~ h i s p a p e r n o t t o b e c i t e d w i t h o u t p r i o r r e f e r e n c e t o t h e a u t h o r s .
Symposium on
A c o u s t i c Methods i n F i s h e r i e s R e s e a r c h
No. 2 4
TARGET STRENGTH MEASUREMENTS OF FISH
Odd Nakken and K j e l l O l s e n I n s t i t u t e o f M a r i n e R e s e a r c h
B e r g e n , Norway
INTRODUCTION
Methods o f f i s h s i z i n g u s i n g t h e r e f l e c t e d sound s i g n a l f r o m i n d i - v i d u a l f i s h a r e d e v e l o p e d (CUSHING 1 9 6 8 , C R A I G and F.0RBES 1 9 6 9 , M I D T T U N 1 9 6 6 ) . I t h a s a l s o b e e n s u g g e s t e d t h a t d i s c r i m i n a t i o n b e t w e e n s p e c i e s m i g h t be p o s s i b l e by s t u d y i n g t h e c h a n g e i n t a r g e t s t r e n g t h when i n d i v i d u a l f i s h e s p a s s t h r o u g h t h e sound beam
(MIDTTUN a n d NAKKEN 1 9 7 1 ) . S e v e r a l w o r k e r s h a v e s t u d i e d t h e £ r e - q u e n c y r e s p o n s e s o f , f i s h i n o r d e r t o f i n d methods o f i d e n t i f i c a t i o n and s i z i n g (McCARTNEY a n d STUBBS 1 9 7 1 , HOLLIDAY 1 9 7 2 ) . So f a r , t h e r e l a t i o n s b e t w e e n t a r g e t s t r e n g t h , and f i s h s p e c i e s and s i z e a r e n o t s a t i s f a c t o r l y known t o e n a b l e u s d o i n g a c c u r a t e s i z i n g and r e l i a b l e i d e n t i f i c a t i o n a s r o u t i n e work a t s e a .
Due t o t h e c o m p l e x i t y o f f a c t o r s g o v e r n i n g t h e r e f l e c t i o n o f sound from f i s h i t i s i m p o s s i b l e t o c a l c u l a t e t h e s c a t t e r e d sound f i e l d . Thus t h e r e l a t i o n s b e t w e e n t a r g e t s t r e n g t h a n d f i s h p a r a m e t e r s
( i . e . s p e c i e s a n d s i z e ) h a v e t o b e e s t a b l i s h e d e m p i r i c a l l y a n d t h e r e a r e two ways o f a p p r o a c h . F i r s t , series o f t a r g e t s t r e n g t h
measurements c a n be made w i t h c a l i b r a t e d e c h o s o u n d e r s a t s e a p r o v i d e d t h a t t h e f i s h u n d e r o b s e r v a t i o n w i t h i n e a c h s e r i e s b e l o n g s t o o n l y one s p e c i e s and o f e q u a l s i z e , and t h a t r e p r e - s e n t a t i v e f i s h samples c a n b e c a u g h t . The t a r g e t s t r e n g t h
o b s e r v e d by t h i s method w i l l b e a n " a v e r a g e d o r s a l a s p e c t t a r g e t s t r e n g t h " d e p e n d i n g on t h e unknown a v e r a g e i n c l i n a t i o n d i s t r i - b u t i o n of t h e f i s h e s u n d e r o b s e r v a t i o n . Second, t h e t a r g e t s t r e n g t h c a n be measured u n d e r f u l l y c o n t r o l l e d c o n d i t i o n s i n l a b o r a t o r y e x p e r i m e n t s and s e v e r a l works o f t h i s k i n d have been r e p o r t e d (LOVE 1969 and 1 9 7 1 , HASLETT.1969, McCARTNEY and STUBBS 1971, M I D T T U N and HOFF 1962, SHIBATA 1 9 7 0 ) . But a s t h e a v e r a g e i n c l i n a t i o n o f t h e f i s h i n t h e f i e l d h a s n o t been con- s i d e r e d , t h e r e s u l t s from s u c h e x p e r i m e n t s m i g h t b i a s t h e
e s t i m a t e s of f i s h l e n g t h s made a t s e a ( M I D T T U N and NAKKEN 1 9 7 1 ) . To s u p p o r t s u c h measurements o b s e r v a t i o n s of f i s h i n c l i n a t i o n s s h o u l d t h e r e f o r e be a v a i l a b l e (OLSEN 1971, BARHAM 1970, BELTESTAD 1 9 7 3 ) .
