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Daily rings in otoliths of sockeye salmon (Oncorhynchus nerka) and their relationship to growth Wilson, Kenneth H. 1981

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D A I L Y R I N G S I N O T O L I T H S OF S O C K E Y E AND  SALMON ( O n c o r h y n c h u s n e r k a )  T H E I R R E L A T I O N S H I P TO GROWTH  by  K e n n e t h H. W i l s o n B.Sc,  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 , 1977  A THESIS SUBMITTED  IN P A R T I A L F U L F I L L M E N T OF  T H E R E Q U I R E M E N T S FOR THE DEGREE MASTER OF  OF  SCIENCE  in the Department of Zoology  We  accept this  t h e s i s as c o n f o r m i n g t o the  required standard  T H E U N I V E R S I T Y OF B R I T I S H COLUMBIA March (c)  1981  K e n n e t h H. W i l s o n  In p r e s e n t i n g  this  thesis i n partial  f u l f i l m e n t of the  r e q u i r e m e n t s f o r an a d v a n c e d d e g r e e a t t h e of B r i t i s h Columbia, I agree that it  freely  the L i b r a r y s h a l l  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 .  agree t h a t p e r m i s s i o n for  University  f o r extensive  s c h o l a r l y p u r p o s e s may  for  financial  shall  of  The U n i v e r s i t y o f B r i t i s h 2075 W e s b r o o k P l a c e V a n c o u v e r , Canada V6T 1W5 Date  Columbia  my  It is thesis  n o t be a l l o w e d w i t h o u t my  permission.  Department  thesis  be g r a n t e d by t h e h e a d o f  copying or p u b l i c a t i o n of this  gain  further  copying of t h i s  d e p a r t m e n t o r by h i s o r h e r r e p r e s e n t a t i v e s . understood that  I  make  written  ii  ABSTRACT This study reports the occurrence o f d a i l y r i n g s i n the o t o l i t h s o f Oncorhynchus  nerka f r y and examines  t h e i r r e l a t i o n s h i p t o growth.  In e x p e r i m e n t 1, sockeye salmon f r y were c o l l e c t e d from t h e F u l t o n R i v e r spawning channel a t Babine Lake, B r i t i s h Columbia i n May 1978.  The f i s h  were r e a r e d f o r 26 days i n e n c l o s u r e s i n t h e spawning channel and were sampled e v e r y seven t o t e n days.  S a g i t t a e were removed from 25 f i s h from  each sample, and t h e growth r i n g s i n one o t o l i t h from each f i s h were counted.  A r e g r e s s i o n o f t h e number o f r i n g s on t h e number o f days  since  c a p t u r e showed t h a t t h e s e r i n g s a r e , on a v e r a g e , formed d a i l y , b e g i n n i n g a t t h e time o f emergence.  A number o f p o s s i b l e t e c h n i c a l and b i o l o g i c a l  causes o f v a r i a t i o n i n r i n g counts w i t h i n and between samples a r e c o n s i d e r e d .  In Experiment 2, sockeye salmon f r y were r e a r e d i n t h e l a b o r a t o r y from f e r t i l i z e d eggs taken i n t h e f a l l  o f 1978 a t the Weaver Creek  spawning channel near M i s s i o n , B r i t i s h Columbia.  A random sample o f 64  o f t h e s e f r y was marked t o e n a b l e i d e n t i f i c a t i o n o f i n d i v i d u a l s . i n d i v i d u a l was weighed  Each  i n i t i a l l y on June 6 o r 8, a g a i n on J u l y 6, and  s u r v i v i n g f i s h were weighed a t h i r d t i m e on J u l y 20.  After a final  w e i g h i n g , s a g i t t a e were removed and a s t a n d a r d o t o l i t h r a d i u s was d e t e r mined by c o u n t i n g back t h e a p p r o p r i a t e number o f d a i l y r i n g s which c o r r e s ponded t o each w e i g h t .  The r e g r e s s i o n o f £n o t o l i t h r a d i u s on  w e i g h t was l i n e a r , and had an R  2  £n f i s h  o f 0.92, which demonstrates a r e l a t i o n -  s h i p between the mean w i d t h o f a d a i l y r i n g i n sockeye salmon f r y s a g i t t a e , and a mean d a i l y change i n t h e w e i g h t o f t h e f r y .  Using t h i s  regression  l i n e , we b a c k - c a l c u l a t e d t h e p r e v i o u s w e i g h t o f t h e i n d i v i d u a l f i s h from  i ii  the c o r r e s p o n d i n g and  o t o l i t h r a d i u s and a l a t t e r f i s h w e i g h t and o t o l i t h  found the e r r o r s to be r e l a t i v e l y s m a l l —  i n the o r d e r o f 15 per  radius cent.  iv  TABLE OF CONTENTS Page T I T L E PAGE  .  ABSTRACT  1 i i  TABLE OF CONTENTS  iv  L I S T OF FIGURES  v i  L I S T OF TABLES  v i i  L I S T OF APPENDICES  viii  ACKNOWLEDGEMENTS  ix  INTRODUCTION  1  GENERAL METHODS  9  Fish Preservation  9  Otolith  Removal  9  Otolith  Storage  11  Otolith  Preparation  EXPERIMENT  and O b s e r v a t i o n  1: DAILY GROWTH RINGS IN THE OTOLITHS  11 OF  J U V E N I L E SOCKEYE SALMON ( O n c o r h y n c h u s n e r k a )  15  Introduction  15  Methods  16  Results  IV  Discussion EXPERIMENT 2: THE RELATIONSHIP BETWEEN WIDTuH OF DAILY GROWTH RINGS IN SAGITTAE AND CHANGE IN BODY S I Z E OF SOCKEYE SALMON ( O n c o r h y n c h u s n e r k a )  19 23  Introduction  23  Methods  23  V  Results  30  Discussion  36  SUMMARY  42  LITERATURE CITED  43  APPENDIX 1  APPENDIX 2  The back c a l c u l a t i o n o f p r e v i o u s f i s h weight from c o r r e s p o n d i n g p r e v i o u s o t o l i t h r a d i u s and a l a t t e r o t o l i t h r a d i u s and f i s h weight..  45  A n n o t a t e d p h o t o g r a p h s o f O. n e r k a o t o l i t h s as an a i d t o age and g r o w t h a n a l y s i s  48  vi  L I S T OF Figure  Figure  1.  2.  F i g u r e 3.  F i g u r e 4. F i g u r e 5. Figure  6.  FIGURES  The l a b y r i n t h o f t h e c o d , Gadus morhua ( L . ) showing the r e l a t i o n s h i p between the semic i r c u l a r c a n a l s and t h e o t o l i t h chambers S a g i t t a e o f 0.  nerka f r y and and  S a g i t t a e f r o m a 0. n e r k a f r y removed t h e e n c l o s u r e on J u n e 9  S a g i t t a e o f an 0. the  enclosure  count versus  from 18  age  in  from  13  Cold  Figure  8.  S a g i t t a e o f an 0. n e r k a f r y s h o w i n g r o s t r u m , p o s t - r o s t r u m , n u c l e i and l o c a t i o n of p r e f e r r e d radius S a g i t t a e o f 0. n e r k a f r y removed f r o m t a n k on J u l y 2  29  £n f i s h w e i g h t v e r s u s in o t o l i t h r a d i u s f o r i n d i v i d u a l f i s h (No's 1-29) a t t h e i n i t i a l ( J u n e 6 o r 8 ) , s e c o n d ( J u l y 6) and, where a p p l i c a b l e , t h i r d ( J u l y 20) w e i g h i n g  31  P l o t of r e s i d u a l s from l i n e a r r e g r e s s i o n of f i s h w e i g h t on £n o t o l i t h r a d i u s as shown i n F i g u r e 10  32  S a g i t t a e o f O. n e r k a f r y s h o w i n g p o s s i b l e t y p e s of back c a l c u l a t i o n of p r e v i o u s f i s h w e i g h t u s i n g the c o r r e s p o n d i n g previous o t o l i t h r a d i u s and a l a t t e r f i s h w e i g h t and o t o l i t h r a d i u s as d e s c r i b e d i n t h e t e x t . . . .  35  R e s i d u a l s o f b a c k c a l c u l a t e d w e i g h t {Un b a c k c a l c u l a t e d f i s h w e i g h t - Zn o b s e r v e d f i s h w e i g h t ) v e r s u s p r e d i c t e d Jin w e i g h t  37  Figure  Figure  Figure  11.  12.  13.  i n an 0.  22  7.  F i g u r e 10.  sites  10  20  n e r k a f r y removed May  after gills  Figure  F i g u r e 9.  branding  on  2 3  A p p e a r a n c e o f 0. n e r k a f r y b e f o r e r e m o v a l o f l o w e r jaw, g i l l c o v e r s to r e v e a l s a g i t t a e  Graph of o t o l i t h r i n g d a y s s i n c e emergence  Page  nerka f r y  25  27  L I S T OF  TABLES Page  Table I  an o t o l i t h r a d i u s ( AnR) and in f i s h w e i g h t (JlnW) f o r e x p e r i m e n t a l f r y a t t h e i n i t i a l ( J u n e 6 o r 8 ) , s e c o n d ( J u l y 6 ) , and t h i r d ( J u l y 20) w e i g h i n g s  33  vi i i  L I S T OF Appendix  Appendix  1  2  APPENDICES  The back c a l c u l a t i o n o f p r e v i o u s f i s h w e i g h t from c o r r e s p o n d i n g p r e v i o u s o t o l i t h r a d i u s and a l a t t e r o t o l i t h r a d i u s and f i s h weight  45  A n n o t a t e d p h o t o g r a p h s o f 0. n e r k a o t o l i t h s as an a i d t o age and g r o w t h a n a l y s i s  48  ix  ACKNOWLEDGEMENTS I am d e e p l y g r a t e f u l  t o D r . P. A. L a r k i n  f o r h i s guidance,  e n c o u r a g e m e n t and s u p p o r t .  I a l s o w i s h t o t h a n k H e l e n Hahn, G a i l  Sugden, M o i r a Greaven, B i l l  G a z e y , J o h n N e i l s o n , Cameron West and  many o t h e r s f o r t h e i r h e l p and a d v i c e , and g r a t e f u l l y financial  s u p p o r t p r o v i d e d by t h e M a r i n e R e s o u r c e s  M i n i s t r y o f Environment  acknowledge  Branch,  o f t h e Government o f B r i t i s h  C o l u m b i a and  t h e N a t u r a l S c i e n c e s and E n g i n e e r i n g R e s e a r c h C o u n c i l o f C a n a d a . I a l s o thank t h e Department staff  o f F i s h e r i e s and O c e a n s , C a n a d a , t h e  o f t h e F u l t o n R i v e r spawning  Pacific  Salmon f i s h e r i e s commission  c h a n n e l , and t h e I n t e r n a t i o n a l for their  cooperation.  