f u r t h e r a f f e c t t h e f o o d c o n s u m p t i o n ( A l l e e , 1933; L a r k i n , 1956). The w o r k o f S u r b e r (1930) a n d B a j k o v (1935) p r o v i d e d a b a s i c f i e l d m e t h o d f o r e s t i m a t i n g t h e d a i l y , s e a s o n a l a n d a n n u a l f o o d c o n s u m p t i o n o f f i s h e s , b u t r e q u i r e d t h a t t h e r a t e o f p a s s a g e o f f o o d f r o m t h e s t o m a c h t o t h e i n t e s t i n e be k n o w n . i One a p p r o a c h t o d e t e r m i n i n g t h e r a t e s o f p a s s a g e o f f o o d t h r o u g h t h e s t o m a c h a n d i n t e s t i n e h a s b e e n t o s u b - s a m p l e t h e f i s h p o p u l a t i o n a t r e g u l a r i n t e r v a l s o v e r a 2 4 h r p e r i o d a n d e x a m i n e e i t h e r t h e r a t e o f p a s s a g e o f c e r t a i n " m a r k e r " o r g a n i s m s ( W e b s t e r , 1942; D a r n e l l & M e i e r o t t o , 1962; H a m i l t o n , M S , I965), o r t h e r a t e a t w h i c h t h e s t o m a c h s a r e e m p t i e d ( B a t t l e e t a l , 1936; R i c k e r , 1937b). 4 6 A s e c o n d a p p r o a c h i n v o l v e s t h e a c t u a l p u m p i n g o u t o f t h e s t o m a c h c o n t e n t s b y w a t e r d i s p l a c e m e n t i n o r d e r t o m e a s u r e t h e r a t e o f d i g e s t i o n o f a p a r t i c u l a r f o o d o r g a n i s m w h i c h has b e e n i n t h e s t o m a c h f o r a known l e n g t h o f t i m e ( S e a b u r g , 1957; S e a b u r g & M o y l e , 1964). R a t e s o f d i g e s t i o n i n p r e d a t o r y s p e c i e s h a v e a l s o b e e n m e a s u r e d b y s e q u e n t i a l X - r a y ( M o l n a r e t a l , I967). I n t h e p r e s e n t s t u d y , t h e d a i l y r a t i o n o f f o o d i n t h e r a i n b o w a n d k o k a n e e p o p u l a t i o n s i s b a s e d o n c a l c u l a t i o n s d e r i v e d f r o m t w o ( s e t s o f e x p e r i m e n t s o n r a i n b o w t r o u t f r o m M a r i o n L a k e . H a m i l t o n ( M S , 1965) d e s c r i b e d a n e x p e r i m e n t c o n d u c t e d i n A u g u s t , I963, a t M a r i o n L a k e i n w h i c h t h e r a t e o f p a s s a g e o f p a r t i c u l a r " m a r k e r " o r g a n i s m s was s t u d i e d o v e r a 2 4 h r p e r i o d . He c a l c u l a t e d t h a t t h e g u t was c l e a r e d e v e r y 8 h r ( 3 x i n 2 4 h r ) a t a t e m p e r a t u r e a b o v e 17°C. The mean m o n t h l y t e m p e r a t u r e f o r A u g u s t (18.8°C) a t 3 m d e p t h h a s b e e n assumed i n t h e p r e s e n t c a l c u l a t i o n s . H a m i l t o n a l s o f o u n d t h a t the-t o t a l number o f o r g a n i s m s a c t u a l l y consumed w i t h i n t h e l a s t d i g e s t i v e p e r i o d ( i . e . 8 h r ) c o u l d be c a l c u l a t e d f r o m t h e r a t i o o f t h e number o f p a r t i c u l a r m a r k e r o r g a n i s m s f o u n d i n t h e i n t e s t i n e d i v i d e d b y t h e number o f t h e s e same o r g a n i s m s f o u n d i n t h e s t o m a c h ( r a t i o f o r c a d d i s l a r v a e 1.70, c h i r o n o m i d l a r v a e 1.58, c h i r o n o m i d p u p a e 1.41, c h i r o n o m i d a d u l t s 1.18, m o l l u s c s 1 .48, a m p h i p o d s 1.08). The o r g a n i s m s w i t h t h e m o s t c o n s p i c u o u s i n d i g e s t i b l e p a r t s ; a n d h e n c e t h e b e s t " m a r k e r " o r g a n i s m s h a d t h e h i g h e s t r a t i o s . The mean r a t i o f o r t h e c a d d i s l a r v a e , c h i r o n o m i d l a r v a e a n d m o l l u s c s was 1.6. 47 Thus f o r e a c h i n d i v i d u a l o b s e r v e d i n t h e s t o m a c h t h e r e w e r e e s t i m a t e d t o b e 1.6 i n t h e i n t e s t i n e o r a t o t a l o f 2.6 i n t h e w h o l e d i g e s t i v e t r a c t . H a m i l t o n t h u s c o n c l u d e d t h a t , " f o r s o f t b o d i e d o r e a s i l y -f r a g m e n t e d f o r m s , m u l t i p l y i n g t h e number o f o r g a n i s m s i n t h e s t o m a c h s b y 2.6 p r o v i d e s a n e s t i m a t e o f t h e number o f o r g a n i s m s e a t e n d u r i n g t h e same p e r i o d as t h e more d u r a b l e o r g a n i s m s s t i l l p r e s e n t i n t h e i n t e s t i n e " . B y d e t e r m i n i n g t h e r e l a t i v e numbers o f v a r i o u s o r g a n i s m s i n d i f f e r e n t p a r t s o f t h e d i g e s t i v e t r a c t a n d r a n k i n g t h e m i n o r d e r o f d e c r e a s i n g s i z e h e f o u n d t h a t e a c h o f t h e f o o d i t e m s consumed b y t h e r a i n b o w t r o u t moved t h r o u g h t h e d i g e s t i v e t r a c t a t a u n i f o r m r a t e . T h i s i s c o n t r a r y t o t h e g e n e r a l l y a c c e p t e d v i e w t h a t f o o d i s h e l d a t t h e p y l o r u s f o r l o n g e r p e r i o d s . However i t h a s b e e n shown b y W i n d e l l (I967) i n f e e d i n g e x p e r i m e n t s u s i n g m e a l worms w i t h h i g h f a t l e v e l s , t h a t t h e r a t e o f p a s s a g e may d e p e n d more o n t h e q u a l i t y o f t h e f o o d r a t h e r t h a n o n t h e s i z e . H i g h e r f a t c o n t e n t o f t h e f o o d i s c o r r e l a t e d w i t h a d e c r e a s i n g r a t e o f p a s s a g e . D u r i n g t h e summer o f I968, D . M . Ware ( p e r s . comm.) c o n d u c t e d a s e r i e s o f f e e d i n g e x p e r i m e n t s u s i n g M a r i o n L a k e r a i n b o w t r o u t i n w h i c h he d e t e r m i n e d t h a t a t 9°C f o o d w o u l d p a s s t h r o u g h t h e d i g e s t i v e t r a c t i n a b o u t 20 h r . o H a m i l t o n ' s t u r n o v e r r a t e o f Jx. a t 18.8 C and W a r e ' s r a t e o f 1.2x a t 9°C w e r e p l o t t e d as l o g r a t e a g a i n s t t e m p e r a t u r e i n o r d e r t o o b t a i n e s t i m a t e s o f t h e t u r n o v e r r a t e s a t o t h e r t e m p e r a t u r e s ( F i g . 1 1 ) . F o r c o m p a r i s o n t h e d i g e s t i o n r a t e s i n h e r r i n g a t t e m p e r a t u r e s f r o m 6-20°C as d e t e r m i n e d b y B a t t l e e t a l (1936) h a v e b e e n c o n v e r t e d F i g u r e 1 1 . T u r n o v e r r a t e s i n f o o d ( l o g ) a g a i n s t t e m p e r a t u r e ( ° C ) . TURNOVER RATE 49 t o t u r n o v e r r a t e s b y d i v i d i n g t h e number o f h o u r s r e q u i r e d t o c l e a r t h e e n t i r e d i g e s t i v e t r a c t i n t o 2 4 h r ( F i g . 11). A s i m i l a r s t u d y b y K r o g i u s a n d K r o k h i n (1948) p r o v i d e d some i n f o r m a t i o n o n t h e r a t e o f d i g e s t i o n i n j u v e n i l e s o c k e y e . A l t h o u g h n o t e x p r e s s l y s t a t e d , i t i s assumed t h a t t h e i r " h o u r s f o r d i g e s t i o n " a r e b a s e d o n t h e c l e a r i n g o f t h e e n t i r e d i g e s t i v e t r a c t r a t h e r t h a n t h e s t o m a c h a l o n e , as t h e y compare t h e i r d a t a t o t h e " w h o l e t r a c t " d a t a o f B a t t l e e t a l (1936)0 T u r n o v e r r a t e s w e r e c a l c u l a t e d a n d p l o t t e d ( F i g . 11). The e q u a t i o n s f o r t h e t h r e e l i n e a r r e g r e s s i o n s w e r e d e t e r m i n e d b y m e t h o d o f l e a s t s q u a r e s ( T a b l e 11). A s a f u r t h e r c h e c k o n t h e a p p l i c a b i l i t y o f t h e a b o v e method:, o f e x p r e s s i n g t u r n o v e r r a t e s a n d t h e c a l c u l a t i o n o f d a i l y r a t i o n , t h e d a t a o f R i c k e r a n d F o e r s t e r ( 1 9 4 8 , p . 201) o n t h e f o o d consumed by-C u l t u s L a k e s o c k e y e s a l m o n (1931 y e a r c l a s s i n 1932-1933) w e r e e x a m i n e d . T h e i r t u r n o v e r r a t e s w e r e b a s e d o n t h e f a c t t h a t a " c o n s e r v a t i v e e s t i m a t e o f s o c k e y e f e e d i n g i n summer w o u l d be t h a t t h e y consume t h e i r a v e r a g e o b s e r v e d s t o m a c h c o n t e n t s e v e r y d a y " ( R i c k e r , 1937b), b u t t h a t a " m o s t p r o b a b l e e s t i m a t e " was b e t t e r a n d t h i s was c o n s i d e r e d t o be " f o u r t i m e s t h e o b s e r v e d c o n t e n t s , i n summer" ( R i c k e r & F o e r s t e r , 1948). D u r i n g autumn a n d w i n t e r t h e s e r a t e s w e r e r e d u c e d t o 3 X a n d 2x r e s p e c t i v e l y . A p p l y i n g t h e s e r a t e s t o t h e s t o m a c h c o n t e n t s ( e x p r e s s e d as $ o f b o d y w e i g h t ) t h e y c a l c u l a t e d d a i l y r a t i o n , w h i c h when m u l t i p l i e d b y p o p u l a t i o n w e i g h t y i e l d s t h e amount o f f o o d consumed f o r one t i m e • i n t e r v a l (•§• m o n t h ) . Summing t h e s e v a l u e s , t h e t o t a l e s t i m a t e d f o o d 50 T a b l e 11. R e l a t i o n o f l o g t u r n o v e r r a t e o f f o o d t o t e m p e r a t u r e ( C ) . A t l a n t i c h e r r i n g y = .462 e , o 9 1 x B a t t l e e t a l (1936) ( C l u p e a h a r e n g u s ) »125x S o c k e y e s a l m o n y = .638 e* K r o g i u s & K r o k h i n (1948) ( O n c o r h y n c h u s n e r k a ) R a i n b o w t r o u t y = .545 e* H a m i l t o n ( M S , 1965), a n d ( S a l m o g a i r d n e r i ) Ware ( p e r s . comm.) 51 c o n s u m p t i o n b y t h e 1931 y e a r c l a s s was 103 m e t r i c t o n s w e t w e i g h t . T h i s v a l u e t h e y r e g a r d as l o w a n d o n t h e b a s i s o f f o o d c o n v e r s i o n e f f i c i e n c i e s , s u g g e s t t h a t t h e b e s t e s t i m a t e o f p l a n k t o n c o n s u m p t i o n was 154 m e t r i c t o n s . H o w e v e r , e v e n t h i s v a l u e was c o n s i d e r e d l o w b e c a u s e a minimum f o o d c o n s u m p t i o n o f 154 m e t r i c t o n s was " a c h i e v e d i n a l a t e r y e a r w i t h a much s m a l l e r i n i t i a l p o p u l a t i o n " . I n o r d e r t h a t t h e t u r n o v e r r a t e s as c a l c u l a t e d f o r t h e M a r i o n L a k e f i s h be a p p l i e d t o t h e C u l t u s L a k e d a t a i t was n e c e s s a r y t o assume t h e same mean m o n t h l y t e m p e r a t u r e s . The t e m p e r a t u r e d a t a g i v e n f o r C u l t u s L a k e b y R i c k e r (1937a) s u p p o r t s t h i s a s s u m p t i o n . R e c a l c u l a t i n g t h e f o o d c o n s u m p t i o n o f C u l t u s L a k e f i s h , u s i n g t h e p o p u l a t i o n w e i g h t a n d s t o m a c h c o n t e n t d a t a ( m u l t i p l i e d b y 2.6) as g i v e n b y R i c k e r & F o e r s t e r , b y s u b s t i t u t i n g t h e M a r i o n L a k e t u r n o v e r r a t e s ( o f f i s h f o o d ) , g i v e s t h e t o t a l c o n s u m p t i o n f o r t h e 1931 y e a r c l a s s as I67 m e t r i c t o n s w e t w e i g h t . The e s t i m a t e o f f o o d c o n s u m e d , c a l c u l a t e d b y t h i s m e a n s , i s g r e a t e r t h a n t h e 154 m e t r i c t o n s c o n s i d e r e d b y R i c k e r & F o e r s t e r t o be t h e minimum e s t i m a t e a n d f a r g r e a t e r t h a n t h e 103 m e t r i c t o n s c a l c u l a t e d b y t h e m . T h e s e r e s u l t s h e l p t o s u b s t a n t i a t e t h e m e t h o d o f c a l c u l a t i n g d a i l y f o o d r a t i o n s i n t h e p r e s e n t p a p e r . The s e a s o n a l c h a n g e i n t u r n o v e r r a t e i n M a r i o n L a k e f i s h ( T a b l e 12) was c a l c u l a t e d b y i n s e r t i n g t h e v a l u e s f o r t h e mean m o n t h l y t e m p e r a t u r e i n t h e H a m i l t o n - W a r e r e g r e s s i o n ( T a b l e 11). I n t h e a b s e n c e o f a n y e x p e r i m e n t a l d a t a o n t h e d i g e s t i v e r a t e s i n M a r i o n L a k e k o k a n e e , t h e t u r n o v e r r a t e s as c a l c u l a t e d f o r r a i n b o w 52 T a b l e 1 2 . C a l c u l a t e d d a i l y t u r n o v e r r a t e o f f o o d a t d i f f e r e n t t e m p e r a t u r e s . M o n t h Mean t e m p , a t C a l c u l a t e d 3 m d e p t h t u r n o v e r r a t e ( ° C ) J a n . 3.0 0.72 F e b . 3.0 0.72 M a r . 4 .3 0 . 8 1 A p r . 6 .0 0 .94 May U . 5 1.55 J u n e 1 4 . 1 I .96 J u l y 16 .2 2.37 A u g . 18 .8 3.00 S e p . 14 .8 2.09 O c t . 9.3 I.27 N o v . 6.5 0.98 D e c . 4 .0 0.78 53 w e r e u t i l i z e d . D a i l y r a t i o n s f o r e a c h m o n t h w e r e c a l c u l a t e d f r o m t h e f o l l o w i n g e q u a t i o n : W t . o f f o o d i n d i g e s t i v e t r a c t ( g ) D a i l y r a t i o n = • • • • • <• x T u r n o v e r r a t e x 100$ W t . o f f i s h ( g ) -w h e r e , w e i g h t o f f o o d i n d i g e s t i v e t r a c t = w e i g h t o f s t o m a c h c o n t e n t s x 2 . 6 . One p o s s i b l e s o u r c e o f e r r o r n o t a c c o u n t e d f o r i n c a l c u l a t i o n o f t h e d a i l y r a t i o n i s t h e c h a n g e i n f o o d i n t a k e w i t h s i z e . P a n d i a n (1967), i n h i s s t u d y o f t w o s p e c i e s o f I n d i a n e u r y h a l i n e f i s h e s , d e m o n s t r a t e d t h a t as w e i g h t i n c r e a s e d u p t o 20 g t h e r e was a s h a r p d e c r e a s e i n f o o d i n t a k e a n d t h a t f o r f i s h l a r g e r t h a n 20 g t h e d e c l i n e c o n t i n u e d b u t a t a s l o w e r r a t e . F o r M a r i o n L a k e f i s h i t i s assumed f o r s i m p l i f i c a t i o n t h a t t h e d a i l y r a t i o n i s t h e same f o r a l l w e i g h t s o f f i s h . The amount o f f o o d i n t h e s t o m a c h s e x a m i n e d was m e a s u r e d b y ; d i s p l a c e m e n t v o l u m e ( m l ) . H o w e v e r , i t h a s b e e n shown b y B a l l (19^8) a n d H u n t (i960) t h a t 1 gram i s a r e a s o n a b l e a p p r o x i m a t i o n f o r 1 m l v o l u m e d i s p l a c e m e n t . Thus t h e v o l u m e o f f o o d i n e a c h s t o m a c h was e x p r e s s e d as grams f o r c a l c u l a t i o n ( T a b l e 13). D u r i n g t h e w i n t e r t h e d a i l y r a t i o n p r o b a b l y r e m a i n s c l o s e t o 1$ o f t h e b o d y w e i g h t ( F i g . 12). I n summer t h e r e a r e two p e a k p e r i o d s (May a n d A u g u s t ) when t h e d a i l y r a t i o n a p p r o a c h e s maximum v a l u e s i n b o t h s p e c i e s . B r e t t e t a l (1969) i n a l a b o r a t o r y s t u d y o f t h e f e e d i n g r a t e s 54 T a b l e 13- D a i l y f o o d r a t i o n e x p r e s s e d a s % o f t o t a l b o d y w e i g h t f o r M a r i o n L a k e r a i n b o w t r o u t (mean w e i g h t o f a l l t r o u t e x a m i n e d 91*1 g ) a n d k o k a n e e (mean w e i g h t 45.1 g) (number o f f i s h e x a m i n e d i n b r a c k e t s ) . R a i n b o w t r o u t M o n t h F o o d i n s t o m a c h D a i l y r a t i o n (# o f b o d y w t . ) (# o f b o d y w t . ) J a n . - a 1 . 0 0 F e b . - a 1 . 0 0 M a r . - a 1 . 0 0 A p r . - a 2 . 0 0 May- 1 . 3 ^ 5 . 4 2 ( 9 ) J u n e 0 . 7 7 3 . 9 3 ( 1 1 ) J u l y 0 . 4 9 3 . 0 2 ( 1 0 ) A u g . 0 . 8 3 6 . 4 5 (?) S e p . - 3 . 0 0 a O c t . 0 . 4 5 1 . 4 9 ( 9 ) N o v . 0 . 4 8 1 . 2 2 ( 4 ) D e c . 0.55 1 .12 ( 4 ) T o t a l number o f f i s h ( 5 4 ) a E s t i m a t e d v a l u e 55 T a b l e 13. ( c o n t i n u e d ) . D a i l y f o o d r a t i o n e x p r e s s e d as $ o f t o t a l b o d y w e i g h t f o r M a r i o n L a k e r a i n b o w t r o u t (mean w e i g h t o f a l l t r o u t e x a m i n e d 91.1 g) a n d k o k a n e e (mean w e i g h t 45.1 g) (number o f f i s h e x a m i n e d i n b r a c k e t s ) . M o n t h J a n . F e b . M a r . A p r . May J u n e J u l y A u g . S e p . O c t . N o v . D e c . K o k a n e e F o o d i n s t o m a c h ( $ o f b o d y w t . ) 0 .84 1.03 0 .44 0 .49 0 .48 0.25 0 .38 0 .47 0 .68 D a i l y r a t i o n ( $ o f b o d y w t . ) 1.00a i.ooa i.ooa 2.06 ( 7 ) 4 . 1 6 (12) 2 .24 ( 9 ) 3.02 (12) 3 .74 (12) 1.36 ( 1 4 ) 1.25 (8) 1.20 ( 7 ) 1.38 (6) T o t a l number o f f i s h (87) E s t i m a t e d v a l u e F i g u r e 1 2 , S e a s o n a l c h a n g e i n d a i l y r a t i o n as a p e r c e n t a g e o f b o d y w e i g h t . PERCENT O F BODY W E I G H T 5? a n d g r o w t h o f j u v e n i l e s o c k e y e s a l m o n f o u n d t h a t t h e maximum d a i l y r a t i o n was a b o u t 5«7$ ( o n a w e t b a s i s ^ ) a t opt imum t e m p e r a t u r e s a n d 7 o t h a t t h i s f e e d i n g l e v e l f e l l t o 0.94$ a t 1 C . F o r M a r i o n L a k e f i s h i t i s o f i n t e r e s t t o n o t e t h a t t h e p e r c e n t d a i l y r a t i o n i n t h e r a i n b o w t r o u t i s g e n e r a l l y e q u a l t o o r g r e a t e r t h a n t h e k o k a n e e , w h i c h i s t h e same r e l a t i o n s h i p as was f o u n d i n t h e s e a s o n a l c h a n g e i n e n e r g y c o n t e n t o f t h e f o o d . Thus i t w o u l d a p p e a r t h a t t h e r a i n b o w t r o u t h a v e a s e l e c t i v e a d v a n t a g e i n t h e i r f o o d h a b i t s ; n o t o n l y do t h e y i n d i v i d u a l l y g e t more f o o d t h a n a k o k a n e e b u t t h e n u t r i t i v e v a l u e o f t h a t f o o d i s g r e a t e r . A S S I M I L A T I O N E F F I C I E N C I E S , A s a p a r t o f t h e f e e d i n g e x p e r i m e n t s c o n d u c t e d b y D . M . Ware ( p e r s . comm.) a t M a r i o n L a k e i n 1968, s m a l l r a i n b o w t r o u t (mean 13 cm) w e r e h e l d i n a q u a r i a (212 1) w i t h o u t f o o d f o r a t l e a s t 55 h r i n o r d e r t o empty t h e s t o m a c h a n d t h e g u t . A t t h e s t a r t o f a n e x p e r i m e n t e a c h f i s h was f e d a known number o f o r g a n i s m s o f u n i f o r m s i z e ( T a b l e 1 4 ) . I n some c a s e s t h e s i z e o f t h e r a t i o n was c l o s e t o t h e s a t i a t i o n l e v e l o f t h e f i s h f o r a s i n g l e f e e d i n g . F e c a l m a t e r i a l was c o l l e c t e d r e g u l a r l y b e t w e e n t h e t i m e o f f i r s t 6 . C a l c u l a t e d o n b a s i s o f d a i l y r a t i o n 7.5$ ( d r y ) ; m o i s t u r e o f f i s h 72$, o f f o o d 63$. 7. C a l c u l a t e d o n b a s i s o f d a i l y r a t i o n 1.5$ ( d r y ) ; m o i s t u r e o f f i s h 77$, o f f o o d 63$. T a b l e 1 4 . A s s i m i l a t i o n e f f i c i e n c y i n r a i n b o w t r o u t b a s e d o n n a t u r a l f o o d s ( H y a l e l l a 3740 c a l / g d r y w e i g h t , C r a n g o n y x 3890 c a l / g d r y w e i g h t ) . R a i n b o w t r o u t N o . 1 D a t e W a t e r F o o d D r y w t . E n e r g y i n t a k e E n e r g y l o s s E n e r g y t e m p . (mg) ( c a l c u l a t e d ) ( e g e s t i o n o n l y ) a s s i m i l a t e d <°« ( c a l o r i e s ) ( c a l o r i e s ) ( * ) 12 V I 68 8 88 H y a l e l l a 70.5 264 37.0 86.0 19 V I 68 9 16 C r a n g o n y x 91.2 355 36.9 89.6 2 4 V I 68 11 19 C r a n g o n y x 107 416 54.4 86.9 17 V I I 68 9 193 H y a l e l l a 135 505 98.9 80.4 T o t a l 1540 T o t a l 227 mean 85.4 T a b l e 1 4 . ( c o n t i n u e d ) . A s s i m i l a t i o n e f f i c i e n c y i n r a i n b o w t r o u t b a s e d o n n a t u r a l f o o d s ( H y a l e l l a 3740 c a l / g d r y w e i g h t , C r a n g o n y x 3890 c a l / g d r y w e i g h t ) . R a i n b o w t r o u t N o . 2 D a t e W a t e r F o o d D r y w t . E n e r g y i n t a k e E n e r g y l o s s E n e r g y t e m p . (mg. ) ( c a l c u l a t e d ) ( e g e s t i o n o n l y ) a s s i m i l a t e d ( ° C ) ( c a l o r i e s ) ( c a l o r i e s ) (*> 1 2 V I 6 8 8 15 C r a n g o n y x 8 1 . 7 318 4 0 . 4 8 7 . 3 14 V I 6 8 10 49 H y a l e l l a 3 9 . 2 147 2 7 . 0 8 1 . 6 19 V I 68 9 100 H y a l e l l a 7 0 . 0 262 4 5 . 8 8 2 . 5 2 1 V I 68 11 67 H y a l e l l a 4 0 . 2 150 2 0 . 0 8 6 . 7 2 4 V I 68 11 19 H y a l e l l a 13.3 50 6 . 9 8 6 . 2 15 V I I 68 9 198 H y a l e l l a 178 6 6 6 85.8 8 7 . 1 T o t a l 1593 T o t a l 2 2 6 mean 85.8 F o r b o t h f i s h T o t a l 3133 T o t a l 4 5 3 mean 85.5 60 a p p e a r a n c e ( a b o u t 12 h r ) u n t i l t h e d i g e s t i v e t r a c t was empty ( n o t l o n g e r t h a n 55 h r ) . The m a t e r i a l c o l l e c t e d was d r i e d a n d w e i g h e d . The e n e r g y c o n t e n t o f t h e f o o d a n d f e c a l s a m p l e s was d e t e r m i n e d b y c o m b u s t i o n i n a P a r r c a l o r i m e t e r . P e r c e n t a s s i m i l a t i o n v a l u e s a r e a n e x p r e s s i o n o f t h e c h a n g e i n e n e r g y c o n t e n t o f t h e f o o d a f t e r p a s s a g e t h r o u g h t h e d i g e s t i v e t r a c t ( T a b l e 1 4 ) . The a v e r a g e amount o f f o o d a s s i m i l a t e d , b a s e d o n t h e d i f f e r e n c e b e t w e e n t h e c a l o r i f i c c o n t e n t o f t h e f o o d a n d t h e f e c a l m a t e r i a l i s 85.5$ u n d e r t h e e x p e r i m e n t a l c o n d i t i o n s d e s c r i b e d . However t h i s v a l u e i s u n d o u b t e d l y a n o v e r e s t i m a t e s i n c e t h e c a l o r i f i c c o n t e n t o f t h e m a t e r i a l e x c r e t e d i s u n k n o w n . W i n b e r g (1956) s t a t e s t h a t t h e e n e r g y l o s s a t t r i b u t a b l e t o e x c r e t i o n may r a n g e f r o m l e s s t h a n 3-5$ "to a. maximum o f 14$ d e p e n d i n g o n t h e r a t e o f g r o w t h o f t h e f i s h a n d t h e p r o d u c t s o f e x c r e t i o n . I t h a s b e e n p r o p o s e d b y W i n b e r g (1956) t h a t a n 8 0 $ e f f i c i e n c y ( o r 20$ l o s s t h r o u g h e g e s t i o n a n d e x c r e t i o n ) be c o n s i d e r e d as a n a v e r a g e f o r a l l f i s h . T h i s v a l u e , r e p r e s e n t i n g t h e p h y s i o l o g i c a l l y u s e f u l p o r t i o n o f t h e f o o d , has , b e e n u s e d b y Mann (1965). P a l o h e i m o & D i c k i e (I966) a n d o t h e r s t o c a l c u l a t e p o p u l a t i o n e n e r g y b u d g e t s o f f i s h c o n s u m i n g n a t u r a l f o o d s . T h e r e f o r e , i t h a s b e e n assumed t h a t t h e a s s i m i l a t i o n e f f i c i e n c y i n b o t h t h e r a i n b o w t r o u t a n d k o k a n e e p o p u l a t i o n s ' o f M a r i o n L a k e i s 8 0 $ . T h e r e a r e h o w e v e r , a t l e a s t two p o s s i b l e s o u r c e s o f e r r o r i n t h i s , a s s u m p t i o n . F i r s t i t i s h i g h l y p r o b a b l e t h a t a s s i m i l a t i o n e f f i c i e n c i e s d e c r e a s e a t t e m p e r a t u r e s a b o v e o r b e l o w w h a t i s n o r m a l l y c o n s i d e r e d t h e opt imum r a n g e , a n d s e c o n d i t h a s b e e n d e m o n s t r a t e d a l r e a d y t h a t " a b s o r p t i o n a n d c o n v e r s i o n " e f f i c i e n c e s d e c r e a s e w i t h i n c r e a s i n g s i z e ( P a n d i a n , 1967). 