I n o r d e r t o o b t a i n more knowledge o f t h e back s c a t t e r i n g prop- e r t i e s o f t h e f i s h s p e c i e s which a r e most commonly r e c o r d e d i n t h e n o r t h - e a s t e r n , A t l a n t i c , s t u d i e s of t a r g e t s t r e n g t h of i n d i - v i d u a l f i s h were c a r r i e d o u t d u r i n g summer 1971. The e x p e r i m e n t s were c a r r i e d o u t a t two f r e q u e n c i e s which a r e commonly used i n f i e l d work. I n t h i s p a p e r r e s u l t s o f t h e s e s t u d i e s a r e r e p o r t e d . E s t i m a t e s of t a r g e t s t r e n g t h which a r e t o be e x p e c t e d a t s e a a r e o b t a i n e d by combining t h e e x p e r i m e n t a l r e s u l t s w i t h f i e l d o b s e r - v a t i o n s of f i s h i n c l i n a t i o n .
MATERIAL AND METHODS
E x p e r i m e n t a l s e t - u p
The e x p e r i m e n t s were c a r r i e d o u t i n a s h e l t e r e d i n l e t which i s 2 0 0 m a c r o s s , 12
-
1 4 m d e e p w i t h s o f t bottom. The e x p e r i m e n t a l s e t - u p i s shown i n F i g . 1. An anchored r a f t c a r r i e d b o t h t h e l a b o r a t o r y and t h e accomodations f o r t h e s t a f f .The upward l o o k i n g t r a n s d u c e r s were mounted i n a h e a v i l y l o a d e d
s t e e l frame submerged from t h e r a f t i n a d j u s t a b l e w i r e s . The f i s h w e r e k e i t i n a n L p s i d e down p o s i t i o n i n t h e c e n t r a l p a r t of t h e sound beam by a frame o f t h i n n y l o n g u t . A s p e c i a l h o i s t i n g d e v i c e made i t p o s s i b l e t o hook t h e f i s h t o t h e frame a t t h e s u r f a c e and t h e n lower i t t o t h e m e a s u r i n g p o s i t i o n a t
The a s p e c t o f t h e f i s h c o u l d be c o n t i n u o u s l y changed i n two ways, t i l t i n g and r o l l i n g , w i t h o u t any change of hooking.
The f i s h was t i l t e d by o p e r a t i o n of t h e a u t o m a t i c " t i l t i n g b a r "
between -45O and +45O of h o r i z o n t a l p o s i t i o n w i t h
-
+ l a c c u r a c y . OThe t i l t i n g s p e e d was lo p e r second. When o n l y t i l t v a r i a t i o n s were
I wanted a s t a b l e u p s i d e down p o s i t i o n was o b t a i n e d by s m a l l f l o a t s a t t a c h e d t o t h e f i s h b e l l y . When r o l l v a r i a t i o n s a l s o were wanted t h i n n y l o n g u t s from t h e f i s h s i d e s t o a s m a l l wheel r e p l a c e d
t h e f l o a t s . The wheel was o p e r a t e d manually and worked normal t o t h e " t i l t i n g b a r " . For c o m p l e t e change of a s p e c t t h e f i s h was h a u l e d t o t h e s u r f a c e and t h e p o i n t s of hookinq changed.
I n s t r u m e n t a t i o n and d a t a r e c o r d i n g
I A b l o c k d i a g r a m of t h e i n s t r u m e n t a t i o n i s shown i n F i g . 2 . Two e c h o s o u n d e r working a t f r e q u e n c i e s 38 kHz and 120 kHz (Simrad E k 38 A and.Ek 120 A ) and w i t h t r a n s d u c e r s 1 0 x 10 c m and 5 cm i n d i a m e t e r , were u s e d . The t r a n s m i t t e d p u l s e l e n g t h s , measured a t h a l f t h e a m p l i t u d e , w e r e 0.6 m i l l i s e c o n d f o r b o t h s o u n d e r s .