1  INTRODUCTION  The system.  o t o l i t h s of teleost The t y p i c a l  teleost  each l a b y r i n t h ; a s a g i t t a lagena,  and a l a p i l l u s  invariably  f i s h e s are p a r t s o f the l a b y r i n t h fish  has s i x o t o l i t h s ,  three i n  i n e a c h s a c c u l u s , an a s t e r i s c u s i n e a c h  i n each u t r i c u l u s .  the l a r g e s t o t o l i t h  The s a g i t t a  i n non-osteriophysids.  i s almost Figure 1  ( a f t e r B l a c k e r 1 9 7 4 ) shows t h e l a b y r i n t h o f a c o d (Gadus morhua) and  the associated o t o l i t h s .  1977)  represents  F i g u r e 2 ( a f t e r T a u b e r t and C o b l e  t h e s a g i t t a o f a sockeye salmon  (Oncorhyncus  nerka). Each o t o l i t h nite  i s composed p r i m a r i l y  (calcium carbonate),  called  otolin  and a p r o t e i n r e l a t e d  ( Degens e_t a _ l . 1 9 6 9 ) .  p h o s p h a t e and c a l c i u m s u l p h a t e  of the l i t e r a t u r e  on o t o l i t h s  sented  by R.W.  (1974).  D a i l y g r o w t h r i n g s were f i r s t  occurrence  amounts o f c a l c i u m  (1971).  An e x c e l l e n t  reported  to occur  i n the oto-  c h u s s and Gadus morhua  S u b s e q u e n t r e s e a r c h has c o n f i r m e d  i n at least  seven s p e c i e s —  a n c h o v y , E n g r a u l i s mordax a d u l t nehu, S t e l a p h o r u s  salts  and t h e i r g r o w t h i s p r e -  of Merluccius b i l i n e a r i s , Utrophycis  by P a n n e l l a  to collagen  as w e l l as c o m p a r a b l e s o d i u m  review  liths  Small  ( M o r r i s and K i t t l e m a n 1 9 6 7 ) .  Blacker  of argo-  O t o l i n c o n s t i t u t e s between  0.25 and 10 p e r c e n t o f t h e o t o l i t h .  are a l s o present  (85-90 p e r c e n t )  their  j u v e n i l e northern  ( B r o t h e r s e t a _ l . 1976 ); j u v e n i l e and  purpurens  ( S t r u s a k e r and U c h i y a m a 1 9 7 6 ) ;  j u v e n i l e s o f the green s u n f i s h , Lepomis c y a n e l l u s , pumpkinseed,  2  F i g u r e 1.  The l a b y r i n t h  o f t h e c o d , Gadus morhua  the  relationship  the  otolith  the  outer side  labyrinth  (L.) showing  between t h e s e m i c i r c u l a r  chambers.  (a) L e f t  and (b) d o r s a l  (after Blacker  labyrinth  c a n a l s and viewed  view of the l e f t  1974).  from  Semi-circular canals (a)  Nerve Ampullae  Lapillus Utriculus Sagitta  Sacculus  Blood vessel Macula  acustica  (b)  Nerve  Asteriscus  Utriculus Lapillus  Macula acustica  Ampullae  Semi-circular canals  3  F i g u r e 2.  S a g i t t a e o f 0. 1977)  n e r k a f r y ( a f t e r T a u b e r t and  Coble  3a  SAGITTAL  SECTION  Dorsal  Edge  Anterior  Posterior  Edge  Edge  TRANSVERSE SECTION Dorsal Edge  LATERAL VIEW  Ventral  Nucleus  Edge  Distal Proximal  Surface  Surface  Proximal  Surface  Nucleus  Posterior Edge  Anterior Edge Distal  Surface  DORSAL VIEW  FRONTAL  SECTION  Ventral Edge  4  L . G i b b o s u s , Mozambique m o u t h b r o o d e r , M e i d i a m e n i d i a  ( Barkman  1 9 7 8 ) ; and j u v e n i l e s o c k e y e s a l m o n , O n c o r h y n c h u s n e r k a and  L a r k i n 1980).  daily  Wilson  ring patterns  (Wilson  and L a r k i n (1980) have a l s o o b s e r v e d  i n the s a g i t t a e of Chinook  (Oncorhynchus tshawytscha),  chum s a l m o n  (0.  (0. k i s u t c h ) , rainbow t r o u t  gorbuscha),  coho s a l m o n  gairdneri), Pacific starry  flounder  (Hippoglossus  staghorn  (0.  salmon  k e t a ) , pink  s c u l p i n (Leptocottus  (Platichthys stellatus),  Pacific  (Ptychocheilus  otolith  d a i l y g r o w t h r i n g s were f i r s t first  described  as c o n c r e t e  recognized  by P a n n e l l a ,  and c o n n e c t e d by s t o u t r a d i a l  micron wide, which are secreted through the l a m e l l a e .  as i n t e r l o c k i n g  Under " o p t i c a l "  zone o f w i d t h  fibers.  spaced  fibers. needle-like  f i b e r s and p a s s  The c o n c e n t r i c l a m e l l a e d e s c r i b e d (1971).  t h a t t h e s m a l l e s t bands i n t h e o t o l i t h  microns wide, with  merluccius)  l e n g t h and l e s s t h a n one  among t h e r a d i a l  H i c k l i n g were e x a m i n e d by P a n n e l l a  distinct parts.  who  s h e l l s or lamellae of organic material  c r y s t a l s a b o u t 40 m i c r o n s i n maximum  organic  squawfish  by H i c k l i n g ( 1 9 3 1 ) ,  i n o r g a n i c component was d e s c r i b e d  a light  halibut  t h e m i c r o s t r u c t u r e s o f t h e hake ( M e r l u c c i u s  a b o u t two m i c r o n s a p a r t The  armatus),  oregonensis).  t h e y were p r o b a b l y described  (Salmo  s t e n o l e p s i s ) , and O l y m p i c mud minnow (Novumbra  h u b b s i ) and i n t h e a s t e r i s c u s o f t h e n o r t h e r n  Although  salmon  observed  were made up o f two  examination  0.5 t o 1 m i c r o n ,  Pannella  by  t h e bands c o n s i s t o f  and a d a r k zone 0.5 t o 2.5  t h e d a r k zone c o n t a i n i n g a much d e n s e r mesh o f  P a n n e l l a s t a t e s t h a t t h e d a r k zone  corresponds  5  t o p e r i o d s o f r a p i d p r o t e i n d e p o s i t i o n , and t h a t " d u r i n g deposition  the production  of organic  fibers  i s h i g h , but c a l c i f i -  cation  i s e v e n h i g h e r , so t h a t t h e r a t i o o f o r g a n i c  fibers  i s overwhelmingly  90 p e r c e n t )  (Sic).  slow  are produced, but the r a t i o  resulted  "optical"  examination  reflected  light.  T h i s s t a t e m e n t may have otoliths  and a c e t a t e  and i t i s n o t c l e a r w h e t h e r t h e  i n v o l v e d t h e use o f t r a n s m i t t e d o r  Another i n t e r p r e t a t i o n o f these position  i s possible.  observations  In the formation  of o t o l i t h  of the annulus,  f a s t g r o w t h zone w h i c h i s r e l a t i v e l y h i g h  I n temperate f i s h e s ,  i n t h e summer. higher  i n c a l c i u m c o n t e n t and  appears transparent  lamellae, ring,  ( B l a c k e r 1 9 7 1 , C h r i s t e n s e n 1 9 6 4 ) , and  under o b s e r v a t i o n w i t h t r a n s m i t t e d  light)  i s high  light.  (observed  with  i n p r o t e i n , analagous to H i c k l i n g ' s  i s t h e t h i n n e s t o f t h e two z o n e s w h i c h c o m p r i s e a d a i l y  and i s p r o b a b l y  otolith  t h e f a s t g r o w t h zone u s u a l l y f o r m s  the h y a l i n e p o r t i o n of a d a i l y r i n g  transmitted  (Blacker  The h y a l i n e zone u s u a l l y f o r m s i n w i n t e r , i s  i n p r o t e i n content  Similarly,  com-  i t i s the  w h i c h i s opaque u n d e r o b s e r v a t i o n w i t h t r a n s m i t t e d l i g h t 1971).  fibers  i s i n favour of the organic p o r t i o n ,  i s almost n i l . "  of otoliths,  (over  d e p o s i t i o n fewer o r g a n i c  from the o b s e r v a t i o n o f both  impressions  to inorganic  i n favour o f the o r g a n i c p o r t i o n  During  since c a l c i f i c a t i o n  fast  growth.  transmitted  The t h i c k e r  light,  formed d u r i n g  formed d u r i n g  the hours of slow  f i s h and  z o n e , w h i c h a p p e a r s opaque u n d e r  i s composed p r i m a r i l y o f c a l c i u m s a l t s  t h e h o u r s o f more r a p i d g r o w t h .  I speculate  and i s that  6  the rate o f calcium s a l t d e p o s i t i o n most r a p i d o t o l i t h formed  by a r a p i d  growth,  i s highest during period of  but whether the p r o t e i n  lamellae are  increase i n p r o t e i n d e p o s i t i o n , a decrease i n  c a l c i u m s a l t d e p o s i t i o n , o r a c o m b i n a t i o n o f b o t h , c a n n o t be determined  by o p t i c a l o b s e r v a t i o n o f p r e s e r v e d  Pannella ring  (1974) o b s e r v e d  4, 8, 1 4 , and 2 8 - d a y c y c l e s  thickness i n a variety of t r o p i c a l  attributed  t o l u n a r and t i d a l  (unpub ms.) o b s e r v e d  effects.  collected  T h e i r o b s e r v a t i o n s suggest  d a i l y growth  ring  and C o b l e  zone i s formed  Another  f e d a t t h r e e hour hours.  A third  times of the  i s formed  during the  d u r i n g the day.  (1977) w o r k e d w i t h t h r e e s p e c i e s o f L e p o m i s  One t a n k was k e p t  intervals.  (Haemulon  that the dark p o r t i o n of the  to d i f f e r e n t i a t e  e f f e c t s o f p h o t o p e r i o d and f e e d i n g r e g i m e  hour  o t o l i t h s , w h i c h he  during different  and T i l a p i a m o s a m b i c a , and a t t e m p t e d  formation.  