61 I n t h e a b s e n c e o f a n y d a t a o n how t h e s e f a c t o r s m i g h t a f f e c t t h e M a r i o n L a k e f i s h t h e one v a l u e f o r a s s i m i l a t i o n e f f i c i e n c y h a s b e e n a p p l i e d t o a l l f i s h i n a l l s e a s o n s , ROUTINE METABOLISM The e n e r g y consumed b y m e t a b o l i c p r o c e s s e s i n f i s h i s n o r m a l l y d e t e r m i n e d i n d i r e c t l y b y m e a s u r i n g t h e r a t e o f o x y g e n u p t a k e . F r y (1957) d e f i n e d t h r e e l e v e l s o f o x y g e n c o n s u m p t i o n , s t a n d a r d , r o u t i n e a n d a c t i v e . M e a s u r e m e n t o f t h e s t a n d a r d a n d a c t i v e r a t e s o f o x y g e n c o n s u m p t i o n i n f i s h e s was g i v e n g r e a t i m p e t u s b y F r y (1947) a n d a s s o c i a t e s . Numerous p a p e r s h a v e b e e n p u b l i s h e d o n m e t a b o l i c r a t e s i n a v a r i e t y o f f i s h e s ( s u m m a r i z e d b y W i n b e r g , 1956, 1961) i n c l u d i n g s a l m o n i d s ( F r y , 1 9 4 7 , 1957 ; G r a h a m , 1949; G i b s o n & F r y , 1954; J o b , 1955; B e a m i s h , 1964a, 1964b; B r e t t , 1964),; The e x p e r i m e n t a l a p p r o a c h t o d e t e r m i n a t i o n o f o x y g e n c o n s u m p t i o n r a t e s i n s a l m o n i d s , i n a l m o s t e v e r y c a s e , h a s b e e n f r o m a p h y s i o l o g i c a l p o i n t o f v i e w . T h i s i s p a r t i c u l a r l y t r u e i n t h e measurement o f s t a n d a r d r a t e s . The f i s h u s e d i n t h e s e e x p e r i m e n t s u s u a l l y a r e o b t a i n e d f r o m c u l t u r e d ( h a t c h e r y ) s t o c k , m a i n t a i n e d o n a n a r t i f i c i a l d i e t ( B e a m i s h , 1 9 6 4 b ) , h e l d i n t a n k s a t c o n s t a n t ( a c c l i m a t i o n ) t e m p e r a t u r e s , a n d s t a r v e d f o r 36 h o u r s b e f o r e commencing t h e e x p e r i m e n t ( B r e t t , 1 9 6 4 ) , I n e a r l i e r s t u d i e s , f i s h w h i c h h a d b e e n p r e c o n d i t i o n e d i n t h i s way w e r e c o n s i d e r e d t o y i e l d s t a n d a r d r a t e s o f o x y g e n c o n s u m p t i o n . H o w e v e r , B e a m i s h a n d M o o k h e r j i i ; ( 1 9 6 4 ) p o i n t e d o u t t h a t m o s t e a r l i e r v a l u e s o f s t a n d a r d r a t e w e r e i n f a c t , r o u t i n e r a t e s s i n c e some s p o n t a n e o u s a c t i v i t y d i d o c c u r . M o r e r e c e n t l y , s t a n d a r d r a t e s h a v e b e e n b a c k - c a l c u l a t e d f r o m r o u t i n e r a t e s b y m e a s u r i n g 62 t h e amount o f s p o n t a n e o u s a c t i v i t y ( B e a m i s h , 1964b), o r b y m e a s u r i n g t h e o x y g e n c o n s u m p t i o n a t v a r i o u s l e v e l s o f f o r c e d a c t i v i t y a n d b a c k c a l c u l a t i n g t o z e r o a c t i v i t y ( B r e t t , 1964). T h e s e s t u d i e s d e m o n s t r a t e a h i g h d e g r e e o f r e p r o d u c i b i l i t y , c o n s i d e r i n g t h e v a r i a t i o n t h a t e x i s t s w i t h i n b i o l o g i c a l s y s t e m s . However f r o m a n e c o l o g i c a l p o i n t o f v i e w , t h e s t a n d a r d a n d a c t i v e r a t e s d e t e r m i n e d b y t h e s e m e t h o d s , r e f l e c t o n l y t h e maximum u p p e r a n d l o w e r l i m i t s w i t h i n w h i c h t h e f i s h o p e r a t e . I n t h e M a r i o n L a k e f i s h , r o u t i n e r a t e s o f o x y g e n c o n s u m p t i o n w e r e m e a s u r e d i n o r d e r t o e s t i m a t e t h e n o r m a l m e t a b o l i c r e q u i r e m e n t s o f t h e t w o p o p u l a t i o n s . F o r t h i s r e a s o n , t h e f i s h w e r e t r a n s f e r r e d as q u i c k l y as p o s s i b l e f r o m t h e i r n a t u r a l e n v i r o n m e n t t o t h e r e s p i r o m e t e r ( l a r g e enough t o a l l o w s p o n t a n e o u s a c t i v i t y ) , a n d n o t s t a r v e d f o r a p r e l i m i n a r y p e r i o d . A t t e m p t s t o m a i n t a i n t h e f o o d i n t a k e ( n a t u r a l f o o d s ) o f f i s h i n t h e ; r e s p i r o m e t e r w e r e u n s u c c e s s f u l . j The c a l c u l a t e d r o u t i n e r a t e was b a s e d on t h e a v e r a g e r a t e o f o x y g e n c o n s u m p t i o n o c c u r r i n g b e t w e e n t h e 8th a n d 16th h r f r o m t h e s t a r t o f t h e e x p e r i m e n t . D u r i n g t h e f i r s t 8 h r i n t h e r e s p i r o m e t e r e a c h f i s h e x h i b i t e d a n i n i t i a l h i g h r a t e o f o x y g e n c o n s u m p t i o n w h i c h g r a d u a l l y d e c l i n e d t o a f a i r l y s t e a d y r a t e d u r i n g t h e n e x t 8 h r . T h e s e h i g h i n i t i a l r a t e s a r e p r o b a b l y due t o t h e e f f e c t s o f h a n d l i n g t h e f i s h , w h i c h i m p o s e s a m e t a b o l i c l o a d i n t h e f o r m o f i n c r e a s e d l a c t i c a c i d l e v e l s . B l a c k e t a l (1962) h a v e o shown t h a t h a t c h e r y - r e a r e d r a i n b o w t r o u t a c c l i m a t e d t o 11.5 C r e t u r n t o n o r m a l l e v e l s o f m u s c l e l a c t a t e w i t h i n 8 h r f o l l o w i n g f a t i g u e . R e c o v e r y o f t h e n o r m a l l e v e l s o f b l o o d a n d l i v e r l a c t a t e a n d p y r u v a t e t o o k a b o u t 12 h r . M u s c l e p y r u v a t e r e t u r n e d t o n o r m a l l e v e l s w i t h i n 4 h r o f f a t i g u e . H o c h a c h k a (1961) d e m o n s t r a t e d t h a t r e s p i r a t i o n r a t e s r e p r e s e n t i n g 63 r o u t i n e m e t a b o l i s m w e r e r e a c h e d w i t h i n 4-6 h r f o l l o w i n g f a t i g u e . He c o n c l u d e d t h a t n o t a l l o f t h e a c c u m u l a t e d l a c t a t e was o x i d i z e d b e f o r e t h e f i s h moved o u t o f a s t a t e o f o x y g e n d e b t , d e s p i t e t h e f a c t t h a t m u s c l e g l y c o g e n a n d l a c t i c a c i d l e v e l s d i d n o t r e t u r n t o n o r m a l l e v e l s f o r a much l o n g e r t i m e . T h e r e f o r e t h e d a t a o n o x y g e n c o n s u m p t i o n c o l l e c t e d d u r i n g t h e f i r s t 8 h r o f t h e e x p e r i m e n t was d i s r e g a r d e d i n t h e c a l c u l a t i o n o f t h e a v e r a g e r o u t i n e m e t a b o l i c r a t e . D u r i n g 1967-1968 t h e r o u t i n e m e t a b o l i c r a t e o f 43 M a r i o n L a k e f i s h was m e a s u r e d . T h e s e e x p e r i m e n t s w e r e c o n d u c t e d o n 39 r a i n b o w t r o u t r a n g i n g i n s i z e f r o m 57 g t o 369 g ( m e a n , 121 g) a t t e m p e r a t u r e s f r o m , o o 4.5-20.5 C a n d 4 k o k a n e e , 46 g t o 60 g a t t e m p e r a t u r e s f r o m 6-16 C . A v e r a g e r a t e o f o x y g e n c o n s u m p t i o n (8th-l6th h r ) was e x p r e s s e d i n m l / h r f o r a g i v e n f i s h a t a g i v e n t e m p e r a t u r e . No c o n s i s t e n t p a t t e r n o f i n c r e a s e i n o x y g e n c o n s u m p t i o n a s s o c i a t e d w i t h dawn o r d u s k was a p p a r e n t . B y c o n f i n i n g t h e e x p e r i m e n t a l p e r i o d t o a n 8 h r i n t e r v a l a t n i g h t t h e e f f e c t o f a n y d i e l r e s p o n s e was e l i m i n a t e d . I n o r d e r t o d e t e r m i n e t h e r e l a t i o n s h i p b e t w e e n t h e r o u t i n e m e t a b o l i s m a n d w e i g h t , as e x p r e s s e d b y t h e e q u a t i o n , Q = a W k w h e r e £ i s t h e r a t e o f m e t a b o l i s m , W i s b o d y w e i g h t ( k g w e t ) , a n d a a n d k a r e c o n s t a n t s , i t was n e c e s s a r y t o c o n v e r t t h e o x y g e n c o n s u m p t i o n r a t e s a t v a r i o u s t e m p e r a t u r e s t o a s i n g l e t e m p e r a t u r e (15°C). W i n b e r g (1956) a p p l i e d t h e same t e c h n i q u e t o h i s summary o f m e t a b o l i c r a t e s a n d f o r 6 4 c o n v e n i e n c e c a l c u l a t e d a t a b l e o f m u l t i p l i e r s b a s e d o n t h e s o - c a l l e d " n o r m a l " c u r v e (Ege & K r o g h , 1914). Mann (1965) h a s shown t h a t f o r c y p r i n i d s , a t l e a s t , t h a t t h i s i s a r e a s o n a b l e a p p r o x i m a t i o n . P a l o h e i m o & D i c k i e (1966) c o n c l u d e d t h a t t h e " K r o g h c o r r e c t i o n f o r t e m p e r a t u r e " c o u l d b e u s e d e x c e p t " w h e r e a c c l i m a t i o n o r s e a s o n a l m e t a b o l i c phenomena a r e i n v o l v e d " . A l t h o u g h t h e " n o r m a l " o r c o n v e x c u r v e h a s l o n g b e e n u s e d t o p r e d i c t t h e r a t e o f r e a c t i o n a t d i f f e r e n t t e m p e r a t u r e s f o r a g r e a t v a r i e t y o f o r g a n i s m s , i t w o u l d a p p e a r t h a t t h e r e a r e some m a r k e d e x c e p t i o n s . , M o s t i m p o r t a n t , f r o m t h e p o i n t o f v i e w o f t h e p r e s e n t s t u d y , a r e t h e d i f f e r e n t c u r v e s f o u n d f o r s a l m o n i d s . B o t h Graham (1949) a n d B e a m i s h (1964b) f o u n d t h e r e l a t i o n s h i p b e t w e e n l o g o x y g e n c o n s u m p t i o n a n d t e m p e r a t u r e (10-20°C) t o be l i n e a r f o r S a l v e l i n u s f o n t i n a l i s . B r e t t (1964) i n h i s c o m p r e h e n s i v e s t u d y o f t h e o x y g e n c o n s u m p t i o n o f y o u n g s o c k e y e , f o u n d t h a t t h e c u r v e ; f o r r a t e o f o x y g e n u p t a k e p l o t t e d a g a i n s t t e m p e r a t u r e ( s e m i - l o g ) was c o n c a v e o v e r t h e r a n g e 0-25°C, L a c k i n g d e t a i l e d i n f o r m a t i o n o n t h e t e m p e r a t u r e r e l a t i o n s i n M a r i o n L a k e f i s h i t was d e c i d e d t o c a l c u l a t e a new s e t o f m u l t i p l i e r s ( c f . W i n b e r g , 1956) b a s e d o n t h e c o n c a v e c u r v e o f B r e t t (1964). The r a t e o f o x y g e n c o n s u m p t i o n a t v a r i o u s t e m p e r a t u r e s was 0 t r a n s f o r m e d t o t h e r a t e a t 15 C b y m u l t i p l y i n g t h e o b s e r v e d v a l u e b y 15 - , t 0 ( Q 1 0 ) 10 w h e r e Q i s d e t e r m i n e d f r o m B r e t t ' s c u r v e a n d t Q i s t h e t e m p e r a t u r e o f 65 t h e e x p e r i m e n t . The v a l u e o f t h e m u l t i p l i e r a t t e m p e r a t u r e s o f 6 C a n d l e s s was d e t e r m i n e d b y e x t r a p o l a t i o n . When a l l t h e v a l u e s h a d b e e n t r a n s f o r m e d t o 1 5 ° C , l o g o x y g e n c o n s u m p t i o n was p l o t t e d a g a i n s t l o g w e i g h t , a n d a r e g r e s s i o n l i n e c a l c u l a t e d f o r t h e r a i n b o w t r o u t ; "jfi o Q = 62.6 W m l / h r a t 15 C w h e r e W i s i n k g ( s t a n d a r d e r r o r o f a i s - 1.19)• T h i s e q u a t i o n i s r e m a r k a b l y s i m i l a r t o t h a t f o u n d b y W i n b e r g (1956) f o r s a l m o n i d s i n g e n e r a l : , , • 7 6 . o Q = 60.4 W m l / h r a t 15 C ( t r a n s f o r m e d u s i n g h i s t a b l e o f m u l t i p l i e r s ) . I n p r e p a r a t i o n f o r c a l c u l a t i n g t h e p o p u l a t i o n e n e r g y b u d g e t s , t h e e q u a t i o n f o r t h e M a r i o n L a k e f i s h was c o n v e r t e d f r o m m l / h r t o K c a l / d a y a s s u m i n g a r e s p i r a t o r y q u o t i e n t (RQ) o f 0 .82 a n d t h u s 1 l i t r e 0£ consumed e q u a l s 4.825 K c a l e n e r g y u t i l i z e d . The e q u a t i o n i s now o f t h e f o r m Q = 7.25 W * 7 6 K c a l / d a y a t 15°C w h e r e W i s w e i g h t i n k g . The s e a s o n a l change i n r o u t i n e m e t a b o l i s m was • c a l c u l a t e d u s i n g t h e a b o v e e q u a t i o n a d j u s t e d t o t h e mean m o n t h l y t e m p e r a t u r e ( T a b l e 15). D e t e r m i n a t i o n o f r o u t i n e m e t a b o l i c r a t e i n k o k a n e e p r o v e d v e r y d i f f i c u l t . D e s p i t e c a r e f u l h a n d l i n g t h e m o r t a l i t y r a t e was h i g h e v e n a t l o w t e m p e r a t u r e s . P r i o r t o t h e o n s e t o f m a t u r i t y t h e s c a l e s o f t h e s e f i s h 66 T a b l e 15. S e a s o n a l c h a n g e i n r o u t i n e m e t a b o l i c r a t e b a s e d o n mean m o n t h l y t e m p e r a t u r e . M o n t h Mean t e m p . ( ° « M u l t i p l i e r R o u t i n e m e t a b o l i s m K c a l / d a y , W i n k g B a s i c e q u a t i o n J a n . F e b . M a r . A p r . May J u n e J u l y A u g . S e p . O c t . N o v . D e c . 15.0 3.0 3.0 4 .3 6 . 0 11.5 1 4 . 1 1 6 . 2 1 8 . 8 1 4 . 8 9.3 5.5 4 . 0 1 . 0 0 0 1 . 4 2 0 1 . 4 2 0 1 .415 1 . 4 0 4 1.236 1 . 0 6 9 0 . 9 0 3 0.691 1 . 0 1 6 1 . 3 4 0 1 . 4 0 7 1 . 4 1 6 Q = 7.25 W .76 Q = 5.11 W Q a 5.11 W Q a 5.12 W Q = 5.16 W Q = 5.87 W Q » 6.78 W' Q » 8.03 W .76 .76 .76 .76 .76 ,76 .76 .76 Q = 1 0 o 4 9 w Q = 7 . 1 4 w * 7 6 Q a. 5.41 w Q = 5.15 W Q = 5.12 W' .76 .76 .76 67 a r e h i g h l y d e c i d u o u s a n d i t was f o u n d t h a t t h e l o s s o f e v e n a f e w s c a l e s r e s u l t e d i n d e a t h . A l t h o u g h numerous e x p e r i m e n t s w e r e t r i e d u s i n g k o k a n e e , o n l y f o u r w e r e s u c c e s s f u l a n d i n e a c h c a s e t h e f i s h a p p e a r e d t o b e i n p e r f e c t c o n d i t i o n . B e c a u s e o f t h e l i m i t e d s a m p l e s i z e a n d l i m i t e d s i z e r a n g e no r e g r e s s i o n c a l c u l a t i o n was a t t e m p t e d . H o w e v e r , t h e v a l u e s when t r a n s f o r m e d t o 15°C, a r e w i t h i n t h e 95$ c o n f i d e n c e l i m i t s o f t h e r e g r e s s i o n c a l c u l a t e d f o r t h e r a i n b o w t r o u t . I n c a l c u l a t i n g p o p u l a t i o n e n e r g y r e q u i r e m e n t s , t h e r o u t i n e m e t a b o l i c r a t e o f t h e r a i n b o w t r o u t a n d k o k a n e e w e r e c o n s i d e r e d i d e n t i c a l . A C T I V E METABOLISM A n a t t e m p t was made t o d e t e r m i n e t h e n o r m a l a c t i v i t y o f f i s h i n a n a t u r a l e n v i r o n m e n t b y e q u i p p i n g i n d i v i d u a l f i s h w i t h s m a l l u l t r a s o n i c t r a n s m i t t e r s a n d t r a c k i n g t h e i r m o v e m e n t s . T e c h n i c a l d i f f i c u l t i e s a s s o c i a t e d w i t h t h e s i g n a l r a n g e h a v e n o t b e e n overcome a t t h e p r e s e n t t i m e , h o w e v e r t h i s a s p e c t o f t h e p r o g r a m i s t o be c o n t i n u e d . From i n v e s t i g a t i o n s i n w h i c h b o t h t h e s t a n d a r d ( o r r o u t i n e ) a n d a c t i v e m e t a b o l i c r a t e s h a v e b e e n d e t e r m i n e d ( F r y , 1947; G r a h a m , 194-9: J o b , 1955), i t w o u l d a p p e a r t h a t t h e g r e a t e s t d i f f e r e n c e b e t w e e n t h e s e t w o r a t e s i s a b o u t 12 t i m e s ( B r e t t , 1964). However t h e d i f f e r e n c e b e t w e e n t h e n o r m a l r o u t i n e m e t a b o l i c r a t e a n d t h e d a i l y a c t i v e r a t e ( a s o p p o s e d t o t h e maximum s u s t a i n a b l e r a t e ) i s much l e s s . I t was p r o p o s e d b y W i n b e r g (1956) t h a t t h e m e t a b o l i c r a t e o f f i s h u n d e r -68 n a t u r a l c o n d i t i o n s b e c o n s i d e r e d as t w i c e t h e r o u t i n e r a t e . Mann (1965) h a s shown t h a t r e s u l t s o f f e e d i n g e x p e r i m e n t s w i t h n a t u r a l f o o d o f t e n show c l o s e a g r e e m e n t w i t h W i n b e r g ' s p r e d i c t i o n s , a n d s o t h e m e t a b o l i c r e q u i r e m e n t o f t h e Thames f i s h was c o n s i d e r e d t o be t w i c e t h e r o u t i n e r a t e . I n t h e p r e s e n t s t u d y , t h e a c t i v e m e t a b o l i c r a t e was n o t a s s i g n e d a p a r t i c u l a r v a l u e b u t a n e s t i m a t e c o u l d be made o n t h e b a s i s o f t h e a v a i l a b l e e n e r g y r e m a i n i n g a f t e r a l l o t h e r l o s s e s h a d b e e n s u b t r a c t e d f r o m t h e t o t a l i n t a k e e n e r g y . MOISTURE CONTENT i Wunder (1936) o b s e r v e d t h a t when p e r c h w e r e s t a r v e d , t h e m o i s t u r e c o n t e n t i n c r e a s e d f r o m 73.5$ t o 82 .1$ . I d l e r a n d Clemens (1959) f o u n d t h a t s o c k e y e s a l m o n d u r i n g t h e i r u p s t r e a m m i g r a t i o n i n c r e a s e d i n w a t e r c o n t e n t . S e a s o n a l c h a n g e i n m o i s t u r e c o n t e n t was o b s e r v e d b y M c B r i d e e t a l (1959) i n t h e P a c i f i c h e r r i n g . From t h e s e s t u d i e s , a n d o t h e r s ( S w i f t , 1955: B r e t t , 1964: B r e t t e t a l , I969), w h e r e t h e f a t c o n t e n t was a l s o d e t e r m i n e d , i t c a n be shown t h a t t h e r e i s a n i n v e r s e r e l a t i o n s h i p b e t w e e n t h e m o i s t u r e c o n t e n t a n d t h e f a t c o n t e n t . M e a s u r e m e n t o f t h e s e a s o n a l c h a n g e i n m o i s t u r e c o n t e n t may p r o v i d e a n i n d i r e c t m e a s u r e o f t h e s e a s o n a l c h a n g e i n e n e r g y c o n t e n t . F I S H EXCLUDING GONADS I n t h e r a i n b o w t r o u t a n d k o k a n e e f r o m M a r i o n L a k e , t h e s e a s o n a l 69 c h a n g e i n m o i s t u r e c o n t e n t o f t h e w h o l e f i s h e x c l u d i n g gonads was e x a m i n e d as w e l l as t h e c h a n g e i n t h e g o n a d m a t e r i a l . The amount o f m o i s t u r e i s e x p r e s s e d as a p e r c e n t a g e o f l i v e w e i g h t ( F i g . 13, T a b l e 16). D u r i n g t h e s p r i n g , b o t h s p e c i e s e x h i b i t a n i n c r e a s e i n w a t e r c o n t e n t , b u t b y e a r l y summer t h e r e i s a s h a r p d e c l i n e . R a i n b o w t r o u t a n d i m m a t u r e k o k a n e e r e m a i n a t a l e v e l o f a b o u t 76$ w a t e r c o n t e n t u n t i l t h e f o l l o w i n g s p r i n g . I n S e p t e m b e r , a d u l t k o k a n e e b e g i n i n c r e a s i n g t h e i r m o i s t u r e c o n t e n t w h i c h r e a c h e s a maximum l e v e l o f 88.3$ i n November when t h e f i s h a r e i n a s p e n t c o n d i t i o n a n d d y i n g . W i t h t h e e x c e p t i o n o f t h e s e a d u l t k o k a n e e , i n p r e - a n d p o s t - s p a w n i n g c o n d i t i o n , t h e r e m a i n d e r o f t h e k o k a n e e p o p u l a t i o n e x h i b i t a d i s t i n c t l y l o w e r m o i s t u r e c o n t e n t t h a n t h e r a i n b o w t r o u t t h r o u g h o u t t h e y e a r . GONADS The s e a s o n a l c h a n g e i n m o i s t u r e c o n t e n t o f t h e gonads i s b a s e d o n r a t h e r s m a l l s a m p l e s i z e s ( F i g . 14, T a b l e 17). The o v e r a l l p a t t e r n o f c h a n g e i n m o i s t u r e c o n t e n t o f t h e w h o l e f i s h a n d t h e gonads i s s i m i l a r , h o w e v e r t h e t i m e s e q u e n c e a n d t h e m a g n i t u d e o f t h e c h a n g e s a r e d i f f e r e n t . ENERGY CONTENT U n t i l q u i t e r e c e n t l y m o s t s t u d i e s i n v o l v i n g m e t a b o l i s m o r p r o d u c t i o n o f f i s h e s h a v e assumed t h a t t h e e n e r g y c o n t e n t o f f i s h ( f l e s h ) i s e q u a l t o 1 K c a l / g w e t w e i g h t ( W i n b e r g , 1956; M a n n , 1965; H o r t o n e t a l , I968) d e s p i t e t h e g e n e r a l l y a c c e p t e d f a c t t h a t f i s h e s i n n o r t h t e m p e r a t e a r e a s i n c r e a s e t h e i r f a t c o n t e n t d u r i n g t h e summer months a n d u t i l i z e t h e s e F i g u r e 13. S e a s o n a l change i n p e r c e n t a g e m o i s t u r e c o n t e n t o f t h e b o d y ( l e s s g o n a d ) . r I 71 T a b l e 16. S e a s o n a l c h a n g e i n p e r c e n t a g e m o i s t u r e c o n t e n t o f r a i n b o w t r o u t a n d k o k a n e e ( e x c l u d i n g g o n a d s ) (number o f f i s h i n b r a c k e t s ) „ M o n t h M a r . A p r . May-J u n e J u l y A u g . S e p , O c t . N o v . D e c . R a i n b o w t r o u t $ m o i s t u r e m a t u r e i m m a t u r e m a t u r e i m m a t u r e m a t u r e i m m a t u r e m a t u r e i m m a t u r e m a t u r e i m m a t u r e m a t u r e i m m a t u r e m a t u r e i m m a t u r e m a t u r e i m m a t u r e m a t u r e i m m a t u r e m a t u r e i m m a t u r e 78.6 79.6 82.1 80.6 81.2 79.2 77.8 76.7 76,4 W (10) (2) (1) (13) (1) (12) 76.3 (12) 76.5 (10) 75.7 (5) (10) (9) 77.0 (6) K o k a n e e % m o i s t u r e 77.4 (6) 78.7 (6) 78.2 (13) 74.0 (9) 74.8 (12) 76.8 (12) 78.8 (10) 76.0 (4) 84.9 (14) 88.3 (1) 76o4 (8) 75o5 (6) F i g u r e Ik, S e a s o n a l c h a n g e i n p e r c e n t a g e m o i s t u r e c o n t e n t o f t h e g o n a d s . 