i
I l The r e p e t i o n r a t e o f t h e s o u n d e r s were i n c r e a s e d t o 4 p u l s e s p e r I l second. For m e a s u r i n g and r e c o r d i n g of d a t a a two c h a n n e l
o s c i l l o s c o p e ( H e w l e t t P a c k a r d , 1 4 1 A ) , a n echo i n t e g r a t o r (Simrad e c h o i n t e g r a t o r , QM) w i t h a two c h a n n e l r e c o r d e r ( H e w l e t t P a c k a r d , 7702 B ) and a p o l a r p l o t l e v e l r e c o r d e r
r ru el
and Kjær 2304) were u s e d . One o f t h e c h a n n e l s o f b o t h t h e o s c i l l o s c o p e and t h ei n t e g r a t o r r e c o r d e r was u s e d f o r p r e s e n t a t i o n o f e c h o a m p l i t u d e s , on t h e o t h e r c h a n n e l t h e c o r r e s p o n d i n g t i l t a n g l e s were r e c o r d e d . A f i l m camera t r i g g e r e d by t h e e c h o s o u n d e r s was a t t a c h e d t o t h e
I
o s c i l l o s c o p e .A hydrophone (LC 32, A t l a n t i c R e s . ) was u s e d f o r c a l i b r a t i o n o f t h e
equipment. In-add-ition, a d a i l y c a l i b r a t i o n was c a r r i e d o u t by m e a s u r i n g t h e t a r g e t s t r e n g t h o f a r i g i d s t e e l s p h e r e , 5 cm i n d i a m e t e r , which was lowered i n t o t h e m e a s u r i n g p o s i t i o n .
During one m e a s u r i n g program t h e t i l t i n g b a r s t a r t e d from h o r i - z o n t a l p o s i t i o n , moved t o +45O, back a g a i n t h r o u g h t h e h o r i z o n t a l t o -45' and t h e n back t o O . During t h e f i r s t q u a r t e r of t h i c c y c l e s u i t a b l e g a i n s e t t i n g s were s e l e c t e d . The d a t a c o l l e c t e d d u r i n g t h e c o m p l e t e h a l f c y c l e between +45O and -45O was u s e d f o r f u r t h e r t r e a t m e n t . F i g . 3 shows examples o f r e c o r d i n g s .
The f i s h was s t u n n e d o r k i l l e d by h i t t i n g t h e f r o n t a l p a r t of t h e b r a i n by a s h a r p t o o l . When s u s p e n d i n g t h e f i s h , c a r e was t a k e n t o a v o i d a i r i n g i l l s and stomach. The measurements were s t a r t e d i m m e d i a t e l y a f t e r t h e f i s h had been lowered i n t o t h e m e a s u r i n g p o s i t i o n . I n o r d e r t o o b t a i n n e c e s s a r y i n f o r m a t i o n a b o u t t h e r e l a t i o n between t a r g e t s t r e n g t h c h a r a c t e r i s t i c s o f dead and l i v e f i s h , a few f i s h were a l s o measured a l i v e . They were t h e n a l l o w e d t o move t h e i r t a i l and body w i t h o u t c h a n g i n g t h e i r p o s i t i o n s w i t h i n t h e sound beam.
Data p r o c e s s i n g
The r e c o r d e d d a t a c o n s i s t i n g o f c o r r e s p o n d i n q v a l u e s o f v o l t a g e , V , and tilt a n g l e , q , ( F i g . 3 ) were t r a n s f e r r e d t o p u n c h c a r d s .
The amount o f d a t a punched from e a c h o b s e r v a t i o n s e r i e s were
1
l a r g e enough t o e n s u r e s u f f i c i e n t r e p r o d u c t i o n of t h e d i a g r a m s . The c a l c u l a t i o n s of t a r g e t s t r e n g t h , TS, were done by computer from e q u a t i o n
where V i s t h e o b s e r v e d v o l t a g e , V, i s t h e v o l t a g e from t h e
r e f e r e n c e s p h e r e and TSr i s t h e t a r g e t s t r e n g t h of t h e r e f e r e n c e s p h e r e i n d e c i b e l ( d B ) . The t h e o r e t i c a l v a l u e o f T S r i s - 3 8 . 1 dB w h i l e t h e measured v a l u e s u s i n g t h e d a t a o b t a i n e d by hydrophone c a l i b r a t i o n , were -38.0 dB and -38.5 dB f o r t h e 38 kHz and t h e 120 kHz e c n o s o u n d e r s r e s p e c t i v e l y . When computing T S , t h e measured v a l u e s o f TSr were u s e d .
A s a f i r s t s t e p i n t h e a n a l y s i s o f t h e m a t e r i a 1 , ~ o u t p r i n t s o f t h e f o l l o w i n g p a r a m e t e r s from e a c h f i s h , s p e c i e and a s p e c t were made:
No : F i s h r e f e r e n c e number L : F i s h l e n g h t ( i n cm)
TSmax : Maximum o b s e r v e d t a r g e t s t r e n g t h ( d ~ )
e/
: T i l t a n g l e ,q ,
( i n d e g r e e s ),
a t TSmaxCf) i s n e g a t i v e f o r head down, p o s i t i v e f o r head up.