in daily  B r o t h e r s and M c F a r l a n d  ( using transmitted light)  n i g h t , and t h e l i g h t Taubert  fish  a s m a l l number o f F r e n c h g r u n t  flavolineatum) otoliths day.  otoliths.  i n daily  i n constant l i g h t  between t h e  ring  and f e d a t t h r e e  was a l s o k e p t u n d e r c o n s t a n t l i g h t and  i n t e r v a l s w i t h a n i n e h o u r h i a t u s e v e r y 24  was k e p t u n d e r a 2 4 - h o u r t o 1 2 - h o u r  light-dark  c y c l e and f e d e v e r y s i x h o u r s , and a f o u r t h was m a i n t a i n e d  under  a 1 5 - h o u r t o 9-hour l i g h t - d a r k c y c l e and f e d e v e r y s i x h o u r s . Fish  i n groups  one and two d e m o n s t r a t e d  no d a i l y r i n g f o r m a t i o n ,  but  i n some c a s e s had as many as f o u r t i m e s as many r i n g s as t h e  age  i n days.  These presumably  t h r e e d i d not produce  a ring  represent feeding rings.  e v e r y 36 h o u r s .  Taubert  Group  and C o b l e  concluded  that a diurnal  cause d a i l y r i n g  feeding  r e g i m e was n o t s u f f i c i e n t t o  f o r m a t i o n , and t h a t a d i u r n a l  p e r i o d was s u f f i c i e n t t o p r o d u c e d a i l y culated  t h a t t h e f i s h under  rings.  the 24-hour  p h o t o p e r i o d d i d n o t produce 36-hour  change i n p h o t o They f u r t h e r  l i g h t t o 12-hour  r i n g s because  spe-  dark  the l i g h t  cycl  was t o o a b n o r m a l  to entrain a biological  damping e f f e c t .  T a u b e r t and C o b l e were a l s o a b l e t o s t o p d a i l y  ring  c l o c k , and i n f a c t had  f o r m a t i o n by r e d u c i n g w a t e r t e m p e r a t u r e , p r o d u c i n g an a r t i -  ficial  annulus which resembled the annulus i n w i l d  They c o n c l u d e d t h a t d a i l y growth r i n g  fish  otoliths  f o r m a t i o n c e a s e s when f i s h  growth ceases. The growth  relationship i s largely  relationship  between  unexplored.  between  sagitta  B r o t h e r s and M c F a r l a n d between  T a u b e r t and C o b l e f o u n d a l i n e a r r a d i u s and f i s h l e n g t h  changes  relationship  i n d a i l y growth r i n g  summary, t h e r e  formation  i s much s p e c u l a t i o n and l i t t l e  in particular.  finest v i s i b l e rings ticularly  width to  transitions.  e v i d e n c e r e g a r d i n g o t o l i t h growth ring  (r=0.9936).  l o g o t o l i t h r a d i u s and l o g f i s h l e n g t h f o r F r e n c h g r u n t s  history In  w i d t h and f i s h  (unpub ms.) f o u n d a l i n e a r  and a t t e m p t e d t o r e l a t e life  d a i l y growth r i n g  documented  i n g e n e r a l , and d a i l y  growth  I t i s commonly assumed t h a t t h e  i n any f i s h o t o l i t h a r e formed  i f they occur i n the f a s t e s t  growth  zone.  daily,  par-  I t was  the purpose of t h i s s t u d y :  to demonstrate the o c c u r r e n c e of d a i l y growth r i n g s sagittae  o f 0.  to examine  nerka.  the r e l a t i o n s h i p  growth r i n g  between  fish  g r o w t h and  daily  width.  t o d e v e l o p t e c h n i q u e s f o r t h e o b s e r v a t i o n and d a i l y growth  i n the  rings.  measurement  9  GENERAL METHODS  Fish  Preservation For  For  this  long  study,  f r y were p r e s e r v e d  term storage  Otolith Of The  the  fish.  We  The  (1977). stage  b r i g h t dots  For  acidic  small f i s h  head i s p l a c e d  or pressed  up  forceps.  were removed by h o l d i n g  d i s s e c t i n g microscope with opercular  seen under top  The  of  were and glass  appear  then  the  as  be  otoliths  o t o l i t h s of l a r g e r  f r y on  the  t h e d o r s a l s i d e down and  plates (Fig.3).  illumination  the  cm.)  otoliths  head can  The  size  a piece of  u s i n g drawn g l a s s p r o b e s and  using probes or f i n e  the  by T a u b e r t  The  The  (1-2  on  bet-  are  d e p e n d i n g on  to that described  under s i d e i l l u m i n a t i o n .  picked  and  techniques  the  o f t h e head  techniques  of a d i s s e c t i n g microscope.  gently apart  (>_ 2cm)  these  o t o l i t h s of extremely  The  used.  t h r o u g h t h e head b e h i n d  t h r o u g h the c e n t e r  small f i s h  e m p l o y e d two  teased  gills  since  to  u s u a l l y removed by m a k i n g a t r a n s -  o f t h e way  removed i n a manner s i m i l a r  fry  are  o r b i t s , or l o n g i t u d i n a l l y  the  7,  taken  to d e c a l c i f y o t o l i t h s .  cut three-quarters  Coble  was  t h r e e p a i r s o f o t o l i t h s , o n l y t h e s a g i t t a e were  inefficient.  on  a pH o f a p p r o x i m a t e l y  ethanol.  Removal  ween t h e o r b i t s .  the  had  s a g i t t a e of l a r g e f i s h  verse eye  tend  cent  ( s e v e r a l weeks o r more) c a r e  ensure t h a t the e t h a n o l preservatives  i n 70 p e r  stage  of  the  removing  s a g i t t a e could  j u s t p o s t e r i o r t o t h e eye  then  orbits,  the be and  10  g u r e 3.  A p p e a r a n c e o f 0^ n e r k a f r y b e f o r e and o f l o w e r jaw, g i l l sag i t t a e .  c o v e r s and  gills  after  removal  to r e v e a l  lower  jaw,  gil.l c o v e r s ,  gills  removed  11  were removed by t e a s i n g a p a r t t h e Otolith  cartilage.  storage  O t o l i t h s were s t o r e d d r y i n t h e d e p r e s s i o n s o f M i c r o T e s t tissue culture plates. p l a t e was  numbered and  Each of the s i x t y d e p r e s s i o n s lettered  f o r convenience,  and  3034  i n every the  otoliths  were h e l d i n the d e p r e s s i o n s w i t h a sheet o f l / 1 6 t h i n c h plexiglass  cut to f i t i n s i d e the p l a t e .  The  p l e x i g l a s s was  held  a g a i n s t t h e d e p r e s s i o n s by a p i e c e o f sponge between t h e l i d and the p l e x i g l a s s , elastic  and  the e n t i r e  As  h e l d t o g e t h e r by s e v e r a l  bands.  O t o l i t h P r e p a r a t i o n and The  u n i t was  Observation  o t o l i t h s of very s m a l l sockeye  t h e f r y grow, t h e o t o l i t h s  becomes o b s c u r e d .  f r y were e x a m i n e d  t h i c k e n and  the  internal  T h i c k e r o t o l i t h s were g r o u n d  into  whole.  structure  thin  sec-  t i o n s by a t t a c h i n g them t o t h e c e n t e r o f a g l a s s m i c r o s c o p e  slide  using a c r y l i c  by  adhesive.  Initially  h a n d , u s i n g C a r b o r u n d u m No. water  on a s i n t e r e d  rounding  50 a l u m i n u m o x i d e powder m i x e d  glass plate.  keep the s l i d e p a r a l l e l  t h e o t o l i t h s were g r o u n d  E x t r e m e c a r e had  t o t h e g r i n d i n g s u r f a c e , and  the ground s u r f a c e o f the o t o l i t h .  g r i n d i n g methods d e v e l o p e d were found  t o be  thus  avoid  by J o h n N e i l s o n ( N e i l s o n , I_n P r e s s )  s u p e r i o r and  were used f o r l a t t e r  to the g r i n d i n g s u r f a c e to ensure  m a i n t a i n s c o n s t a n t and  taken  Subsequently,  N e i l s o n ' s method e m p l o y s a g r i n d i n g j i g w h i c h parallel  t o be  with  h o l d s the  a flat  controllable pressure.  work.  grind,  slide and  Mylar grinding  to  12  sheets as had  (available  f r o m 3M  i n a v a r i e t y of g r i t  the g r i n d i n g s u r f a c e . a particle  otoliths.  s i z e o f 0.5  Rapid  by  was  m i c r o n s and  r e s i n as an a d h e s i v e  The  was  critical.  remove r i n g s f r o m t h e o t o l i t h ,  otolith fairly  otoliths for We  and  use  the ground The  sur-  grinding sensitive  found t h a t  this  U c h i y a m a 1976)  acetate  greatly simplified. can  insufficient grinding  can  d u r i n g g r i n d i n g , and  T a u b e r t and  have suggested  Coble  The a  impressions,  i n e t c h i n g r a t e w i t h i n and  and  1977;  e t c h i n g the either  found  ground  in preparation  i m p r e s s i o n , o r t o augment d i r e c t  viewing.  considerable  between o t o l i t h s e c t i o n s .  This variation  r e s u l t e d i n o c c a s i o n a l l o s s e s of m a t e r i a l i n  t h e c e n t e r and  edges o f the o t o l i t h .  improvement i n the v i s i b i l i t y a potential  and  is required.  ( P a n n e l l a 1971;  t a k i n g an a c e t a t e  variation  was  f o r s e v e r a l seconds i n d i l u t e HCl,  d i d not  was  Excessive grinding  numerous t i m e s  high l e v e l of s k i l l  Strusaker  glue  by an o v e r b u r d e n o f c a l c i u m c a r b o n a t e .  must be o b s e r v e d  Several authors  while  We  the  t o remove t h e o t o l i t h s  the g r i n d i n g p r o c e s s  leave r i n g s obscured  and  paper,  paper.  