73 T a b l e 17. S e a s o n a l c h a n g e i n p e r c e n t a g e m o i s t u r e c o n t e n t o f r a i n b o w t r o u t a n d k o k a n e e gonads (number o f f i s h i n b r a c k e t s ) . R a i n b o w t r o u t K o k a n e e M o n t h % m o i s t u r e % m o i s t u r e May f e m a l e 8 1 . 1 (3) 74.7 (7) m a l e 8 2 . 2 (1) -J u n e f e m a l e 8 0 . 6 (5) 76.8 (3) m a l e - 79.3 (5) J u l y f e m a l e 79.4 (2) 72.8 (1) m a l e mm 80 .0 (3) A u g . f e m a l e 68.1 (5) 58.0 (3) m a l e 8 2 . 1 C D 74.9 (5) S e p . f e m a l e - -m a l e 79o0 (3) 75.3 C7) O c t . f e m a l e 57.3 (6) 84 .0 G O m a l e 71.8 (2) 8 4 .3 (9) N o v . f e m a l e 58.7 (5) 88.4 (1) m a l e 70.1 (2) -D e c . f e m a l e 59.3 (4) 70.5 (3) m a l e 70.3 (2) mm 74 r e s e r v e s w h i l e o v e r w i n t e r i n g . The b a s i s o f t h i s a s s u m p t i o n w o u l d a p p e a r t o t o be v a r i o u s b i o c h e m i c a l a n a l y s e s o f f i s h w h i c h d e t e r m i n e d t h e r e l a t i v e p r o p o r t i o n s o f w a t e r , f a t a n d p r o t e i n , a n d t o w h i c h e n e r g y u n i t s w e r e a s s i g n e d . H o w e v e r , d e t a i l e d b i o c h e m i c a l s t u d i e s s u c h as t h a t o f I d l e r & Clemens (1959) o n m i g r a t i n g s o c k e y e s a l m o n i l l u s t r a t e t h e p o t e n t i a l h a z a r d o f a s s u m i n g a c o n s t a n t e n e r g y v a l u e f o r f i s h o D e t e r m i n a t i o n o f t h e e n e r g y c o n t e n t o f a v a r i e t y o f b i o l o g i c a l m a t e r i a l s b y bomb c a l o r i m e t r y h a s g r a d u a l l y become more w i d e s p r e a d as a r e s u l t o f t h e w o r k b y ; R i c h m a n (1958), S l o b o d k i n & R i c h m a n (1961), G o l l e y (1961), S l o b o d k i n (1962) a n d Cummins ( M S , I967). The n e c e s s i t y o f p r o v i d i n g d e t a i l e d i n f o r m a t i o n o n t h e n a t u r e o f t h e m a t e r i a l b u r n e d a n d t h e e x p r e s s i o n o f t h e r e s u l t s h a s b e e n ; e m p h a s i z e d b y Cummins ( M S , I967). I n h i s summary o f a l l c a l o r i f i c v a l u e s a v a i l a b l e u p t o 1967, h e l i s t e d f i v e s p e c i e s o f f i s h (no s a l m o n i d s ) . S i n c e t h e n , B r o c k s e n e t a l (I968) h a v e p u b l i s h e d t h e ,• r e s u l t s o f t h e i r bomb c a l o r i m e t r y o f t h e s c u l p i n , C o t t u s p e r p l e x u s (17 s a m p l e s ) , a n d o f t h e c u t t h r o a t t r o u t , S a l m o c l a r k i ( 2 8 s a m p l e s ) . These t r o u t w e r e f e d a t t h r e e d i f f e r e n t r a t i o n l e v e l s w h i c h was r e f l e c t e d i n t h e i r e n e r g y c o n t e n t . The v a l u e s w h i c h r a n g e d f r o m 4136 c a l / g t o 5174 c a l / g w o u l d a p p e a r t o b e l o w . However as Cummins ( M S , I967) h a s p o i n t e d o u t , i t i s v e r y d i f f i c u l t t o make c o m p a r i s o n s b e t w e e n d i f f e r e n t s e t s o f d a t a when d e t a i l s a r e l a c k i n g . F o r e x a m p l e B r o c k s e n e t a l d i d n o t i n c l u d e a n y i n f o r m a t i o n o n t h e s i z e o f t h e f i s h e x a m i n e d , t h e s e a s o n , t h e m a t e r i a l u s e d i n t h e s a m p l e s ( f o r e x a m p l e i f t h e w h o l e f i s h was u s e d w h e t h e r i t i n c l u d e d o r e x c l u d e d t h e gonads o r g u t c o n t e n t s ) , 75 a n d f i n a l l y , o n t h e v a r i a b i l i t y o f t h e a s h c o n t e n t . L a c k i n g t h i s i n f o r m a t i o n , c o m p a r i s o n w i t h t h e p r e s e n t s t u d y i s n o t p o s s i b l e 0 C A L O R I F I C CONTENT OF THE F I S H EXCLUDING GONADS M a t u r e a n d i m m a t u r e r a i n b o w t r o u t i n M a r i o n L a k e o v e r w i n t e r w i t h a n a v e r a g e b o d y c o n t e n t o f 6100 c a l / a s h - f r e e g . B y m i d - F e b r u a r y , t h o s e f i s h a p p r o a c h i n g s p a w n i n g c o n d i t i o n b e g i n a g r a d u a l d e c l i n e i n e n e r g y c o n t e n t w h i c h c o n t i n u e s u n t i l c o m p l e t i o n o f s p a w n i n g ( F i g . 15). D u r i n g t h e p r e - a n d p o s t - s p a w n i n g p e r i o d , t h e i m m a t u r e t r o u t m a i n t a i n a r e l a t i v e l y c o n s t a n t e n e r g y l e v e l c l o s e t o t h a t o b s e r v e d d u r i n g t h e w i n t e r m o n t h s . Once t h e m a t u r e f i s h h a v e c o m p l e t e d s p a w n i n g t h e r e i s a s h a r p i n c r e a s e i n t h e i r a v e r a g e e n e r g y c o n t e n t t o a s e a s o n a l maximum i n A u g u s t w h i c h r e f l e c t s t h e i n c r e a s e i n f a t c o n t e n t . From t h e A u g u s t l e v e l u n t i l t h e f o l l o w i n g s p r i n g , t h e r e i s no s i g n i f i c a n t d i f f e r e n c e b e t w e e n t h e e n e r g y l e v e l s o f m a t u r e a n d i m m a t u r e f i s h . K o k a n e e i n M a r i o n L a k e a p p e a r t o o v e r w i n t e r a t a h i g h e r a v e r a g e b o d y e n e r g y c o n t e n t t h a n t h e t r o u t (6225 c a l / a s h - f r e e g) ( F i g . 16). The s e a s o n a l maximum e n e r g y l e v e l i s r e a c h e d b y J u l y . A t t h i s t i m e i t i s p o s s i b l e t o s e p a r a t e t h o s e f i s h w h i c h w i l l s p a w n i n t h e c u r r e n t y e a r f r o m t h o s e s t i l l i n a n i m m a t u r e s t a t e . The m a t u r e k o k a n e e b e g i n * a d e c l i n e i n t h e a v e r a g e e n e r g y c o n t e n t i n A u g u s t , w h i c h i n t e n s i f i e s i n t h e 4-6 week p e r i o d p r i o r t o s p a w n i n g . The g r e a t l o s s o f e n e r g y p r i o r t o s p a w n i n g i n b o t h r a i n b o w t r o u t a n d k o k a n e e was t h o u g h t a t f i r s t t o r e f l e c t t h e p r o c e s s o f f i n a l m a t u r a t i o n F i g u r e 15. S e a s o n a l c h a n g e i n e n e r g y c o n t e n t ( c a l / a s h - f r e e g , mean * s t a n d a r d d e v i a t i o n ) o f t h e f i s h e x c l u d i n g gonad i n r a i n b o w t r o u t . o> (1) <D (_ c_ d) a ro u 6 4 0 0 6 3 0 0 6 2 0 0 6100 • 6 0 0 0 5 9 0 0 5 8 0 0 5 7 0 0 5 6 0 0 -5 5 0 0 5 4 0 0 •«:::«-J A N F E B M A R APR S A M P L E , A , ci7p (ma tu re ) O - 4 S I Z E RAINBOW TROUT ( i m m a t u r e ) J 2 8 MAY 8 5 1 JUN 2 10 J U L 6 6 A U G 7 3 SEP 2 3 O C T - 10 NOV DEC F i g u r e 16. S e a s o n a l c h a n g e i n e n e r g y c o n t e n t ( c a l / a s h - f r e e g , mean -s t a n d a r d d e v i a t i o n ) o f t h e f i s h e x c l u d i n g gonad i n k o k a n e e . 6400 O) <D <D t_ M— i _C if) CD (_ <D a To u 6300 6200 6100 6000 5900 5800 5700 5600 5500 5400 • 5300 SAMPLE S'ZE KOKANEE [™™nL A ( immature) A 7 8 o f t h e g o n a d s , h o w e v e r t h i s was n o t t h e c a s e , as i s shown l a t e r . C A L O R I F I C CONTENT OF THE GONADS The e n e r g y c o n t e n t o f t h e r a i n b o w t r o u t o v a r y i s c o n s i s t e n t l y h i g h e r t h a n t h a t o f t h e r e m a i n d e r o f t h e b o d y , b u t u n e x p e c t e d l y e x h i b i t s t h e same p a t t e r n o f s e a s o n a l c h a n g e as t h e b o d y ( F i g . 17). A t a n y g i v e n t i m e o f y e a r t h e e n e r g y c o n t e n t o f t h e r a i n b o w t r o u t t e s t e s i s l o w e r t h a n t h a t o f t h e m a l e b o d y . I n t h e p e r i o d p r i o r t o s p a w n i n g t h e p a t t e r n o f c h a n g e i n e n e r g y i s u n l i k e t h a t o f , t h e o v a r y o r t h e b o d y . I n o r d e r t o c a l c u l a t e t h e amount o f e n e r g y a s s o c i a t e d w i t h t h e gonads o f i m m a t u r e r a i n b o w t r o u t f o r t h e p o p u l a t i o n e n e r g y b u d g e t , .. t h e b e s t e s t i m a t e i s t a k e n t o be t h a t e n e r g y l e v e l r e a c h e d b y t h e m a t u r e gonads one month a f t e r s p a w n i n g ( o v a r i e s - 6100 c a l / a s h - f r e e g , t e s t e s 5600 c a l / a s h - f r e e g ) . 1 The o v e r w i n t e r i n g e n e r g y c o n t e n t o f k o k a n e e gonads i s assumed t o b e a b o u t 6300 c a l / a s h - f r e e g b a s e d o n i m m a t u r e f i s h c a p t u r e d i n December ( F i g . 1 8 ) . T h i s v a l u e i s u s e d a l s o i n t h e c a l c u l a t i o n o f t h e t o t a l amount o f e n e r g y c o n t a i n e d i n i m m a t u r e gonads d u r i n g t h e y e a r . , As i n t h e r a i n b o w t r o u t , t h e p a t t e r n o f s e a s o n a l c h a n g e i n t h e e n e r g y c o n t e n t o f t h e o v a r i e s i s s i m i l a r t o t h a t o f t h e b o d y b u t a t a somewhat h i g h e r l e v e l . 1 K o k a n e e t e s t e s d u r i n g m a t u r a t i o n a p p e a r t o u n d e r g o a s t e a d y d e c r e a s e i n e n e r g y c o n t e n t f r o m a w i n t e r l e v e l o f a b o u t 6300 c a l / a s h -f r e e g t o a p r e s p a w n i n g l o w o f 5200-5300 c a l / a s h - f r e e g . A f t e r s p a w n i n g F i g u r e 17. S e a s o n a l c h a n g e i n e n e r g y c o n t e n t ( c a l / a s h - f r e e g , mean -s t a n d a r d d e v i a t i o n ) o f t h e gonads i n r a i n b o w t r o u t . 7 0 0 0 F i g u r e 1 8 . S e a s o n a l c h a n g e i n e n e r g y c o n t e n t ( c a l / a s h - f r e e g , mean -s t a n d a r d d e v i a t i o n ) o f t h e gonads i n k o k a n e e . D ) 0) CD £_ *7 n CO ro £_ CD CL ro u 7 0 0 0 6 8 0 0 6 6 0 0 6 4 0 0 6 2 0 0 6 0 0 0 5 8 0 0 5 6 0 0 5 4 0 0 5 2 0 0 5 0 0 0 SAMPLE SIZE JAN KOKANEE FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC (ovaries) X 7 3 1 3 2 1 1 ( immature) ( testes) A - 5 3 4 5 6 • 2 8 1 t h e r e i s a m a r k e d i n c r e a s e i n t h e e n e r g y c o n t e n t . The d i s t i n c t i v e p a t t e r n o f s e a s o n a l . c h a n g e i n e n e r g y c o n t e n t o f t e s t e s , i n b o t h r a i n b o w t r o u t a n d k o k a n e e , i s t h o u g h t t o be d u e t o a d i l u t i o n e f f e c t o f t h e s p e r m . I n b o t h s p e c i e s t h e t i m i n g o f t h e d e c l i n e i n e n e r g y c o n t e n t c o r r e s p o n d s e x a c t l y t o t h a t o f t h e i n c r e a s e i n w e i g h t o f t h e t e s t e s . A s s u m i n g t h a t t h e t e s t i c u l a r t i s s u e h a s a n e n e r g y c o n t e n t s i m i l a r t o t h a t o f t h e a v e r a g e b o d y t i s s \ i e , a n d t h e s p e r m h a s l i t t l e o r no e n e r g y c o n t e n t , t h i s w o u l d e x p l a i n t h e s t e a d y d e c l i n e i n e n e r g y c o n t e n t o f t h e k o k a n e e t e s t e s a n d t h e l o w w i n t e r l e v e l i n t h e m a t u r e r a i n b o w t r o u t . The d i s c h a r g e o f l o w e n e r g y s p e r m a t s p a w n i n g t i m e i s t h e o n l y o b v i o u s e x p l a n a t i o n f o r t h e s h a r p i n c r e a s e i n a v e r a g e e n e r g y c o n t e n t i n t h e t e s t e s o f s p e n t f i s h . ASH CONTENT F o l l o w i n g t h e c o m b u s t i o n o f e a c h s a m p l e i n t h e c a l o r i m e t e r , t h e amount o f d r i e d a s h m a t e r i a l r e m a i n i n g i n t h e c r u c i b l e was w e i g h e d . Cummins ( M S , 1967) p r o p o s e d t h a t c a l o r i f i c d e t e r m i n a t i o n s b e e x p r e s s e d b o t h i n t e r m s o f c a l / g d r y w e i g h t a n d c a l / a s h - f r e e g b e c a u s e o f t h e v a r i a b i l i t y i n a s h v a l u e s b a s e d o n r e s i d u e w e i g h t s o b t a i n e d i n bomb c a l o r i m e t r y . Much o f t h i s v a r i a b i l i t y a p p e a r s t o be r e l a t e d t o t h e l o s s o f m a t e r i a l f r o m s h a l l o w c r u c i b l e s d u r i n g c o m b u s t i o n . The c u p -s h a p e d c r u c i b l e o f t h e G a l l e n k a m p c a l o r i m e t e r l a r g e l y e l i m i n a t e s t h i s l o s s . I n o n l y t h r e e s a m p l e s was m a t e r i a l o b s e r v e d t o b e l o s t f r o m ' t h e c r u c i b l e . I n e a c h c a s e t h e s a m p l e c o n s i s t e d o f a l a r g e amount (1-1.5 g) o f h i g h e n e r g y m a t e r i a l ( f i s h e g g s ) w h i c h o n i g n i t i o n a p p a r e n t l y 82 d e t o n a t e d r a t h e r t h a n b u r n e d , r e s u l t i n g i n p a r t i c l e s o f i n c o m p l e t e l y c o m b u s t e d m a t e r i a l b e i n g d i s t r i b u t e d o v e r t h e i n t e r i o r o f t h e bomb. The p e r c e n t a s h w i t h i n a s e r i e s o f s a m p l e s f r o m one f i s h r e m a i n e d c o n s t a n t . However t h e p e r c e n t a s h b e t w e e n d i f f e r e n t f i s h was v a r i a b l e . The amount o f a s h was i n v e r s e l y r e l a t e d t o t h e e n e r g y c o n t e n t o f t h e f i s h a n d c o u l d be e x p r e s s e d b y t h e e q u a t i o n 1 — _ = b x + c y w h e r e x e q u a l s $ a s h , y_ e q u a l s e n e r g y c o n t e n t o f s a m p l e ( c a l / a s h - f r e e g ) , a n d b a n d £ a r e c o n s t a n t s . The e q u a t i o n s f o r r a i n b o w t r o u t a n d k o k a n e e a r e as f o l l o w s : R a i n b o w t r o u t 1 = (3.363 x 10~ 6 ) x + (I.375 x 10"4) ( n = 95) y K o k a n e e 1 = (3.489 x 10" 6 )x + (1.391 x 10" 4) ( n = 102) y T h i s t y p e o f r e l a t i o n s h i p i s n o t a p p a r e n t i n t h e d a t a o f M c B r i d e e t a l (1959) o n t h e P a c i f i c h e r r i n g . A l t h o u g h t h e amount o f f a t a n d p r o t e i n v a r i e d s e a s o n a l l y t h e a s h c o n t e n t r e m a i n e d c o n s t a n t . I t w o u l d a p p e a r t h a t t h e i n c r e a s e i n a s h c o n t e n t w i t h a d e c r e a s e i n e n e r g y may r e f l e c t i n p a r t t h e d i f f e r e n c e s i n t h e r e l a t i v e ' p r o p o r t i o n s o f s k e l e t a l m a t e r i a l , p r o t e i n a n d f a t . However i t h a s b e e n 83 o b s e r v e d i n t h e c a l o r i f i c d e t e r m i n a t i o n o f t h e t e s t e s a n d o v a r i e s ( w h i c h c o n s i s t o f w a t e r , f a t a n d p r o t e i n o n l y ) , t h a t t h e amount o f a s h p r e s e n t r e m a i n s f a i r l y c o n s t a n t o v e r a m o d e r a t e r a n g e o f e n e r g y v a l u e s , b u t t h a t t h e r e a r e c o n s i s t e n t d i f f e r e n c e s b e t w e e n t e s t e s ( r a i n b o w t r o u t 8.7$ a s h , k o k a n e e 8.4$) a n d o v a r i e s ( r a i n b o w t r o u t 5.5$ a s h , k o k a n e e 4.4$). S i n c e t h e t e s t e s a r e g e n e r a l l y l o w i n e n e r g y a n d t h e o v a r i e s h i g h , t h e r e l a t i o n s h i p b e t w e e n e n e r g y a n d a s h c o n t e n t i s s t i l l i n v e r s e . ENERGY CHANGE I N MARION L A K E F I S H COMPARED WITH MIGRATORY SOCKEYE SALMON I d l e r a n d Clemens (1959) p u b l i s h e d a d a t a - f i l l e d a c c o u n t o f t h e e n e r g y e x p e n d i t u r e s o f two s t o c k s o f F r a s e r R i v e r s o c k e y e s a l m o n d u r i n g t h e i r m i g r a t i o n . A d u l t f i s h w e r e s a m p l e d a t v a r i o u s p o i n t s a l o n g t h e i r f r e s h w a t e r m i g r a t i o n r o u t e . The c h a n g e i n m o i s t u r e c o n t e n t , amount o f f a t a n d amount o f p r o t e i n i n t h e b o d y a n d v i s c e r a w e r e d e t e r m i n e d a n d e x p r e s s e d i n grams a n d p e r c e n t a g e s , c o r r e c t e d t o a , " s t a n d a r d " s i z e f i s h . U s i n g t h e c o n v e r s i o n 1 g f a t e q u a l s 9*3 K c a l a n d l g p r o t e i n e q u a l s 4.1 K c a l t h e r a t e o f l o s s o f e n e r g y i n t h e b o d y d u r i n g t h e m i g r a t i o n was c a l c u l a t e d b y t h e m a n d e x p r e s s e d as K c a l / k g / k m . From t h e d a t a g i v e n i n t h e i r t a b l e s (59-62) i t was p o s s i b l e t o c a l c u l a t e t h e e n e r g y c o n t e n t o f t h e w h o l e f i s h e x c l u d i n g g o n a d s , o n a d r y w e i g h t b a s i s . I n o r d e r t h a t t h e s e v a l u e s b e c o m p a r a b l e t o t h o s e d e t e r m i n e d f o r M a r i o n L a k e f i s h , t h e c o n v e r s i o n v a l u e f o r p r o t e i n was s e t a t 1 g p r o t e i n e q u a l s 5»7 K c a l , w h i c h more c l o s e l y r e p r e s e n t s 8 4 t h e e n e r g y r e l e a s e d o n c o m b u s t i o n o f p r o t e i n as o p p o s e d t o t h e l e s s e r amount o f e n e r g y (4.1 K c a l / g ) r e l e a s e d b y m e t a b o l i c p r o c e s s e s ( B r o d y , 1945). The c o n v e r s i o n v a l u e u s e d f o r f a t was 9.5 K c a l / g ( l o c . c i t . ) . A n a d d i t i o n a l c o r r e c t i o n f a c t o r was a p p l i e d t o t h e w e i g h t i n grams o f p r o t e i n d a t a p u b l i s h e d b y I d l e r a n d Clemens (1959). T h e i r p r o c e d u r e i n d e t e r m i n i n g t h e amount o f p r o t e i n i n a g i v e n s a m p l e was b y " c o n v e n t i o n a l m a c r o K j e l d a h l m e t h o d " . E x a m i n a t i o n o f t h e i r n i t r o g e n t o p r o t e i n c o n v e r s i o n i n d i c a t e s t h a t t h e y m u l t i p l i e d t h e amount o f ; n i t r o g e n b y a f a c t o r o f 6.25 t o g i v e t h e amount o f p r o t e i n i n g r a m s . However i t was shown b y L o v e (1957) t h a t t h e m o s t l i k e l y c o n v e r s i o n f a c t o r f o r f i s h p r o t e i n i s 6.025. Thus a l l w e i g h t s o f p r o t e i n u s e d i n r e - c a l c u l a t i o n o f d a t a f r o m I d l e r & Clemens (1959) h a v e b e e n m u l t i p l i e d b y 6.025/6.25 = 0.964. The t i m e p e r i o d , o v e r w h i c h t h e e n e r g y l o s s o c c u r s , i s a p p r o x i m a t e l y t h e same i n e a c h p o p u l a t i o n . F o r t h e S t u a r t r u n t h e e l a p s e d t i m e , f o r m i g r a t i o n a n d s p a w n i n g , b e t w e e n A l b i o n a n d d e a t h o n t h e s p a w n i n g g r o u n d s , i s 36 d a y s , w h e r e a s i n t h e C h i l k o r u n t h e t i m e i s 43 d a y s • I n M a r i o n L a k e k o k a n e e , t h e t o t a l t i m e s p a n b e t w e e n t h e mid-summer maximum e n e r g y c o n t e n t ( p r o b a b l y c o m p a r a b l e t o t h a t o f t h e s o c k e y e s p r i o r t o t h e i r o c e a n m i g r a t i o n ) a n d p o s t - s p a w n i n g d e a t h i s a b o u t 4 months ( F i g . 16). H o w e v e r , i n c o m p a r i n g t h e e n e r g y l o s s o f t h e s o c k e y e s a n d k o k a n e e s , o n l y l o s s e s w h i c h o c c u r 45 d a y s p r i o r t o d e a t h a r e u s e d f o r t h e l a t t e r s p e c i e s . The e n e r g y l o s s i n m a t u r e r a i n b o w t r o u t a l s o o c c u r s o v e r a 45 d a y p e r i o d ( F i g . 15). The s i m i l a r i t y b e t w e e n t h e s m a l l k o k a n e e a n d t h e much l a r g e r 85 s o c k e y e s a l m o n a t s p a w n i n g t i m e i s e v i d e n t i n t h e amount o f g o n a d m a t e r i a l p r e s e n t , e x p r e s s e d as a p e r c e n t a g e o f b o d y w e i g h t ( k o k a n e e t e s t e s 3*9$, s o c k e y e t e s t e s 3*3$, k o k a n e e o v a r i e s 13.2$, s o c k e y e o v a r i e s 13.7$) ( K i z e v e t t e r , 19^8; I d l e r & C l e m e n s , 1959). I n M a r i o n L a k e f i s h t h e e n e r g y c o n t e n t o f t h e w h o l e f i s h l e s s gonads was n o t s i g n i f i c a n t l y d i f f e r e n t f o r m a l e s a n d f e m a l e s s o a l l d a t a w e r e c o m b i n e d - As s h o w n i n T a b l e 18 t h e e n e r g y l o s s e s o c c u r r i n g i n m i g r a t o r y a n d n o n - m i g r a t o r y f i s h a r e q u i t e s i m i l a r . The e n e r g y l o s t b y t h e M a r i o n L a k e f i s h m i g h t b e g r e a t e r i f i t w e r e n o t f o r t h e f a c t t h a t b o t h t h e r a i n b o w t r o u t a n d k o k a n e e c o n t i n u e t o f e e d , e v e n t h o u g h a t a r e d u c e d l e v e l , d u r i n g t h i s p e r i o d . The l o s s o f e n e r g y f r o m t h e w h o l e f i s h e x c l u d i n g gonads was; t h o u g h t a t f i r s t t o be a r e d i s t r i b u t i o n p r o c e s s w h e r e b y e n e r g y s t o r e d i n t h e b o d y was u t i l i z e d i n t h e f i n a l g o n a d m a t u r a t i o n p r o c e s s o r t h e g e n e r a l i n c r e a s e i n t h e mass o f t h e gonads p r i o r t o s p a w n i n g . T h i s e x p l a n a t i o n was r e j e c t e d f o r t w o r e a s o n s . F i r s t , when t h e e n e r g y c o n t e n t o f t h e t e s t e s a n d o v a r i e s was e x a m i n e d f o r s e a s o n a l d i f f e r e n c e s , i i t was f o u n d u n e x p e c t e d l y t h a t t h e c h a n g e i n e n e r g y c o n t e n t o f t h e • o v a r i e s was a n a l m o s t p e r f e c t r e f l e c t i o n o f t h e s e a s o n a l c h a n g e i n t h e e n e r g y c o n t e n t o f t h e f i s h e x c l u d i n g g o n a d s , a n d t h a t t h e a v e r a g e e n e r g y c o n t e n t o f t h e t e s t e s was l o w ( r a i n b o w t r o u t ) o r d e c r e a s i n g , ( k o k a n e e ) i n t h e p e r i o d p r i o r t o s p a w n i n g ( F i g s . 