N e i l s o n a l s o employed heat  instead of a c r y l i c .  g r i n d i n g process  the a c r y l i c  to the s l i d e .  method i n v o l v e d o n l y s l i g h t h e a t i n g r e a t t a c h them, and  used t o f i n i s h  i n acetone,  reattached  repeated.  lapping  p e r f o r m e d on 25 m i c r o n  immersing the s l i d e  then  called  was  s i d e of the o t o l i t h ,  f a c e o f t h e o t o l i t h was process  finest grit,  g r i n d i n g was  A f t e r p o l i s h i n g one softened  The  s i z e s ) were used  and  t h e r e was  was  not used f o r r o u t i n e work.  of the  Since  t h e r e was  both  o n l y minor  internal otolith structure,  for introducing error,  this  technique  13  The p r e p a r e d o t o l i t h was c l a r i f i e d or immersion otolith  o i l prior  becomes  absorbed,  to viewing.  by i m m e r s i o n  The i n t e r n a l  increasingly visible  obscures  r i n g s by m a k i n g t h e opaque p o r t i o n o f t h e r i n g serial  cess of c l a r i f i c a t i o n s l o w l y decreased.  rapidly  stable  f o r s e v e r a l days.  It  that f o rsmall o t o l i t h s  s i x t o e i g h t hours o f c l a r i f i c a t i o n .  s e c t i o n s and l a r g e r o t o l i t h s  stable  thin  i n c r e a s e d f o r up t o  f o r s e v e r a l d a y s , and d e c r e a s e d  f o r a week o r  O b s e r v a t i o n s were made on t h e s e o t o l i t h s on t h e d a y  following placing  and t h e n  i n c r e a s e d f o r up t o s i x h o u r s , r e m a i n e d  c o u n t s were made a f t e r  more.  growth  transparent. For  i n c r e a s e d , then s t a b i l i z e d  was a c c o r d i n g l y made s t a n d a r d p r a c t i c e  a day, remained  the d a i l y  counts of d a i l y r i n g s during the pro-  f o r s i x t o t w e l v e h o u r s , and t h e n d e c r e a s e d  on t h i c k e r  agent i s  The c o u n t s on v e r y s m a l l o t o l i t h s o r v e r y  sections of otoliths  Counts  s t r u c t u r e o f the  as t h e c l a r i f y i n g  but excessive c l a r i f i c a t i o n  salmon f r y o t o l i t h s ,  i n glycerine  immersion.  Overclarified  o t o l i t h s were r e c o v e r e d by  them i n e t h a n o l t o remove t h e c l a r i f y i n g  a g e n t , and  reclarified. The c l a r i f i e d  o t o l i t h s were p h o t o g r a p h e d  t r i n o c u l a r compound  microscope  and t r a n s m i t t e d l i g h t ,  w i t h a W i l d 35mm a u t o m a t i c c a m e r a . 50X and 200X negative).  u s i n g an Olympus and f i t t e d  O t o l i t h s were p h o t o g r a p h e d  ( b a s e d on t h e m a g n i f i c a t i o n  at  f a c t o r f o r the f i l m  A p r o j e c t o r s l i d e was p r e p a r e d d i r e c t l y  from t h e f i l m  n e g a t i v e and p r o j e c t e d o n t o a w h i t e t a b l e t o p f o r c o u n t s and measurements. A s e r i e s of annotated photographs been prepared  o f sockeye  o t o l i t h s has  t o a s s i s t o t h e r s i n i n t e r p r e t i n g d a i l y growth p a t -  14  t e r n s and liths  understanding  examined  A p p e n d i x 1. lysing  in this  The  t h e methods used study.  the  oto-  These p h o t o g r a p h s are p r e s e n t e d  methods e m p l o y e d  the data are presented  in interpreting  i n c o u n t i n g , measuring  i n the s p e c i f i c  methods  and  in  ana-  sections.  15  EXPERIMENT 1: DAILY GROWTH RINGS IN THE OTOLITHS OF JUVENILE SOCKEYE SALMON ( O n c o r h y n c h u s  nerka)  Introduction. The  early  life  h i s t o r y o f s a l m o n makes t h e i n t e r p r e t a t i o n o f  growth r i n g data d i f f i c u l t . s t r e a m b e d s between occur u n t i l  Sockeye  eggs h a t c h i n t h e g r a v e l o f  December and J a n u a r y , b u t emergence may n o t  March, A p r i l  o r May.  F r y g e n e r a l l y move d i r e c t l y t o  a l a k e upon e m e r g e n c e , b u t s h o r t - t e r m s t r e a m r e s i d e n c y uncommon. (Dill  t o e m e r g e n c e , t h e f r y e x i s t on y o l k  1 9 6 8 ) , and e x o g e n o u s  exhausted. ring  Prior  T a u b e r t and C o b l e  (1977) s u g g e s t t h a t d a i l y i s prompted  i n t e n s i t y and i n f l u e n c e d  changes  emergence.  in light  Salmon f r y may  and may be e x p o s e d  Some s o c k e y e s a l m o n  f r y do n o t b e g i n f e e d i n g Such  f a c t o r s may a l l  first  suspect the a p p l i c a t i o n  i n the time  p r o d u c e s g r o w t h r i n g s and make  to wild populations of results  from l a b o r a t o r y s t u d i e s .  To make t h i s  study of o t o l i t h  a p p l i c a b l e t o w i l d p o p u l a t i o n s , we c o l l e c t e d their natural  facilities.  serve to increase the v a r i a t i o n  w h i c h an i n d i v i d u a l  environment.  even  f r yare c a l l e d  " p i n h e a d s " and a r e commonly o b s e r v e d i n r e a r i n g  at  to diur-  i n t e n s i t y and t e m p e r a t u r e p r i o r t o  a f t e r completely absorbing their yolks.  These  growth  by f l u c t u a t i o n s i n  by t e m p e r a t u r e .  occur a t v a r i o u s depths i n the g r a v e l nal  reserves  f e e d i n g b e g i n s as y o l k r e s e r v e s a r e  formation i n the s a g i t t a  light  i s not  and r e a r e d  obtained growth fry in  16  Methods S o c k e y e f r y were c o l l e c t e d  from  converging  t h e mouth o f F u l t o n R i v e r s p a w n i n g c h a n n e l British  Columbia,  i n 1978.  To d e t e r m i n e  emergence and c a p t u r e a t t h e c h a n n e l  throat traps at  2 a t Babine  Lake,  t h e maximum t i m e  between  mouth, 10,000 f r y were dyed  w i t h n e u t r a l r e d and r e l e a s e d a t t h e u p s t r e a m end o f t h e c h a n n e l . Experimental  f r y were c a p t u r e d  i n the l a t e evening  m o r n i n g o f May 12 and 1 3 , and r e a r e d channel. and  were  enclosures i n the  S a m p l e s were t a k e n on May 1 3 , 2 3 , and 3 1 , and J u n e 9,  were p r e s e r v e d  examined from and  in  and e a r l y  i n 70 p e r c e n t e t h a n o l .  each sample.  any o t o l i t h s which  Twenty-five  fish  One o t o l i t h was used f r o m e a c h  were c r y s t a l l i n e o r o t h e r w i s e  were fish,  uncountable  rejected. Sagittae  were removed  g l a s s microscope upward.  i n t h e l a b o r a t o r y and a f f i x e d t o  s l i d e s w i t h t h e s u l c u s (_i.e. p r o x i m a l  T h i c k e r specimens ( p r i m a r i l y  those  from  surface)  the June 9  s a m p l e ) were g r o u n d l i g h t l y on t h e p r o x i m a l s u r f a c e , u s i n g a l u m i n i u m o x i d e on a s i n t e r e d clarified  glass plate.  w i t h a 35 mm p h o t o g r a p h i c prepared  directly  Ring counts defined  and a s t a n d a r d  attachment.  compound  film  negative.  were made on t h e p r o j e c t e d i m a g e s .  as a p a i r o f d i s t i n c t  microscope  A 35 mm p r o j e c t o r s l i d e  using the developed  b a n d s , one l i g h t  d i s c e r n i b l e r i n g s were c o u n t e d . lith  were  w i t h g l y c e r i n e and p h o t o g r a p h e d a t a m a g n i f i c a t i o n o f  200X u s i n g t r a n s m i t t e d l i g h t  was  A l l otoliths  A ring  was  and one d a r k . A l l  F i g u r e 4 shows a p r e p a r e d  ( f r o m t h e J u n e 9 s a m p l e ) and t h e a s s o c i a t e d  count.  oto-  17  Results Of  t h e 10,000 f r y r e l e a s e d  a t t h e u p s t r e a m end o f t h e  c h a n n e l , 340 were r e c a p t u r e d i n s a m p l i n g a t t h e mouth; 339  were  c a p t u r e d w i t h i n 5 h o u r s , and o n l y one  the  second evening a f t e r  release.  subsequently, during  With such a s h o r t passage  t h r o u g h the c h a n n e l , the date o f c a p t u r e o f the f r y from o t o l i t h s were s a m p l e d If  i n days  (after  which  be t a k e n as t h e d a t e o f e m e r g e n c e .  r i n g s a r e b e i n g formed  least-squares linear  ring  may  of time  daily  i n sockeye salmon  r e g r e s s i o n of r i n g  sagittae,  c o u n t a g a i n s t known  c a p t u r e ) s h o u l d have a s l o p e o f 1.  a  age  Further, i f  f o r m a t i o n began on t h e a v e r a g e a t t h e t i m e o f c a p t u r e , t h e  regression should s t i l l  have a s l o p e o f 1 when c o n s t r a i n e d  through the o r i g i n . The  mean and v a r i a n c e o f d a i l y  sockeye salmon British  ring  counts i n samples  from F u l t o n R i v e r spawning  C o l u m b i a , 1978  SAMPLE  were as  2  Mean  3.9565  10.000  17.850  25.668  Variance  53.316  58.000  27.503  19.101  test  of homogeneity  ces o f the f o u r samples  23  May  4  May  1  May  3  DATE  Bartlett s  of variance  31  June  indicates  9  the  are not s i g n i f i c a n t l y d i f f e r e n t .  2, 3, and 4 a r e d i s t r i b u t e d  Lake,  follows:  1 13  c h a n n e l , Babine  o f 25  around  a mean w h i c h  varianSamples  i n each case i s  c l o s e t o a l i n e p a s s i n g t h r o u g h t h e o r i g i n w i t h a s l o p e o f 1.  18  g u r e 4.  Sagittae  f r o m an 0. n e r k a f r y removed  e n c l o s u r e on J u n e 9.  from the  19  The  counts below  the  first  t h e mean i n t h e s e s a m p l e s  s a m p l e by z e r o c o u n t s b e c a u s e  mean i s n o t due t o c o u n t i n g e r r o r . impossible,  the f i r s t  was  t o a l l t h e d a t a , and a s e c o n d  t h r o u g h t h e o r i g i n was f i t t e d (Fig.5).  regression.  One r e g r e s s i o n  regression  line  constrained  of regression of d a i l y  three ring  i s as f o l l o w s :  Intercept  (all  negative counts are  t o t h e d a t a from t h e l a s t  The s t a t i s t i c s  c o u n t a g a i n s t t i m e i n days  Standard  below the  s a m p l e c a n n o t have a mean o f z e r o and was  from t h e c o n s t r a i n e d  samples  the v a r i a t i o n  Because  eliminated fitted  must be r e p r e s e n t e d i n  3.4869  regression  Slope  SD o f s l o p e  0.81660  0.10020  0.97584  0.033899  data)  Regression  constrained  through the o r i g i n ( d a t a f o r May 2 3 , 31) Discussion The  r e s u l t s suggest t h a t growth  s a g i t t a e a r e formed  daily.  Because  r i n g s i n sockeye  salmon  the vast majority of f r y  m i g r a t e d r a p i d l y and hence were c a p t u r e d a l m o s t  i m m e d i a t e l y upon  emergence, the r e s u l t s  further  first  f o r m a t i o n i s t h e date o f emergence.  discernible ring  apparent reduction the  suggest that the average date of  i n v a r i a n c e w i t h age c a n be a t t r i b u t e d  The both to  d e a t h o f t h e f i s h w h i c h do n o t grow, and t o t h e n a t u r e o f  pre-emergent  growth  r i n g s which  a r e formed  w h i l e the f i s h  are i n  20  F i g u r e 5.  Graph o f o t o l i t h emergence.  ring  c o u n t v e r s u s age i n days  since  AO  36-  32-  DAYS  SINCE  EMERGENCE  21  the  g r a v e l of the  s t r u c t u r e and  channel.  T h e s e r i n g s have an e x t r e m e l y  m i g h t be o b s c u r e d as  Another p o s s i b l e explanation variance  of r i n g counts with  r i n g s at e a r l i e r  ages.  r i n g s , even i n f i s h  age  f o r the  thicken  r e d u c t i o n of  i s the p r e s e n c e o f  H o w e v e r , we  r a i s e d i n the  s u b s t a n t i a l l y higher  the o t o l i t h s  have n o t  observed  i n our  (Fig.6). the  subdaily subdaily  l a b o r a t o r y at growth  than those o c c u r r i n g  fine  rates  experimental  enclosures. Finally,  there  is a possibility  of  s o u r c e s of e r r o r t h a t account f o r the age.  R i n g s can  g r i n d i n g , and  can  where g r i n d i n g t o be  be  removed f r o m t h e be  o b s c u r e d by  is insufficient.  the  faulty  techniques  r e d u c t i o n of v a r i a n c e  otoliths  through  l a y e r of calcium  Such e r r o r s are  not  s e r i o u s f o r s o c k e y e f r y a t e a r l y a g e s as h e r e  t h o u g h t h e y may  p r o v e t o be  at l a t e r  stages  of  the  as with  excessive overburden considered reported,  life history.  22  F i g u r e 6.  S a g i t t a e o f an 0. on  May  13.  n e r k a f r y removed f r o m t h e  enclosure  22a  23  EXPERIMENT 2: THE RELATIONSHIP BETWEEN WIDTH OF DAILY GROWTH RINGS IN SAGITTAE AND CHANGE IN BODY S I Z E OF SOCKEYE SALMON ( O n c o r h y n c h u s  nerka)FRY  Introduction O t o l i t h s have been used as a method o f a g i n g f i s h turn of this lizing  c e n t u r y (Graham 1 9 2 9 ) .  otoliths  are often confined  W h i l e growth  studies  t o those species  fish  growth  and o t o l i t h  ( J o n s s o n and S t e n s e t h 1 9 7 7 ) .  Until  size  the r e l a -  i n salmonids  recently, only techniques  utilizing  a n n u l a r marks were a v a i l a b l e  studies.  The o c c u r r e n c e o f d a i l y g r o w t h r i n g s i n t e l e o s t  liths  ( P a n e l l a 1971) i n c l u d i n g  uti-  with  u n r e a d a b l e s c a l e s , a t l e a s t one r e c e n t s t u d y examined t i o n s h i p between  since the  f o r age and g r o w t h oto-  t h o s e o f sockeye salmon  (Wilson  and L a r k i n 1 9 8 0 ) p r o v i d e s an i m p o r t a n t new m e t h o d o l o g y  f o r age  and g r o w t h The between  studies.  purpose o f t h i s  e x p e r i m e n t i s t o examine  f i s h w e i g h t and o t o l i t h  f o r sockeye salmon as t h e t i m e  the r e l a t i o n s h i p  s i z e over short time  fry, utilizing  daily rings  intervals  instead of annuli  marker.  Methods Sockeye tilized spawning  salmon  f r y were r e a r e d  eggs t a k e n i n t h e f a l l  i n t h e l a b o r a t o r y from  fer-  o f 1978 a t t h e Weaver c r e e k  channel near M i s s i o n , B r i t i s h  Columbia.  A random  sample  24  o f 64 f r y was b r a n d e d The  to enable i d e n t i f i c a t i o n of i n d i v i d u a l s .  marks c o n s i s t e d o f t h e p r e s e n c e o r a b s e n c e  any one o f f i v e d i s t i n c t  l o c a t i o n s on t h e body.  a n a e s t h e t i z e d w i t h MS222 and b r a n d e d in liquid  nitrogen.  the p e c t o r a l  fin,  The f r y were  The b r a n d s i t e s w e r e : on t h e l e f t  below  the d o r s a l  brand i n  w i t h a c o p p e r w i r e immersed  fin,  side  above  and a b o v e t h e a n a l f i n ,  and on t h e r i g h t s i d e above t h e p e c t o r a l fin  of a cold  f i n and b e l o w  the d o r s a l  ( s e e F i g . 7) .  T h e s e v a r i o u s p a t t e r n s e n a b l e d 32 f r y t o be i d e n t i f i e d individuals  i n e a c h o f two t a n k s .  and  before being p l a c e d i n a tank ( 1 2 i n x 1 2 i n x 2 4 i n ) .  branded  One t a n k was s t o c k e d on J u n e later.  6, 1 9 7 9 , and a s e c o n d  The f i s h were n o t f e d f o r 24 h o u r s p r i o r  order to minimize v a r i a t i o n fish  E a c h o f t h e f r y was  as  were f e d a t one h o u r  i n stomach  to weighing i n  c o n t e n t s ; o t h e r w i s e the  o f a M e t t l e r a u t o m a t i c pan b a l a n c e e n a b l e d t h e f i s h  t a k e n by r e m o v i n g  the f i s h  two days  i n t e r v a l s e i g h t times a day.  on t h e p a n , a w e i g h t t a k e n , t h e s c a l e  weighed  tared  from the pan.  The use  t o be p l a c e d  and a s e c o n d  weight  The two w e i g h t s were  compared t o e n s u r e a g r e e m e n t w i t h i n 0.01 g r a m s , and a v e r a g e d . The  f i s h were r e w e i g h e d  carried  out during  i f necessary.  The w e i g h i n g s were a l l  t h e same p e r i o d o f t h e d a y , d u r i n g  the l a t e  afternoon. T h i r t y days  after  the f i r s t  b o t h t a n k s were a n a e s t h e t i z e d  t a n k was s e t up, t h e f i s h i n  and r e w e i g h e d .  s e c o n d w e i g h i n g were r e a r e d f o r an a d d i t i o n a l  F r y s u r v i v i n g the two weeks.  Data  25  F i g u r e 7.  Cold branding  s i t e s on 0.  nerka f r y .  Brand marks  Brand marks  cn Co  26  were r e c o r d e d f o r t h o s e f r y w h i c h d i e d  i m m e d i a t e l y as a r e s u l t  of  h a n d l i n g s t r e s s d u r i n g t h e s e c o n d w e i g h i n g as w e l l as f o r t h o s e still  r e m a i n i n g a t t h e end o f t h e e x p e r i m e n t .  died during  t h e c o u r s e o f t h e e x p e r i m e n t were  S a g i t t a e were removed and g r o u n d Methods).  One  otolith  Kodak p a n a t o m i c scope.  Projector  negatives.  S l i d e s were p r o j e c t e d  f a c t o r 382X.  changes  is oval  usually  and m o r p h o l o g y  as i t g r o w s .  radial  sockeye  The  otolith. foci,  s a g i t t a o f the y o l k sac f r y becomes  typi-  t h e r o s t r u m and p o s t r o s t r u m e x h i b i t i n g  growth.  i n our samples  These a r e a s o f g r e a t e s t  was  l e s s prone  p r e p a r a t i o n , more u n i f o r m l y s h a p e d , easily discernible daily  of  t y p i c a l l y has s e v e r a l  show t h e most d i s t i n c t d a i l y r i n g  postrostrum  focus,  sockeye  i n s h a p e , b u t as t h e f r y grows t h e s a g i t t a  t h e most r a p i d  a  a d i s c u s s i o n o f t h e measurement  the s a g i t t a  c a l l y heart-shaped with  top using  M e a s u r e m e n t s were t a k e n  Figure 8 represents a generalized  shape  film  image.  