17 & 1 8 ) . Thus t h e l o s s i n e n e r g y f r o m t h e b o d y was n o t b e i n g t r a n s f e r r e d t o t h e g o n a d s . S e c o n d , a l t h o u g h t h e i n c r e a s e i n t h e w e i g h t o f t h e gonads i n k o k a n e e c o r r e s p o n d e d i n t i m e t o t h e g e n e r a l d e c r e a s e i n b o d y e n e r g y i 86 T a b l e 1 8 . E n e r g y l o s s ( c a l / g d r y w e i g h t ) d u r i n g m a t u r a t i o n i n M a r i o n L a k e r a i n b o w t r o u t a n d k o k a n e e , a n d m i g r a t o r y F r a s e r R i v e r s o c k e y e s a l m o n . S o c k e y e s a l m o n C h i l k o r u n A l b i o n t o C h i l k o R . ( s p e n t ) (43 d a y s , 596 km) b S t u a r t r u n A l b i o n t o F o r f a r C r . ( s p e n t ) (36 d a y s , 1023 km) E n e r g y l o s s m a l e s & f e m a l e s * 965 c a l / g d r y 1309 c a l / g d r y m a l e s f e m a l e s , 1269 c a l / g d r y 1504 c a l / g d r y K o k a n e e M a r i o n L a k e S e p . t o m i d - O c t . ( s p e n t ) (45 d a y s ) m a l e s a n d f e m a l e s 960 c a l / g d r y ° R a i n b o w t r o u t M a r i o n L a k e M a r . t o m i d - A p r i l ( s p e n t ) (45 d a y s ) m a l e s a n d f e m a l e s 820 c a l / g d r y C V a l u e s b a s e d o n c h a n g e i n e n e r g y c o n t e n t o f f i s h e x c l u d i n g gonads ( u s i n g c o n v e r s i o n f a c t o r s 1 g f a t = 9*5 K c a l , 1 g p r o t e i n = 5»7 K c a l ) , E n e r g y o f v i s c e r a b a s e d o n f> c o m p o s i t i o n o f v i s c e r a o f C h i l k o R u n . I n c l u d e s a s h c o n t e n t . 87 ( F i g . 6 ) , t h i s was n o t t r u e f o r t h e r a i n b o w t r o u t ( F i g . 7)» I n t h e t r o u t t h e t e s t e s a n d o v a r i e s b e g i n i n c r e a s i n g i n w e i g h t b y l a t e summer a n d e a r l y a u t u m n , r e a c h i n g a l e v e l c l o s e t o maximum b y F e b r u a r y a l m o s t a month p r i o r t o t h e g e n e r a l d e c l i n e i n e n e r g y . E n e r g y l o s s i n t h e k o k a n e e a n d s o c k e y e s a l m o n m i g h t w e l l b e a t t r i b u t e d t o t h e g e n e t i c s o f t h e genus O n c o r h y n c h u s a n d t h e phenomenon o f c o m p l e t e p o s t - s p a w n i n g m o r t a l i t y i f i t w e r e n o t f o r t h e f a c t t h a t t h e r a i n b o w t r o u t e x h i b i t t h e same l o s s . The a v a i l a b i l i t y a n d c o n s u m p t i o n o f f o o d o r t h e l a c k o f i t m u s t a l s o b e r e j e c t e d b e c a u s e t h e r a i n b o w t r o u t a n d k o k a n e e c o n t i n u e r e d u c e d f e e d i n g w h i l e t h e s a l m o n t e r m i n a t e f e e d i n g . The e n e r g y c o s t f o r m i g r a t i o n i s o b v i o u s l y n o t as g r e a t as s u g g e s t e d b y I d l e r & C lemens (1959) b e c a u s e o f t h e s i m i l a r l o s s e s i n t h e s e n o n - m i g r a t o r y s a l m o n i d s . T h i s h a s b e e n f u r t h e r s u b s t a n t i a t e d b y R o b e r t s o n e t a l (196l) who f o u n d t h a t t h e d e g e n e r a t i v e c h a n g e s o c c u r r i n g i n s t e e l h e a d t h a t m i g r a t e d 5 m i l e s u p s t r e a m t o s p a w n w e r e as g r e a t as i n t h o s e w h i c h h a d m i g r a t e d 160 m i l e s . A n i n c r e a s e i n a c t i v i t y p r i o r t o a n d d u r i n g s p a w n i n g p r o b a b l y a c c o u n t s f o r some o f t h e e n e r g y l o s s b u t i t i s u n l i k e l y t h a t l o s s e s e x h i b i t e d b y t h e f i s h e s i n M a r i o n L a k e w o u l d be s o s i m i l a r t o t h o s e i n f i s h m i g r a t i n g g r e a t d i s t a n c e s i f a c t i v i t y l e v e l s w e r e t h e s o l e c a u s e o f t h i s l o s s . T h e r e r e m a i n s h o w e v e r a t l e a s t one f a c t o r , common t o b o t h t h e M a r i o n L a k e f i s h a n d t h e m i g r a t i n g s o c k e y e s a l m o n , f o r w h i c h t h e e n e r g y r e q u i r e m e n t s a r e n o t k n o w n , a n d t h a t i s t h e c h a n g e i n p h y s i o l o g i c a l 8 8 s t a t e . P r o b a b l y t h e m o s t d r a m a t i c c h a n g e s o c c u r i n t h e s o c k e y e . The b r i g h t s i l v e r y s i d e s a n d b l u e - b l a c k c o l o u r o f t h e b a c k a r e r e p l a c e d b y b r i g h t r e d o n t h e b o d y a n d o l i v e g r e e n c o l o u r a t i o n o n t h e h e a d . T h e r e i s a l s o a c o n s p i c u o u s t h i c k e n i n g o f t h e s k i n a n d a b s o r p t i o n o f t h e s c a l e s i n b o t h s e x e s w h i c h may s e r v e t o p r o t e c t t h e f i s h f r o m i n j u r y o r t h e i n v a s i o n o f f u n g u s a n d b a c t e r i a , e t c . ( V a n O o s t e n , 1957)• M a r i o n L a k e k o k a n e e do n o t e x h i b i t a d e v e l o p m e n t o f t h e r e d c o l o u r a t i o n o r a n y o f t h e a b o v e s e c o n d a r y s e x u a l c h a r a c t e r s e x c e p t f o r a m o d e s t e n l a r g e m e n t o f t h e t e e t h . However t h e r e i s a c o n s p i c u o u s c h a n g e i n t h e c o n d i t i o n o f t h e s k i n . D u r i n g t h e summer t h e s i d e s . a n d b e l l y o f m a t u r e a n d i m m a t u r e k o k a n e e a r e c o v e r e d w i t h b r i g h t , s i l v e r y and h i g h l y d e c i d u o u s s c a l e s . B y e a r l y a u t u m n , a t a t i m e c o r r e s p o n d i n g t o t h e l o s s o f e n e r g y i n t h e m a t u r e k o k a n e e , t h e s e s c a l e s a r e r e s p r b e d o r m o d i f i e d i n t o a t o u g h " i m p e r v i o u s " l a y e r , a c c o m p a n i e d b y a g e n e r a l t h i c k e n i n g o f t h e s k i n w h i c h becomes b l a c k i n c o l o u r . R o b e r t s o n (19&1) i n h i s s t u d y o f t h e e f f e c t s o f c a s t r a t i o n o n k o k a n e e , o b s e r v e d t h a t t h e r e s o r p t i o n o f s c a l e s a n d t h i c k e n i n g o f t h e s k i n o c c u r r e d i n t h e f i n a l s t a g e s o f m a t u r a t i o n o f n o r m a l f i s h as w e l l as i n t h o s e w h i c h h a d r e g e n e r a t e d s m a l l amounts^ (200 mg o r l e s s ) o f g o n a d a l t i s s u e , . Those f i s h w h i c h h a d b e e n c o m p l e t e l y c a s t r a t e d e x h i b i t e d none o f t h e c h a n g e s i n s e c o n d a r y s e x u a l c h a r a c t e r s a n d d i d n o t d i e i n t h e i r 4th y e a r as t h e n o r m a l f i s h d i d , b u t s u r v i v e d up t o t h e i r 9th y e a r . S i m i l a r c h a n g e s i n t h e s k i n a l s o o c c u r i n t h e r a i n b o w t r o u t p r i o r t o s p a w n i n g h o w e v e r t h e e f f e c t i s somewhat l e s s c o n s p i c u o u s , b e c a u s e t h e r a i n b o w s a r e g e n e r a l l y d a r k i n c o l o u r t h r o u g h o u t t h e w i n t e r . 89 N e v e r t h e l e s s t h e r e i s some i n t e n s i f i c a t i o n o f c o l o u r p r i o r t o s p a w n i n g a n d a n a p p a r e n t c h a n g e i n s k i n f o r m . R o b e r t s o n e t a l (1961) h a v e d e m o n s t r a t e d t h a t i n t h e p e r i o d p r i o r t o s p a w n i n g t h e r e i s a w h o l e s e r i e s o f i n t e r n a l c h a n g e s t a k i n g p l a c e . D e g e n e r a t i o n o f t h e p i t u i t a r y , a d r e n a l , s t o m a c h , l i v e r , k i d n e y a n d c a r d i o v a s c u l a r s y s t e m o c c u r s i n . b o t h t r o u t a n d s a l m o n a l t h o u g h t h e e x t e n t o f t h e s e c h a n g e s i s much l e s s i n t h e t r o u t . T h e r e a r e a l s o m a j o r p h y s i o l o g i c a l c h a n g e s o c c u r r i n g i n t h e b l o o d . K i z e v e t t e r (1948) r e c o g n i z e d t h a t some e n e r g y was r e q u i r e d t o p r o v i d e f o r t h e m o r p h o l o g i c a l c h a n g e s b u t made no e s t i m a t e o f t h e amount u s e d f o r t h i s p u r p o s e . The s i m i l a r i t y i n t h e e n e r g y l o s s p r i o r t o s p a w n i n g i n b o t h m i g r a t o r y a n d n o n - m i g r a t o r y s a l m o n i d s w o u l d seem t o be d u e l a r g e l y ; t o t h e m e t a b o l i c c o s t o f t h e s e p h y s i o l o g i c a l c h a n g e s , a n d i t w o u l d : , a p p e a r t h a t t h e e n e r g y r e q u i r e m e n t s f o r m i g r a t i o n h a v e b e e n o v e r e s t i m a t e d . U n t i l s u c h t i m e as t h i s p r o b l e m c a n b e r e - e x a m i n e d e x p e r i m e n t a l l y , i t may be p o s s i b l e t o make a c r u d e e s t i m a t e o f t h e p e r c e n t a g e o f t h e t o t a l e n e r g y l o s t w h i c h r e s u l t s f r o m m i g r a t o r y a c t i v i t y . B r e t t (1965a) i n h i s s t u d y o f t h e F r a s e r R i v e r s o c k e y e s a l m o n ( S t u a r t r u n ) c o n c l u d e d t h a t t h e r a t e o f u p s t r e a m m i g r a t i o n was 2.65 mph (4.26 k m / h r o r 118 c m / s e c ) . The a v e r a g e t e m p e r a t u r e o f t h e r i v e r d u r i n g t h i s r u n i s 15.6°C ( I d l e r & C l e m e n s , 1959). A swimming r a t e o f 118 cm/sec ( a t 15°C) i s assumed t o be a b o u t 3/4 maximum s u s t a i n e d r a t e f o r l a r g e s o c k e y e s ( B r e t t , 1965b). The r a t e o f o x y g e n u p t a k e (y_) i n mg/hr a t t h e 3/^  maximum a c t i v i t y l e v e l c a n be c a l c u l a t e d f r o m t h e f o l l o w i n g e q u a t i o n w h e r e x i s t h e w e t w e i g h t o f t h e f i s h i n g r a m s . 90 l o g £ = -.223 + 0.926 l o g x ( B r e t t , 1965b) The a v e r a g e w e i g h t s o f t h e S t u a r t r u n s o c k e y e s a r e f o r m a l e s , 24-75 g a n d f o r f e m a l e s , 2200 g ( I d l e r & B i t n e r s , 1959). S u b s t i t u t i n g t h e s e v a l u e s i n t h e a b o v e e q u a t i o n g i v e s t h e r a t e o f o x y g e n c o n s u m p t i o n a n d b y f u r t h e r c a l c u l a t i o n t h e e n e r g y c o s t f o r swimming u p s t r e a m ( T a b l e 19). T h i s c a l c u l a t e d r a t e o f e n e r g y e x p e n d i t u r e f o r m i g r a t i o n i n b o t h m a l e a n d f e m a l e s o c k e y e s i s a b o u t 0.35 K c a l / k g / k m , w h i c h i s i n c o n t r a s t t o t h e v a l u e s g i v e n b y I d l e r & B i t n e r s (1959) f o r m a l e s 0 . 8 4 K c a l / k g / k m a n d f o r f e m a l e s 1.1 K c a l / k g / k m . T h e s e l a t t e r v a l u e s a r e b a s e d o n t h e d i f f e r e n c e b e t w e e n t h e t o t a l e n e r g y c o n t e n t o f t h e f i s h a t t h e b e g i n n i n g a n d e n d o f t h e m i g r a t i o n . From t h e s e f i g u r e s i t w o u l d a p p e a r t h a t t h e a c t u a l c o s t o f m i g r a t i o n f o r f i s h e s o f t h e S t u a r t r u n i s a b o u t 32-42$ o f t h e t o t a l e n e r g y l o s t . T h i s p e r c e n t a g e w o u l d b e e x p e c t e d t o d e c r e a s e i n f i s h , m i g r a t i n g o v e r s h o r t e r d i s t a n c e s o r i n s l o w e r f l o w i n g r i v e r s . Thus t h e s h o r t e r t h e m i g r a t i o n , t h e l a r g e r t h e e n e r g y r e s e r v e i n t h e e v e n t o f d e l a y s . I n t h e c a s e o f t h e n o n - m i g r a t o r y k o k a n e e , a n y r e s e r v e j w h i c h r e m a i n s i s p r o b a b l y u s e d i n i n c r e a s e d s u r v i v a l t i m e p o s t -s p a w n i n g , h o w e v e r t h e r e m a i n i n g e n e r g y i s i n s u f f i c i e n t t o s u s t a i n t h e s e f i s h o v e r w i n t e r a n d b y s p r i n g m o r t a l i t y i s c o m p l e t e . 91 T a b l e 19. C a l c u l a t e d m e t a b o l i c c o s t o f m i g r a t i o n i n F r a s e r R i v e r s o c k e y e s a l m o n ( S t u a r t r u n ) . M a l e s F e m a l e s A v e r a g e w e i g h t ( g ) 2475 2200** R a t e o f o x y g e n c o n s u m p t i o n " * (mg O g / h r ) 1111 996 ( m l 0 2 / h r ) 777 697 M e t a b o l i c c o s t ( K c a l / h r ° ) 3,69 3.31 ( K c a l / k m d ) .866 .777 ( K c a l / k g / k m ) .350 .353 M e t a b o l i c c o s t a c c o r d i n g t o I d l e r & B i t n e r s (1959) ( K c a l / k g / k m ) .84 1.1 S o u r c e - I d l e r & B i t n e r s (1959) 3 l o g v_ = -.223 + .926 l o g x , w h e r e v_ i s mg 0 , , / h r a n d x i s w e i g h t i n g . ' A s s u m i n g o x y c a l o r i f i c e q u i v a l e n t o f 4.75 K c a l / l 0 ( B r e t t , 1964). A s s u m i n g a swimming r a t e o f 4.26 k m / h r . 92 DISCUSSION U T I L I Z A T I O N OF ENERGY I n o r d e r t o i n t e g r a t e t h e e n e r g y c h a n g e s o u t l i n e d i n t h e p r e v i o u s s e c t i o n s , t h e d a t a w e r e e x p r e s s e d as m o n t h l y r a t e s a n d t h e c a l c u l a t i o n s made b y means o f a n I B M 1130 c o m p u t e r . The l o g i c o f t h e p r o g r a m u s e d i n t h i s a n a l y s i s was as f o l l o w s . F o r t h e f i r s t month t h e t o t a l number o f f i s h p r e s e n t i n e a c h age c l a s s a n d t h e i r a v e r a g e w e t w e i g h t ( k g ) , d e t e r m i n e d p r e v i o u s l y , p r o v i d e d a s t a r t i n g p o i n t . The f o l l o w i n g c a l c u l a t i o n s w e r e made f o r e a c h age g r o u p a t t h e b e g i n n i n g o f e a c h m o n t h . (1) t o t a l w e i g h t ( k g ) f o r .each age g r o u p , (2) t o t a l amount o f e n e r g y ( K c a l ) u s e d i n r o u t i n e m e t a b o l i s m , (3) t o t a l e n e r g y ( K c a l ) c o n t e n t o f t h e f o o d i n t a k e , (4) t o t a l e n e r g y ( K c a l ) l o s t t h r o u g h e g e s t i o n a n d e x c r e t i o n , (5) r a t i o o f m a l e s t o f e m a l e s , (6) number o f m a t u r e a n d i m m a t u r e w i t h i n e a c h age g r o u p , (7) e n e r g y c o n t e n t ( K c a l ) o f t h e b o d y b a s e d o n m o n t h l y v a l u e s o f e n e r g y ( K c a l / a s h - f r e e g ) , $ m o i s t u r e , $ a s h , (8) e n e r g y c o n t e n t ( K c a l ) o f g o n a d s . Once t h i s p h a s e was c o m p l e t e d t h e m o n t h l y , g r o w t h r a t e was a p p l i e d t o e a c h age g r o u p . A t t h e e n d o f t h e m o n t h , t h e new w e i g h t , b o d y e n e r g y a n d g o n a d e n e r g y w e r e r e c a l c u l a t e d . A t t h i s p o i n t t h e m o n t h l y m o r t a l i t y r a t e was a p p l i e d t h u s b e g i n n i n g t h e n e x t m o n t h w i t h f e w e r f i s h . The number o f r e c r u i t s t o t h e p o p u l a t i o n a n d t h e i r a v e r a g e w e i g h t w e r e i n s e r t e d i n t h e a p p r o p r i a t e m o n t h s . On c o m p l e t i o n o f t h e a n a l y s i s f o r a 12 month p e r i o d t h e d a t a ( n u m b e r , w e i g h t , i n t a k e , e g e s t i o n - e x c r e t i o n , r o u t i n e m e t a b o l i s m , b o d y 93 e n e r g y , g o n a d e n e r g y , c h a n g e i n w e i g h t , c h a n g e i n b o d y e n e r g y a n d c h a n g e i n gonad e n e r g y ) w e r e summed b y (1) age g r o u p , ( 2 ) s e x a n d s t a t e o f m a t u r i t y a n d ( 3 ) c o m b i n e d t o t a l f o r e a c h month ( A p p e n d i x t a b l e s ) . EGG SURVIVAL C a l c u l a t i o n o f t h e number o f m a t u r e f e m a l e s o f e a c h age g r o u p p r e s e n t i n t h e l a k e p r i o r t o s p a w n i n g made i t p o s s i b l e t o e s t i m a t e t h e t o t a l a n n u a l egg d e p o s i t i o n ( T a b l e 2 0 ) . The c a l c u l a t e d p e r c e n t s u r v i v a l f r o m egg t o f r y ( e n t e r i n g t h e l a k e ) i s 1.9$ f o r r a i n b o w t r o u t a n d 4.9$ f o r k o k a n e e . M c F a d d e n (19&9) s u g g e s t e d t h a t i n g e n e r a l , t h e egg t o f r y s u r v i v a l i n s a l m o n i d s i s l e s s t h a n 10$. Low s u r v i v a l r a t e s s u c h as t h e s e i n M a r i o n L a k e w e r e a n t i c i p a t e d i n v i e w o f t h e r e s t r i c t e d a r e a a v a i l a b l e f o r s p a w n i n g a n d t h e h i g h l y v a r i a b l e w a t e r - f l o w r a t e s w h i c h o c c u r w h i l e eggs a n d f r y a r e i n t h e g r a v e l . D u r i n g 1968, t h e o n l y y e a r f o r w h i c h c o m p l e t e d a t a a r e a v a i l a b l e , a c o n t i n u o u s m o n i t o r i n g o f t h e f l o w r a t e s o f t h e m a i n i n l e t streams t o M a r i o n L a k e ( b y Canada D e p t . o f E n e r g y , M i n e s & R e s o u r c e s , I n l a n d W a t e r s B r a n c h ) r e v e a l e d d i f f e r e n c e s r a n g i n g f r o m 1.5 t o 867 c u b i c f e e t p e r s e c (.043-24.6 k l / s e c ) . I n c r e a s e s i n f l o w r a t e c a n be v e r y r a p i d d u r i n g h e a v y r a i n s , f o r e x a m p l e t h e l a k e l e v e l h a s b e e n o b s e r v e d t o r i s e o v e r 1.5 m i n a m a t t e r o f a f e w h o u r s . F l o o d i n g s u c h as i t h i s h a s l o n g b e e n r e c o g n i z e d as a m a j o r c a u s e o f m o r t a l i t y i n eggs a n d f r y ( W i c k e t t , 1958; M c N e i l , 1966). I t i s p o s s i b l e t h a t a s i g n i f i c a n t p o r t i o n o f t h e f r y o c c u p y i n g t h e o u t l e t s t r e a m a r e s w e p t d o w n s t r e a m a n d o v e r i m p a s s a b l e f a l l s . T h e y may s u r v i v e t h e f a l l s b u t 94 T a b l e 20. E s t i m a t e d a n n u a l egg d e p o s i t i o n i n M a r i o n L a k e r a i n b o w t r o u t a n d k o k a n e e a n d p e r c e n t s u r v i v a l t o f r y s t a g e . R a i n b o w t r o u t K o k a n e e A g e g r o u p 3 + ^ + 2 3 + a N o . m a t u r e f e m a l e s 269 87 696 196 T o t a l w t . ( k g ) 23.4 17.1 34 .I 17.0 A v e r a g e w t . ( k g ) 87 197 "49 88 b C a l c u l a t e d l e n g t h (cm) 19.7 25.8 16.4 19.7 c A v e r a g e n o . e g g s / f e m a l e 326 422 148 192 T o t a l n o . e g g s / a g e g r o u p x 103 87.7 36.7 103 37.3 3 T o t a l n o . e g g s / s p e c i e s x 10 124 140 A v e r a g e n o . f r y p r o d u c e d 2350 6850 $ s u r v i v a l 1.9$ 4.9$ i a A p r i l 1st f o r r a i n b o w t r o u t , O c t o b e r 1st f o r k o k a n e e . b 3 0 3.6 3 2 32 From W = .0109 L f o r r a i n b o w t r o u t , f r o m W = .0058 L f o r k o k a n e e . c 0.959 , 1 .414 From F = 1 8 . 7 L f o r r a i n b o w t r o u t , f r o m F « 2 . 8 4 L f o r k o k a n e e . 95 a r e c o m p l e t e l y l o s t t o t h e M a r i o n L a k e s y s t e m . I f t h e egg t o f r y s u r v i v a l r a t e i s h i g h e r t h a n h a s b e e n c a l c u l a t e d t h e e f f e c t o n t h e o v e r a l l p r o d u c t i o n f i g u r e s w o u l d b e t o r a i s e t h e m s l i g h t l y , h o w e v e r t h e d i f f e r e n c e i f a n y , w o u l d p r o b a b l y b e s m a l l . ANNUAL ENERGY BUDGET From t h e a p p e n d i x t a b l e s , t h e a n n u a l e n e r g y b u d g e t f o r M a r i o n L a k e f i s h h a s b e e n c a l c u l a t e d ( T a b l e 21). I t s h o u l d be n o t e d t h a t a l l v a l u e s e x p r e s s e d i n K c a l a r e s l i g h t l y h i g h ( e x c e p t m e t a b o l i s m ) s i n c e t h e y a r e b a s e d o n t h e amount o f e n e r g y r e l e a s e d o n c o m b u s t i o n a n d n o t o n m e t a b o l i z a b l e e n e r g y . No c o r r e c t i o n was a t t e m p t e d s i n c e t h e p e r c e n t a g e p r o t e i n c o m p o s i t i o n u n d o u b t e d l y v a r i e d . The e n e r g y r e m a i n i n g a f t e r e g e s t i o n - e x c r e t i o n , g r o w t h ( p r o d u c t i o n ) a n d r o u t i n e m e t a b o l i s m h a v e b e e n s u b t r a c t e d f r o m t o t a l e n e r g y i n t a k e , p r o v i d e s some i n d i c a t i o n o f t h e amount o f e n e r g y u s e d i n a c t i v i t y o r o t h e r w o r k ( T a b l e 21). W i n b e r g (1956), Mann (1965) a n d o t h e r s h a v e a s s u m e d t h a t t h e m e t a b o l i c r a t e i n n a t u r e i s t w i c e t h e : r e s t i n g r a t e * F o r s a l m o n i d s t h i s e s t i m a t e may b e l o w . Mann (1965) i n h i s r e c a l c u l a t i o n o f t h e d a t a f r o m P e n t e l o w (1939) o n b r o w n t r o u t ( S a l m o t r u t t a ) d e m o n s t r a t e d r e m a r k a b l y c l o s e a g r e e m e n t t o t h e v a l u e s p r e d i c t e d b y W i n b e r g . H o w e v e r , P e n t e l o w ' s f i s h were; h e l d i n t r o u g h s a n d p r o b a b l y d i d n o t e x h i b i t t h e amount o f a c t i v i t y t h a t f i s h i n a n a t u r a l s t a t e w o u l d u n d e r g o i n t h e p r o c e s s o f f e e d i n g , d e f e n d i n g t e r r i t o r i e s , a v o i d i n g p r e d a t o r s , e t c . ( C h a p m a n , 1967). M T a b l e 21 . A n n u a l e n e r g y b u d g e t f o r M a r i o n L a k e r a i n b o w t r o u t a n d k o k a n e e , K c a l / m ( x 10 = K c a l / h a ) . R a i n b o w t r o u t K o k a n e e A v e r a g e b i o m a s s B T o t a l f o o d E g e s t i o n - P r o d u c t i o n R o u t i n e i n t a k e e x c r e t i o n m e t a b o l i s m 1.53 (12.3 k g / h a ) 10.72 2.23 ( 1 6 . 6 k g / h a ) 9.67 2 . 1 4 1 . 9 4 1.23 1.58 M 1.43 1.72 R e m a i n i n g e n e r g y F - (E+ P + M) 5.92 4.43 ON 3.76 ( 2 8 . 9 k g / h a ) 20 .39 4 . 0 8 2 . 8 1 3.15 P r o d u c t i o n ( n e t e n e r g y g a i n i n b o d y a n d g o n a d s ) . 97 I n p u b l i c a t i o n s as r e c e n t as t h a t o f F r y (1957), i t was g e n e r a l l y b e l i e v e d t h a t t h e maximum d i f f e r e n c e b e t w e e n r a t e s o f s t a n d a r d m e t a b o l i s m a n d a c t i v e m e t a b o l i s m was 4 t i m e s . S i n c e t h e s t a n d a r d r a t e was c o n s i d e r e d t o b e t h e l o w e s t m e a s u r a b l e r a t e a n d t h e a c t i v e r a t e a c h i e v e d a t t h e h i g h e s t swimming s p e e d , i t was p r o b a b l y n o t a n u n r e a s o n a b l e a s s u m p t i o n t h a t i n n a t u r e n o r m a l a c t i v i t y was a t some i n t e r m e d i a t e l e v e l . M o r e r e c e n t l y i t h a s b e e n shown b y B r e t t (1964) i n h i s s t u d y , o o f y o u n g s o c k e y e , t h a t a t t e m p e r a t u r e s up t o 15 C t h e a c t i v e r a t e s ; w e r e a b o u t 10-12 t i m e s t h e s t a n d a r d r a t e . O n l y as t h e t e m p e r a t u r e , a p p r o a c h e d t h e u p p e r l e t h a l (25°C) d i d t h e a c t i v e r a t e a p p r o a c h 4 t i m e s t h e s t a n d a r d r a t e . B r e t t c o n c l u d e d t h a t e x c i t a b i l i t y h a d n o t b e e n a d e q u a t e l y a c c o u n t e d f o r i n measurement o f s t a n d a r d r a t e s i n p r e v i o u s s t u d i e s a n d a l s o t h a t a c t i v e l e v e l s h a d n o t b e e n b a s e d o n maximum s u s t a i n a b l e r a t e s . Thus t h e " s c o p e f o r a c t i v i t y " ( F r y , 1947) i s p r o b a b l y much g r e a t e r , a t l e a s t i n s a l m o n i d s , t h a n was p r e v i o u s l y e x p e c t e d . T h e r e w o u l d a l s o a p p e a r t o b e a n o t h e r p o t e n t i a l s o u r c e o f e n e r g y u t i l i z a t i o n - t h e c o s t o f h a n d l i n g t h e f o o d . S e p a r a t i o n o f t h i s e n e r g y r e q u i r e m e n t f r o m t h e c o l l e c t i v e m e t a b o l i c c o s t s , r e m a i n s a p r o b l e m . F o r e x a m p l e I v l e v ( i n W i n b e r g , 1956) i n c l u d e d as a p a r t o f t h e p h y s i o l o g i c a l l y u s e f u l e n e r g y a f a c t o r t e r m e d " p r i m a r y h e a t " w h i c h i n one s t u d y amounted t o 45$ o f t h e a v a i l a b l e e n e r g y . W i n b e r g b e l i e v e s . t h a t r e s p i r a t i o n r a t e s u s e d b y I v l e v a r e t o o l o w a n d when e l e v a t e d t o t h e c o r r e c t l e v e l t h e v a l u e o f p r i m a r y h e a t p r o d u c t i o n becomes z e r o . J o b ( i n F r y , 1957) a l s o o b s e r v e d t h a t when f i s h a r e f e d , t h e y 98 h a v e a h i g h e r ( r o u t i n e ) m e t a b o l i c r a t e t h a n f i s h w h i c h a r e f a s t i n g . B r o c k s e n e t a l (1968) h a v e c a l c u l a t e d t h e e n e r g y r e q u i r e m e n t s f o r f o o d h a n d l i n g ( " s p e c i f i c d y n a m i c a c t i o n " ) i n a s e r i e s o f e x p e r i m e n t s u s i n g c u t t h r o a t t r o u t o n a d i e t o f n a t u r a l f o o d s . T h e y f o u n d t h a t t h e r e was a d i r e c t c o r r e l a t i o n b e t w e e n t h e amount o f f o o d consumed a n d t h e c o s t o f f o o d h a n d l i n g . U n f o r t u n a t e l y no i n f o r m a t i o n was p r o v i d e d o n t h e t e m p e r a t u r e o r d u r a t i o n o f t h e e x p e r i m e n t , o r t h e ',. w e i g h t o f t h e f i s h u s e d , a l t h o u g h t h e t o t a l e n e r g y c o n t e n t o f e a c h f i s h i s g i v e n . I n summary, some e n e r g y i s u n d o u b t e d l y l o s t as h e a t i n t h e m e t a b o l i s m o f t h e f o o d , b u t i t w o u l d a p p e a r t h a t t h i s l o s s i n c o m p a r i s o n t o t h a t e n e r g y u s e d i n t h e a c t u a l f e e d i n g p r o c e s s i s v e r y s m a l l ( J . R . B r e t t , p e r s . c o m m . ) . BIOMASS AND PRODUCTION W i n b e r g (1956), P a l o h e i m o & D i c k i e (1966) a n d o t h e r s h a v e p o i n t e d o u t t h e n e e d t o e x p r e s s t h e c h a n g e s i n p o p u l a t i o n s i n e n e r g y u n i t s ( K c a l ) . H o w e v e r , now t h a t t h i s s t a t e h a s b e e n r e a c h e d i t i s d i f f i c u l t t o make m e a n i n g f u l c o m p a r i s o n s w i t h o t h e r p u b l i s h e d d a t a . F o r s a l m o n i d s t h e a s s u m p t i o n t h a t 1 g o f f i s h ( w e t ) e q u a l s 1 K c a l o f e n e r g y i s p r o b a b l y i n c o r r e c t f o r m o s t e x a m p l e s ( W i n b e r g , 1956; H o r t o n e t a l , 1968). The s e a s o n a l v a r i a t i o n s a r e l a r g e ( F i g s . 15 & 16) a n d e n e r g y e q u i v a l e n t s f o r 1 g o f f i s h ( w e t ) i n r a i n b o w t r o u t c a n r a n g e f r o m 0.90-1.35 K c a l a n d i n k o k a n e e f r o m 0.73-1*55 K c a l b a s e d o n a v e r a g e s o f m a t u r e age g r o u p s . 99 V a r i a t i o n i n i n d i v i d u a l f i s h g r e a t l y e x c e e d t h e s e r a n g e s . A p r o b a b l e a v e r a g e v a l u e c a n b e c a l c u l a t e d f r o m t h e b i o m a s s e s t i m a t e s g i v e n i n T a b l e 21 i n u n i t s o f K c a l / h a a n d k g / h a . The e n e r g y e q u i v a l e n t f o r r a i n b o w t r o u t s o c a l c u l a t e d i s 1.25 K c a l / g a n d f o r k o k a n e e 1.35 K c a l / g , mean f o r b o t h s p e c i e s i s 1.30 K c a l / g . S t a n d i n g c r o p o r b i o m a s s has b e e n e s t i m a t e d f o r a l a r g e number o f f i s h p o p u l a t i o n s ( C a r l a n d e r , 1955; I s a a c & B o n d , 1963). I t w o u l d a p p e a r t h a t t h e n o r m a l r a n g e o f b i o m a s s v a l u e s i n t r o u t l a k e s ir> f r o m 9-32 k g / h a ( C a r l a n d e r , 1955) a l t h o u g h t h e b i o m a s s i n t r o u t s t r e a m s may e x c e e d t h a t o f l a k e s b y 10 o r more t i m e s ( A l l e n , 1951; M c F a d d e n & C o o p e r , 1962). if. M a r i o n L a k e , i r i / t h a t o t a l f i s h b i o m a s s o f 28.9 k g / h a (3.76 x 10 K c a l / h a ) i s p r o b a b l y t y p i c a l o f s m a l l , r e l a t i v e l y s h a l l o w l a k e s w h i c h a r e c o o l enough t o s u p p o r t a t r o u t p o p u l a t i o n . H o w e v e r i t s h o u l d be n o t e d t h a t M a r i o n L a k e a l s o s u p p o r t s ( d u r i n g t h e summer m o n t h s ) a n a v e r a g e b i o m a s s o f s a l a m a n d e r s (5$ T a r i c h a , 95$ Ambystoma) e q u i v a l e n t t o 27.8 k g / h a ( I . N e i s h , p e r s . c o m m . ) . S t a n d i n g c r o p s o f T a r i c h a  g r a n u l o s a as h i g h as 220 k g / h a h a v e b e e n r e c o r d e d f o r O r e g o n ponds ( I s a a c & B o n d , I963). The t e r m p r o d u c t i o n h a s b e e n a s s i g n e d a v a r i e t y o f m e a n i n g s i n f i s h e r i e s management, some o f w h i c h h a v e b e e n i n t e r p r e t e d i n t h e r e v i e w b y Chapman (I967). The d e f i n i t i o n o f p r o d u c t i o n a s u s e d i n t h e p r e s e n t s t u d y i s b a s e d o n t h a t o f I v l e v (19^5) a n d may b e d e f i n e d a s t h e amount o f e n e r g y e l a b o r a t e d i n e a c h p o p u l a t i o n i n a s p e c i f i e d p e r i o d o f t i m e ( m o n t h ) , w h e t h e r o r n o t a l l o f i t s u r v i v e s t o t h e ; e n d 100 o f t h a t t i m e . B y d e t e r m i n i n g t h e c h a n g e s i n e n e r g y c o n t e n t o f t h e b o d y a n d g o n a d o f t h e f i s h s e p a r a t e l y i t was p o s s i b l e f o r t h e f i r s t t i m e t o d e t e r m i n e t h e p r o d u c t i o n r a t e ( i n e n e r g y u n i t s ) i n d i f f e r e n t m o n t h s . L o s s o f e n e r g y ( n e g a t i v e p r o d u c t i o n ) o c c u r s i n t h e b o d i e s o f b o t h s p e c i e s i n t h e w i n t e r m o n t h s , b u t c h a n g e s i n t h e t o t a l e n e r g y c o n t e n t o f t h e gonads a r e l a r g e l y d e t e r m i n e d b y t h e t i m e o f y e a r a t w h i c h . s p a w n i n g o c c u r s . A n n u a l p r o d u c t i o n i n M a r i o n L a k e r a i n b o w t r o u t i s 1.23 x ID*1" K c a l / h a / y r a n d i n k o k a n e e i s 1.58 x 10^  K c a l / h a / y r . The r a t i o o f p r o d u c t i o n t o a v e r a g e b i o m a s s ( P / B ) i s 0.80 f o r r a i n b o w t r o u t a n d 0.71 f o r k o k a n e e o r 0.75 f o r b o t h s p e c i e s c o m b i n e d . The r a t i o s ( P / B ) g i v e n b y Chapman (1967) f o r s a l m o n i d s r a n g e f r o m 1.0-2.5. Mann (I965) a n d H o r t o n e t a l (I968) h a v e shown t h a t t h e P / B r a t i o s f o r s a l m o n i d s and o t h e r s p e c i e s may b e l e s s t h a n 1. Odum (1959) a n d G e r k i n g (1962) h a v e d e f i n e d f i v e e c o l o g i c a l e f f i c i e n c i e s , t h r e e w i t h i n a g i v e n t r o p h i c l e v e l a n d t w o b e t w e e n t r o p h i c l e v e l s . W i t h i n t h e t r o p h i c l e v e l t h e f i r s t e f f i c i e n c y i s t h a t o f a s s i m i l a t i o n (amount a s s i m i l a t e d / i n t a k e ) w h i c h i n t h e p r e s e n t s t u d y has b e e n assumed t o be 80$ a t a l l i n t a k e l e v e l s . The s e c o n d a n d t h i r d e f f i c i e n c i e s w i t h i n t h e t r o p h i c l e v e l a r e t h e " e c o l o g i c a l g r o w t h e f f i c i e n c y " ( p r o d u c t i o n / t o t a l i n t a k e ) a n d t h e " t i s s u e g r o w t h e f f i c i e n c y " ( p r o d u c t i o n / a s s i m i l a t e d e n e r g y ) w h i c h a r e e q u i v a l e n t t o K-^  a n d K 2 as d e f i n e d b y I v l e v (1945). The v a l u e o f t h e s e l a t t e r ' e f f i c i e n c i e s , f o r a l l age g r o u p s o f M a r i o n L a k e f i s h c o m b i n e d , a r e f o r r a i n b o w t r o u t K n = 11.5$, = 14.3$ a n d f o r k o k a n e e = 16.3$, 101 K 2 = 2 0 . 4 $ . S l o b o d k i n ( I960) p r e d i c t e d t h a t t h e r a t i o o f y i e l d t o i n t a k e ( o r K g ) i n s t a b l e a q u a t i c s y s t e m s i s i n t h e r a n g e o f 5 t o 1 5 $ , w h e r e " y i e l d " i s i n t e r p r e t e d as t o t a l p r o d u c t i o n consumed b y p r e d a t o r s ( M a n n , 1 9 6 5 ) » On t h i s b a s i s t h e M a r i o n L a k e r a i n b o w t r o u t a r e w i t h i n t h e p r e d i c t e d r a n g e b u t t h e v a l u e f o r k o k a n e e i s h i g h e r t h a n p r e d i c t e d , T h e s e h i g h v a l u e s may b e a f u r t h e r r e f l e c t i o n o f t h e u s e o f e n e r g y e q u i v a l e n t s r a t h e r t h a n w e i g h t . A t t h e p r e s e n t t i m e a d e t a i l e d a n a l y s i s o f t h e e n e r g y e f f i c i e n c i e s b e t w e e n t r o p h i c l e v e l s i n M a r i o n L a k e i s n o t p o s s i b l e a l t h o u g h the.^ u l t i m a t e a i m o f t h i s I B P p r o j e c t i s t o p r o v i d e a n e n e r g y b u d g e t f o r a l l s y s t e m s w i t h i n t h e l a k e . U n t i l s u c h t i m e as t h i s i n f o r m a t i o n i s a v a i l a b l e t h e f o l l o w i n g c r u d e e s t i m a t e c a n be made o f t h e " e c o t r o p h i c c o e f f i c i e n t " ( r a t i o o f t o t a l i n t a k e / p r o d u c t i o n i n l o w e r t r o p h i c l e v e l ) , as d e f i n e d b y G e r k i n g ( 1 9 6 2 ) . H a r g r a v e ( M S , I969) e s t i m a t e d t h a t t h e t o t a l b i o m a s s o f b e n t h i c i n v e r t e b r a t e s was 62 k g / h a d r y w e i g h t . U s i n g t h e p e r c e n t m o i s t u r e . : c o n t e n t d a t a g i v e n b y K a r z i n k i n & K r i v o b o k (1964) f o r e a c h g r o u p o f i n v e r t e b r a t e s , t h e t o t a l b i o m a s s l i v e w e i g h t was e s t i m a t e d a t 380 k g / h a . U s i n g t h e c o n v e r s i o n 1 k g = 1000 K c a l ( i n t h e a b s e n c e o f a n y o t h e r , 4 i n f o r m a t i o n ) t h e t o t a l e n e r g y o f t h e b i o m a s s i s assumed t o be 38 x 10 K c a l / h a . I t i s f u r t h e r a s s u m e d t h a t t h e P / B r a t i o f o r t h e b e n t h i c i n v e r t e b r a t e s i s 2 ( E f f o r d - p e r s . c o m m . ) , a n d t h u s t h e t o t a l w o u l d ; 4 be e q u i v a l e n t t o 76 x 10 K c a l / h a . 4 ! .• T o t a l e n e r g y i n t a k e b y b o t h s p e c i e s o f f i s h i s 2 0 . 4 x 10 K c a l / h a , 102 h o w e v e r f r o m t h e s t o m a c h a n a l y s i s i t c a n be shown t h a t t h i s i n c l u d e s a b o u t 10$ m a t e r i a l o f t e r r e s t r i a l o r i g i n a n d a b o u t 5$ o b t a i n e d f r o m t h e w a t e r c o l u m n . The t o t a l i n t a k e d e r i v e d f r o m t h e b e n t h i c i n v e r t e b r a t e s 4 i s a b o u t 1?.3 x 10 K c a l / h a 0 I f t h e amount consumed b y t h e s a l a m a n d e r s i s e q u a l t o t h a t o f t h e f i s h t h e e c o t r o p h i c c o e f f i c i e n t f o r t h e v e r t e b r a t e s o f M a r i o n L a k e i s e q u i v a l e n t t o 0 . 4 6 . P r o d u c t i o n i n M a r i o n L a k e a p p e a r s t o b e l o w a t a l l t r o p h i c l e v e l s ( E f f o r d , 1967)» E f f o r d f o u n d t h a t t h e l o w p r i m a r y p r o d u c t i v i t y was d u e m a i n l y t o t h e v e r y l o w n u t r i e n t l e v e l s i n t h e l a k e i t s e l f a n d t h e n e g l i g i b l e amount o f n u t r i e n t m a t e r i a l e n t e r i n g t h e l a k e f r o m t h e s u r r o u n d i n g a r e a . The m o s t i m p o r t a n t s i n g l e f a c t o r a f f e c t i n g t h e n u t r i e n t c o n t e n t w o u l d a p p e a r t o be t h e h e a v y r a i n f a l l ( 2 4 0 c m / y r a v e r a g e ) w h i c h r e s u l t s i n a h i g h f l u s h i n g r a t e i n t h e l a k e . E f f o r d (1967), a n d D i c k m a n (1969) e x p l o r e o t h e r p o s s i b l e e x p l a n a t i o n s f o r t h e l o w p r i m a r y p r o d u c t i v i t y , b u t t h e f i n a l c o n c l u s i o n w h i c h must be d r a w n i s t h a t t h e s e l o w p r o d u c t i o n l e v e l s p a r t i c u l a r l y o f p h y t o p l a n k t o n a r e r e f l e c t e d a t a l l h i g h e r t r o p h i c l e v e l s up t o t h e f i s h . 103 LITERATURE CITED A L L E E , W. C . 1933* E f f e c t s o f c r o w d i n g - e f f e c t s o f numbers o n amount o f f o o d c o n s u m e d . T r a n s . A m . F i s h . S o c . 63: 25-27. 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B l a c k w e l l S c i e n t i f i c P u b l i c a t i o n s , O x f o r d a n d E d i n b u r g h . W I 3 S I N G , T . E . AND A . D . H A S L E R . I968. C a l o r i f i c v a l u e s o f some i n v e r t e b r a t e s i n L a k e M e n d o t a , W i s c o n s i n . J . F i s h . R e s . B d . C a n a d a 25(11): 2515-2518. WUNDER, W, 1936. [ P h y s i o l o g y o f t h e f r e s h - w a t e r f i s h e s o f C e n t r a l E u r o p e . ] I n R . Demo 11 a n d H . N . M a i e r [ e d j ] H a n d b u c h d e r B i n n e n f i s c h e r e i M i t t e l e u r o p a s . 2(B) E . S c h w e i z e r b a r t , S t u t t g a r t . 115 A p p e n d i x T a b l e 1 . E n e r g y b u d g e t o f m a t u r e a n d i m m a t u r e r a i n b o w t r o u t o f b o t h s e x e s b y m o n t h . K e y t o t a b l e : F o r e a c h age g r o u p Number p r e s e n t a t b e g i n n i n g o f m o n t h m a l e i m m a t u r e f e m a l e i m m a t u r e m a l e m a t u r e f e m a l e m a t u r e W e i g h t ( k g ) p r e s e n t a t b e g i n n i n g o f month I n t a k e e n e r g y ( K c a l ) E g e s t i o n - e x c r e t i o n ( K c a l ) R o u t i n e m e t a b o l i s m ( K c a l ) B o d y e n e r g y ( K c a l ) Gonad e n e r g y ( K c a l ) W e i g h t ( k g ) a t e n d o f month Change i n b o d y e n e r g y ( K c a l ) Change i n g o n a d e n e r g y ( K c a l ) 116 JANUARY 0 + 1 + 2 + 3 + 4+, 0 7 3 3 3 2 2 1 8 3 0 0 1 0 6 4 6 9 2 1 2 2 0 0 0 1 8 7 1 4 8 7 1 0 0 C 3 2 8 9 7 Q < .OOOOOOE 0 0 0 . 1 3 9 3 7 4 E 02 0 . 1 3 9 7 4 8 E 0 2 0 . 1 4 4 5 7 0 E 02 0 .OOOOCOE 0 0 p. 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0 . 1 7 9 6 1 8 E 0 4 0 . 1 1 0 2 7 1 E 0 4 0 .OOOOOOE 00 0 . 2 2 3 8 8 3 E 0 4 0 . 3 1 6 3 2 6 E 0 4 0 . 1 3 1 5 0 1 E 0 4 0 . 8 0 3 2 7 1 E 03 0 .OOOOOOE 00 OoCOOOOOE 0 0 0 . 1 1 6 8 6 3 E 0 4 0 . 1 5 2 7 0 6 E 0 4 0 • 9 4 6 7 3 7 E 0 3 Oi .GOOCOOE CO O.OOOOOOE 0 0 0 .OOOOOOE 00 0 . 2 7 9 9 7 6 E 0 4 0 • 1 7 3 6 9 5 E 0 4 0 .OOOOOOE 00 0 . 2 1 2 4 7 9 E 05 0 . 2 1 1 7 6 4 E 05 0 . 2 1 6 0 4 1 E 0 5 0 . 1 1 3 6 9 4 E 05 Oi .OOOOOOE 00 0 . 2 8 8 2 4 9 E 05 0 . 4 5 4 2 3 9 E 05 0 . 1 4 3 1 2 0 E 0 5 0 . 7 4 8 2 2 2 E 0 4 0 . » OOCOOOE 00 O.OOOOOOE 0 0 c • 1 2 0 3 2 5 E 05 0 . 1 7 1 0 8 9 E 0 5 0 . 9 1 2 0 6 7 E 0 4 0 . .OOOOOOE 00 O.OOOOOOE 0 0 0 .OOOOOOE 00 0 . 3 7 7 1 4 4 E 0 5 0 . 2 0 1 2 3 3 E 05 0 . . 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GOOCOOE 00 0 . 1 1 0 2 6 8 E - 0 5 0 . 5 0 6 6 3 9 E - 0 6 0 . 4 7 6 8 3 7 E - 0 6 0 . 1 0 4 3 0 8 E - 0 6 Oi .OOOOOCE 00 0 • 1 9 0 7 3 4 E - 0 5 0 • 5 0 0 6 7 9 E - 0 5 0 . 8 3 4 4 6 5 E - 0 6 0 . 1 7 5 8 3 3 E - 0 5 Oi . 0 0 0 0 , 0 0 E 00 0.OOOOOOE 0 0 0 . 1 5 9 9 7 1 E 0 3 0 . 2 2 7 4 6 1 E 0 3 0 . 1 2 1 2 5 8 E 03 Oi .OOOOOOE 00 O.OOOOOOE 0 0 0 •OOOOOOE 00 © • 1 6 2 4 2 9 E 0 4 0 . 8 6 6 6 7 7 E 03 0 .OOOOOOE 00 125 OCTOBER 0 + 1+ 2 + 3 + 4 + 8 7 7 3 5 8 2 0 1 4 3 0 1 1 9 2 7 6 9 1 3 4 2 8 0 0 2 0 7 1 6 3 34 0 0 0 3 6 1 7 7 0 0 . 1 5 5 7 3 6 E 02 0 o l 5 5 2 3 8 E 02 0 . 1 5 8 6 0 1 E 0 2 0 . 8 4 4 0 9 2 E 0 1 0 oOOOOOOE 00 0 o 2 i l 6 7 3 E 0 2 0 o 3 3 3 4 5 8 E 0 2 0 o l 0 5 7 3 4 E 0 2 0 . 5 4 9 6 4 1 E 0 1 0 0 OOOOOOE 00 Oo OCOOCOE 00 0 o 8 9 7 6 0 6 E 0 1 0 8 1 2 8 6 1 7 E 0 2 0 o 6 6 7 4 2 2 E 0 1 0 0 OOOOOOE 00 OoOOOOOOE 0 0 0 oOOOOOOE 00 0 . 2 8 4 8 5 1 E 0 2 0 • 1 5 1 1 5 1 E 02 0 oOOOOOOE o'o 0 o 7 3 5 l 7 0 E 0 4 0 e 7 3 2 8 2 0 E 0 4 0 o 7 4 8 6 9 6 E 0 4 0 o 3 9 8 4 6 3 E 0 4 0 oOOOOOOE 00 0 . 9 9 9 2 2 8 E 0 4 0 o l 5 7 4 1 3 E 05 0 . 4 9 9 1 3 0 E 0 4 0 0 2 5 9 4 6 4 E 0 4 0 o OOOOOOE CO OoOOOOOOE 0 0 0 9 4 2 3 7 2 5 E 0 4 0 . 6 0 7 1 5 1 E 0 4 0 0 3 1 5 0 6 4 E 0 4 0 oOOOOOOE 00 OoOOOOOOE 00 0 .OOOOOOE 0 0 0 . 1 3 4 4 6 7 E 0 5 0 „ 7 1 3 5 2 8 E 0 4 0 oOOOOOOE CO 0 o1 4 7 0 3 4 E 0 4 0 o 1 4 6 5 5 4 E 0 4 0 0 1 4 9 7 3 9 E 0 4 0 o 7 9 6 9 2 7 E 0 3 0 0 OOOOOOE 00 Oo 1 9 9 8 4 5 E 0 4 0 . 3 1 4 8 2 6 E 0 4 0 o 9 9 8 2 6 1 E 0 3 0 0 5 1 8 9 2 9 E 0 3 0 oOOOOOOE 00 OoOOOOOOE CO 0 o 8 4 7 4 5 1 E 0 3 0 . 1 2 1 4 3 0 E 0 4 0 o 6 3 0 1 2 8 E 03 0 0 OOOOOOE 00 OoOOOOOOE 00 0 .OOOOOOE 00 0 o 2 6 8 9 3 4 E 0 4 0 0 1 4 2 7 0 5 E 0 4 0 s OOOOOOE 00 0 o l 3 5 l 3 7 E 0 4 0 o l 3 4 8 0 8 E 0 4 0 0 1 3 7 0 2 2 E 0 4 0 o 8 4 8 4 2 8 E 0 3 0 oOOOOOOE 00 C o l 7 0 6 3 3 E 0 4 0 . 2 4 1 0 2 9 E 0 4 0 . 1 0 0 6 8 4 E 0 4 0 o 6 1 2 3 7 7 E 03 0 oOOOOOOE 00 OoOOOOOOE 0 0 e 8 8 9 0 0 5 E 0 3 0 0 1 1 6 8 4 9 E 0 4 0 . 7 0 9 7 4 6 E 03 0 oOOOOOOE 00 OoOOOOOOE 00 0 oOOOOOOE 0 0 0 6 2 1 3 8 2 9 E 0 4 0 o l 3 2 1 0 2 E 0 4 0 e OOOOCOE 00 Oo 2 0 0 2 1 1 E 05 0 o l 9 9 5 7 0 E 05 0 o 2 0 3 8 9 4 E 0 5 0 0 108 5 14 E 0 5 0 oOOOOOOE 00 0 o 2 7 1 7 1 3 E 05 0 o 4 2 8 0 4 3 E 05 0 . 1 3 5 7 2 5 E 0 5 0 o 7 0 5 5 4 6 E 0 4 0 e OOOOOOE 00 OoOOOOOOE 0 0 0 . 1 1 2 2 5 2 E 0 5 0 o l 6 0 8 4 5 E 0 5 n e 8 3 4 6 6 l E 0 4 0 oOOOOOOE 00 OoOOOOOOE 0 0 0 oOOOOOOE 00 0 . 3 4 8 6 0 3 E 0 5 0 0 1 8 4 9 8 0 E 05 0 oOOOOOOE' 00 0 e l 4 3 3 2 4 E 0 2 0 . 1 4 2 8 6 6 E 0 2 0 . 1 4 5 9 6 1 E 0 2 0 . 7 7 6 8 2 1 E 0 1 0 0 OOOOOOE 00 0 . 5 7 9 5 9 0 E 02 0 • 9 1 3 0 5 5 E 02 0 o 2 8 9 5 l 4 E 0 2 0 o 1 5 0 4 9 9 E 02 0 e OOOCOOE 00 OoOOOOOOE 0 0 0 . 3 4 3 1 9 0 E 03 C . 4 9 1 7 5 4 E 0 3 • 0 . 2 5 5 1 8 2 E 03 0 oOOOOOOE 00 OoOOOOOOE 0 0 0 oOOOOOOE 00 0 . 3 5 4 2 6 1 E 0 4 C 0 1 8 7 9 8 3 E 0 4 0 oOOOOOOE 00 O o l 5 5 7 36E 02 0 o l 5 5 2 3 8 E 0 2 0 • 1 5 8 6 0 1 E 0 2 0 o 8 4 4 0 9 2 E 0 1 0 oOOOOOOE 00 0 o 2 1 1 6 7 3 E 02 0 o 3 3 3 4 5 8 E 0 2 0 . 1 0 5 7 3 4 E 0 2 0 o 549 6 4 1 E 0 1 0 oOOOOOOE 00 OoOOOOOOE 0 0 0 o 8 9 7 6 0 6 E 0 1 0 o l 2 8 6 l 7 E 0 2 0 . 6 6 7 4 2 2 E 01 0 oOOOOOOE 00 OoOOCOOOE 0 0 0 oOOOOOOE 00 0 . 2 8 4 8 5 1 E 0 2 0 • 1 5 1 1 5 1 E 02 0 0 OOOOOOE 00 0 o 3 1 7 1 1 8 E 03 . - 0 o 3 1 6 1 0 5 E 0 3 -o . 3 2 2 9 4 9 E 0 3 -0 . 1 7 1 8 7 7 E 0 3 0 oOOOOOOE 00 0 . 4 3 0 3 6 5 E 0 3 - 0 • 6 7 7 9 9 2 E 0 3 -o o 2 1 4 9 7 6 E 0 3 - 0 . 1 1 1 7 5 3 E 03 0 0 OOOOOOE 00 OoOOCOOOE 0 0 - 0 o 6 5 2 5 6 3 E 0 2 - 0 . 9 3 5 0 1 7 E 0 2 -0 o 4 8 5 1 9 7 E 02 0 oOOOOOOE 00 OoOOOOOOE 0 0 0 .OOOOOOE 00 - 0 . 1 4 3 6 0 2 E 0 4 -0 o 7 6 1 9 9 5 E 0 3 0 oOOOOOOE 00 0 . 9 5 3 6 7 4 E - 0 6 0 o 8 6 4 2 6 7 E ' - 0 6 0 o 9 5 3 6 7 4 E --0 6 0 o 7 8 9 7 6 l E - 0 6 0 oOOOOOOE 00 O o 6 1 9 8 8 8 E ~ 0 5 0 . 4 8 8 7 5 8 E ' - 0 5 0 o 2 2 6 4 9 7 E ' -0 5 0 • 8 3 4 4 6 5 E - 0 6 0 oOOOOOOE 00 OoOOOOOOE 0 0 -o « 1 0 8 8 9 4 E 03 - 0 • 1 5 6 0 3 3 E 0 3 -0 0 8 0 9 6 9 4 E 02 0 oOOOOOOE 00 J.OOOOOCE 0 0 0 .OOOOOOE 00 0 • 1 5 7 2 5 6 E 0 4 0 . 8 3 4 4 5 7 E 0 3 0 oOOOOOOE 00 126 0 + 8 4 5 1 1 4 9 0 0 0 . 1 5 0 0 5 3 E 02 0 . 2 0 4 0 3 7 E 02 OoOOOOOOE 0 0 OoOOOOOOE 00 0 o 6 5 4 6 4 1 E 0 4 0 . 8 9 C 1 5 8 E 0 4 OoOOOOOOE 0 0 OoOOCOOOE 0 0 0 . 1 3 0 9 2 8 E 0 4 0 o l 7 8 0 3 l E 0 4 OoOOOOOOE 0 0 OoOOOOOOE 0 0 O o l 2 1 C 2 5 E 0 4 O o l 5 2 8 6 4 E 0 4 OoOOOOOOE CO OoOOOOOOE 0 0 0 o l 8 9 8 5 0 E 05 0 . 2 5 7 7 6 3 E 05 OoOOOOOOE 00 OoOOOOOOE 00 0 o l 3 8 0 9 4 E 02 0 o 5 5 8 6 8 2 E 0 2 OoOOOOOOE CO OoOOOOOOE CO i O , 1 5 0 0 5 3 E 0 2 0 . 