facilitate  I n sockeye salmon and  using  angle, with a fixed  A b r i e f d e s c r i p t i o n of the growth otoliths will  from  on a w h i t e t a b l e  h e i g h t and  from the p r o j e c t e d  system used.  photographed  s l i d e s were p r e p a r e d d i r e c t l y  to achieve a magnification  salmon  (see G e n e r a l  i n a 35mm camera mounted on a compound  p r o j e c t o r held at a fixed  directly  which  discarded.  on b o t h s i d e s  f r o m e a c h f i s h was  X film  Those f i s h  ring  growth  formations.  The  to breakage  during  and c o n s i s t e n t l y had  t h e most  patterns.  To e n s u r e t h a t m e a s u r e m e n t s were t a k e n on t h e same r a d i u s f o r each o t o l i t h ,  a l i n e was  drawn t o t h e r o s t r u m t h r o u g h t h e  27  F i g u r e 8.  S a g i t t a e o f an 0. post-rostrum, radius.  nerka  nucleii  f r y showing  and  rostrum,  l o c a t i o n of p r e f e r r e d  28  focus immediately a d j a c e n t to the p o s t - r o s t r u m . drawn i n a s t r a i g h t bulge at a fixed  line  from t h i s  focus to the  a n g l e o f 40 d e g r e e s  For the f i s h  from the f i r s t  of the experiment,  t h e r a d i u s was  r i n g 14 r i n g s b a c k  f o c u s to the growth radii  tank which  s u r v i v e d to the  determined  Similarly,  (2) from the f o c u s  from the edge;  those f i s h which  the experiment  from the second  (3) from  initial  to the edge,  back  The  and  the  These weights  s u r v i v e d t o t h e end  t a n k ( s t o c k e d two d a y s  f i r s t ) were measured  end  for three distances:  c o r r e s p o n d e d t o t h e f i n a l , m i d d l e , and  from the edge.  post-rostral  r i n g 44 r i n g s b a c k f r o m t h e e d g e .  respectively.  was  (Fig. 8).  (1) f r o m t h e f o c u s t o t h e edge o f the o t o l i t h ; to the growth  A radius  after  the  t o t h e 1 4 t h and 42nd  o t o l i t h s of those f i s h  from the  of  rings first  t a n k w h i c h d i e d as a r e s u l t o f h a n d l i n g s t r e s s were measured  to  t h e edge and  from  t o t h e r i n g 30 r i n g s b a c k  the second  t a n k were measured  r i n g s back  from the edge ( F i g . 9 ) .  cess l e f t  t o t h e edge and  The f r y was  b r a n d i n g and w e i g h i n g  measured  and a v e r a g e v a l u e s were c a l c u l a t e d . determined  28  from the  pro-  edge  (Fig. 9).  I n e a c h c a s e t h e r a d i u s was  otolith  The  Those  to the r i n g  a c l e a r s t r e s s m a r k 28 o r 30 r i n g s b a c k  in every otolith  was  from the edge.  three independent  Thus,  f o r each f i s h  a s e r i e s o f two o r t h r e e w e i g h t s  r a d i u s corresponding to each  times  there  and a s t a n d a r d  weight.  r e l a t i o n s h i p b e t w e e n o t o l i t h m e a s u r e m e n t s and w e i g h t s e v i d e n t l y c u r v i l i n e a r , as would  of  be e x p e c t e d c o n s i d e r i n g  29  F i g u r e 9.  S a g i t t a e o f 0. n e r k a f r y removed 6.  f r o m Tank 2 on J u l y  30  that the r e l a t i o n measurements. to  i s between l i n e a r  Accordingly a standard  the l o g o f the weight  Additionally,  and e s s e n t i a l l y v o l u m e t r i c regression  l i n e was  fitted  and t h e l o g o f t h e r a d i u s .  t h e d a t a i n raw f o r m were f i t t e d  by o r t h o g o n a l  polynomials. The l o g r e g r e s s i o n l i n e was e m p l o y e d t o b a c k c a l c u l a t e w e i g h t o f each  individual  using a l a t e r weight pared w i t h those  f i s h at the time o f p r e v i o u s w e i g h i n g s ,  and r a d i u s .  w e i g h t s were com-  b e t w e e n o t o l i t h measurements  f r y was e v i d e n t l y c u r v i l i n e a r , that the r e l a t i o n  to  The p r e d i c t e d  observed.  The r e l a t i o n s h i p  measurements.  the  and w e i g h t s o f  as w o u l d be e x p e c t e d  i s between l i n e a r  considering  and e s s e n t i a l l y v o l u m e t r i c  Accordingly a standard  regression  l i n e was  fitted  t h e l o g o f t h e w e i g h t and t h e l o g o f t h e r a d i u s .  Additionally,  t h e d a t a i n raw f o r m were f i t t e d  by o r t h o g o n a l  polynomials. Results Fish  weights  and c o r r e s p o n d i n g r a d i i a r e g i v e n i n T a b l e I .  The s t a n d a r d l i n e a r  regression line fitted  t o t h e l o g g e d d a t a had  a l i n e o f b e s t f i t £nW = -11.27 + 4.409* & n  R  0.9230 ( s e e F i g . 1 0 ) .  o f the r e s i d u a l s  A visual  inspection  ( F i g . 1 1 ) showed  no marked non-random  points  f i s h are l i n k e d ,  f o r each  made t h a t a l l d a t a p o i n t s were  trends.  and an r  2  of  A l t h o u g h the data  the c o n s e r v a t i v e assumption independent.  was  31  F i g u r e 10.  in fish dual  fish  second 20)  w e i g h t v e r s u s in (No's  1-29)  otolith  radius for i n d i v i -  at the i n i t i a l  (June 6 o r 8 ) ,  ( J u l y 6) a n d , where a p p l i c a b l e ,  weighing.  third  (July  In Otolith Radius  32  F i g u r e 11.  P l o t o f r e s i d u a l s from l i n e a r on  r e g r e s s i o n o f In w e i g h t  Jin r a d i u s as shown i n F i g u r e  10.  32a  Residual  o  O  o 4*  "i  1  p 1  o  r  o  i  r  a •  > o c •o  >• O •  Table I  Jin o t o l i t h  radius  experimental  f r y at the i n i t i a l  ( J u l y 6) and t h i r d  Fish# 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29  F i r s t Weighing JlnR JlnW 2. 5014 2. 5479 2. 5408 2. 6174 2. 4874 2. 5088 2. 4973 2. 4681 2. 5518 2. 5233 2. 4423 2. 5080 2. 4510 2. 4891 2. 4553 2. 3842 2. 5392 2. 4441 2. 4570 2. 4380 2. 4380 2. 4204 2. 5031 2. 4562 2. 5337 2. 3979 2. 4998 2. 5193 2. 4105  -0. -0. -0. -0. -0. -0. -0. -0. -0. -0. -0. -0. -0. -0. -0. -0. -0. -0. -0. -0. -0. -0. -0. -0. -0. -0. -0. -0. -0.  31608 18995 20702 22941 21567 36962 27839 30517 25618 20949 51584 45256 52256 39304 45413 63488 45099 47160 67139 52763 56212 38566 43232 39156 02020 66943 21691 25489 27181  ( JlnR) and Zn f i s h w e i g h t ( Jin) f o r (June 6 o r 8 ) , s e c o n d  ( J u l y 20) w e i g h i n g s .  Second Weighing JlnR JlnW 2. 7246 2. 7428 2. 7447 2. 7402 2. 7020 2. 7324 2. 7147 2. 6797 2. 7606 2. 7498 2. 6589 2. 7114 2. 6426 2. 6831 2. 6672 2. 6064 2. 7441 2. 5665 2. 6844 2. 6469 2. 6123 2. 6525 2. 6940 2. 6617 2. 7376 2. 6851 2. 7074 2. 7279 2. 6797  0. 63340 0. 80648 0. 99990 0. 92861 0. 81802 0. 60704 0. 86752 0. 68057 0. 78800 0. 83291 0. 49042 0. 58667 0. 30822 0. 73573 0. 57774 0. 30895 0. 65389 0. 40213 0. 61842 0. 51462 0. 18482 0. 67905 0. 66783 0. 72368 0. 88830 0. 61896 0. 68914 0. 84243 0. 79706  T h i r d Weighing JlnPl JlnW  2. 7350 2. 6946 2. 7344 2. 7763 2. 7376 2. 8232 2. 7763 2. 7955 2. 8214 2. 7850  0. 70804 0. 60103 0. 87338 0. 91228 0. 97305 1. 0435 0. 87838 0. 75565 1. 1105 1. 0282  34  The  orthogonal polynomial regression yielded  an r ^ o f  0.9157, b u t t h e r e s i d u a l s showed a marked non-random t r e n d i n that the weights of the f i s h  with  the s m a l l e s t r a d i i  were  underestimated. The  w e i g h t o f each  individual  w e i g h i n g s was b a c k c a l c u l a t e d £nW,. = £nW  2  InR^)  t o be e s t i m a t e d  W2 i s some l a t e r Rl  using the formula:  - bUnR^ -  where Wl i s t h e w e i g h t  at the time of previous  measured  weight  i s the r a d i u s corresponding t o the weight  t o be e s t i m a t e d  R2 i s t h e r a d i u s c o r r e s p o n d i n g t o W2 A simple d e r i v a t i o n  f o r this  I t s h o u l d be n o t e d  procedure  that four d i f f e r e n t  c a l c u l a t i o n s are possible with died  a t t h e second  i s g i v e n i n Appendix  these d a t a .  1.  s e t s o f back For those f i s h  weighing, o n l y the i n i t i a l  which  w e i g h t s c a n be  estimated  ( g r o u p A, F i g . 1 2 ) . F o r t h o s e f i s h w h i c h s u r v i v e d t o  the  weighing, the i n i t i a l  third  w e i g h t c a n be c a l c u l a t e d  w e i g h t and r a d i u s c o r r e s p o n d i n g t o e i t h e r weighings  u s i n g t h e f i n a l w e i g h t and r a d i u s  I t s h o u l d be n o t e d  initial  the middle or f i n a l  ( g r o u p s B and C, F i g . 1 2 ) , and t h e m i d d l e w e i g h t c a n be  back c a l c u l a t e d 12).  from t h e  w e i g h t based  further  ( g r o u p D, F i g .  that o n l y the e s t i m a t e o f the  on t h e m i d d l e w e i g h t and r a d i u s  (Group  B)  d o e s n o t u t i l i z e a r a d i u s m e a s u r e d t o t h e edge o f t h e o t o l i t h t h a t back c a l c u l a t i n g  the weight a t the second  weighing  and  utilizes  35  F i g u r e 12.  S a g i t t a e o f 0.  