2 0 4 0 3 7 E 02 OoOOOOOOE 00 OoOOOOOOE 0 0 -0 . 3 5 2 7 7 2 E 0 3 0 o 4 7 8 9 6 9 E 03 OoOOOOOOE CO 0 ft vy 0 0 0 V "<J 0 C 0 o 8 6 4 2 6 7 E - 0 6 0 . 1 6 6 3 9 3 E - 0 5 OoOOCOOOE 0 0 OoOOOOOOE OC 1+ 3 4 5 7 4 1 2 0 0 0 0 e l 4 9 6 0 1 E 0 2 0 . 3 2 1 3 1 7 E 02 0 . 8 6 7 2 5.2E 0 1 OoOOOOOOE 00 0 . 6 5 2 6 6 7 E - 0 4 0 o l 4 0 1 8 1 E 05 0 . 3 7 8 3 5 7 E 0 4 OoOOOQpOE 0 0 O o l 3 0 5 3 3 E 0 4 0 . 2 8 0 3 6 3 E 0 4 0 o 7 5 6 7 1 5 E 03 OoOOOOOOE 0 0 0 o l 2 0 7 4 7 E 0 4 0 . 2 1 5 8 7 2 E 0 4 0 . 7 9 7 8 4 5 E 03 OoOOOOOOE 00 O o l 3 9 2 7 7 E 05 0 . 4 0 5 9 2 5 E 05 O o l 0 7 8 2 5 E 05 O.OOOOOOE 00 0 . 1 3 7 6 7 8 E C2 0 . 8 7 9 8 1 0 E 02 0 . 2 2 6 3 7 3 E 0 3 O.OOOOOOE 0 0 0 o l 4 9 6 0 1 E 02 0 . 3 2 1 3 1 7 E 0 2 0 . 8 6 7 2 5 2 E 0 1 O.OOOOOOE 0 0 0 . 3 5 1 7 1 1 E 0 3 0 o 7 5 4 2 9 4 E 0 3 0 o l 9 7 1 2 7 E C3 O.OOOOOOE 00 0 . 1 2 5 1 6 9 E - C 5 0 . 2 0 2 6 5 5 E - 0 5 O o 6 5 5 4 9 5 E 0 1 O.OOOOOOE 00 1 9 5 1 2 9 1 5 7 3 4 9 0 o l 5 3 8 6 6 E 0 2 0 o 1 0 1 7 8 3 E 0 2 0 o 1 2 3 8 8 2 E 0 2 0 o 2 7 5 3 8 2 E 0 2 0 . 6 7 1 2 7 8 E 0 4 0 o 4 4 4 0 7 6 E 0 4 0 o 5 4 0 4 6 5 E 0 4 O o l 2 0 1 4 1 E 0 5 0 o l 3 4 2 5 5 E 0 4 0 o 8 8 6 l 5 2 E 0 3 0 . 1 0 8 0 9 3 E 0 4 0 o 2 4 0 2 8 3 E 0 4 0 o 1 2 3 3 5 5 E 0 4 0 o 9 0 1 1 l 5 E 0 3 0 o l 0 4 6 2 0 E 0 4 0 o l 9 1 9 9 0 E 0 4 0 e l 9 4 6 7 4 E 0 5 0 o l 2 8 5 9 1 E 0 5 0 o l 5 4 0 2 3 E 0 5 0 o 3 2 3 1 3 2 E 0 5 0 . 1 4 1 6 0 4 E 0 2 0 • 2 7 8 7 1 1 E 0 2 0 o 3 2 3 3 6 2 E 0 3 0 . 4 9 4 5 1 5 E 0 4 0 • 1 5 3 8 6 6 E 0 2 0 o 1 0 1 7 8 S E 0 2 0 . 1 2 3 8 8 2 E 0 2 0 . 2 7 5 3 8 2 E 0 2 0 . 3 6 1 7 40E 0 3 0 . 2 3 8 9 4 8 E 0 3 0 • 2 8 1 5 8 5 E 0 3 0 . 1 2 5 0 7 8 E 0 4 0 . 1 4 0 0 7 0 E - 0 5 0 . 1 9 0 7 3 4 E - 0 5 0 . 9 3 6 3 4 0 E 0 1 0 • 1 1 3 9 9 1 E 0 4 3 + 4 1 27 33 7 4 0 o 6 0 4 8 3 2 E 0 1 0 . 5 3 0 0 1 1 E 0 1 0 . 6 4 7 7 9 2 E 0 1 0 o l 4 5 2 6 2 E 0 2 O o 3 5 l l 2 5 E 0 4 0 o 2 3 1 2 2 9 E 0 4 0 . 2 8 2 6 1 3 E 0 4 0 o 6 3 3 7 3 9 E 0 4 0 . 7 0 2 2 5 1 E 03 0 o 4 6 2 4 5 8 E 03 0 . 5 6 5 2 2 6 E 03 O o l 2 6 7 4 7 E 0 4 0 . 7 5 3 8 1 4 E 0 3 O o 5 4 8 7 6 3 E 03 0 . 6 3 9 1 7 4 E 0 3 0 o l l 8 0 7 6 E 04 0 o l O l 8 2 8 E C5 0 . 6 6 9 5 7 2 E 0 4 0 o 8 0 5 4 0 3 E 0 4 0 e l 7 0 4 5 0 E 0 5 0 . 7 4 0 6 9 0 E 0 1 0 . 1 4 5 1 2 4 E 02 Q o l 6 9 0 8 8 E 03 0 o 2 6 0 8 5 3 E 0 4 0 o 8 0 4 8 3 2 E 0 1 O o 5 3 0 0 l l E 0 1 0 o 6 4 7 7 9 2 E 0 1 0 . 1 4 5 2 6 2 E 0 2 - 0 o l 8 9 2 l 7 E 03 - 0 o l 2 4 4 l 9 E 03 - O o l 4 7 2 4 2 E 03 - 0 o 6 5 9 7 7 8 E 03 C o 2 8 3 1 2 2 E - 0 6 0 . 2 0 8 6 1 6 E - 0 6 - 0 . 4 8 9 6 1 6 E 0 1 O o 6 0 1 3 0 0 E 03 NOVEMBER 4 + 0 0 0 0 OoOOOOOOE 00 O.OOOOOOE CO OoOOOOOOE 00 OoOOOOOOE GO OoOOOOOOE 00 OoOOOOOOE 00 OoOOOOOOE 00 OoOOOOOOE 00 OoOOOOOOE 00 0 0 OOOOOOE 00 O.OOOOOOE 00 O.OOOOQOE 00 0 0 OOOOOOE 00 OoOOOOOOE 00 OoOOOOOOE 00 OoOOOOOOE 00 OoOOOOOOE CO OoOOOOOOE 00 O.OOOOOOE 00 OoOOOOOOE 00 OoOOOOOOE CO OoOOOOOOE 00 OoOOOOOOE 00 0.OOOOCOE CO OoOOOOOOE 00 OoOOOOOOE 00 OoOOOOOOE 00 O.OOOOOOE CO OoOOOOOOE 00 O.OOOOOOE 00 OoOOOOOOE 00 OoOOOOOOE 00 OoOOOOOOE 00 OoOOOOOOE 00 OoOOOOOOE 00 OoOOOOOOE 00 127 DECEMBER 0 + 1 + 2 + 3 + 4 + 3 1 6 3 3 3 1 8 8 4 0 0 1 1 0 8 7 1 5 1 2 5 26 0 0 1 9 3 1 5 2 3 2 0 0 0 3 3 8 7 1 0 0 . 1 4 4 9 0 3 E 0 2 0 . 1 4 4 3 9 7 E 0 2 0 . 1 4 8 3 4 3 E 0 2 0 . 7 8 5 2 0 2 E 0 1 0 oOOGOOOE 00 0 s 1 9 6 7 5 6 E 02 0 . 3 1 0 0 4 2 E 0 2 0 . 9 8 6 3 2 6 E 0 1 0 . 5 1 0 3 8 1 E 0 1 0 .OOOOOOE 00 Co OOOOOOE 0 0 0 o 8 3 6 8 9 8 E 0 1 0 . 1 1 9 9 3 7 E 0 2 0 . 6 2 8 1 6 2 E 0 1 0 .OOOOOOE 00 0 9 OOOOOOE 0 0 O.OOOOOOE 00 0 . 2 6 6 7 0 2 E 0 2 0 . 1 3 9 3 7 3 E 02 0 •OOOOOOE do 0 . 5 8 9 6 4 0 E 0 4 0 . 5 8 7 5 8 0 E 0 4 0 . 6 0 3 6 3 7 E 0 4 0 . 3 1 9 5 1 3 E 0 4 0 .OOOOOOE 00 Oo 8 C 0 6 3 9 E 0 4 0 . 1 2 6 1 6 2 E 0 5 0 . 4 0 1 3 5 4 E 0 4 0 . 2 0 7 6 8 3 E 0 4 0 •OOOOOOE 00 OoCOOOOOE 0 0 C . 3 4 0 5 4 9 E 0 4 0 . 4 8 8 0 4 6 E 0 4 0 . 2 5 5 6 1 0 E 0 4 0 .OOOOOOE CO 0« OOOOOOE 0 0 O.OOOOOOE 00 0 . 1 0 8 5 2 6 E 0 5 0 . 5 6 7 1 3 6 E 0 4 0 .OOOOOOE 00 0 . 1 1 7 9 2 8 E 0 4 0 . 1 1 7 5 1 6 E 0 4 0 . 1 2 0 7 2 7 E 0 4 0 . 6 3 9 0 2 6 E 03 0 .OOOOOOE 00 0 o l 6 0 1 2 7 E 0 4 0 . 2 5 2 3 2 4 E 0 4 r\ V • S 0 2 7 0 8 E 0 3 0 . 4 1 5 3 6 7 E 0 3 0 •OOOOOOE 00 Oo OOOOOCE CO 0 . 6 8 1 0 9 9 E 03 0 . 9 7 6 0 9 3 E 0 3 0 . 5 1 1 2 2 1 E 0 3 0 eOOOOOOE 00 OoCOOOOOE r, n O.OOOOCOE 00 0 . 2 1 7 0 5 2 E 0 4 0 . 1 1 3 4 2 7 E 0 4 0 .OOOOOOE 00 0 . 1 2 1 Q 7 4 E 0 4 0 . 1 2 0 7 5 2 E 0 4 0 . 1 2 3 2 5 2 E 0 4 0 . 7 6 0 0 0 7 E 03 0 e OOOOOOE 00 0 . 1 5 2 7 6 2 E 0 4 0 . 2 1 5 8 3 0 E 0 4 0 . 9 0 3 8 3 1 E 0 3 0 . 5 4 7 8 1 2 E 0 3 0 .OOOOOOE 00 CoOOOOOOE 0 0 0 . 7 9 7 7 4 2 E 03 0 o 1 0 4 8 6 6 E 0 4 0 . 6 4 1 4 5 4 E 0 3 0 •OOOOOOE CO Oo OOCOOOE 0 0 O.OOOOOOE 00 0 . 1 9 2 4 9 1 E 0 4 0 . 1 1 7 5 4 6 E 0 4 0 .OOOOOOE 00 0 . 1 7 9 9 2 7 E 0 5 0 . 1 7 9 2 9 9 E 0 5 0 8 1 8 4 1 9 8 E 0 5 0 . 9 7 4 9 9 0 E 0 4 0 . 0 0 0 0 0 0 E 00 0 . 2 4 3 9 4 7 E 05 0 . 3 8 4 4 0 3 E 05 0 . 1 2 2 2 8 8 E 0 5 0 . 6 3 2 7 9 2 E 0 4 0 o OOOOOOE 00 OoCOOOOOE 0 0 C . 1 0 2 1 4 9 E 05 0 . 1 4 6 3 9 2 E 0 5 0 . 7 6 6 7 1 9 E 0 4 0 e OOCOOOE 00 QoOCOCOOE 0 0 O.OOOOOOE 00 0 . 3 0 0 8 3 4 E 0 5 0 . 1 5 7 2 0 9 E 0 5 0 o OOOOOOE 00 0 . 1 3 3 3 5 5 E 0 2 C . 1 3 2 8 8 9 E 0 2 0 . 1 3 6 5 2 1 E 0 2 0 . 7 2 2 6 2 4 E 0 1 0 eOOOOOOE 00 0 o 5 3 8 7 4 6 E 0 2 0 . 8 4 8 9 3 9 E 0 2 0 . 2 7 0 0 6 9 E 0 2 0 . 1 3 9 7 4 9 E 02 0 eOOOOOOE 00 OoCOOOOOE 0 0 G . 2 1 2 1 2 4 E 0 3 0 . 3 0 3 9 9 9 E 0 3 0 . 1 5 9 2 1 7 E 03 0 eOOOOOOE 00 OoOOOOOOE 0 0 O.OOOOOOE 0 0 0 . 5 8 9 3 2 7 E 0 4 0 . 3 0 7 9 7 0 E 0 4 0 eOOOOOOE 00 0 o 1 4 4 9 0 3 E 0 2 0 . 1 4 4 3 9 7 E 02 0 . 1 4 8 3 4 3 E 0 2 0 . 7 8 5 2 0 2 E 0 1 0 .OOOOOOE 00 0 . 1 9 6 7 5 6 E 02 0 . 3 1 0 0 4 2 E 02 0 . 9 8 6 3 2 6 E 0 1 0 . 5 1 0 3 8 1 E 0 1 0 eOOOOOOE 00 OoCOOOOOE 00 0 . 8 3 6 8 9 8 E 0 1 0 o l l 9 9 3 7 E 0 2 0 . 6 2 8 1 6 2 E 0 1 0 .OOOOOOE 00 OoCOOOOOE 0 0 O.OOOOOOE 0 0 0 . 2 6 6 7 0 2 E 0 2 0 . 1 3 9 3 7 3 E 02 0 eOOOOOOE 00 0 o 2 4 4 2 5 0 E 03 - 0 . 2 4 3 3 9 4 E 0 3 - 0 . 2 5 0 0 4 2 E 0 3 - 0 . 1 3 2 3 5 2 E 03 0 •OOOOOOE 00 0 . 3 3 1 1 5 4 E 03 - 0 . 5 2 1 8 1 8 E 0 3 - 0 . 1 6 6 0 0 2 E 0 3 - 0 . 8 5 8 9 9 2 E 02 0 eOOOOOOE 00 Oo OOOOOOE 0 0 - 0 . 1 3 8 6 6 6 E 03 -o . 1 9 8 7 2 0 E 0 3 - 0 . 1 0 4 0 8 0 E 03 0 •OOOOOOE 00 OoOOOOOOE 0 0 • O.OOOOOOE 00 - 0 • 9 3 1 5 6 4 E 0 3 - 0 . 4 8 6 8 1 6 E 03 0 •OOOOOOE 00 0 . 8 3 4 4 6 5 E - 0 6 0 . 9 2 3 8 7 2 E - 0 6 0 .OOOOOOE 0 0 0 . 1 4 9 0 1 1 E - Q 7 0 •OOOOOOE 00 0 . 1 5 4 9 7 2 E - 0 5 0 . 1 9 0 7 3 4 E - 0 5 0 • 3 6 3 5 8 8 E -0 5 C . 1 5 1 9 9 1 E - 0 5 0 •OOOOOOE 00 OoOOOOOOE 0 0 - 0 . 6 4 2 9 2 5 E 0 1 -0 . 9 2 1 3 9 3 E 0 1 - 0 . 4 8 2 5 6 9 E 0 1 0 .OOOOOOE CO 0.OOOOOOE 0 0 O.OOOOOOE CO 0 . 8 8 8 4 5 2 E 0 3 0 . 4 6 4 2 8 7 E 03 0 •OOOOOOE 00 128 A p p e n d i x T a b l e 2 . S u m s , b y m o n t h , a g e , s e x , m a t u r i t y a n d t o t a l , o f e a c h component o f t h e e n e r g y b u d g e t f o r r a i n b o w t r o u t . K e y t o t a b l e : F e b r u a r y M a r c h A p r i l M a y J u n e J u l y A u g u s t S e p t e m b e r O c t o b e r November December J a n u a r y sum o f e a c h age g r o u p sum m a l e i m m a t u r e sum m a l e m a t u r e sum t o t a l sum f e m a l e i m m a t u r e sum f e m a l e m a t u r e 0 + O.OOOOOOE 0 0 0 . 1 2 8 8 0 0 E 0 4 0 . 4 0 6 0 0 0 E 03 0 . 3 9 9 7 0 0 E 0 4 OoOOOOOOE 0 0 0 . 1 2 4 3 0 0 E 0 4 0 8 3 9 2 0 0 0 E 03 0 . 3 8 5 5 0 0 E 0 4 OoOOOOOOE 0 0 0 . 1 2 C 2 0 Q E 0 4 0 0 3 7 9 0 0 0 E 03 0 o 3 7 3 2 0 0 E 0 4 OoOOOOOOE 0 0 0 a l l 6 0 0 0 E 0 4 0 o 3 6 6 0 0 0 E 0 3 O o 3 6 0 1 0 0 E 0 4 O.OOOOOOE 0 0 O0IO68OOE 0 4 0 o 2 6 2 0 0 0 E 03 0 o 3 l 3 9 0 0 E 0 4 0 . 2 3 4 9 0 0 E 0 4 0 o l 9 9 0 0 0 E 0 4 0 o 2 1 4 0 0 O E 0 3 0 o 5 1 9 8 0 0 E 0 4 . 0 . 2 3 0 7 0 0 E 0 4 0 o l 6 4 9 0 C E 0 4 0 o 4 5 0 0 0 0 E 0 3 0 . 4 9 5 1 0 0 E 0 4 0 . 2 2 2 4 0 0 E 0 4 0 o l 5 8 9 0 0 E 0 4 O o 4 3 4 0 0 0 E 0 3 .0 . 4 7 7 3 0 0 E 0 4 • O o 2 1 4 4 0 0 E 0 4 -0 . 1 5 3 2 0 0 E 0 4 O o 4 l 9 0 0 0 E 0 3 0 . 4 6 0 3 0 0 E 0 4 0 o 2 0 6 9 C 0 E 0 4 O o l 4 7 9 0 0 E 0 4 0 . 4 0 4 O O O E 03 J o 4 4 4 4 0 0 E 0 4 . D . 1 9 9 4 0 0 E 0 4 O o l 4 2 6 0 0 E 0 4 D . 3 9 0 0 0 0 E 0 3 J o 4 2 8 5 0 0 E 0 4 J o l 9 2 4 0 0 E 0 4 3 o l 3 7 7 0 0 E 0 4 3 . 3 7 7 0 0 0 E 0 3 : o 4 ! 3 7 0 0 E 0 4 1 + 0 o l 8 4 7 0 0 E 0 4 0 o l 8 7 8 0 0 E 0 4 0 0 4 2 5 0 0 0 E 0 3 0 e 1 7 8 0 0 0 E 0 4 0 a l 8 1 0 0 0 E 0 4 0 . 4 1 0 0 0 0 E 0 3 C 1 7 2 2 0 0 E 0 4 0 . 1 7 5 3 0 0 E 0 4 0 . 3 9 8 0 0 0 E 0 3 0 . 1 6 6 0 0 0 E 0 4 0 . 1 6 9 2 0 0 E 0 4 0 o 3 8 3 0 0 0 E 0 3 0 . 1 6 0 2 0 0 E 0 4 0 o l 5 9 5 0 0 E 0 4 0 . 2 1 4 0 0 0 E 0 3 0 , 1 5 4 4 0 0 E 0 4 0 . 2 8 6 3 0 0 E 0 4 0 o l 3 1 0 0 0 E 0 3 0 o l 4 9 0 0 0 E 0 4 0 o 2 3 6 6 0 0 E 0 4 0 . 4 8 6 0 0 0 E 0 3 0 « 1 4 3 5 0 0 E 0 4 0 . 2 2 8 1 0 0 E 0 4 0 o 4 6 9 0 0 0 E 0 3 0 . 1 3 8 3 0 0 E 0 4 0 o 2 1 9 9 0 0 E 0 4 0 o 4 5 3 0 0 0 E 03 0 o l 3 3 4 0 0 E 0 4 O o 2 1 2 3 0 0 E 0 4 0 . 4 3 8 0 0 0 E 0 3 0 o l 2 8 6 0 0 E 0 4 0 . 2 0 4 6 0 0 E 0 4 0 . 4 2 3 0 0 0 E 0 3 0 o l 2 4 l 0 0 E 0 4 0 . 1 9 7 4 0 0 E 0 4 0 , 4 0 9 0 0 0 E 0 3 129 2 + 0 . 1 2 0 1 0 0 E 0 4 0 . 1 1 5 9 0 0 E 0 4 0 . 1 1 2 3 0 0 E 0 4 0 . 1 0 8 6 0 0 E 0 4 0 . 1 0 3 9 0 0 E 0 4 0 . 9 9 4 0 0 0 E 0 3 0 . 9 5 1 0 0 0 E 0 3 0 e 9 1 8 0 0 0 E 0 3 0 . 8 8 7 0 0 0 E 0 3 0 . 8 5 9 0 0 0 E 0 3 0 . 8 3 0 0 0 0 E 0 3 0 o 8 0 3 0 0 0 E 0 3 3+ 0 . 7 8 1 0 0 0 E 0 3 0 o 7 5 4 0 0 0 E 03 O o 7 3 0 0 0 0 E 03 0 . 7 0 4 0 0 0 E 03 0 0 4 6 9 0 0 0 E 03 0 o 3 0 7 0 0 0 E 0 3 0 « 2 0 3 0 0 0 E 0 3 0 . 1 9 6 0 0 0 E 0 3 0 o l 8 9 0 0 0 E 0 3 0 o l 8 2 0 0 0 E 0 3 0 o l 7 5 0 0 0 E 03 0 . 1 6 9 0 0 0 E 03 NUMBER 4+ O 9 I 6 8 O O O E 03 0 o l 6 2 0 0 0 E 03 0 . 1 5 7 0 C 0 E 03 0 s l 5 l O C 0 E 03 0 @ 2 9 0 0 0 0 E 02 0 o 4 0 0 0 0 0 E 0 1 OoOOOOOOE 00 OoOOOOOOE 00 OoOOOOOOE 00 O.OOOOOOE 00 OoOOOOOOE 00 O.OOOOOOE 00 0 + O.OOOOCOE 0 0 0 . 4 2 3 6 9 2 E 02 0 . 3 3 7 5 2 1 E 0 2 0 . 1 7 9 6 9 4 E 03 O.OOOOOOE 0 0 Q . 4 C 9 1 9 4 E 0 2 0 . 3 2 6 6 0 5 E 0 2 0 . 1 7 3 3 6 7 E 03 O.OOOOOOE 0 0 0 . 3 9 5 6 6 3 E 02 0 . 3 1 4 5 9 4 E 0 2 0 . 1 6 7 8 9 4 E 0'3 O.OOOOCOE 0 0 0 . 3 8 1 9 5 5 E 02 0 . 3 0 4 1 1 1 E 02 0 . 1 6 1 9 5 2 E 03 O.OOOOOOE 0 0 0 . 3 2 8 0 4 8 E 0 2 0 . 1 6 3 7 5 1 E 02 0 . 1 1 6 3 5 4 E 03 0 . 7 0 4 6 9 9 E 0 0 0 . 3 5 4 9 2 4 E 02 0 . 1 3 9 1 2 1 E 02 0 . 1 1 6 4 5 4 E 0 3 0 . 1 5 3 2 3 0 E 0 1 0 . 3 0 6 7 9 5 E 0 2 0 . 2 1 6 3 1 5 E 0 2 0 . 1 2 3 4 1 1 E 0 3 0 . 7 6 3 7 9 8 E 0 1 0 . 3 8 3 3 9 1 E 0 2 0 . 2 5 3 0 2 3 E 0 2 0 . 1 5 0 2 0 8 E 0 3 0 . 3 8 0 7 2 T E 0 2 0 . 5 7 2 7 9 . 1 E . 0 2 0 . 2 9 6 4 6 0 E 02 0 . 2 0 5 1 3 3 E 03 0 . 3 6 7 4 0 9 E 0 2 0 . 5 5 3 9 8 4 E 0 2 0 . 2 8 5 1 1 9 E 02 0 . 1 9 8 0 9 3 E 03 0 . 3 5 4 0 9 1 E 02 0 . 5 3 4 0 0 4 E 02 0 . 2 7 5 3 8 7 E 0 2 0 . 1 9 1 0 1 8 E 03 0 . 3 4 1 6 6 0 E 0 2 0 . 5 1 6 1 6 5 E 0 2 0 . 2 6 6 4 4 3 E 0 2 0 . 1 8 4 5 1 5 E 03 1 + 0 . 3 2 8 7 6 6 E 0 2 0 . 5 8 6 0 9 9 E 02 0 . 4 4 9 6 2 7 E 0 2 0 0 3 1 6 8 4 0 E 02 C . 5 6 4 7 9 1 E 0 2 0 . 4 3 3 0 8 1 E 0 2 0 . 3 0 6 5 1 5 E 02 0 . 5 4 7 4 3 3 E 02 0 . 4 2 1 2 5 4 E 0 2 0 . 2 9 5 4 7 9 E 0 2 0 . 5 2 8 7 5 1 E 0 2 0 . 4 0 4 7 0 8 E 02 0 . 2 8 5 1 5 5 E 02 0 . 4 8 1 9 3 0 E 0 2 0 . 1 8 9 0 1 7 E 0 2 0 . 3 4 3 9 7 1 E 02 0 . 5 3 8 3 8 8 E 0 2 0 . 1 3 2 1 1 4 E 0 2 0 . 4 1 2 4 6 8 E 0 2 0 . 3 7 1 0 9 3 E 02 Q . 3 3 9 9 1 3 E 0 2 0 . 4 9 7 1 7 7 E 02 0 . 4 7 4 3 3 1 E 0 2 0 . 3 9 1 3 4 2 E 0 2 0 . 5 9 9 7 0 5 E 0 2 0 . 7 3 0 7 2 7 E 02 0 . 4 5 1 3 6 0 E 0 2 0 . 5 7 8 4 5 7 E 0 2 0 . 7 0 5 8 3 0 E 02 0 . 4 3 6 0 0 2 E 02 0 . 5 5 7 6 4 3 E 02 0 . 6 8 0 1 4 4 E 02 0 . 4 2 0 6 4 4 E 02 0 . 5 3 8 1 3 0 E 02 Q . 6 5 6 4 7 0 E 02 0 . 4 0 6 0 7 6 E 02 130 2+ 0 . 5 2 1 2 3 3 E 0 2 0 . 5 0 3 0 0 5 E 0 2 0 . 4 8 7 3 8 1 E 0 2 . 0 . 4 7 1 3 2 4 E 0 2 0 . 4 5 0 9 2 5 E 0 2 0 . 5 0 0 6 0 9 E 0 2 0 . 5 5 4 4 6 4 E 0 2 0 . 6 2 2 6 5 0 E 0 2 0 . 6 9 9 8 9 7 E 0 2 0 . 6 7 7 8 0 3 E 0 2 0 . 6 5 4 9 2 0 E 0 2 0 . 6 3 3 6 1 6 E 0 2 3 + 0 . 6 1 6 9 8 9 E 02 0 . 5 9 5 6 5 9 E 02 0 . 5 7 6 6 9 9 E 02 0 . 5 5 6 1 5 9 E 02 0 . 3 7 0 5 0 9 E 02 0 . 3 0 5 0 6 5 E 0 2 0 . 2 5 1 8 6 2 E 0 2 0 . 3 0 5 8 7 9 E 0 2 0 . 3 7 1 0 0 8 E 0 2 0 . 3 5 7 2 6 7 E 0 2 0 . 3 4 3 5 2 6 E 02 0 . 3 3 1 7 4 8 E 02 WEIGHT ( K G ) 4 + 0 . 3 2 9 9 5 2 E 02 0 . 3 1 8 1 6 7 E 02 0 . 3 0 8 3 4 8 E . 02 ' 0 . 2 9 6 5 6 3 E 02 0 . 5 6 9 5 5 9 E O l 0 . 7 8 5 6 0 0 E 00 O.OOOOOOE 00 OoOOOOOOE 00 O.OOOOOOE 00 O.OOOOOOE 00 O.OOOOOOE 00 O.OOOOOOE 00 0 + 0 oOOOOOOE 0 0 0 9 1 4 8 4 1 9 E 05 0 • 1 1 8 2 3 3 E 0 5 0 e 6 2 9 4 6 8 E 05 0 « OOOOOOE CO 0 . 1 2 5 0 0 0 E 0 5 0 o 9 9 7 7 1 5 E 0 4 0 o 5 2 9 6 0 2 E 0 5 0 oOOOOOOE 0 0 0 o l 2 8 9 1 1 E 0 5 0 o 1 0 2 4 9 7 E 05 0 o 5 4 7 0 1 7 E 05 0 o OOOOOOE 0 0 0 o 2 1 3 5 8 9 E 05 r> . 1 7 0 0 5 9 E 05 0 o 9 0 5 6 3 9 E 05 0 oOOOOOOE 00 0 • 5 3 4 8 4 9 E 05 0 o 2 6 6 3 3 1 E 0 5 0 o l 8 9 2 4 3 E 06 0 o 9 4 7 l 5 8 E 0 3 0 o 4 7 7 0 3 9 E 0 5 0 o1 8 6 9 8 7 E 0 5 0 o1 5 6 5 2 2 E 0 6 0 o l 5 1 6 3 1 E 0 4 0 o 3 0 3 5 9 3 E 05 0 o 2 1 4 0 5 8 E 05 0 o l 2 2 1 2 3 E 0 6 0 o l 4 2 3 3 6 E 0 5 c . 7 1 4 4 6 2 E 05 0 . 4 7 1 5 1 7 E 05 0 • 2 7 9 9 1 9 E 0 6 0 o 3 0 7 7 0 4 E 05 0 o 4 6 2 9 2 9 E 05 0 o 2 3 9 5 9 9 E 05 0 o l 6 5 7 8 9 E 0 6 0 o l 7 3 4 3 9 E 05 0 o 2 6 1 5 1 5 E 0 5 0 • 1 3 4 5 9 4 E 05 0 o 9 3 5 l 2 4 E 0 5 0 o l 5 4 4 8 0 E 05 0 o 2 3 2 9 7 1 E 05 0 o 1 2 0 1 4 3 E 05 0 e 8 3 3 3 5 8 E 05 0 . 1 3 9 0 2 7 E 05 0 o 2 1 0 0 3 7 E 0 5 0 o 1 0 8 4 2 0 E 0 5 0 o 7 5 0 8 2 7 E 05 1+ 0 o ! 1 5 1 6 6 E 05 0 o 2 0 5 3 1 0 E 05 r\ \j o l 5 7 5 0 4 E 05 0 o 9 6 7 8 8 2 E 0 4 0 • 1 7 2 5 3 2 E 0 5 0 o l 3 2 2 9 7 E 05 C e 9 9 8 6 5 9 E 0 4 0 o l 7 8 3 5 9 E 05 0 o l 3 7 2 4 8 E 05 0 e l 6 5 2 3 2 E 0 5 0 . 2 9 5 6 7 8 E 05 C • 2 2 6 3 1 2 E 05 0 o 4 6 3 7 8 7 E 05 0 o 7 8 3 8 2 6 E 05 0 o 3 0 7 4 2 5 E 05 0 . 4 6 2 3 1 8 E 05 0 o 7 2 3 6 2 5 E 05 0 o l 7 7 5 7 0 E 05 0 o 4 0 8 l 6 3 E 05 0 o 3 6 7 2 2 0 E 05 0 B 3 3 6 3 6 6 E 05 0 o 9 2 6 5 0 7 E 05 0 « 8 8 3 9 3 3 E 05 0 o 7 2 9 2 8 0 E 05 0 o 4 8 4 6 8 l E 05 0 o 5 9 0 5 7 3 E 05 0 8 3 6 4 7 8 9 E 05 0 o 2 7 3 0 6 7 E 05 0 • 3 3 3 1 9 5 E 0 5 0 o 2 0 5 8 2 0 E 05 0 o 2 4 3 2 8 4 E 05 0 o 2 9 6 7 2 7 E 05 0 o l 8 3 5 1 5 E 05 0 . 2 1 8 9 7 5 E 05 0 . 2 6 7 1 2 9 E 05 0 o l 6 5 2 3 9 E 05 2 + 0 o l 8 2 5 8 8 E 0 5 0 . . 1 5 3 6 5 8 E 0 5 O o l 5 8 7 9 3 E 0 5 0 e 2 6 3 5 6 4 E 0 5 O o 7 3 3 4 0 0 E 0 5 0 o 6 7 2 8 4 9 E 0 5 0 e 5 4 8 6 7 7 E 0 5 0 o l l 6 0 3 3 E 0 6 0 o 5 6 5 6 5 7 E 0 5 O o 3 l 9 9 6 5 E 0 5 0 o 2 6 5 7 2 3 E 0 5 0 o 2 5 7 8 3 0 E 0 5 I N T A K E 3 + 0 o 2 1 6 1 3 1 E 0 5 0 . 1 8 1 9 6 2 E 0 5 0 o l 6 7 8 9 4 E 0 5 0 o 3 1 l 0 0 4 E 05 0 e 6 0 2 6 0 9 E 0 5 0 o 4 l 0 0 2 5 E 0 5 C o 2 4 9 2 3 3 E 05 0 o 5 7 0 0 1 7 E 05 0 o 2 9 9 8 4 8 E 0 5 0 . 1 6 8 6 5 2 E 0 5 0 o l 4 9 8 7 0 E 0 5 0 o l 3 4 9 9 4 E 05 ENERGY ( K C A L ) 4 + O o l l 5 5 8 2 E 05 O o 9 7 l 9 3 9 E 0 4 0 e l 0 0 4 6 2 E 05 0 o l 6 5 8 3 8 E 05 0 o 9 2 6 3 5 l E 04 0 o l 0 5 5 8 9 E 04 OoOOOOOOE 00 OoOOOOOOE 00 OoOOOOOOE 00 OoOOOOOOE CO OoOOOOOOE 00 OoOOOOOOE 00 E G E S T I O N - E X C R E T I O N ( K C A L ) OO- 1+ 2 + 3 + 4 + Oo OOOOOOE 0 0 0' . 2 3 0 3 3 3 E 0 4 0 . 3 6 5 1 7 6 E 0 4 0 o 4 3 2 2 6 3 E 0 4 0 . 2 3 1 1 6 4 E 0 4 0 . 2 9 6 8 3 8 E 0 4 0 o 4 1 0 6 2 l E 0 4 0 0 2 3 6 4 6 7 E 0 4 0 o 3 1 5 0 0 9 E 0 4 0 0 1 2 5 8 9 3 E 05 0 . OOOOOOE 0 0 0 o l 9 3 5 7 6 E 0 4 0 . 3 0 7 3 1 6 E 0 4 0 • 3 6 3 9 2 4 E 0 4 0 o l 9 4 3 8 7 E 04 0 * 2 5 0 Q C 1 E 0 4 0 o 3 4 5 C 6 5 E 0 4 0 . 1 9 9 5 4 3 E 0 4 0 . 2 6 4 5 9 5 E 04 0 . 1 0 5 9 2 0 E 05 0 . OOOOOOE 00 0 o l 9 9 7 3 1 E 0 4 0 o 3 1 7 5 8 7 E 0 4 0 • 3 7 5 7 8 9 E 0 4 0 o 2 0 0 9 2 5 E 0 4 Oa 2 5 7 8 2 2 E 0 4 0 o 3 5 6 7 1 8 E 0 4 0 . 2 0 4 9 9 5 E 0 4 0 • 2 7 4 4 9 7 E 0 4 c . 1 C 9 4 0 3 E 05 0 . OOOOOOE 0 0 0 o 3 3 0 4 6 4 E 04 O o 5 2 7 1 2 8 E 0 4 0 o 6 2 2 0 0 9 E 0 4 0 . 3 3 1 6 7 7 E 04 0 a 4 2 7 1 7 9 E 0 4 0 . 5 9 1 3 5 5 E 0 4 0 a 3 4 0 1 1 8 E 0 4 0 o 4 5 2 6 2 5 E 0 4 0 a 1 8 1 1 2 7 E 05 Oo OOOOOOE 0 0 0 . 9 2 7 5 7 4 E 0 4 O o 1 4 6 6 8 0 E 0 5 0 o 1 2 0 5 2 1 E 0 5 0 • 1 8 5 2 7 0 E 04 Oo 1 0 6 9 6 9 E 05 0 • 1 5 6 7 6 5 E 05 0 0 5 3 2 6 6 3 E 0 4 0 o 6 1 4 8 5 0 E 0 4 0 o 3 7 8 4 8 6 E 05 0 0 1 8 9 4 3 1 E 03 0 o 9 2 4 6 3 7 E 0 4 0 o l 3 4 5 6 9 E 0 5 0 o 8 2 0 0 5 1 E 0 4 0 • 2 1 1 1 7 8 E 03 0 0 9 5 4 0 7 9 E 0 4 0 o l 4 4 7 2 5 E 05 0 o 3 7 3 9 7 5 E 0 4 0 o 3 5 5 1 4 0 E 0 4 Oo 3 1 3 0 4 4 E 05 Oo 3 0 3 2 6 3 E 03 0 o S l 6 3 2 6 E 0 4 0 o l 0 9 7 3 5 E 0 5 . 0 o 4 9 S 4 6 7 E 0 4 0 oOOOOOOE 00 Oo 6 0 7 1 6 7 E 0 4 0 o 7 3 4 4 4 0 E 0 4 0 0 4 2 8 1 1 6 E 0 4 0 o 6 7 2 7 3 2 E 0 4 0 0 2 4 4 2 4 7 E 05 0 o 2 8 4 6 7 2 E 0 4 0 . 1 8 5 3 0 1 E 05 0 . 2 3 2 0 6 6 E 0 5 0 a 1 1 4 0 0 3 E 0 5 0 •OOOOOOE CO 0 0 1 4 2 8 9 2 E 05 0 o l 7 6 7 8 6 E 05 Oo 9 4 3 0 3 4 E 0 4 0 o l 4 5 8 5 5 E 05 0 . 5 5 9 8 3 8 E 05 0 0 6 1 5 4 0 8 E 0 4 0 o 9 6 9 3 6 3 E 0 4 0 . 1 1 3 1 3 1 E 0 5 0 o 5 9 9 6 9 7 E 0 4 0 oOOOOOOE 00 Oo 9 2 5 6 5 9 E 0 4 0 • 1 1 8 1 1 4 E 05 0 o 4 7 9 1 9 8 E 0 4 0 o 7 2 9 5 7 3 E 0 4 0 0 3 3 1 5 7 S E 05 0 o 3 4 6 8 7 9 E 0 4 0 . 5 4 6 1 3 5 E 0 4 0 « 6 3 9 9 3 0 E 0 4 0 . 3 3 7 3 C 4 E 0 4 0 oOOOOOOE 00 0 o 5 2 3 0 3 0 E 0 4 0 • 6 6 6 3 9 0 E 0 4 0 0 2 6 9 1 8 8 E 0 4 o 4 1 1 6 4 0 E 0 4 0 0 1 8 7 0 2 4 E .05 Oo 3 0 8 9 5 9 E 0 4 0 . 4 8 6 5 6 3 E 0 4 0 o 5 7 1 4 4 7 E 0 4 0 o 2 9 9 7 4 1 E 04 o oOOOOOOE 00 0 o 4 6 5 9 4 2 E 04 0 o 5 9 3 4 5 5 E 0 4 Oo 2 4 0 2 8 7 E 0 4 0 • 3 6 7 0 3 1 E 0 4 0 o 1 6 6 6 7 1 E . 05 0 0 2 7 8 0 5 5 E 0 4 0 . 4 3 7 9 5 0 E 0 4 C o 5 1 5 6 6 0 E 0 4 0 • 2 6 9 9 8 8 E 0 4 0 o 0 0 0 0 0 0 u 00 Oo 4 2 0 0 7 4 E 0 4 0 . 5 3 4 2 5 9 E 0 4 0 o 2 1 6 8 4 1 E 0 4 0 . 3 3 0 4 7 9 E 0 4 Oo 1 5 0 1 6 5 E 0 5 133 ROUTINE METABOLISM (KCAL) 0 + 1 + 2 + 3 + 4 + 0.OOOOOOE 0 0 0 . 2 6 5 4 2 4 E 0 4 0 . 4 0 5 5 9 6 E 0 4 0 . 4 9 9 8 6 7 E 0 4 0 o 2 6 6 l 3 2 E 0 4 0 . 3 5 5 4 9 9 E 0 4 0 o 4 4 7 0 0 3 E 0 4 0 . 2 9 7 7 9 7 E 0 4 0 . 3 3 6 7 2 0 E 0 4 . 0 o 1 4 3 7 0 2 E 05 OeOOOOOOE 0 0 0 . 2 3 3 1 0 2 E 0 4 0 . 3 5 6 5 4 2 E 0 4 0 e 4 3 9 5 2 6 E 0 4 0 . 2 3 3 8 6 7 E 04 0 e 3 l 2 7 0 9 E 0 4 0 . 3 9 2 4 6 8 E 0 4 0 . 2 6 2 3 0 2 E 0 4 0 . 2 9 5 5 5 7 E 0 4 0 e 1 2 6 3 0 3 E 0 5 OoOOOOOOE GO 0 . 2 5 2 1 5 0 E 0 4 0 e 3 8 6 l 7 8 E 0 4 0 . 4 7 5 7 5 C E 0 4 0 . 2 5 3 2 4 3 E 0 4 0 . 3 3 8 1 4 0 E 0 4 0 0 4 2 5 2 2 5 E 0 4 0 . 2 8 2 8 7 4 E 0 4 0 . 3 2 1 0 8 3 E 0 4 0 . 1 3 6 7 3 2 E 0 5 O.OOOOOOE 0 0 0 . 2 3 9 1 6 6 E 0 4 0 . 3 6 7 1 4 8 E 0 4 0 o 4 5 1 3 9 1 E 0 4 0 . 2 3 9 8 2 9 E 04 0 . 3 2 1 G 6 8 E 0 4 0 . 4 0 3 8 8 3 E 0 4 0 . 2 6 8 8 5 8 E 0 4 0 . 3 0 3 7 2 4 E 0 4 O o 1 2 9 7 5 3 E 05 Oo OOOOOOE CO 0 . 2 7 3 6 4 3 E 0 4 0 . 4 1 7 2 8 1 E 0 4 . 0 . 3 8 9 7 0 0 E 0 4 0 o 8 0 4 0 2 5 E 03 0 e 3 3 6 l 4 7 E 0 4 0 . 4 3 8 0 6 7 E 0 4 C o 1 9 4 7 7 0 E 0 4 0 . 1 9 2 0 4 2 E 0 4 0 . 1 1 6 1 0 2 E 05 0 • 1 3 3 7 2 1 E 0 3 0 . 3 5 2 7 2 7 E 0 4 0 . 5 0 4 9 8 8 E 0 4 0 . 3 7 5 7 8 1 E 0 4 0 . 1 9 9 9 6 3 E 03 0 . 4 0 1 4 4 3 E 0 4 0 . 5 3 5 5 S 7 E 0 4 0 o 1 8 3 4 8 2 E 0 4 . 0 . 1 5 1 3 5 3 E .04 0 « 1 2 7 1 8 6 E 0 5 0 . 4 0 5 7 5 5 E 0 3 0 . 5 3 3 5 3 3 E 0 4 0 . 7 2 4 1 5 1 E 0 4 0 . 3 9 7 2 7 3 E 0 4 OeOOOOOOE 00 0 o 4 4 4 1 9 2 E 0 4 0 . 4 9 6 4 3 3 E 0 4 0 o 3 3 1 7 9 9 E 0 4 0 . 4 2 3 1 0 8 E 0 4 0 . 1 6 9 5 5 3 E 05 0 . 1 7 9 6 9 1 E 0 4 0 . 8 0 3 2 1 2 E 0 4 o . 1 0 3 3 2 0 E 0 5 C . 6 0 1 6 3 7 E G4 OoOOOOOOE 00 Oo 7 0 7 0 8 5 E 0 4 0 . 8 0 4 6 7 2 E 0 4 0 . 4 8 9 3 2 3 E 0 4 0 . 6 1 6 6 6 6 E 0 4 0 . 2 6 1 7 7 4 E 05 0 a 4 0 l 2 4 7 E 0 4 0 e 6 l 0 0 9 9 E 0 4 0 . 7 4 3 8 0 4 E 0 4 Oo 4 5 8 9 6 7 E 0 4 OeOOOOOOE 00 0 . 6 4 4 1 6 3 E 0 4 0 . 7 5 2 0 3 8 E 0 4 0 . 3 6 4 2 4 4 E 0 4 0 e 4 5 3 6 7 2 E 0 4 0 . 2 2 1 4 1 1 E 05 0 . 3 Q 5 7 7 Q E 0 4 0 . 4 6 4 7 3 8 E 0 4 0 • 5 6 S 3 8 6 E 0 4 0 . 3 4 9 1 5 8 E 0 4 OoOOOOOOE 00 0 . 4 9 1 8 1 1 E 0 4 0 . 5 7 3 5 8 5 E 0 4 0 . 2 7 6 7 2 4 E 0 4 0 e 3 4 5 9 3 2 E 0 4 0 . 1 6 8 8 0 5 E 05 0 o 2 7 3 8 9 0 E 0 4 0 . 4 1 6 4 0 4 E 0 4 0 . 5 1 0 0 7 8 E 0 4 0 . 3 1 2 2 5 2 E 0 4 O.OOOOOOE 00 0 . 4 4 0 5 0 9 E 0 4 0 . 5 1 3 7 2 5 E 0 4 0 . 2 4 8 3 2 2 E 0 4 0 o 3 1 0 0 6 7 E 0 4 0 . 1 5 1 2 6 2 E 0 5 0 . 2 7 3 8 3 7 E 0 4 0 e 4 1 6 3 5 7 E 0 4 0 • 5 1 0 9 9 3 E 0 4 0 . 3 1 2 4 7 3 E 0 4 O.OOOOOOE 00 0 o 4 4 1 0 8 0 E 0 4 0 . 5 1 3 7 5 7 E 0 4 0 . 2 4 8 7 8 6 E 0 4 0 . 3 1 0 0 3 7 E 0 4 0 . 1 5 1 3 6 6 E 05 134 BODY, ENERGY. ( K C A L ) c + 1 + 2 + . 3 + 4 + U © O \) -0 w \> \j L 0 0 0 . 4 0 2 3 4 0 E 0 5 0 • 6 3 6 1 8 9 E 0 5 0 8 7 1 8 9 5 2 E 0 5 . 0 • 3 7 6 1 2 5 E 05 0 . 5 1 8 9 5 8 E 0 5 0 e 7 1 6 8 0 6 E 05 0 . 4 0 6 3 7 8 E 0 5 0 e 4 9 1 4 6 4 E 0 5 0 . 2 1 3 3 6 0 E 0 6 O t O O O O O C E 0 0 0 . 3 8 0 3 2 9 E 0 5 0 8 & 0 2 1 8 3 E 0 5 0 • 6 7 6 3 9 1 E 0 5 0 • 3 5 2 6 5 1 E 05 0 . 4 9 1 6 1 4 E 0 5 0 8 6 7 7 5 3 4 E 05 G o 38 56 35E 0 5 0 e 4 5 6 7 7 1 E 05 0 . 2 0 1 1 5 5 E 0 6 f> ;~\ n ry -"\ -~ JT w 0 vy <J Vv v> v/ w i-. 00 0 e 3 4 8 1 5 8 E 05 0 a 5 5 0 1 5 4 E 0 5 .. o 8 6 C 9 3 8 4 E 0 5 - 0 8 3 1 4 3 5 3 E 05 C - . 4 4 9 8 0 7 E 0 5 0 e 6 2 1 4 1 1 E 05 0 e 3 4 3 3 8 5 E 05 0 . 4 0 7 4 4 7 E 05 0 o 18 2 2 C 5 E 0 6 0 e OOOOOOE 0 0 0 . 3 1 9 1 7 1 E 05 0 a 4 9 1 9 4 2 E . 0 5 0 . 5 2 1 3 9 2 E 0 5 0 o 2 5 0 9 0 0 E 05 0 . 4 1 2 9 3 7 E 0 5 0 a 5 7 0 7 8 2 E 0 5 0 . 2 5 75 84E 05 G a 3 4 2 1 0 2 E 05 0 a l 5 8 3 4 C E 0 6 . - . - , ^ £• \.j A U w ;J w C. 0 0 0 . 3 2 4 1 6 3 E 05 0 • 4 9 9 0 3 8 E 0 5 . 0' 8 3 7 7 2 8 4 E 0 5 0 a 5 3 6 7 8 4 E 0 4 0 . 3 7 4 1 5 5 E 05 0 a 5 4 7 5 0 5 E 05 0 . 1 5 4 5 1 0 E 0 5 0 « 1 7 7 9 9 2 E 05 0 a 12 5 4 1 6 E 0 6 0 » 9 1 0 6 2 0 E 0 3 0 8 4 4 4 4 8 3 E 05 0 • 6 3 6 7 2 2 E 0 5 0 8 3 6 9 7 9 3 E 0 5 " 0 8 9 1 2 1 1 6 E 03 0 a 4 5 9 0 3 3 E 0 5 C 6 9 5 2 6 7 E 05 0 a l 6 1 6 8 8 E 05 0 a l 5 3 2 3 7 E 05 0 . 1 4 6 9 2 2 E 0 6 0 • 2 0 7 8 8 3 E 0 4 0 . 5 5 8 7 6 7 E 05 0 • 7 4 7 7 7 3 E 0 5 0 a 3 3 9 6 6 8 E 0 5 0 8OOOOOOE 00 0 a 4 1 6 5 8 1 L 0 5 0 0 5 0 3 1 3 0 E 05 0 . 