n e r k a f r y showing p o s s i b l e types  back c a l c u l a t i o n s of p r e v i o u s corresponding fish text.  previous  w e i g h t and  otolith  otolith  f i s h weight using r a d i u s and  of the  a  latter  r a d i u s as d e s c r i b e d  i n the  36  a change i n r a d i u s f o r a p e r i o d 28-30 and  42-44 d a y s f o r t h e o t h e r  The lation  o f o n l y 14  d a y s (compared  estimates).  d i f f e r e n c e between the w e i g h t s e s t i m a t e d  and  by  back c a l c u -  t h e w e i g h t s o b s e r v e d , when p l o t t e d a g a i n s t  t h e p r e d i c t e d w e i g h t shows a non-random d i s t r i b u t i o n G r o u p s A,  B,  and  C,  show a s i m i l a r d e c l i n i n g t r e n d  from p o s i t i v e to n e g a t i v e  as  fish  the d i f f e r e n c e s i n group D are  with  size increases.  the  n  of  (Fig.13).  of  differences  A l l but  one  of  positive.  Discussion Because the the  r a d i u s m e a s u r e m e n t s were d e t e r m i n e d p r i m a r i l y on  b a s i s of d a i l y r i n g counts,  otolith  r a d i u s and  r^  value  f o r the  m i n e d by J o n s s e n and  fish  of a d a i l y r i n g  regression  Stenseth  back c a l c u l a t e d weights of  fry.  to t h a t length  The  deterversus  sample s i z e  and  study.  individual the  fish  i s presumably  r e g r e s s i o n model as  of growth, a r e s u l t of  and an a  measurement  both.  A b r u p t changes i n the o t o l i t h w e i g h t may  the  d i f f e r e n c e between o b s e r v e d  o f some i n a d e q u a c y o f  r e f l e c t i o n of the p a t t e r n s e r r o r , or  is identical  (1978) f o r l o g f i s h  i n our  non-randomness o f the  expression  i n sockeye salmon f r y  r a d i u s , even t h o u g h a much s m a l l e r  s i z e r a n g e were used The  shows a r e l a t i o n s h i p  a mean d a i l y change i n t h e w e i g h t o f  o f 0.92  log o t o l i t h  c l o s e r e l a t i o n s h i p between  w e i g h t as h e r e d e s c r i b e d  b e t w e e n t h e mean w i d t h s a g i t t a e , and  the  occur.  F i s h w e i g h t was  growth p a t t e r n or the  fishes  u s e d h e r e as a measure o f s i z e  37  Figure  13.  R e s i d u a l s o f back lated  fish  predicted  calculated  w e i g h t - Un &n  weight.  weights  observed  fish  ( I n back weight)  calcuversus  0.4 0.3 +  •  0.2  © 9 ®  9  0.1 o ?  © 0  (/)  "9  0  cr  9  •  0.1 1  0.2 0.3 0.4 0.5  10  20  30  40  Sample Number  50  60  70  38  because  f o r very small  accurately  than l e n g t h .  f i s h , w e i g h t c a n be measured much more However, i f the o t o l i t h  i s growing i n  d i r e c t p r o p o r t i o n t o o t h e r h a r d p a r t s , a b r u p t changes form r e s u l t i n g  from changes  i n growth  r e s u l t o f s t r e s s , c o u l d cause  lity  o f time dependent  o r t h a t m i g h t o c c u r as a  the weight to o t o l i t h  t i o n s h i p t o d e v i a t e from a l i n e a r changes  log-form.  in otolith  ( p e r s . comm.) h a s n o t e d t h a t f o r s a m p l e s suffering remained  i n body  radius  There  rela-  is a possibi-  radius.  David  Levy  o f c h i n o o k salmon f r y  f r o m h a n d l i n g s t r e s s , w e i g h t d e c r e a s e d and l e n g t h c o n s t a n t ( o v e r a two week p e r i o d ) , b u t o t o l i t h  r i n g s were s t i l l  formed  growth  daily.  A n o t h e r p o s s i b l e cause o f the p a t t e r n o f d e p a r t u r e o f observed the  from p r e d i c t e d  course of the experiment.  relatively larger,  I f smaller  s m a l l e r and l a r g e r f i s h  fish  tend t o remain  tend t o remain  relatively  then s i z e - s e l e c t i v e m o r t a l i t y a t the second  c o u l d have c o n t r i b u t e d the  v a l u e s c o u l d be s e l e c t i v e m o r t a l i t y i n  not only to the r e l a t i v e l y  weighing  large value of  s l o p e (4.4) but a l s o t o the p a t t e r n o f d e p a r t u r e s of observed  from b a c k - c a l c u l a t e d factor  in this  vived  less  T h i s does n o t a p p e a r  a n a l y s i s , because  f i s h which died ficantly  sizes.  t h e mean i n i t i a l  a t the second w e i g h i n g  t h a n t h e mean i n i t i a l  t o t h e end o f t h e e x p e r i m e n t  weight of those  (0.693 gms) was n o t s i g n i -  weight o f the f i s h which  sur-  (0.700 g m s ) .  M e a s u r e m e n t e r r o r may be a f a c t o r b e c a u s e p r o c e d u r e o f back c a l c u l a t i o n  t o have been a  the s t a t i s t i c a l  assumes a c o n s t a n t s l o p e .  measurement e r r o r o r o t h e r random e f f e c t s  If  are a s i g n i f i c a n t  com-  39  ponent i n the calculation of  variation  will  f i s h with  o f w e i g h t and  always tend  a fortuitously  underestimate those with calculation constant  formula  rank order  relationship.  to overestimate  assumes t h a t t h e with  respect  fish will  t o t h e w e i g h t and could  ments f o r the  B,  t h e w e i g h t s and  individual  fish  G r o u p D ( F i g . 12 and of  fish  two  weeks p r i o r  mated f r o m f i n a l  w e i g h t and  end  r a d i u s and  The  p r e c i s i o n of the w e i g h t e s t i m a t e s  are  as good as  vals  ( g r o u p s A and  underestimation  B and  Stress  r a d i u s r a t i o at the  discussed) otolith  by  f o r the  the  i n the r e s i d u a l s .  radius  the  28-30 and  (as  weight  esti-  from the  i n t e r v a l of  related reductions  shorter  measure-  the p r e d i c t e d  radius  f o r an  weighing  the  i t is  42-44 day The  focus  edge. two  weeks  inter-  consistent  w e i g h t s e e n i n g r o u p D has  final  pat-  connecting  r i n g s back from the  a number  i n weight to  oto-  previously with  time i n t e r v a l  s m a l l change i n o t o l i t h r a d i u s ) , c o u l d deviation  By  e x p e r i m e n t , as  o r measurement e r r o r s a s s o c i a t e d  accentuated  C.  group C r e s p e c t i v e l y ) .  of previous  of p o s s i b l e causes. lith  estimates  radius  occur.  of the  o f t h e o t o l i t h t o t h e g r o w t h r i n g 14  the  and  Fig.13) represents  to the  a  otolith  (as shown i n F i g . 1 0 ) , did  back  thus g e n e r a t e the  corresponding  a p p a r e n t t h a t changes i n rank o r d e r  and  The  maintain  back  weight  ratio,  a f o r t u i t o u s l y small r a t i o .  Changes i n r a n k o r d e r  representing  the p r e v i o u s  l a r g e weight to r a d i u s  t e r n o f r e s i d u a l s s e e n f o r g r o u p s A, points  r a d i u s , t h i s method o f  cause the  the (and  edge o f hence  observed  the  40  M e a s u r e m e n t e r r o r i n w e i g h t w o u l d a p p e a r t o be a m i n o r c o n t r i b u t o r , because c o n s e c u t i v e  w e i g h t s on e a c h f i s h were  seen  t o a g r e e w i t h i n 0.01 gms.  Measurement e r r o r i s , h o w e v e r , a m a j o r  component o f t h e v a r i a t i o n  i n radius values.  minus 5 p e r cent uncommon. mining nical  o v e r t h r e e m e a s u r e m e n t s o f a r a d i u s were n o t  Much o f t h e v a r i a t i o n  the exact  Errors of plus or  r e s u l t e d from d i f f i c u l t y  l o c a t i o n of the a p p r o p r i a t e  problems c o n t r i b u t e d to v a r i a t i o n  focus.  rostrum  or post-rostrum,  m a t e r i a l removed d u r i n g Variation  during  grinding  t o the o t o l i t h can  and v a r i a t i o n  i n t h e amount o f  g r i n d i n g c a n change a p p a r e n t r i n g  i n t h e shape o f t h e o t o l i t h w i t h  i n d i v i d u a l s o f t h e same s i z e a l s o i n t r o d u c e s dard  method i s e m p l o y e d .  width.  s i z e , and b e t w e e n  variation  i n stan-  r a d i u s m e a s u r e m e n t s , p a r t i c u l a r l y when a r e l a t i v e l y  standard  tech-  the apparent l o c a t i o n of the  membranes a d h e r i n g  o b s c u r e t h e edge o f t h e o t o l i t h ,  Other  i n the radius  measurement; c h i p p i n g o r c r a c k i n g o f the o t o l i t h or p r e p a r a t i o n can obscure or s h i f t  deter-  simple  F o r example, because the o t o l i t h  c h a n g e s shape as i t g r o w s , a s t r a i g h t l i n e drawn f r o m t h e f o c u s t o t h e o u t s i d e edge o f t h e o t o l i t h pendicular  n o t n e c e s s a r i l y be p e r -  t o any o r a l l o f t h e r i n g s i t i n t e r s e c t s , nor w i l l i t  necessarily use  will  i n t e r s e c t a l l o f t h e r i n g s a t t h e same a n g l e .  o f a curved  radius, perpendicular  t o each r i n g  i t intersects,  was c o n s i d e r e d .  Unfortunately,  proved d i f f i c u l t  t o measure a c c u r a t e l y and d i r e c t l y ,  resulting  the l e n g t h o f a curved  r a d i u s d i d not always leave  to the l o n g i t u d i n a l a x i s .  