2 9 0 9 7 3 E 05 0 e 4 5 6 3 1 2 E 05 0 a 1 6 6 6 9 9 E 0 6 Oa 1 0 1 9 5 5 E 05 0 e 6 6 3 0 7 0 E 05 0 6 8 2 7 4 4 6 E 0 5 0 e 4 0 6 4 7 7 E 05 0 oOOGOOOE GO 0 a 5 12 2 1 3 E 05 0 . 6 3 2 7 5 9 E 05 0 . 3 3 6 1 1 2 E 05 0 a 17 8 64E 05 0 . 1 9 9 t ; 9 4 E 0 6 Oa 5 0 U 7 2 8 E 05 0 »7 8 6 3 2 9 E 05 0 a 9 0 7 3 9 5 E 0 5 0 9 4 8 0 9 5 6 E 0 5 0 aOOOOOOE 00 0 o75 3 9 8 0E 0 5 0 B 9 6 0 4 3 0 E 05 0 . 3 8 2 6 2 I E 05 0 . 5 7 8 3 7 7 E 05 0 8 2 6 7 5 4 0 E 0 6 0 . 4 7 1 9 2 4 E 0 5 0 « 7 3 9 8 6 6 E 05 0 8 8 4 9 0 6 8 E 0 5 0 8 4 4 7 5 1 6 E 0 5 ' c •OOOOOOE 00 0 . 7 1 2 1 9 0 E 05 0 « 9 0 6 0 3 7 E 05 0 e 3 5 6 5 6 3 E 05 0 . 5 3 3 5 8 3 E 05 0 a 2 5 G 8 3 7 E G6 0 a 4 4 7 6 1 3 E 0 5 C a 7 0 3 0 2 8 E 05 0 8 8 0 0 4 2 2 E 0 5 0 a 4 1 9 7 7 6 E 0 5 0 •GOOCOOE 00 0 a 6 7 3 6 3 1 E 0 5 0 . 8 5 9 2 3 7 E 05 0 o 3 4 2 3 90E 05 0 . 4 9 3 5 8 2 E 05 0 . 2 3 7 0 ; ; ^ E 0 6 C a42 3 8 7 5 E 05 0 . 6 6 5 8 5 2 E 05 0 . 7 5 3 7 1 4 E 0 5 0 • 3 9 4 6 5 9 E 0 5 0 •OOOOOOE '~ 0. 0 « 6 4 0 9 2 5 E 05 0 a 2 1 3 9 1 8 E 05 0 . 3 2 5 2 1 4 E 05 0 . 4 5 8 0 4 3 E 05 135 GONAD ENERGY (KCAL) 0 + 1 + 2 + 3 + 4 + 0. OOOOOOE 0 0 O o 6 4 6 8 4 7 E 0 2 0 o 2 9 4 5 6 7 E 0 3 0 . 6 9 159 8E 0 4 0 . 5 1 8 6 9 7 E 0 4 0 . 3 8 9 9 2 5 E 02 0 o l 6 0 4 8 2 E 0 3 0 o 8 2 9 5 7 0 E 03 0 9 1 1 4 3 3 1 E 05 0 • 1 2 4 6 2 2 E 05 0. OOOOOOE 0 0 Q . 6 2 3 2 5 2 E 02 0 e 2 8 2 3 9 9 E 0 3 0 o 7 3 4 8 5 9 E 0 4 O o 5 4 6 l 4 7 E 0 4 0 o 3 7 6 5 8 2 E 02 0 o l 5 4 6 4 7 E 0 3 0 e 7 9 6 8 4 7 E 03 O o l 2 1 6 5 6 E 05 0 e l 3 l 5 4 7 E 05 OoOOOOOOE 0 0 0 o 6 0 3 0 7 3 E 0 2 0 o 2 6 0 7 2 7 E 0 3 0 . 6 9 8 8 0 0 E 0 4 0 . 5 2 3 3 7 2 E 0 4 0 o 3 6 4 l 3 1 E 0 2 0 » 1 4 9 8 9 4 E 03 0 8 7 l l 6 9 4 E 03 0 o l l 6 4 4 7 E 05 Q . 1 2 5 4 2 7 E 05 OoOOOOOOE 00 0 . 5 8 1 2 6 8 E 02 0 o 9 9 9 2 0 5 E 0 2 0 0 1 6 2 2 3 C E 03 0 0 1 0 1 6 6 5 E 03 0 e 3 5 1 5 1 5 E 02 O o l 4 4 7 7 9 E 0 3 0 o 5 7 6 6 l 8 E 0 2 C o l 8 4 3 5 0 E 0 3 0 o 4 2 1 9 4 2 E 0 3 OoOOOOOOE 00 0 o 5 6 1 0 8 9 E 0 2 0 o 9 5 0 2 8 0 E 02 Oe 1 0 4 3 9 2 E 03 0 e l 8 8 3 4 7 E 02 0 o 3 0 2 6 3 9 E 02 0 o l 3 1 9 5 9 E 03 C o 2 9 7 6 8 l E 02 O o 8 2 3 7 3 3 E 02 0 o 2 7 4 3 6 4 E 03 0 o l 3 8 6 3 9 E 0 1 0 o 6 7 6 5 8 2 E 02 0 o l 0 5 5 3 4 E 0 3 Oe 8 9 0 5 5 4 E 02 0 e 2 5 2 1 1 4 E 01 0 a 3 2 6 6 3 8 E 02 0 e l 4 7 4 l 8 E 03 0 e 2 5 3 6 7 5 E 02 O o 6 0 7 0 6 6 E 02 0 o 2 6 6 l 5 6 E 0 3 0 o 3 0 l 4 8 2 E 0 1 0 o 8 8 6 8 0 5 E 0 2 0 • 1 8 5 9 9 3 E 0 3 0 . 8 4 6 4 2 1 E 02 OoOOOOOOE 00 0 e 2 8 2 3 4 5 E 02 0 o l 0 1 6 1 0 E 03 0 o 4 5 3 1 9 2 E 0 2 0 o l 8 7 1 6 7 E 03 0 e 3 6 2 3 3 1 E 03 0 o 1 5 0 2 6 6 E 02 O o l 3 7 5 6 4 E 03 0 o 6 5 7 6 3 6 E 0 3 0 e 3 2 4 6 1 8 E 03 OeOOOOOOE 00 0 e 3 5 2 8 3 6 E 02 0 9 1 2 9 8 7 8 E 0 3 0 e l 5 3 3 7 5 E 03 O o 8 1 6 3 0 8 E 03 0 e l l 3 4 8 4 E 0 4 O o 7 4 9 C 5 2 E 0 2 O o 3 0 5 9 l 8 E 03 0 o 2 3 6 0 3 7 E 0 4 O e 1 2 5 8 8 6 E 0 4 OoOOOOOOE 00 0 o 5 2 7 1 4 1 E 02 O o 2 C 0 0 8 3 E 03 0 o 6 2 4 7 9 4 E 03 O o 3 1 2 2 4 7 E 0 4 0 . 4 0 0 0 0 6 E 0 4 0 . 7 2 2 9 1 5 E 02 0 . 4 4 8 7 3 3 E 03 0 o 4 0 7 7 9 1 E 0 4 0 o 2 l 5 7 8 3 E 0 4 OeOOOOOOE 00 Oe 5 0 9 8 3 4 E 0 2 0 . 1 9 3 2 6 5 E 0 3 Oo 1 0 9 0 1 2 E 0 4 0 o 5 4 2 2 4 4 E 0 4 O o 6 7 5 6 8 2 E 0 4 0 o 6 9 6 7 7 7 E 02 0 o 3 2 8 l 2 2 E 0 3 0 o 5 3 1 0 5 4 E 0 4 0 o 2 7 9 9 5 4 E 0 4 OoOOOOOOE 00 0 o 4 9 1 4 4 6 E 0 2 0 o l 8 6 2 3 2 E 0 3 0 o 7 l 3 8 2 4 E 03 0 o 7 5 5 3 6 8 E 0 4 0 o 8 5 0 7 8 8 E 0 4 0 . 6 7 2 1 0 . 2 E 0 2 0 e 3 l 0 3 0 7 E 0 3 0 o 6 2 3 7 9 3 E 0 4 Oo 3 2 6 0 1 2 E 0 4 O.OOOOOOE 00 0 o 4 7 5 0 2 8 E 02 0 . 1 7 9 7 5 0 E 0 3 0 o 6 7 5 3 4 0 E 0 3 0 o 8 9 7 2 9 8 E 0 4 0 o 9 8 7 5 5 8 E 0 4 136 F I N A L WEIGHT (KG) 0 + 1+ 2 + 3 + 4 + 0 . OOOOOOE 0 0 ' 0 e 3 2 8 7 6 6 E 02 0 e 5 2 1 2 3 3 E 0 2 Oe 6 1 6 9 8 9 E 02 ' • 0 . 3 2 9 9 5 2 E 02 0 . 4 2 3 6 9 2 E 0 2 0 e 5 8 6 0 9 9 E 0 2 0 . 3 3 7 5 2 1 E 0 2 0 e 4 4 9 6 2 7 E 0 2 0 . 1 7 9 6 9 4 E 03 i Oo OOOOOOE 0 0 0 e 3 l 6 8 4 0 E 0 2 0 o 5 0 3 0 0 5 E 0 2 Oe 5 9 5 6 5 9 E 02 0 . 3 1 8 1 6 7 E 02 Oe 4 0 9 1 9 4 E 02 0 e 5 6 4 7 9 1 E 02 Oe 3 2 6 6 0 5 E 02 0 . 4 3 3 0 8 1 E 0 2 0 e 1 7 3 3 6 7 E 0 3 0 e OOOCOOE 0 0 0 e 3 0 6 5 1 5 E 02 0 e 4 8 7 3 8 1 E 0 2 Oe 5 7 6 6 9 9 E 02 0 o 3 0 8 3 4 8 E 02 0 . 3 9 5 6 6 3 E 0 2 0 e 5 4 7 4 3 3 E 02 0 e 3 1 4 5 9 4 E 02 0 e 4 2 1 2 5 4 E 0 2 Oe 1 6 7 S 9 4 E 0 3 0 . OOOOOOE 0 0 0 e 2 9 5 4 . 7 9 E 0 2 0 « 4 7 1 3 2 4 E 0 2 0 e 5 5 6 l 5 9 E 02 0 . 2 9 6 5 6 3 E 02 0 e 3 8 1 9 5 5 E 02 0 e 5 2 6 7 5 1 E 02 Oe 3 0 4 1 1 1 E 0 2 0 o 4 0 4 7 0 8 E 0 2 0 e 1 6 1 9 5 2 E 03 0 . OOOCOOE 0 0 0 0 3 5 6 8 9 3 E 02 0 e 5 2 3 2 7 2 E 0 2 Oe 4 6 6 0 4 4 E 02 0 • 5 6 9 5. 59 E 0 1 Oe 4 0 1 0 8 7 E 02 0 o 5 7 9 S 4 0 E 02 Oe 1 9 3 0 9 1 E 02 0 o 2 2 9 1 4 6 E 02 0 . 1 4 0 3 1 6 E 03 0 e 1 5 6 0 2 0 E 0 1 0 e 4 2 7 4 l 6 E 02 0 . 5 7 9 5 3 4 E 0 2 Oe 3 8 0 8 9 5 E 02 0 . 7 8 5 6 0 0 E 00 r\ . 4 3 2 9 3 5 E 02 0 . 6 4 8 7 1 5 E 02 0 e 1 6 5 6 7 5 E 02 0 e 1 6 3 9 7 8 E 02 0 . 1 4 1 1 3 0 E 0 3 Ce 7 9 2 3 0 3 E 0 1 0 e 5 l 6 2 3 3 E 02 0 e 6 4 5 0 3 3 E 0 2 Oe 3 1 6 8 0 4 E 02 • OeOOOOOOE 00 Oe 3 9 3 3 3 7 E 02 0 e 4 9 1 6 7 2 E 02 0 e 2 6 1 7 7 1 E 0 2 0 e 4 0 5 5 2 0 E 02 0 e 1 5 5 7 3 0 E 03 Oe 3 9 4 9 3 4 E 02 0 0 6 2 2 2 5 3 E 02 0 • 7 2 4 3 5 8 E 0 2 0 . 3 8 4 7 4 9 E 02 Oo OOOCOOE 00 ^ e 5 9 4 3 4 8 E 02 0 e 7 5 7 8 l 2 E 0 2 Oe 3 0 & 6 2 6 E 02 0 • 4 6 7 5 0 7 E 0 2 0 e 2 1 2 6 2 9 E 03 C e 3 8 C 7 2 7 E 02 0 o 5 9 9 7 0 5 E 0 2 0 . 6 9 9 3 9 7 E 0 2 Oe 3 7 1 C 0 8 E 02 OoOOCOOOE CO 0 e 5 7 2 7 9 1 E 02 0 . 7 3 0 7 2 7 E 0 2 Oe 2 9 6 4 6 0 E 02 0 # 4 5 1 3 6 0 E 0 2 0 • 2 0 5 1 3 3 E 03 0 e 3 6 7 4 0 9 E 02 0 e 5 7 8 4 5 7 E 0 2 0 e 6 7 7 8 0 3 E 0 2 Oe 3 5 7 2 6 7 E 02 ' 0 • 0 0 0 0 0 0 E 00 Oe 5 5 3 9 8 4 E 0 2 0 e 7 C 5 8 3 0 E 02 0 . 2 8 5 1 1 9 E 02 0 e 4 3 6 0 0 2 E 02 Oe 1 9 8 0 9 3 E 0 3 0 e 3 5 4 0 9 1 E 02 0 . 5 5 7 6 4 3 E 02 0 • 6 5 4 9 2 0 E 0 2 0 . 3 4 3 5 2 6 E 02 OoOOOOOOE 00 0 e 5 3 4 0 C 4 E 02 0 e 6 8 0 1 4 4 E 02 Oe 2 7 5 3 6 7 E 02 0 . 4 2 0 6 4 4 E 0 2 0 e 1 9 1 0 1 8 E 0 3 Oe 3 4 1 6 6 0 E 02 0 e 5 3 8 l 3 0 E 02 0 . 6 3 3 6 1 6 E 0 2 Oe 3 3 1 7 4 8 . E 02 . OoOOCOOOE 00 0 e 5 1 6 1 6 5 E 02 0 . 6 5 6 4 7 0 E 02 0 e 2 6 6 4 4 3 E 02 0 e 4 0 6 0 7 6 E 02 0 e 1 8 4 5 1 5 E 03 137 CHANGE I N 30DY ENERGY ( K C A L ) 0 + 1 + 2 + 3 + 4+ 0 .OOOOCOE 0 0 - 0 . 7 6 9 5 7 1 E 0 3 - 0 . 1 2 1 8 8 1 E 0 4 - Q . 1 8 2 9 8 4 E 0 4 - 0 • 1 0 5 4 9 9 E 04 0 o 9 9 2 6 3 7 E 0 3 - 0 . 1 3 7 1 0 6 E 0 4 0 o 7 8 5 4 1 6 E 03 - Q . 1 7 2 4 1 0 E 0 4 G o 4 8 7 3 2 2 E 0 4 0 oOOOOOOE 0 0 - 0 8 2 0 4 4 3 6 E 0 4 - 0 . 3 4 3 8 0 4 E 0 4 - 0 e 4 6 9 7 5 6 E 0 4 - 0 . 2 8 0 6 7 9 E 04 0 . 2 6 4 2 5 4 E 0 4 - Q . 3 6 4 1 9 2 E 0 4 0 • 2 9 1 3 8 . 7 E 0 4 » 0 e 3 7 8 8 4 1 E 0 4 0 o 1 2 9 8 6 7 E 05 Oo GOGOGGE oc - G . 1 7 C 6 6 6 E 0 4 - 0 . 4 1 4 7 5 4 E 0 4 - 0 . 6 8 7 6 5 1 E 0 4 - 0 . 5 3 4 8 4 5 E 04 0 o 2 2 0 4 9 4 E 0 4 - 0 . 3 0 4 6 1 2 E 0 4 0 o 7 6 9 2 2 7 E 0 4 ~ O o 5 1 3 5 8 3 E 0 4 0 o 1 8 0 7 9 1 E 0 5 0 eOOOOOOE 0 0 0 . 1 6 7 2 8 6 E 0 4 0 . 2 9 7 4 8 7 E 0 4 0 o 4 5 l 8 0 8 E 0 4 0 o 2 8 8 4 0 3 E 04 0 . 2 1 6 4 3 1 E 0 4 0 . 2 9 9 1 6 2 E 0 4 0 e 2 9 9 3 S 5 E 0 4 0 . 3 8 9 9 9 5 E 0 4 0 o 1 2 C 4 9 8 E 05 0 •OOOOOOE 0 0 - 0 . 1 5 4 9 2 0 E 0 9 - 0 . 1 9 1 9 1 4 E 0 9 - 0 e 7 8 9 4 1 7 E 0 8 O o 1 9 8 6 6 6 E 0 4 0 e l 7 4 2 5 6 E 0 9 - 0 . 2 5 1 5 3 9 E 09 0 e 9 5 1 l 4 7 E 0 4 0 . 1 1 7 6 4 9 E 05 0 e 4 2 5 7 7 4 E 0 9 0 o l 2 0 6 0 6 E 0 4 0 . 1 3 5 3 8 0 E 05 O o 1 4 8 3 7 4 E 0 5 0 . 1 4 3 9 0 1 E 0 5 0 o l 4 3 5 6 0 E 03 0 0 1 2 8 8 2 6 E 05 0 . 1 8 4 2 6 5 E 05 0 . 6 1 1 6 7 6 E 0 4 0 . 6 6 8 9 3 3 E 0 4 0 o 4 4 1 1 5 2 E 05 0 . 8 4 9 7 2 4 E 04 0 . 1 2 9 7 1 5 E 0 5 0 e 1 0 9 4 1 7 E 0 5 0 . 8 1 3 2 9 8 E 0 4 OoOOOOOOE 00 0 . 1 1 5 6 0 C E 05 0 . 1 5 2 7 6 1 E 05 0 . 5 6 7 5 9 8 E 0 4 0 o 8 0 3 l 3 4 E 0 4 0 • 4 0 5 4 3 5 t 05 0 e 4 l 7 4 5 6 E 05 0 . 1 5 4 8 0 0 E 05 0 . 1 1 4 4 6 4 E 0 5 0 . 9 3 7 8 9 4 E 0 4 OoOOOOOOE 00 0 . 2 7 0 1 4 3 E 05 0 . 3 6 3 2 7 1 E 05 0 . 6 5 8 9 2 4 E 0 4 0 . 8 1 2 0 3 1 E 0 4 0 . 7 8 C 5 1 1 E 05 0 . 1 1 6 9 6 7 E 0 4 - 0 . 1 9 2 9 1 9 E 0 4 - 0 e 3 0 6 5 2 8 E 0 4 - 0 . 1 6 2 8 0 0 E 0 4 OeOOOOOOE 00 0 • 1 7 6 1 2 5 E 0 4 - 0 . 2 2 4 3 5 0 E 0 4 0 . 1 1 8 7 53E 0 4 - 0 . 2 5 9 9 8 5 E 0 4 0 • 7 7 9 2 1 6 E 0 4 0 . 7 4 7 4 8 4 E 0 3 - 0 . 1 0 5 9 3 5 E 0 4 - 0 . 2 0 6 7 4 5 E 0 4 - 0 . 109 4 1 4 E 0 4 OoOOOOOOE 00 0 . 1 1 2 8 0 5 E 0 4 - 0 . 1 4 3 5 0 8 E 0 4 0 . 2 0 7 2 7 7 E 03 - 0 . 2 1 9 8 0 1 E 0 4 0 . 4 9 6 8 4 3 E 0 4 0 • 8 3 1 7 4 1 E 03 - 0 . 1 3 0 3 1 3 E 0 4 - 0 . 2 1 3 3 0 5 E 0 4 - 0 . 1 1 2 0 6 5 E 0 4 OoOOOOOOE 00 0 • 1 2 5 5 4 4 E 0 4 - 0 . 1 5 9 6 6 3 E 0 4 0 • 6 2 5 9 5 5 E 03 - 0 . 1 9 1 0 5 6 E 0 4 0 . 5 3 8 8 5 9 E 0 4 A • 5 7 5 4 0 4 E 0 3 - 0 . 9 0 3 8 7 9 E 03 - 0 . 1 5 4 6 3 3 E 0 4 - 0 . 8 0 9 1 4 9 E 03 • •• -OeOOOOOOE 00 A • 8 7 0 0 3 9 E 03 - 0 . 1 1 0 4 8 7 E 0 4 0 • 4 4 1 4 6 8 E 0 3 - 0 . 1 4 1 8 3 8 E 0 4 0 • 3 8 3 4 7 6 E 0 4 138 CHANGE I N GONAD ENERGY ( K C A L ) 0 + 1 + 2+ 3 + 4 + 0 .OOOOOOE 0 0 0 . 2 1 7 5 5 6 E - 0 5 - 0 . 1 4 6 4 5 4 E 0 1 0 . 6 8 7 8 7 5 E 03 0 . 5 0 4 7 7 0 E 03 U • 3 3 6 7 6 6 E - 0 5 0 . 5 4 2 4 0 2 E - 0 5 0 . 6 0 9 0 7 6 E 0 1 0 e l l 9 7 2 7 E 0 4 0 . 1 1 9 1 1 8 E 0 4 0.OOOOOOE 0 0 Q e 2 7 7 l 6 l E - 0 5 - 0 . 1 3 8 6 7 7 E 0 2 - 0 . 1 3 2 1 8 9 E 0 3 - 0 . 1 0 5 8 4 1 E 03 0 . 1 0 7 2 8 8 E - 0 5 O e 1 4 0 C 7 0 E - 0 4 0 . 5 7 9 7 8 5 E 0 2 - 0 e l 9 3 9 2 Q E 03 0 . 2 5 1 8 9 8 E 03 0 eOOOOOOE 0 0 0 . 5 2 7 5 0 1 E - 0 5 - 0 . 1 5 7 4 1 8 E 0 3 - 0 . 6 8 1 9 7 5 E 0 4 - 0 . 5 1 2 7 7 3 E 0 4 0 . 1 4 6 0 3 1 E - 0 5 0 . 1 4 3 0 5 1 E - 0 4 0 . 6 5 2 0 4 5 E 03 - 0 . 1 1 4 5 2 8 E 05 0 . 1 2 1 G 4 9 E 0 5 0 . OOOOOOE 0 0 0 . 3 8 7 4 3 0 E - 0 5 . - 0 . 1 3 9 0 6 9 E 0 2 ' - 0 . 2 4 5 3 2 8 E 02 - 0 . 2 7 2 0 2 0 E 02 0 e 2 7 7 1 6 1 E - 0 5 0 . 1 3 3 8 1 2 E - 0 4 0 . 5 7 6 6 1 8 E 02 - 0 . 7 9 7 9 9 9 E 0 1 o . 6 5 6 4 1 7 E 02 0 .OOOOOOE 00 0 . 1 4 1 1 5 4 E 0 2 0 . 1 5 2 9 1 1 E 0 2 0 . 3 1 6 0 5 3 E 02 0 . 8 0 4 3 8 9 E 00 0 . 6 6 4 8 2 0 E 0 1 0 . 2 6 8 0 9 3 E 0 2 0 . 5 4 4 0 3 0 E 0 1 0 . 2 2 9 1 9 0 E 0 2 0 . 6 1 8 1 6 8 E 02 0 e l 6 8 3 0 8 E 0 1 0 . 1 6 4 1 3 3 E 0 2 0 . 1 9 0 7 7 5 E 0 2 0 8 3 8 6 7 7 3 E 02 0 . 4 6 4 6 7 3 E 00 0 . 7 1 7 9 4 1 E 0 1 0 . 3 0 2 0 9 0 E 0 2 . 9 3 4 2 3 5 E 0 1 0 . 2 9 5 8 5 2 E 02 0 . 7 6 3 1 5 9 E 02 0 • 1 2 5 7 3 8 E 0 2 0 . 5 4 1 9 3 4 E 0 2 0 . 4 9 5 3 4 4 E 0 3 0 e 2 5 1 4 0 2 E 03 O.OOOOOOE 00 0 . 8 4 2 4 6 2 E 0 1 C . 3 3 0 1 6 2 E 0 2 0 . 1 1 3 3 5 8 E 0 3 0 . 6 5 8 7 1 4 E 03 0 • 8 1 3 5 1 4 E 0 3 0 8 6 2 6 7 0 9 E 0 2 0 e 2 5 6 7 8 7 E 02 0 . 1 5 6 6 4 6 E 0 4 o . 8 6 4 2 7 1 E 03 OeOOOOOOE 00 0 e l 9 4 l 4 5 E 0 2 0 . 7 7 6 2 0 9 E 0 2 r> . 4 1 8 0 6 5 E 0 1 0 e 2 4 1 7 8 7 E 0 4 0 . 2 5 1 9 0 8 E 0 4 0 . 3 0 1 Q 0 3 E - 05 0 . 1 5 9 9 7 1 E 0 3 0 . 1 8 5 1 7 5 E 0 4 0 . 9 S 7 9 3 5 E 03 O.OOOOOOE 0 . 2 1 9 0 4 7 E - 05 0 . 9 5 0 6 9 4 E - 0 5 0 • 5 0 8 6 9 2 E 03 0 . 2 4 9 0 9 7 E 0 4 0 . 2 9 9 9 6 6 E 0 4 0 • 7 1 5 2 5 5 E - 05 - 0 . 1 0 8 S 9 4 E 03 0 . 1 4 1 6 5 3 E 0 4 0 • 7 5 3 4 8 7 E 03 OeOOOOOOE 00 0 . 3 5 6 1 3 7 E - 05 0 . 1 4 1 8 5 9 E - 0 4 0 • 3 4 5 8 9 8 E 0 3 0 . 2 4 0 7 0 2 E 0 4 0 . 2 0 6 1 1 2 E 0 4 0 . 2 5 3 3 1 9 E - 05 - 0 . 6 5 5 4 9 5 E 0 1 0 . 1 1 3 0 5 5 E 0 4 0 . 5 9 6 4 0 4 E 0 3 O.OOOOCOE CO 0 . 3 7 9 9 7 9 E - 05 0 . 5 8 1 1 4 5 E - 0 5 r\ . 2 0 8 1 4 5 E 0 2 0 . 1 7 4 1 2 1 E 0 4 0 . 1 7 2 0 4 0 E 0 4 0 . 2 3 8 4 1 8 E - 05 - 0 . 6 4 2 9 2 4 E 0 1 0 . 8 7 9 2 3 8 E 0 3 0 . 4 5 9 4 6 1 . E 03 OeOOOOOOE 00 0 . 1 7 7 3 2 3 E - 0 5 0 . 8 6 1 2 8 7 E - 0 5 0 . 2 0 4 6 8 8 E 02 0 . 1 3 5 2 7 3 E 0 4 0 . 1 3 3 2 2 7 E 0 4 139 A p p e n d i x T a b l e 3. E n e r g y b u d g e t o f m a t u r e a n d i m m a t u r e k o k a n e e o f b o t h s e x e s b y m o n t h . K e y t o t a b l e : F o r e a c h age g r o u p Number p r e s e n t a t b e g i n n i n g o f month m a l e i m m a t u r e f e m a l e i m m a t u r e m a l e m a t u r e f e m a l e m a t u r e W e i g h t ( k g ) p r e s e n t a t b e g i n n i n g o f m o n t h I n t a k e e n e r g y ( K c a l ) E g e s t i o n - e x c r e t i o n ( K c a l ) R o u t i n e m e t a b o l i s m ( K c a l ) B o d y e n e r g y ( K c a l ) Gonad e n e r g y ( K c a l ) W e i g h t ( k g ) a t e n d o f m o n t h Change i n b o d y e n e r g y ( K c a l ) Change i n g o n a d e n e r g y ( K c a l ) 140 JANUARY 0 + 1+ 2 + 3 + 4 + •"• 0 2 5 6 4 6 3 4 0 0 0 2 5 6 4 4 6 3 0 0 0 0 1 0 3 4 2 6 4 0 0 0 1 2 0 4 2 6 4 0 0 •OOOOOOE 0 0 0 . 3 9 9 9 8 4 E 02 0 • 1 7 5 6 1 8 E 0 2 OoOOOOOOE 00 0 i. •OOOOOOE 00 0 . OOOOOOE 0 0 0 e 3 9 9 9 8 4 E 02 0 o l 2 8 2 5 0 E 0 2 OeOOOOOOE 00 0 •OOOOOOE 00 0 .OCOOOOE 0 0 O.OOOOOOE 00 0 o 2 8 6 4 l 7 E 0 2 0 o l 2 9 0 9 5 E 02 • 0 •OOOOOOE 00 0 •OOOOOOE 0 0 0 .OOOOOOE 0 0 0 o 3 3 3 5 0 7 E 02 . 0 . 1 2 9 0 9 5 E 0 2 0 •OOOOOOE do 0 » OOOOOOE 0 0 0 . 1 1 3 4 5 5 E 05 0 • 4 9 8 1 4 0 E 0 4 OeOOOOOOE CO 0 iOOOOOOE 00 r\ \j .OOOOOOE 0 0 0 . 1 1 3 4 5 5 E 0 5 0 o 3 6 3 7 8 3 E 0 4 OoOOOOOOE 00 0 iOOOOOOE 00 0 .OOOOOOE 0 0 OoOOOOOOE 00 0 o 8 1 2 4 2 4 E 0 4 0 . 3 6 6 1 8 0 E 0 4 0 .OOOOOOE CO 0, .OOOOOOE 0 0 "O.OOOOOOE 00 0 • 9 4 5 9 9 5 E 0 4 0 e 3 6 6 1 8 0 E 0 4 ' 0 .OOOOOOE 00 0 < .OOOCOOE 0 0 ' 0 . 2 2 6 9 1 0 E 0 4 • 0 • 9 9 6 2 8 0 E 0 3 O.OOOOOOE 0 0 •'. 0 » 0 0 0 0 0 0 E 00 0 .OOOCOOE 0 0 0 . 2 2 6 9 1 0 E 0 4 0 o 7 2 7 5 6 7 E 0 3 OeOOOOOOE . 0 0 . 0 I O O O O O O E 00 0, , OOOOOOE 0 0 O.OOOOOOE 00 0 o l 6 2 4 8 4 E • 0 4 0 e 7 3 2 3 6 1 E 0 3 • 0 .OOOOOOE' CO 0. . OOOOOCE 0 0 O.OOOOOOE 00 0 • 1 8 9 1 9 9 E 0 4 0 . 7 3 2 3 6 1 E 0 3 0 .OOOOOOE 00 Oi .OOOCOOE 0 0 . 0 . 2 6 1 4 1 7 E 0 4 0 • 1 3 9 8 4 7 E 0 4 OeOOOOOOE 0 0 ; Oi .OOOOOOE 00 Oi .OOOOOOE 0 0 0 . 2 6 1 4 1 7 E 0 4 0 • 1 1 0 1 3 0 E 0 4 ' OeOOOOOOE oo •". • Oi . OOOOOOE 00 0, .OOOOOOE 0 0 . O.OOOOOOE 00 0 • 2 0 2 8 1 6 E 0 4 0 e l l 0 6 8 1 E 0 4 .. . OI .OCOOOOE 00 0 , .OOOOOOE 00.-. O.OOOOOOE 00 0 • 2 2 7 6 8 9 E 0 4 O e 1 1 0 6 8 1 E 0 4 • 0 , OOOOOOE 00 0 , .OOOOOOE 0 0 • 0 . 5 4 8 0 8 0 E 05 0 • 2 4 0 6 4 1 E 0 5 OeOOOOOOE 0 0 -; Oi .OOOOOOE 00 0, .OOOOOOE 0 0 0 . 5 4 6 2 6 9 E 05 0 • 1 7 5 1 5 5 E 0 5 OeOOOOOOE 0 0 OI . OOOOOOE 00 0 , , OOOOOOE 0 0 O.OOOOOOE 00 • 0 o 3 9 2 4 2 6 E . 0 5 0 . 1 7 6 8 7 6 E 0 5 . Oi .OOOOOOE 00 0. .OOOOOOE 0 0 . O.OOOOOOE 00 . 0 o 4 5 3 9 7 0 E 0 5 0 . 1 7 5 7 2 5 E 0 5 . ; o I . OOOOOOE 00 0« .OOOOOOE 00 0 . 9 5 3 2 9 7 E 02 . • b o 4 1 8 5 5 7 E 0 2 . OoOOOOOOE "0 0 ' Oi .OOOOOOE 00 Oi OOOOOOE 0 0 0 . 3 3 4 0 1 0 E 03 - 0 o l 0 7 0 9 7 E 0 3 ' O.OOOOOOE 00 CI .OOOOOOE 00 0. .OOOOOOE 0 0 O.OOOOOOE 00 0 . 7 3 1 3 9 0 E 0 2 0 . 3 2 9 6 5 6 E 0 2 Oi OOOOOOE 00 0, .OOOOOOE 0 0 OoOOOOOOE 00 0 • 4 7 4 0 4 1 E 0 3 • 0 . 1 8 3 4 9 4 E 03 " OI OOOOOOE 00 Oi .OOOOOOE 0 0 ' O o 3 9 9 9 8 4 E 02 . 0 • 1 7 5 6 1 8 E 0 2 OoOOOOOOE 00 . ' 0 * OOOOOOE CO OI OOOOOOE 0 0 0 . 3 9 9 9 8 4 E 02 0 • 1 2 8 2 5 0 E 0 2 O.OOOOOOE .0 0 OI OOOCOOE 00 OI OOOOOOE 0 0 OoOOOOOOE 00 . 0 • 2 8 6 4 1 7 E 0 2 Oe 1 2 9 0 9 5 E 02 OI OOOOOOE CO OI OOOOOOE 0 0 . OoOOOOOOE 0 0 0 • 3 3 3 5 0 7 E 0 2 0 . 1 2 9 0 9 5 E 02 OI OOOOOOE 00 0 i .OOOOOOE 0 0 - O o l 2 6 3 6 6 E 0 4 - 0 • 5 5 4 8 2 5 E 0 3 0 .OOOOOOE 0 0 OI OOOOCOE 00 0. OOOOOOE 0 0 ' - 0 o l 2 5 9 4 9 E 0 4 - 0 • 4 0 3 8 4 1 E 0 3 . O.OOOOOOE 0 0 OI OOOOCOE 00 0, OOOOOOE oo OEOOOOOOE 00 -o • 9 0 4 7 8 2 E 0 3 . - O o 4 0 7 8 l l E 03 OI OOOOOOE 00 0 . OOOOOOE 00 O.OOOOOOE 00 - 0 • 1 0 4 6 6 8 E 0 4 - C e 4 0 5 1 5 5 E 03 , Oe OOOOOOE 00 0 . OOOOOOE 0 0 •'' 0 . 1 2 6 3 6 1 E - 0 4 0 • 7 7 4 8 6 0 E - 0 6 . O.OOOOOOE .0:0. •: Oe OOOOOOE 00 0 . OOOOOOE 0 0 0 . 5 3 8 S 2 6 E - 0 4 0 • 2 8 6 1 0 2 E - 0 5 OeOOOOOOE 0 0 ' • OI O O O O O O E 00 0 . OOOOOOE 0 0 , O.OOOOOOE 00 . 0 • 7 5 1 C 1 8 E - 0 5 0 e 4 6 4 9 l 6 E - 0 5 .'• 0 . OOOOOOE 00 0, OOOOOOE 0 0 O.OOOOOOE 00 : 0 • 1 2 3 9 7 7 E - 0 4 0 e 6 6 7 5 7 2 E - 0 5 . O i OOOOOOE 00 141 FEBRUARY 0 + 1+ 2 + 3+ 4 + 0 2 4 7 2 5 9 5 0 0 0 2 4 7 2 4 3 5 0 0 0 0 97 2 25 5 0 0 0 1 1 3 2 2 5 5 0 OeOOOOOOE 0 0 0 8 3 8 5 6 3 2 E 0 2 0 « 1 6 4 8 1 5 E 0 2 O.OOOOOOE 0 0 OeOOOOOOE 00 OeOOOOOOE 0 0 0 . 3 8 5 6 3 2 E 0 2 0 o l 2 0 4 9 5 E 0 2 OeOOOOOOE 0 0 O.OOOOOOE 00 OoOOOOOOE 0 0 O.OOOOOOE 0 0 0 e 2 6 9 2 4 4 E 0 2 O e 1 2 4 6 9 4 E 0 2 OeOOOOOOE 00 OoOOOOOOE 0 0 OeOOOOOOE 0 0 0 e 3 1 3 5 6 3 E 0 2 0 . 1 2 4 6 9 4 E 02 OeOOOOOOE o'o Oo 0000OOE 0 0 0 o 8 7 6 7 7 2 E 0 4 0 e 3 7 4 7 2 3 E 0 4 O.OOOOOOE 0 0 OeOOOOOOE 00 OoOOOOOOE 0 0 0 9 8 7 6 7 7 2 E 0 4 0 e 2 7 3 9 5 7 E 0 4 OeOOOOOOE 0 0 OeOOOOOOE 00 0 o 0 0 0 0 0 0 E CO O.OOOOOOE 0 0 0 . 6 1 2 1 5 3 E 0 4 0 . 2 8 3 5 0 6 E 0 4 O.OOOOOOE 00 OoOOOOOOE 0 0 OeOOOOOOE 0 0 O o 7 1 2 9 l 9 E 0 4 0 . 2 8 3 5 0 6 E 0 4 OeOOOOOOE 00 Oe OOOOOOE 0 0 0 . 1 7 5 3 5 4 E 0 4 0 e 7 4 9 4 4 6 E 0 3 O.OOOOOOE 0 0 Oe OOOOOOE 00 O e OOOOOOE 0 0 0 . 1 7 5 3 5 4 E 0 4 0 e 5 4 7 9 l 4 E 0 3 O.OOOOOOE 0 0 OeOOOOOOE CO P. 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S 8 3 0 0 0 E 03 0 . 1 2 73 80E 05 0 . 5 9 2 3 0 0 E 0 4 0 . 5 2 78 00E 0 4 . 0 . 8 3 5 0 0 0 E 03 0 . 1 2 2 2 6 0 E 05 0 . 5 7 2 2 0 0 E 0 4 0 . 5 0 8 5 0 0 E 0 4 0 . 7 9 1 0 0 0 E 03 0 . 1 1 7 5 2 0 E 05 0 . 5 5 1 6 0 0 E 0 4 0 . 4 8 5 1 G 0 E 0 4 0 . 7 8 0 C 0 0 E 02 0 a 9 8 5 8 0 0 E 0 4 0 . 5 3 2 4 0 0 E 0 4 0 . 4 6 7 0 0 0 E 0 4 0 . 8 0 0 0 0 0 E 0 1 0 . 9 3 5 5 0 0 E 0 4 1 + 0 . 5 1 2 B 0 0 E 0 4 0 . 3 0 2 7 0 0 E 0 4 0 . 1 4 6 8 0 0 E 0 4 G . 4 9 4 4 0 0 E 0 4 0 . 2 9 0 7 0 0 E 04 0 . 1 3 8 7 0 0 E 0 4 C . 4 7 8 4 0 0 E 0 4 G . 2 8 0 3 0 0 E 0 4 0 . 1 3 1 7 0 0 E 0 4 0 . 4 & 1 2 0 0 E 0 4 0 . 2 6 9 3 C 0 E 04 0 . 1 2 4 4 C 0 E 0 4 0 . 4 4 5 2 0 0 E 0 4 0 . 6 0 1 7 0 0 E 0 4 0 . 1 1 7 7 0 C E 0 4 0 . 4 2 9 2 0 0 E 0 4 0 . 5 7 9 2 0 0 E 0 4 0 . 1 H 2 0 0 E 0 4 0 . 4 1 4 2 0 0 E 0 4 C . 5 5 8 2 0 0 E 0 4 0 . 1 0 5 2 0 0 E 0 4 0 . 3 9 9 4 0 0 E 0 4 0 . 5 3 7 5 0 0 E 0 4 C . 9 9 4 0 0 0 E 0 3 0 . 3 8 5 0 0 0 E 0 4 0 . 5 1 7 3 0 0 E 0 4 0 e 9 4 Q G C 0 E 03 G . 3 7 1 6 0 0 E 0 4 0 . 4 9 8 6 0 0 E 0 4 0 . 8 9 0 0 0 0 E 03 O . 3 5 8 2 0 0 E 0 4 0 . 4 8 4 2 0 0 E 0 4 0 . 8 7 0 0 0 0 E 02 0 e 3 4 5 6 0 0 E 0 4 0 . 4 6 6 9 0 0 E 0 4 C e e O O O O O E 0 1 153 2 + 0 . 3 3 3 5 0 0 E 0 4 0 . 3 1 3 4 0 0 E 0 4 0 . 2 9 6 2 0 0 E 0 4 0 . 2 7 8 4 0 0 E 0 4 0 . 2 6 2 0 0 0 E 0 4 0 . 2 4 6 2 C 0 E 0 4 0 . 2 3 1 8 0 0 E 0 4 0 . 2 1 7 8 0 0 E 0 4 Q . 2 0 4 6 0 0 E 0 4 0 . 1 9 2 6 0 0 E 0 4 0 . 7 1 0 0 0 0 E 0 3 0 . 5 6 9 0 0 0 E 0 3 3 + 0 . 5 2 8 00GE 0 3 0 . 5 1 0 0 0 0 E G3 0 . 4 9 4 0 G C E 0 3 G . 4 7 3 C 0 G E 0 3 G . 4 6 2 0 0 0 E C3 0 . 4 4 6 0 G 0 E 0 3 G . 4 3 0 0 G G E 03 0 . 4 1 6 0 G G E 0 3 0 e 4 G 2 G 0 0 E 0 3 0 . 3 8 8 0 0 0 E 0 3 0 . 5 0 0 G 0 G E 02 ;'\vJi''iEi£k 4 + 0 . 0 C G 0 C 0 E 00 C.OOOOOOE 00 0 . 0 G 0 0 u 0 E G 0 O.OOOOOOE GO O.OOOOOOE 00 OeOOOOOOE GO 0.0 00 G 0 0 E 0 G 0.OOOOOOE CO O.OGOOOGE 00 O.OOOOOOE 00 OeOOOOOOE 00 OeOOOOOOE 00 154 WEIGHT (KG) 0 •+• 1 + 2 + 3 + 4 + 0 . 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GOOCOOE 00 0 . 2 2 6 9 1 0 E 0 5 0 . 2 6 2 0 3 4 E 0 5 0 . 7 3 2 3 6 1 E 0 4 0.OOOOOOE 00 0 « 1 6 3 2 6 9 E 0 5 0 . 1 4 9 8 3 3 E 05 0 . 1 1 7 8 6 G E 0 5 G . 1 3 1 2 1 7 E 0 5 0 . 5 6 21 8 1 E 0 5 O .OOOOOOE 00 0 . 1 7 5 3 5 4 E 05 0 . 1 9 7 3 7 5 E 0 5 0 . 5 6 7 0 1 2 E 04 0 .OOOOOOE o c 0 . 1 2 5 1 4 9 E 0 5 0 . 1 1 5 0 7 3 E 0 5 0.895 6 59E 0 4 0 . 9 9 6 4 2 5 E 04 0 . 4 2 9 4 3 I E 0 5 O .OOOOOOE 00 G . 1 7 5 8 2 9 E 0 5 0 . 1 9 3 3 0 3 E 0 5 0 . 5 6 9 1 2 9 E 0 4 o . o o o q o o E 00 0 . 1 2 4 6 5 6 E 0 5 0 . 1 1 4 7 3 6 E 0 5 0 .88 3 6 6 2 E 04 0 . 9 8 2 8 5 9 E C4 0 . 4 2 6 0 4 5 E 0 5 CO 0 . 3 1 3 0 2 2 E 0 5 0 . 3 3 5 5 1 3 E 05 0 . 1 0 1 6 9 4 E 05 O.OOOOOOE 00 0 . 2 2 0 2 6 3 E 0 5 0 . 2 C 3 1 5 0 E 0 5 0 • 1 5 4 8 5 1 E 0 5 0 . 1 7 1 9 6 4 E 0 5 0 . 7 5 0 2 3 0 E 05 0 . 1 7 3 6 8 4 E 0 4 0 . 5 8 6 6 4 5 E 0 5 0 . 6 1 3 0 2 3 E 0 5 0 • 1 9 0 8 3 0 E 0 5 OeOOOOOOE 00 0 . 4 1 8 5 2 8 E 0 5 0 . 3 8 7 1 7 5 E 0 5 0 . 2 6 5 4 0 5 E 0 5 0 . 3 1 6 7 5 8 E 05 0. 1 4 0 7 8 6 E 06 0 . 3 3 6 7 G 3 E 04 0 . 4 8 5 0 8 9 E 05 0 . 4 9 4 0 8 3 E 0 5 0 . 1 5 8 4 5 1 E 05 O.OOOOOOE 00 0 . 3 5 3 3 0 0 E 0 5 0 . 3 2 8 G 1 4 E 0 5 0 . 2 3 2 3 4 9 E 0 5 0 . 2 5 7 6 3 5 E 0 5 0 . 1 1 7 1 3 0 E 06 0 . 9 9 7 2 0 2 E 0 4 0 . 6 2 9 8 2 1 E 0-5 0 . 6 2 5 3 5 8 E 0 5 0 . 2 0 6 0 8 6 E 0 5 OeOOOOOOE 00 0 . 4 8 3 5 7 0 E 05 0 . 4 5 1 7 1 0 E 0 5 0 . 2 9 7 1 7 2 E 05 0 . 3 2 9 0 3 2 E 0 5 0 . 1 5 6 1 4 8 E 06 0 . 3 6 6 5 7 2 E 05 C . 9 8 7 6 0 4 E 05 0 . 9 5 6 2 8 6 E 0 5 0 . 3 2 5 1 3 0 E 0 5 OeOOOOOOE 00 0 . 8 5 S 8 6 2 E 0 5 0 . 8 1 0 1 2 5 E 0 5 0.4589 3 5 E 0 5 0 . 5 0 7 6 7 1 E 0 5 0 . 2 6 3 5 5 9 E 0 6 0 . 3 2 8 1 9 1 E 05 0 . 3 7 7 1 8 1 E 05 0 . 3 5 5 9 1 8 E 0 5 0 . 1 2 4 8 3 0 E 05 OeOOOOOOE 00 0 . 4 2 0 3 5 6 E 0 5 0 . 4 0 2 0 9 0 E 0 5 0 . 1 7 2 7 0 4 E 0 5 0 . 1 9 0 9 7 0 E 0 5 0 . 1 . 1 8 6 1 2 E 06 0 . 2 9 9 8 3 3 E 0 5 0 . 3 4 4 5 7 1 E 0 5 0 . 3 1 7 1 1 3 E 0 5 0 . 1 1 4 0 3 5 E 05 OeOOOOOOE 00 0 . 3 8 2 4 6 3 E 0 5 0 . 3 6 6 1 6 3 E 0 5 0 . 1 5 5 3 1 2 E 0 5 0 . 1 7 1 6 1 3 E 0 5 0 . 1 0 7 5 5 5 E 06 0 . 2 9 2 9 6 6 E 0 5 0 . 3 3 6 6 5 8 E 05 0 . 1 1 8 4 8 8 E 0 5 0 e 1 4 8 9 4 9 E 04 OeOOOOOOE GO 0 . 3 6 5 2 1 1 E 0 5 0 . 3 6 3 7 0 9 E 0 5 0 . 1 6 2 9 2 4 E 0 4 0 . 1 7 7 9 4 3 E 0 4 0 . 7 6 3 0 0 8 E 0 5 0 . 3 5 9 7 9 6 E 0 5 0 . 4 1 3 2 9 8 E •05 0 . 1 2 0 8 2 5 E 0 5 . 0 . 2 2 7 4 2 9 E 0 3 OeOOOOOOE 00 C . 4 4 6 0 0 4 E 0 5 0 . 4 4 5 7 9 2 E 0 5 C . 2 1 9 S 8 8 E 0 3 0 . 2 1 9 8 8 8 E 03 0 • 8 9 6 1 9 5 E 0 5 156 EGESTION-EXCRETION (KCAL) 0 + 1 + 2+ 3 + 4 + OiOOOCOOE 00 C 4 5 3 8 2 1 E 0 4 0 . 5 2 4 0 6 8 E 0 4 0 o 1 4 6 4 72 E 0 4 OeOOOOOOE 00 O e 3 2 6 5 3 8 E 0 4 0 e 2 9 9 6 6 7 E 0 4 0 . 2 3 5 7 2 1 E 0 4 0 e 2 6 2 4 3 5 E 0 4 0 . 1 1 2 4 3 6 E 05 OeOOOOOOE 0 0 O o 3 5 0 7 0 9 E 0 4 O o 3 9 4 7 5 0 E 0 4 0 e 1 1 3 4 0 2 E 0 4 OeOOCOOOE 00 0 . 2 5 0 2 9 9 E 0 4 0 e 2 3 0 l 4 5 E 0 4 0 o 1 7 9 1 3 1 E 0 4 O e l 9 9 2 8 5 E 0 4 0 e 8 5 8 8 6 2 E 0 4 Oe OOOOOOE 0 0 0 o 3 5 1 6 5 8 E 0 4 0 . 