the focus  The  line  and t h e  a t t h e same  Complex a g e , s i z e and shape  angle  specific  41  measurement s y s t e m s w o u l d a l m o s t c e r t a i n l y r e d u c e v a r i a t i o n i n r a d i u s m e a s u r e m e n t , as w o u l d preparation. describing  Appendix 2 g i v e s  t h e s e and  m e a s u r i n g and  improved  other  techniques  of  otolith  a s e r i e s of annotated  photographs  t e c h n i c a l problems a s s o c i a t e d  interpreting otoliths  and  with  d a i l y growth r i n g s i n  otoliths. Despite t h a t the vides  these s e v e r a l c o n s i d e r a t i o n s  i t i s t o be  r e l a t i o n s h i p b e t w e e n f i s h w e i g h t and  otolith  emphasized radius  a s a t i s f a c t o r y method f o r b a c k c a l c u l a t i o n o f w e i g h t  previous  age.  relatively  The  small  e r r o r s t h a t a r i s e from the (of the o r d e r  i n p r e d i c t i o n e v e n o v e r as  o f 15 p e r  short  a period  techniques  c e n t ) and as  two  pro-  at  used  permissible  weeks.  are  42  SUMMARY Daily  r i n g s a r e found  i n the s a g i t t a e of sockeye  Under the c o n d i t i o n s o f n a t u r a l mation  o f t h e s e r i n g s commenced, on a v e r a g e ,  emergence. of  We  have found  every t e l e o s t  fish  the s i z e of a f i s h , of  incubation observed,  daily rings,  similar  observed.  ring  the f o r -  at the time of  f o r m a t i o n s i n the o t o l i t h s  The c l o s e r e l a t i o n s h i p  between  t h e s i z e o f i t s o t o l i t h s and t h e o c c u r r e n c e  a l l o w s the back c a l c u l a t i o n  t h e r e f o r e , growth  salmon f r y .  over i n t e r v a l s  T h i s shows a r e l a t i o n s h i p  at l e a s t  of f i s h  weight  and,  as s h o r t as two weeks.  b e t w e e n a f i s h ' s change i n w e i g h t  the w i d t h o f the c o r r e s p o n d i n g d a i l y growth  ring.  and  43  LITERATURE CITED B a r k m a n , R.C. 1978. The use o f o t o l i t h g r o w t h r i n g s t o age young A t l a n t i c s i l v e r s i d e s , M e n i d i a m e n i d i a . T r a n s . Am. F i s h . Soc. 1 0 7 ( 6 ) : 790-792 B l a c k e r , R.W. 1974. Recent advances i n o t o l i t h 67-90 I n SEA FISHERIES RESEARCH, F.R. New Y o r k : J o h n W i l e y .  studies. Pages H a r d e n J o n e s , ed.  B r o t h e r s , E.B., C P . Mathews, and B. L a s k e r . 1976. D a i l y g r o w t h i n c r e m e n t s i n o t o l i t h s o f l a r v a l and a d u l t f i s h e s . U.S. F i s h . B u l l . 74. 8 p. B r o t h e r s . E.B. and W.N. M c F a r l a n d . 1979. C o r r e l a t i o n s between o t o l i t h s m i c r o s t r u c t u r e , g r o w t h and l i f e h i s t o r y t r a n s i t i o n s i n n e w l y r e c r u i t e d F r e n c h g r u n t s Haemulon f l o v o l i n e a t u m (Desmarest), Haemulidae. ICES-ELH Symposium on E a r l y L i f e H i s t o r y o f F i s h . A p r i l 1979. Contributed paper. 16p. C h r i s t e n s e n , J.M. 1964. Burning of o t o l i t h s , a technique f o r d e t e r m i n a t i o n o f s o l e and o t h e r f i s h . J . C o n s . i n t . E x p l o r . Mer 26: 73-81  age  D e g e n s , E.T. W.G. Deuser and R.L. H a e d r i c h . 1969. Molecular s t r u c t u r e and c o m p o s i t i o n o f f i s h o t o l i t h s . Man. B i o l . ( B e r l . ) 2: 105-113 Dill,  L.M.  1968. The s u b - g r a v e l b e h a v i o u r o f P a c i f i c s a l m o n l a r v a e . Pp. 89-99 I n SYMPOSIUM ON SALMON AND TROUT I N STREAMS ( T . G . . N o r t h c o t e , e d . ) H.R. M a c m i l l a n l e c t u r e s i n F i s h e r i e s , U n i v e r s i t y of B r i t i s h Columbia, Vancouver, B.C. 388 p.  Graham, M. 1929. S t u d i e s o f Age D e t e r m i n a t i o n i n F i s h P t . I I . A s u r v e y of the l i t e r a t u r e . F i s h e r y I n v e s t . , Land., Ser. 11(3):50p. H i c k l i n g , C F . 1931. The s t r u c t u r e o f t h e o t o l i t h Q u a r t . J . 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SCIENCE 173: 1124-1127  and  S t r u s a k e r , P., a n d J . H . U c h i y a m a . 1 9 7 6 . Age and growth o f the nehu, S t o l e p h o r u s purpurens ( P i s c e s : E n g r a u l i d a e ) , from t h e H a w a i i a n I s l a n d s as i n d i c a t e d by d a i l y g r o w t h i n c r e ments o f s a g i t t a e . F i s h . B u l l . 74: 9-19. T a u b e r t , B., a n d D.W. C o b l e . 1 9 7 7 . Daily rings in otoliths of t h r e e s p e c i e s o f L e p o m i s and T i l a p i a m o s s a m b i c a . J . F i s h . R e s . B o a r d Can. 34: 332-340. W i l s o n , K e n n e t h H. a n d P.A. L a r k i n . 1 9 8 0 . D a i l y g r o w t h r i n g s i n the o t o l i t h s of j u v e n i l e sockeye salmon (Oncorhynchus nerka). Can. J . F i s h . Aquat. S c i . 3 7 ( 1 0 ) : 1495-1498.  APPENDIX 1  THE BACK CALCULATION OF PREVIOUS FISH WEIGHT FROM CORRESPONDING PREVIOUS OTOLITH RADIUS AND A LATTER OTOLITH RADIUS AND FISH WEIGHT  When a s t r a i g h t weight  line  and l o g o t o l i t h  i s f i t t e d to the data  radius,  a relationship  for log fish i s obtained  o f the  form: y = a+bx which provides a p r e d i c t i o n  o f t h e mean v a l u e s o f y f o r a g i v e n  mean v a l u e o f x. To  back c a l c u l a t e  a v a l u e o f y f o r some p r e v i o u s v a l u e o f x  f r o m a g i v e n s e t o f i n d i v i d u a l v a l u e s o f x and y r e q u i r e s an assumption about the constancy If  of either  a i s assumed t o be an i n v a r i a b l e  back c a l c u l a t i o n Algebraically, Y  2  Y  1  The  -a  -a  =  a o r b. q u a n t i t y , the l i n e of  i s g i v e n by t h e d a s h e d l i n e  this X  i n Figure 1-la.  i s equivalent to establishing  the p r o p o r t i o n  2  XI  q u a n t i t y b becomes d i f f e r e n t  A l t e r n a t i v e l y , b may be c o n s i d e r e d  f o r each i n d i v i d u a l  as i n v a r i a b l e ,  case.  i n which  case  X figure  1 - 1 b  47  the dashed l i n e lation  i n Figure  1-lb represents  and t h e p r o p o r t i o n a l i t y  the l i n e  o f back  calcu-  is:  b  =  *2 " X i  or  Y  or  Yi - Y  2  - Y i = b(X - Xi) 2  2  = b(X! - X ) 2  The  value  of a i s different  f o r each i n d i v i d u a l  The  r e l a t i o n between f i s h w e i g h t  ( y ) and o t o l i t h  case. r a d i u s (x)  was f o u n d t o be o f t h e f o r m : £n y = a+b &nx I n as much as t h e r e was no i n c r e a s e increased a constant  i n variance  o f Un y w i t h  &n x, i t f o l l o w s t h a t t h e s e c o n d p r o c e d u r e o f a s s u m i n g slope  i s appropriate.  Hence t h e f o r m u l a  calculation: l nWi = £nW  2  - b (£nRi~  £nR ) 2  f o r back  48  APPENDIX 2  ANNOTATED PHOTOGRAPHS OF ONCORHYNCHUS NERKA OTOLITHS AS AN AID TO AGE AND GROWTH ANALYSIS  Otolith T h i s f i s h was b r a n d e d and w e i g h e d on J u n e 6, w e i g h e d a g a i n 30 d a y s l a t e r , and f i n a l l y and  s a c r i f i c e d 44 d a y s a f t e r  The e f f e c t s ring  initial  weighed  weighing.  of s t r e s s of handling are evident at  44 and 1 4 .  The r i n g s n e a r t h e edge a r e  s l i g h t l y obscured. the d i f f i c u l t i e s  (This problem i s accentuated  of printing  by  a high contrast  negative.) "A" shows an a r e a where two r i n g s come t o g e t h e r along  a f a u l t i n the o t o l i t h .  Finding  the nucleus  i n an o t o l i t h s u c h  as t h i s  involves considerable uncertainty. Prior  to ring  44, t h i s  f i s h was h e l d  populated  t a n k and f e d l e s s  reflected  i n the growth  regularly;  and u n d e r - g r o u n d  Under-ground.  Note the r i n g  by  rostrum  of glue  T h i s c a n be m i s t a k e n  inexperienced readers. o f both  otoliths  this i s  rings.  Over-clarified  the o t o l i t h .  i n a densely  seen  through  as a s t r e s s mark  Note a l s o t h a t the i s crystaline.  49  Under-ground "A" shows r i n g s l o s t due t o e x c e s s i v e in  clarification  glycerine.  The r i n g s a t "B" a r e d a i l y  rings.  "C".  would  The f i r s t  daily  ring  None o c c u r a t likely  near the arrow, a l t h o u g h i t i s d i f f i c u l t on t h i s  t o see i t  print.  Note the r e f r a c t i o n This  be f o u n d  is a fairly  p e r l y ground  and  The p r e f e r r e d  l i n e s near the edge.  typical  sockeye f r y o t o l i t h pro-  clarified.  r a d i u s would  start  a t the n u c l e u s  "A" . Zone "B" shows r i n g s t y p i c a l o f t h o s e f o u n d periods of irregular taken i n counting of  growth; extreme  c a r e must be  r i n g s s u c h as t h e s e b e c a u s e  t h e d a r k e r r i n g s c a n c o n t a i n two o r more  zones. tion  Counts  during  s h o u l d be done a t h i g h e r  t h a n t h e one shown.  each  daily  magnifica-  51  52  53  

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