3 8 6 6 0 7 E 0 4 0 o l l 3 8 2 5 E 0 4 OeOOOOOOE 00 Oe 2 4 9 3 1 3 E 0 4 0 . 2 2 9 4 7 3 E 0 4 0 e 1 7 6 7 3 2 E 0 4 O o 1 9 6 5 7 1 E 0 4 0 , 8 5 2 0 9 1 E 0 4 OeOOOOOOE CO Oo6 2 6 0 4 4 E 0 4 0 e 6 7 l 0 2 6 E 04 0 . 2 0 3 3 8 9 E 0 4 . OeOOOOOOE 00 0 o 4 4 0 5 2 6 E 0 4 0 o 4 0 6 3 0 0 E 0 4 Oe 3 0 9 7 0 3 E 0 4 0 o 3 4 3 9 2 9 E 0 4 0 e 1 5 G 0 4 5 E 05 0 e 3 4 7 3 6 9 E 0 3 O o l l 7 3 2 9 E 05 0 e l 2 2 6 0 4 E 0 5 Oo38 166 0E 0 4 Oe OOOOOOE 00 0 . 8 3 7 0 5 6 E 0 4 0 e 7 7 4 3 5 0 E 0 4 Oo 5 7 0 8 1 1 E 0 4 O o 6 3 3 5 1 7 E 0 4 0 . 2 8 1 5 73E 05 0 . 6 7 3 4 0 6 E 03 0 o 9 7 0 l 7 8 E 0 4 0 e 9 8 8 l 7 7 E 0 4 0 . 3 1 6 9 0 3 E 0 4 OeOOOOOOE 00. Oe 7 0 6 6 0 1 E 0 4 0 o 6 5 6 0 2 8 E 0 4 0 e 4 6 4 6 9 8 E 0 4 0 e 5 l 5 2 7 1 E 0 4 0 . 2 3 4 2 5 9 E 05 0 e 1 9 9 4 4 0 E 0 4 0 e l 2 5 9 6 4 E 05 O o 1 2 5 1 7 1 E 0 5 0 . 4 1 2 1 7 2 E 0 4 OeOOOOOOE 00 0 e 9 6 7 l 4 l E 0 4 0 o 9 0 3 4 2 1 E 0 4 0 o 5 9 43 4 5 E 0 4 O o 6 5 8 0 6 4 E 0 4 0 . 3 1 2 2 9 7 E 05 0 e 7 3 3 l 4 5 E 0 4 0 0 1 9 7 5 2 0 E 05 0 . 1 9 1 2 5 7 E 0 5 0 . 6 5 0 2 6 0 E 0 4 OeOOOOOOE oo 0 o l 7 1 7 7 2 E 05 Oo1 6 2 0 2 5 E 0 5 0 o 9 1 7 8 6 9 E 0 4 0 . 1 0 1 5 3 4 E 05 0 o 5 2 7 1 18 E 05 0 . 6 5 6 3 8 2 E 0 4 O o 7 5 4 3 6 3 E 0 4 0 . 7 1 1 8 3 7 E 0 4 0 o 2 4 9 6 6 1 E 04 0 o OOOOOOE 00 0 o 3 4 C 7 12E C4 0 . 8 0 4 1 8 I E 0 4 0 . 3 4 5 4 0 9 E 0 4 O e 3 8 1 9 4 0 E 0 4 0 o 2 3 7 2 2 4 E 05 0 o 5 9 9 6 6 6 E 0 4 0 e 6 8 9 1 4 l E 0 4 0 o 6 3 4 2 2 6 E 0 4 Oo 2 2 8 0 7 0 E 0 4 Co OOOOOOE 00 0 . 7 6 4 9 2 7 E 0 4 0 « 7 3 2 3 2 6 E 0 4 0 o 310 6 2 5 E 0 4 0 e 3 4 3 2 2 6 E 0 4 0 O215110E 05 0 o 5 85 93 2E 0 4 ' 0 e 6 7 3 3 1 6 E 0 4 0 . 2 3 6 9 7 7 E 0 4 0 o 2 9 7 8 9 8 E 03 OoOOOOOOE oo 0 o 7 3 0 4 2 3 E 0 4 0 . 7 2 7 4 1 9 E 0 4 0 o 3 2 5 8 4 8 E 0 3 0 . 3 5 5 8 8 7 E 03 0 . 1 5 2 6 0 1 E 05 0 . 7 1 9 5 9 3 E 0 4 0 . 3 2 6 5 9 7 E 0 4 0 e 2 4 l 6 5 0 E 0 4 0 o 4 5 4 8 5 8 E 02 0 eOOOOOOE 00 0o 8920 0 9 E 0 4 0 . B 9 1 5 8 4 E 0 4 0 e 4 3 9 7 7 6 E 02 0 e 4 3 9 7 7 6 E 02 O o 1 7 9 2 3 8 E 05 1 157 ROUTINE M E T A B O L I S M ( K C A L ) 0 + 1+ 2 + 3 + 4 + J ( i OOOOOOE 00 0 o522e35E 0 4 0 • 6 8 0 4 8 3 E 0 4 0 . 2 2 1 3 6 3 E 0 4 O.OOOOOOE CO 0 . 401 2 6 5E 04 0 • 3 7 1 5 4 8 E 0 4 0 . 3 1 3 4 9 7 E 0 4 0 • 3 3 8 3 7 1 E 04 c » 1 4 2 4 6 S E 0 5 .OOOOOOE 0 0 C • 4 5 9 3 0 4 E 0 4 w . 5 8 6 2 4 9 E 0 4 0 . 1 9 4 7 3 9 E 0 4 O.OOOOOOE 00 0 .35C015E 04 0 . 3 2 4 518E 0 4 0 . 2 7 2 1 4 8 E 0 4 .0 . 2 9 3 6 0 9 E 0 4 0 . 1 2 4 0 2 9 E 0 5 Oi OOOOCOE GO 0 » 4 9 6 9 2 9 E 0 4 0 . 6 2 3 0 3 1 E 0 4 0 . 2 1 0 8 5 5 E 0 4 O.OOOOOOE 00 Q . 3 7 S 4 9 1 E 0 4 0 e 3 4 9 2 5 8 E 0 4 0 . 2 9 1 0 7 0 E 0 4 0 • 3 1 3 9 9 5 E 0 4 ^ t .1330 8 I E 0 5 0 i OOOOOOE 0 0 • 4 7 1 3 5 5 E 0 4 0 . 5 7 9 7 0 9 E 0 4 0 . 2 0 0 5 6 6 E 0 4 O.OOOOOOE 00 0 . 3 5 4 7 6 9 E 0 4 0 . 3 2 9 5 8 8 E 0 4 0 i . 2 7 3 C 3 5 E 0 4 n . 2 9 4 2 3 8 E 0 4 . 1 2 5 1 6 3 E 05 0 . . 3 7 1 6 8 6 E 0 3 0 . 5 3 9 4 1 5 E 0 4 0 • 6 5 0 7 2 3 E 0 4 0 . 2 2 9 7 4 6 E 0 4 O.OOOOOOE 00 0 < . 4 2 2 0 0 6 E 0 4 0 . 3 9 3 6 6 2 E 0 4 0 •30 3 75 9E 0 4 0 .332625E 0 4 c • , 1 4 5 7 0 5 E 05 Oi . 8 6 1 0 7 6 E 0 3 0 • 6 5 3 9 7 0 E 0 4 0 • 7 7 3 7 1 4 E 0 4 0 . 2 7 9 4 1 8 E 0 4 0 . OOOOOOE 0 0 0 < > 5 2 9 0 3 5 E 0 4 . 4 9 5 3 1 2 E 0 4 Oi 3 7 0 2 3 4 E 0 4 0 . 3 9 8 6 2 3 E 0 4 0 i 1 7 9 3 2 1 E 0 5 0 j 21 3 3 3 3 E 0 4 . 8 6 5 7 4 1 E 0 4 • 1 0 0 5 1 9 E 0 5 0 . 3 7 0 3 9 0 E 0 4 O.OOOOOOE 00 0 . 7 4 5 8 3 3 E 04 0 • 7 0 2 2 9 5 E 04 C i .484694E 04 0 o 5 2 1 5 8 5 E 0 4 0 • 2 4 5 4 6 5 E 05 0 . 5 7 7 6 7 2 E 0 4 0 . 1 2 2 6 3 8 E 05 »139 68 I E 0 5 0 . 5 2 7 3 3 1 E 0 4 OoOOOOOOE 00 0 i . 1 1 S 9 3 0 E 05 0 o l 1 2 8 6 3 E 05 0 i . 6 7 9 83 I E 0 4 0 . 7 3 0 8 5 0 E 0 4 0 • . 3 7 2 8 6 9 E 05 , 7 385 1 0E 0 4 0 . 3 7 6 4 5 4 E 0 4 0 • 9 7 8 4 2 0 E 0 ^ 0* 2 7 8 2 2 9 E 0 4 OoOOOOOOE 00 Oi 1 0 3 3 5 7 E 05 0 . 9 9 C 8 1 2 E 0 4 0 i »48060eE 0 4 . 5 1 6 6 1 4 E 0 4 Ci 3 0 2 1 6 1 E C5 0 i 60099SL C4 0 • 5 6 7 9 9 9 E 0 4 0 . 7 3 1 6 4 0 E 0 4 0 o 2 8 8 2 6 6 E 0 4 0 o OOOCOOE 0 0 0 , 7 3 4 0 2 2 E 0 4 0 • 7 5 2 7 8 4 E 0 4 0 .3& 2 1 6 6 E 04 0 . 3 0 9 1 3 2 E 04 0 . 2 2 8 8 9 C E 0 5 0 .538 4 4 4 E 04 0 e 596444E 0 4 r. « 3 0 4 4 12E 0 4 0.5 59539E 0 3 OoOOOOOOE 00 0 . . 6 3 3 1 0 6 E 0 4 0 • 6 8 5 3 8 6 E 0 4 0 i .598655E 0 3 0 . 6 3 9 C 2 1 E 0 3 0 i . 1 4 9 7 2 5 E 05 0 . 5 3 8 4 5 6 E 0 4 0 .59B2 84E 04 0 . 2 4 2 7 0 2 E 0 4 0 . 1 1 4 7 4 9 E 0-3 0 o OOOOOOE 00 0 . 6 8 4 4 3 3 E 0 4 0 .684118E 0 4 0 •111833E 03 0 o 1118 3 3 E 0 3 Oi 1 3 9 0 9 I E 0 5 158 BODY ENERGY ( K C A L ) 0 + 1 + 2 + 3 + 4 + Oi .OCCOCCE OG Oi . 1 C 9 4 3 4 E 06 0 . 1 2 6 2 1 9 E 0 6 0 . 3 5 2 6 0 2 E 05 OaOOGCGOE 00 0 « . 7 8 B 7 2 1 E 05 Oi . 7 2 1 4 2 4 E 05 0 . . 5 6 9 3 0 3 E 05 0 , > 6 2 9 6 9 5 E 05 r) , 2 7 0 9 1 4 E 06 0 t OOOOOOE CC G ( 1 0 3 C 7 5 E 06 0 o l l 5 8 7 7 E 06 0 . 3 3 2 7 2 8 E G5 Oo OCGOOOE 00 0 ,736862c 05- Oi .675294E 0 5 0 , . 5 2 7 2 9 6E 0 5 0 , .582303E 0 5 o ,252225c 0 6 0 c GOOCOOE 0 0 G i .968227c 05 0 . 1 0 6 3 1 4 E 06 G e 3 1 2 8 6 4 E 05 OoCOOOOOE GO ,637575c 0 5 G i • & 3 0 7 6 9 E 05 0 .48 73 57E 05 0 i • 5 3 8 5 3 7 E 0 5 n ,234423E 0 6 Oi , OOOOOOE GO G i . 8 7 9 7 2 4 E 05 0 . 9 4 1 7 7 9 E 0 5 0.23 5 3 1 7 E 05 OoOOOOOOE 00 0 . 6 2 0 0 5 8 E 05 0 . 5 6 9 9 9 3 E 0 5 0 • 4 3 5 6 7 5 E 0 5 0 , .48Q894E 0 5 0 • 2 10 6 8 2 E 0 6 0 . 2 6 1 4 2 5 E 04 0< . 8 6 3 0 0 5 E 05 0 . 9 2 1 5 7 6 E 0 5 0 . 2 8 6 7 4 3 E 0 5 QoGCCOCOE '•j c 0 • 5 3 1 0 0 1 E 05 0 » 5 S 1 8 0 2 E 05 0 . 4 3 C 2 5 3 E 05 0 . 4 7 4 4 1 0 E 0 5 0 ,211746E 06 0 • S 2 2 5 9 9 E 0 4 G < ,113512c 0 6 0 . 1 2 0 4 3 0 E 06 0 . 3 8 5 8 1 2 E 05 ' OoOGCCCCE 00 0 •364577c 0 5 C< . 8 0 0 0 4 5 E 05 0 » 5 6 7 3 9 6 E 05 0 ,6 2 5 4 8 3c 05 0 . 285 75 0E 06 0 . 1 9 9 7 4 5 E 05 0 , • 1 2 6 1 5 6 E 06 0 o l 2 3 3 0 6 E 06 0 . 4 C 2 8 1 4 E 0 5 0.'GOOCOOE 00 n . 9 7 C 2 2 2 E 0 5 Oi . 9 0 3 3 0 5 E 05 0 , 5B3 6 9 9 E 05 0 . 6 3 9 9 7 G E 05 0 , 3 0 9 7 1 9 E 06 0 . 4 9 5 9 7 7 E C5 0 . 1 3 3 6 2 4 E 06 C . 1 2 3 3 6 3 E 0 6 0 o 4 0 9 75 2 E 0 5 0 * GOGOGGE 00 0 ,116397c 06 Oi 1 1 0 9 4 2 9 E 06 0 .58 6 4 2 8 E 05 G .6 3 0 9 Q 2 E G5 0 » 3 4 7 5 6 0 E 06 Oi i 1 2 9 3 7 0 E 06 0 , 1 4 8 632E 06 0 . 1 1 3 9 6 S E 0 6 0•355 455 E 05 OoOOGOCGE GO 0 ,165975E 06 0 , . 1 5 8 2 3 8 E 06 0 . 5 1 1G55E 05 0 , . 5 2 2 4 7 3 E 0 5 0 •427567E 06 o . 1 2 4 2 5 4 E 06 1 , 142794c 06 0.3 4 8 8 5 2 E 0 5 Ca 2 2 4 3 1 4 E 0 5 OoOOCGOOE CC 0 , 1 5 8 7 6 O E 06 C , 1 5 1 4 9 1 E 0 6 0 . 3 2 4 0 0 6 E 0 5 0 i . 3 1 7 1 3 6 E C5 0 , 3 7 4 3 6 6 E 06 0 . 1 1 9 2 8 0 E 06 0 . , 1 3 7 0 6 9 E 06 0 . 4 3 5 1 2 2 E 0 5 0.239292E 0 4 Go 0GC0GGE 00 0 « , 1 489 4 I E 0 6 0 , .147833E 0 6 Q . 2 6 2 2 6 9 E 04 0 , 2 8 5 3 0 1E 04 0 « . 3 0 2 2 5 5 E 0 6 0 i . 1 1 3 9 4 7 E 06 Oi . 1 3 0 8 9 1 E 0 6 0 . 3 7 8 6 0 9 E 0 5 0 . 2 6 6 9 2 2 E 0 3 0 • GOCGpG E 00 0 , 141483E 0 6 G, . 1 4 0 9 4 9 E 06 0 ,277937c 0 3 Oi . 2 7 6 3 7 9 E 03 0 « , 2 8 2 9 8 7 E 06 159 GONAD ENERGY (KCAL) 0 + 1+ 2+ 3 + 4 + o • OOOOOOE 00 0 • 4 2 9 3 4 0 E 0 3 0 • 6 9 6 1 3 3 E 0 3 0 » 2 1 6 4 6 0 E 0 3 0 • OOOOOOE 00 0 • 13 7 1 8 5 E 03 0 • 4 4 1 1 0 8 E 0 3 p, o 1 0 6 1 0 4 E 0 3 • 6 5 7 5 3 6 E 03 0 o1 3 4 1 9 3 E 0 4 0 •OOOOCOE 0 0 • 4 1 3 9 3 5 E 03 0 • 6 5 4 3 4 8 E 0 3 0 . 2 0 9 0 8 0 E 0 3 OeOOOOOOE 00 0 • 1 3 1 1 9 0 E 0 3 r-> w * 4 2 2 6 4 6 E 03 0 • 1 0 0 5 9 5 E 03 (' ] • 6 2 2 9 3 2 E 03 0 o 1 2 7 7 3 6 E 0 4 0 B OOOOOOE 0 0 0 • 4 0 0 5 3 9 E 03 n \j « 6 16 3 6 6 E 0 3 0 e 2 0 l 6 2 5 E 0 3 OeOOOOOOE 00 0 • 1 2 6 1 0 3 E 0 3 0 • 4 0 6 6 73E 03 0 o 9 ^ S 0 2 8 E 0 2 0 • 5 9 0 9 5 1 E 0 3 •J e 1 2 1 8 53E 0 4 0 •OOOOCOE 0 0 0 • 3 8 6 1 3 8 E 0 3 o 5 6 6 7 2 2 E 0 3 0 « 1 8 9 6 2 2 E 03 0* OOOOOOE, 00 0 • 1 2 0 6 6 1 E 0 3 0 • 3 S 9 9 1 9 E 0 3 0 • 8 5 0 3 3 6 E 02 0 • 5 4 6 8 6 9 E 0 3 0 • 1 1 4 2 4 8 E 0 4 0 • 1 1 0 3 5 5 E 02 0 • 3 7 2 7 4 2 E 0 3 0 • 4 9 7 2 0 7 E 0 3 0' . 1 6 7 0 9 7 F. 0 3 OeOOCOOOE 00 0 • 1 1 8 0 9 0 E 0 3 0 • 3 8 2 7 6 2 E 03 0 • 6 8 2 5 7 9 E 0 2 0 • 4 7 8 9 7 2 E 0 3 0 . 1 0 4 H 0 8 E 0 4 • 2 8 7 9 1 2 E 02 0 • 4 1 4 7 9 6 E 0 3 o • 6 15 9 9 0 E 0 3 0 » 2 2 4 3 4 8 E 0 3 O.OOOOOOE 00 a 1 3 4 1 5 7 E 03 0 • 4 3 6 4 1 C E 0 3 0 . 1 4 8 8 7 8 E 03 0 • 5 6 4 4 8 0 E 0 3 . 1 2 8 3 9 2 E 0 4 0 • 7 2 8 2 4 0 E 0 2 0 • 4 5 9 9 4 7 E 03 0 . 1 9 6 9 7 8 E 0 4 0 . 8 6 9 6 7 6 E 03 OeOOOOOOE GO 0 o 15 6 8 2 2 E 0 3 0 • 5 1 3 2 6 3 E 0 3 o • 6 9 S 3 2 2 E 03 0 • 2 0 0 3 S 2 E 0 4 0 • 3 3 7 2 2 2 E 0 4 0 • 1 9 0 O 2 2 E 0 3 0 • 5 1 1 9 5 0 E 0 3 0 • 6 2 1 3 4 1 E 0 4 0 . 2 9 3 8 7 6 E 0 4 0 e OOOOOOE 00 0 • 1 9 7 7 0 8 E 0 3 0 • 6 5 3 4 1 0 E 03 . 1 6 4 1 9 7 E 0 4 Q • 7 3 6 1 0 5 E 0 4 c • 9 8 5 4 1 5 E 0 4 o . 4 9 5 6 5 3 E 03 0 , 5 6 9 6 4 I E 0 3 0 . 1 0 1 9 3 9 E 0 5 OB 4 9 8 6 6 4 E 0 4 0.OOOOOOE CO 0 • 2 8 1 9 1 9 E 03 0 • 9 4 4 8 5 2 E 03 0 • 2 2 5 3 9 2 E 0 4 0 • 1 2 7 6 5 2 E 05 0 • 1 6 2 4 5 8 E 05 0 • 4 7 8 4 2 9 E 03 0 • 5 4 9 8 1 5 E 03 pi • 4 4 2 3 4 4E 0 4 0 . 2 1 6 6 3 9 E 0 4 OeOOOOOOE 00 0 • 2 7 1 0 Q 9 E 03 0 • 9 0 9 0 7 9 E 03 0 • 7 S 5 7 5 9 E 03 0 • 5 6 5 2 2 3 E 0 4 0 • 7 6 1 8 0 7 E 0 4 0 • 4 6 1 2 0 5 E 0 3 0 • 5 2 9 9 8 8 E 0 3 0 • 1 6 5 8 0 0 E 0 3 O t 7 8 9 7 9 5 E 0 1 0 eOOOOOOE 00 0 • 2 5 5 3 1 6 E 03 0 • 8 9 0 8 8 3 E 03 0 • 1 9 1 9 4 5 E 0 1 0 • 1 6 7 7 4 3 E 02 0 • 1 1 6 4 8 9 E 0 4 0 . 4 4 5 1 5 1 E 03 0 . 5 1 1 3 4 6 E 0 3 0 • 1 4 9 6 4 6 E 0 3 0 . 3 1 3 9 0 8 E 0 1 OeOOOOOOE 00 0 • 2 4 5 0 4 5 E 0 3 0 • 8 5 8 1 6 7 E 03 0 . 1 2 9 4 4 1 E 0 1 0 • 4 7 7 5 5 7 E 0 1 n • 1 1 0 9 2 3 E 0 4 160 FINAL WEIGHT (KG) 0 + 1 + 2+ 3 + 4 + 0.OOOOOOE 0 0 0 o 7 9 9 9 6 8 E 0 2 0 o 9 2 3 7 9 4 E 0 2 0 . 2 5 8 1 9 1 E 02 Oo OOOOOOE 00 0 . 5 7 5 6 0 2 E 02 O o 5 2 8 2 3 5 E 02 0 o 4 l 5 5 1 3 E 0 2 0 o 4 6 2 6 0 3 E 02 Oo 1 9 8 1 9 5 E 0 3 OoOOOOOOE 0 0 O o 7 7 1 2 6 4 E 02 0 . 8 6 8 1 1 7 E 0 2 0 • 2 4 9 3 8 9 E 02 Oo OOOOOOE 00 0 o 5 5 0 4 4 7 E 0 2 0 o 5 0 6 1 2 7 E 0 2 0 o 3 9 3 9 3 8 E 02 O o 4 3 8 2 5 8 E 02 0 o l 8 8 8 7 7 E 03 OoOOOOOOE 0 0 0 o 7 4 6 3 0 4 E 0 2 0 o 8 2 0 4 7 3 E 0 2 0 o 2 4 1 5 6 3 E 02 Oo OOOOOOE 00 0 . 5 2 9 1 0 2 E 02 O o 4 8 6 9 9 8 E 02 0 o 3 7 5 0 6 8 E 02 O o 4 1 7 1 7 3 E 0 2 Oo 1 8 0 8 3 4 E 03 OoCOOOOOE 0 0 O o 7 1 9 4 7 2 E 0 2 O o 7 7 1 1 6 7 E 0 2 0 o 2 3 3 7 4 1 E 02 0 0 OOOOOOE CO 0 o 5 0 6 2 6 8 E 02 O o 4 6 6 9 3 4 E 02 0 o 3 5 5 9 2 2 E 02 0 . 3 9 5 2 5 5 E 02 0 o l 7 2 4 3 8 E 0 3 O o 5 5 6 4 2 3 E 0 1 0 o 8 0 1 6 7 9 E 02 0 » 8 3 7 7 2 6 E 0 2 0 • 2 6 1 5 0 9 E 02 Oo OOOOOOE 00 O o 5 8 7 8 9 2 E 02 0 o 5 4 5 0 4 7 E 02 0 . 3 9 0 3 8 6 E 02 0 o 4 3 3 2 3 1 E 02 Oo 1 9 5 6 5 5 E 0 3 0 o l 4 0 5 8 2 E 02 O o 8 8 8 0 3 2 E 02 0 o 9 0 4 5 0 8 E 0 2 0 . 2 9 0 8 5 5 E 02 Oo OOOOOOE 00 0 o 6 8 6 2 4 4 E 02 0 o 6 3 9 9 5 3 E 02 0 o 4 2 5 7 4 4 E 02 O o 4 7 2 0 3 5 E 02 O o 2 2 2 3 9 7 E 0 3 0 o 3 6 7 l 8 5 E . 02 0 o 9 8 9 2 3 7 E 0-2 0 o 9 8 3 O l 2 E 0 2 0 o 3 2 4 5 9 9 E 02 Oo OOOOOOE CO 0 o 8 6 4 8 0 5 E 0 2 0 o 8 1 4 7 6 4 E 02 0 o 4 6 7 2 1 1 E 02 O o 5 1 7 2 5 2 E 0 2 0 o 2 6 6 4 0 3 E 03 0 . 9 5 8 1 1 1 E 02 O o l 1 0 1 0 8 E 03 0 o 1 0 6 6 1 6 E 0 3 0 o 3 6 3 5 0 4 E 02 0 0 OOOOOOE 00 0 o 1 2 3 2 2 5 E 03 0 . 1 1 7 7 9 1 E 03 0 O 5 1 2 1 7 5 E 0 2 0 o 5 6 5 5 1 I E 02 0 o 3 4 3 8 8 6 E 03 0 o 9 2 3 5 2 5 E 02 0 o l 0 6 l 3 8 E 03 O o 1 C 0 1 5 4 E 0 3 0 8 3 5 1 2 7 1 E 02 0 0 OOOOOOE 00 0 . 1 1 8 2 3 7 E 03 0 . 1 1 3 1 4 7 E 03 0 o 4 8 5 9 8 8 E 02 O o 5 3 7 3 8 7 E 0 2 0 o 3 3 3 7 7 2 E 03 Q o 8 9 1 4 3 2 E 02 0 o l 0 2 4 4 4 E 0 3 0 o 9 4 2 8 0 7 E 0 2 0 o 3 3 9 0 3 7 E 02 Oo OOOOOOE 00 0 o l l 3 7 1 0 E 03 0 o l 0 8 8 6 3 E 03 0 o 4 6 1 7 6 0 E 02 O o 5 1 0 2 2 2 E 02 0 o 3 1 9 7 7 2 E 03 0 0 8 5 9 3 4 0 E 0 2 0 . 9 8 7 5 0 0 E 02 0 o 3 4 7 5 5 6 E 0 2 0 . 4 3 6 9 0 4 E 0 1 0 0 OOOOOOE 00 Oo 1 0 7 1 2 5 E 03 0 o l 0 6 6 8 4 E 0 3 0 o 4 7 7 8 9 5 E 0 1 0 . 5 2 1 9 5 2 E 0 1 0 o 2 2 3 8 0 8 E 0 3 0 o 8 2 9 4 2 8 E 02 0 o 9 5 2 7 6 4 E •02 0 o 2 7 8 5 3 4 E 0 2 0 . 5 2 4 2 8 5 E 0 0 .. Oo OOOOOOE 00 0 . 1 0 2 8 1 6 E 03 0 . 1 0 2 7 6 7 E 03 Oo 5 0 6 9 0 0 E 00 0 . 5 0 6 9 0 0 E 0 0 Oo 2 0 6 5 9 6 E 03 CHANGE I N BODY E N E R G Y ( K C A L ) 0 + 1+ 2 + 3 + 4+ Oi » OOOOOOE CO -0 • 252315E 04 -0.291013E 04 -C.S12967E 0 3 OoOOOOOOE 00 -0 .181849E 04 -Oi .166333E 04 -0 •121259E 04 -0 , 145183E 04 -0 .624525E 04 Oi ,OOOOOOE 00 -0 .3G1461E 04 -0.338901E 04 -0.973121E 03 O.OOOOCOE 00 •an • 2 1 5 5 C e E 04 -0 >197500E 04 -0 ol5421&£ 04 -0 , 170450E 04 —c 3737&75E 04 0. , OOOOOOE GG -0 >556943E 04 -0.611545E 04 -Oa179966E 04 OeOOGOGGE GO -0 S3955C7E G4 -0 .362831E 04 -0 .280338E 04 -0, , 309778E 04 -0 •134645E 05 0, s OOOOOOE GO Gi •350145E 04 0 e 374845E 04 0.113561E 04 OoOOOOOOE GG 0 )246794E 04 0 •226867E 04 0 »173465E 04 Gi , 191405E 04 0 . 8 3 8 5 5 3 E 04 0 .591797E 04 0 ( •3462 95E 05 0a36CG19E 0 5 0.112910E 0 5 O.OOOOOOE CO 0 .271969E 05 0 ,252593E 05 0 ,168096E 05 G i , 185745E 05 0 .378404E C5 0 ,12 468 8E 05 G i >122135E 05 Q.1C5363E 0 5 0.31990 IE 04 0 e G G G 0 0 G t G 0 .147302E 05 0, .140459E 05 0 .471686E 04 G« 492517E 04 •J • 3641P.2E 05 0 i314619E 0 5 0 .124187E 05 0.798154E 04 0 . 2 0 7 2 7 7 E 0 4 Go GCGG.CGE Ob 0 • 243230E 05 0 •236155E 05 0 .34 40 6GE 04 G ,255590E 04 0 .539350E 05 0 .846176E 05 G< .20&191E 05 -0 . 2 02883E 0 4 -0.41917GE 04 0•OOOOOOE GG •J .5&5067E 05 C • 553042E 05 -Q .478353E 04 -o, •801131E 04 . Oi ,99016 IE 05 fO •642303E 03 -Gi .738179E 03 -0.237744E 05 -C•123C47 E 0 5 O.OOOOOOE GO •:-c .824044E 03 -0 .785628E 0 3 '"•0 .17G048E 05 -0 .138451E 0 5 -C •374596E 05 -0 >519892E 03 -0 .597476E 0 3 -0.720236E 0 4 -0.386226E 04 O.OOOOOOE 00 »0 .664283E 0 3 -0 .633852E 03 •Ci •705920E 04 -0 9382466E 04 •0 •121820E 05 •0 •122310E 04 -Oi . 140556E 04 -0.417017E 03 -C a 1919 26E 01 OoOOOOOOE CO •0 .152727E 04 -0 .151596E 04 •0 •236164E 01 -Oi .200614E 01 •Oi i304760E 04 •0 •432843E 03 -0 .554639E 03 0.241316E 03 0.429G72E 0 3 O.OGOCCOE GO •0 > 59 95 2 5 E 03 -0 »597258E 0 3 0 »4l6576E 03 0 •413113E 03 •0 •367G93E 0 3 162 CHANGE I.< GO GAD 72 E 2GY ( KC 0 + 1 + 2 + 3 + 4 + 0. G 0 0 G 0 G E GG G , .6 65137E-04 0.235438E-•0 4 G. 1132487-G 4 0 . GGGG GGE 0 < .13411CE- 04 0 c .567436E- 0 4 0 < 121593E- 04 G ( 190734E- 04 0« 1C13S7E- 03 J 0 G 0 G 0 G E GO G i ,6961827- 04 -0 « 2 209767 01 -0.9245 22 7 0 7 0 e C20C GO E 3 < 115C36E- 04 G f ,6294257- C4 1C2296E CI -0 i ,2111347 01 -0 »313422E 01 2 < >00GG0GE G < ,379-J 8 5E-7 4 -0.132961E 0 2 -0*5G 5 6027 71 0. G G G G G 7 7 o ,1727537- 04 0 ,2753737- G4 -0. ,5194677 01 -0 ,137 5 757 0 2 -0 « >189521E 0 2. , G C J G G 0 E OG G .5 59091E-G4 -0.3841987 02 -0 . 1673352 G2 0. G 0 G G 00 ,.: . IT ~ 3 1 L -04 G< i 5 5 G 7 4 6 E -04 ,13 1316 7 02 -G , 4 2 G 2 6 8 E 02 > S 3 1 G G 3 E G2'J i 1S fi 2 7 5 E 02 6 ,7751677 0.1773077 0 3 0 . 6 5 2 9 9 6 E 2; 2 G. G G G 0 G G E > 2202447 02 Q .7 23 t 5 2£ ,C » 2 2 5 E G2 ,1167797 0 3 •j . 29993 I E G3 v- ,46 65 7 87 G2 G I613C-':4E 02 G.1475707 G i • G.27763 7 r G3 G a ^ G G G 2 0 » 2 9 3 ? 5 1 E 0 2 G , 979 9 03 7 0 2 0 , 58 63 717 0 3 0 , 1542 19E 0 4 0 » ?. 2. 61 9 5 E 04»1242437 0 3 0 ,7097 317 0 2 0.i646 797 G 4 G» 2 16 79 5 2' G4 G . G v 2; G 2 G 2 7 .4229737 02 0 » 1671147 0 3 G < i 10 32 35E C4 G ,7761? >-'•£ 0 4 » ",G0999E G 4 G .324193E C3 G , 78 97 74E G2 G » 4 6 3 G 3 2 c 0 4 0 ,,222 1G4~ G i 2 . GC; GG GG 2 G . •'? 5 v e G1 E 02 G ,3372247 03 D 7 3 " 3 9 G E v.- , 607 2 96 7 04 G , 7 777737 04 2 6 z~ G':* G ,5674367- G4 -0.5496377 G4 -G . 2742GS7 2 '7 G. 2- G' G G G G G 0 ,1126137- 04 Q ,934663E- G 4 <~,' a 1 '•  2 6 7 3 £ 7 4 -G ,6312227 C 4 ;™G I 3 ,? 37 9 5 E 04 •" • 169277L-04 f; , 233418E-05 -0«4I5G36G 0 4 — G o 2 1G 5 12 i.'.. 2 4 G e G... G ~ 2 G L 0 . 1G :•: 7 G 1 E -G4 G , 132 2.3 2 7-04 -0 9 / O •' 1 C~ t-0 3 -G ,242 22 5E C4 -0 • 625f3<-f6E G 4 ;j •25 27 2 3E-0 4 G » 2 2 B 7 8 17 -G 2 » 247223E 0 2 G o 17 261GG 2 2 0 • -;G G G G G 7 c , 370077E-Gi 0 ,2264977- 04 » 1027 39c • v C 7 •3239747 2 2 .4268 34E '• n G C . 236C34E-G4 G ,4553797- Gi G . i 173C5E 01 0« 122 6 :>6 2 G i 0 . C. 0 G G GL . 1 . 13 7 i 5 6 £ -04 0 .6294257- Gi 0 . 14 52S6E-G 7 0 , 2 4 2 9417 GI G » 242 94 9L 01 163 IBM 1 1 3 0 PROGRAM / / JOB / / FOR *ONE WORD INTEGERS * I O C S ( C A R D » 1 1 3 2 PR I N T E R » D I S K ) D I M E N S I O N W T ( 5 » 2 » 2 ) » N R ( 5 ) » W P E R ( 5 ) , P R M ( 2 » 5 ) » S X ( 2 ) » E ( 2 ) » W ( 2 ) » A ( 2 ) . 1 G W I 2 . 2 ) » G E ( 2 »2) » G A Q ( 2 » 2 ) > G A ( 2 ) . G R ( 5 ) > RMR( 5 ) » N ( 5 »2 » 2 ) » E I N ( 5 » 2 » 2 ) » 2EGEX ( 5 »2 » 2 ) » R M ( 5 • 2 » 2 ) » E C ( 5 » 2 . 2 ) » REC ( 5 » 2 »'2 ) . D E L W ( 5 > 2 • 2 ) . D E C ( 5 . 2 , 2 ) » 3 D R E C ( 5 » 2 » 2 ) » H W ( 2 > 2 ) » H E ( 2 » 2 ) » H A Q ( 2 » 2 ) » H A ( 2 ) . R R M ( 2 » 5 ) D I M E N S I O N R N ( 5 » 2 » 2 ) » D A T A ( 2 0 0 ) . S < 5 ) . S S ( 2 » 2 ) D I M E N S I O N X( 2 ) > B ( 2 ) » F ( 2 ) D E F I N E F I L E 1 ( 120 > 40 »U » I D ) 10 W R = 0 . 0 0 0 3 . N R ( 1 ) = 0 W P E R ( 1 ) = 0 . 0 P R M ( 1 » 1 ) = 0 # 0 P R M ( 2 . 1 ) = 0 . 0 P R M ( 1 » 2 ) = 0 . 0 P R M ( 2 . 2 ) = 0 . 0 R M E = 0 . 7 6 E E = 0 . 2 I D= 1 PAUSE 1 R E A D ( 2 » 1 0 0 ) { N R ( I ) » W P E R ( I ) . 1 = 2 . 5 ) >( ( P R M ( J » I ) » J = 1» 2 ) * I = 3 » 5 ) » SX » NREC » 1MR»MS 1 0 0 F O R M A T ( 6 X » 4 ( I 6 » F 6 . 0 ) / 6 X » 6 F 6 e 0 » 6 X , 2 F 6 . 0 , 6 X » I 6 , 2 I 3 ) DO 52 J = 1 . 5 DO 52 K= 1 »2 5 2 R R M ( K . J ) = P R M ( K » J ) DO 1 1 = 1 . 1 2 PAUSE 2 R E A D ( 2 » 1 0 1 ) F D R > ECF » RMK »RMD » E ( 2 ) * W ( 2 ) » A ( 2 ) » E ( 1 ) » W ( 1 ) »A( 1 ) » P 1 » P 2 » 1 ( G W ( J » 2) » G E ( J » 2 ) » G A Q ( J > 2 ) »GW(J . 1 ) » G E ( J » 1 ) » G A Q ( J »1 ) . G A ( J ) » j = 1 , 2 ) » 2GR » R M R * ( H W ( J »2) » H E ( J » 2 ) » H A Q ( J » 2 ) » H W ( J * 1 ) » H E ( J »1 ) • H A Q ( J • 1 ) » H A ( J ) • 3 J = 1 » 2 ) » F ( 2 ) » X ( 2 ) » B ( 2 ) » F { 1 ) » X ( 1 ) » B ( 1 ) 1 0 1 F O R M A T ( 6 X * 3 F 6 . 1 » F 3 . 0 » 3 X » 8 F 6 . 0 / 6 X » 3 F 6 . 1 » F 3 . 0 » 3 F 5 , 0 » F 8 . 0 » . 2 6 F 5 . 0 / 6 X » 1 0 F 6 . 0 / 6 X » 3 F 6 . 1 » F 3 « 0 , 3 F 5 . 0 » F 8 * 0 » 6 F 5 . 0 / 3 0 X » 6 F 6 * 0 ) READt 2 . 9 9 9 ) 9 9 9 F O R M A T ( I X ) I F ( P 1 ) 6 0 . 6 0 . 6 1 6 1 R R M { 1 . 3 ) = P I R R M ( 2 . 3 ) = P2 6 0 I F ( I - M S ) 5 0 . 5 1 . 5 0 5 1 DO 53 K = 1 . 2 DO 53 J = 2 . 4 5 3 R R M ( K . J - l ) = P R M ( K . J ) 5 0 I F ( I - M R ) 5 . 4 . 5 4 N R ( 1 ) = N R E C W P E R ( 1 ) = W R 5 DO 2 J = 1 . 5 DO 2 K = l . 2 N ( J . K . 2 ) = N R ( J ) * S X ( K ) * R R M (K » J ) * 0 . 0 0 0 1 2 N ( J i K » l ) * N R ( J ) * S X U ) * ( 1 0 0 . 0 - R R M ( K i J ) ) * 0 . 0 0 0 1 164 DO 6 J = 1» 5 DO 6 K = l » 2 ' DO 6 L = l » 2 W T ( J » K . L ) = N ( J » K » L ) * W P E R ( J ) E I N ( J » K . L ) = W T ( J . K . L > * F D R * 0 . 0 1 # E C F * R M D E G E X ( J » K » L ) = E I N ( J . K » L ) * E E R M ( J » K » L ) = R M K * ( W T ( J » K» L ) * # R M E ) * R M D E C ( J . K » L ) = W T ( J . K . L ) * ( 1 0 0 . 0 - W ( L ) ) * ( 10 0 . 0 - A ( L ) ) * E ( L ) 1 * ( 1 0 0 . 0 - G W < K » L ) ) # 0 . 0 0 0 0 0 1 R E C ( J » K » L ) = W T ( J . K . L ) *GW ( K » L ) * ( 10 0 • O-GAQ(K * L ) ) * ! 1 0 0 . 0 - G A { K ) ) 1 * 0 . 0 0 0 0 0 1 * G E ( K » L ) DELW ( J »K > U = WT ( J » K » L ) * E X P ( G R U ) ) D E C ( J » K » L ) = D E L W ( J »K» L ) * ( 1 0 0 . 0 - X ( L ) ) * ( 1 0 0 . 0 - B ( L ) ) * F { L ) 1 * ( l O O . O - H W ( K . L ) ) # 0 . 0 0 0 0 0 1 - E C U . K . L ) D R E C ( J »K » L ) =DELW( J . K . . L ) *HW( K. > L ) *< 100 . 0 -HAQ ( K » L ) ) * ( 1 0 0 . 0 - H A (K ) ) * 1 0 . 0 0 0 0 0 1 * H E ( K » D - R E C ( J . K . L ) R N ( J » K » L ) = N ( J • K. > L ! 6 CONTINUE DO 9 J = l . 5 9 N R ( J ) = N R ( J ) * E X P ( - R M R ( J ) ) W R I T E ( 2 * 1 0 3 )N 1 0 3 F O R M A T ( 4 ( 5 I 1 5 / ) ) W R I T E ( 2 * 1 0 2 J W T . E I N . E G E X . R M . E C . R E C . D E L W . D E C . D R E C 1 0 2 F O R M A T ( 4 ( 5 E 1 5 . 6 / ) ) W R I T E ( 1 ' I D ) R N . W T . E I N . E G E X . R M . E C . R E C >DELW.DEC.DREC DO 4 0 0 J = 1 » 5 4 0 0 WPER (J) = W P E R ( J ) * E X P ( G R U J ) 1 CONTINUE DO 3 0 0 L = l » 1 0 I D= 1 DO 30 2 I M = 1 . 1 2 DO 2 0 2 1 = 1 . 5 2 0 2 S ( I ) = 0 . 0 DO 2 0 3 J = 1 . 2 DO 20 3 K = 1 . 2 2 0 3 S S ( J . K ) = 0 . 0 R E A D ( 1 ' I D )DATA DO 3 0 1 1 = 1 . 5 DO 3 0 1 J = l » 2 DO 3 0 1 K = l > 2 M = I + 5 * ( J - l ) + 1 0 * ( K - l ) + 2 0 * ( L - l ) S ( I )= S ( I ) + D A T A ( M ) 3 0 1 S S I J . K ) =SS < J »K l.+DATA (M > SUM = 0 . 0 DO. 3 0 3 1 = 1 . 5 3 0 3 SUM = SUM + S ( I ) 3 0 2 W R I T E ( 2 . 1 0 4 ) S . S S . S U M 1 0 4 F O R M A T ! 5 E 1 5 . 6 / 2 E l 5 . 6 / 2 E 1 5 . 6 / E 1 5 . 6 ) 3 0 0 CONTINUE GO TO 1 0 . END / / XEQ 165 Fortran Program Variable Name Equivalents WT = weight DELW = change i n weight WPER = weight/individual NR = number NREC = number of r e c r u i t s REM = PRM = percent mature SX = sex r a t i o E = body energy W = body water content A = body ash content EC = energy content of f i s h DEC = change i n energy content of f i s h GW = gonad weight f o r month HW = gonad weight f o r following month GE = gonad energy f o r month HE = gonad energy f o r following month GAQ = gonad water content f o r month HAQ = gonad water content f o r following month GA = gonad ash content f o r month HA = gonad ash content f o r following month REC = energy content of gonad DREC = change i n energy content of gonad GR = grox-rth rate RMR = m o r t a l i t y rate EIN = energy intake FDR = percent d a i l y r a t i o n ECF = energy content of food EE = egestion - excretion EGEX = loss through egestion excretion RM = routine metabolism RME = exponent of routine metabolism equation RMK = c o e f f i c i e n t of routine metabolism equation RMD = number of days i n month 

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