FV1 : i n t e r v a l o f 9 w i t h i n which TS
1
TSmax -6 dB FV2 : I n t e r v a l o f q w i t h i n which TS
2
TSmax - 1 0 d ~\ FV3 : I n t e r v a l o f
<p
w i t h i n which TS 1 T S -20 d ~ ~ ~ ~ n 1 : T o t a l number of l o b e s where T S Z TSmax -6 dB n2 : T o t a l number o f l o b e s where TS 2 TSmax -10 dB n 3 : T o t a l number o f l o b e s where TS Z T S m a x -20 dB: Mean a m p l i t u d e w i t h i n FV1 v2 : Mean a m p l i t u d e w i t h i n FV2 v3 : Mean a m p l i t u d e w i t h i n FV3
A : Running mean o f a m p l i t u d e s , c a l c u l a t e d from t h e f ormula
1 i 6
t o +45O. A was p r i n t e d o u t f o r = - 2 1 , -15, - 9 , - 3 , 3 , 9; 1 5 and 21°.
A3 ,: Maximum v a l u e o f A F I : V a l u e o f
q
when A = A 3A l i n e a r r e l a t i o n between maximum d o r s a l a s p e c t t a r g e t s t r e n g t h (Tsmax ) and f i s h l e n g t h ( L )
TSmax = m l o g
10 L + b (11)
was assumed t o e x i s t f o r e a c h s p e c i e s and f r e q u e n c y and t h e c o e f f i - c i e n t s m and b w e r e c a l c u l a t e d by a l e a s t mean s q u a r e r e g r e s s i o n a n a l y s e s .
TS
-
l e n g t h r e l a t i o n s , t a k i n g i n t o a c c o u n t d i s t r i b u t i o n of t h e t i l t a n g l e , Q , which have been o b s e r v e d a t s e a , ( f i g . 4 ) were computed f o r two s p e c i e s , cod and h e r r i n g , a t 38 kHz. The f o l l o w i n g f o r m u l a s were u s e d :s e a
' s e a
and TSsea = 2 0 l o g
- +
TCrv
r1
where k 4 i s t h e f r e q u e n c i e s g i v e n i n F i g . 4 ( i n 6 d e g r e e c l a s s e s o f q ) and A
YJ
i s t h e a m p l i t u d e of t h e J - t h f i s h a t t i l t a n g l eg
b v e r a g e d i n 6 d e g r e e c l a s s e s o f c p ) , n i s t h e number of f i s h i n e a c h i n v e s t i g a t e d l e n g t h g r o u p ( T a b l e 1 ) .
RESULTS
The o b s e r v a t i o n s and t h e r e s u l t s o f t h e l e a s t mean s q u a r e r e g r e s s i o n (TSmax=nlog10 L
+
b ) a r e shown i n T a b l e 2 and Fig: SA-E. I ta p p e a r s t h e - r e g r e s s i o n l i n e s f o r c o d , s a i t h e and p o l l a c k a l m o s t c o i n c i d e , w h i l e t h o s e f o r s p r a t and h e r r i n g a r e d i f f e r e n t . The two l a t t e r s p e c i e s h a v i n g lower maximum d o r s a l a s p e c t t a r g e t s t r e n g t h s t h a n t h e g a d o i d s . The o t h e r measured f i s h were e i t h e r t o o few i n number or t h e l e n g t h r a n g e was t o narrow f o r a p p l y i n g a l e a s t mean s q u a r e r e g r e s s i o n and t h e r e s u l t s f o r t h e s e f i s h a r e shown i n T a b l e 3 and F i g . 5F. I n F i g . SF t h e r e g r e s s i o n l i n e f o r cod i s shown f o r c o m p a r i s o n . The maximum d o r s a l a s p e c t t a r g e t s t r e n g t h of t h e s e s p e c i e s i s a p p r o x i m a t e l y 1-3 dB l e s s t h a n t h a t o f c o d , e x c e p t f o r m a c k e r e l , d o g f i s h and prawns which a l l show c o n s i d e r a b l y lower v a l u e s . The mean v a l u e s o f m a c k e r e l a r e 1 0 - 1 1 dB lower t h a n - t h o s e o f cod, and 3-4 dB lower t h a n f o r h e r r i n g .
The TS-length r e l a t i o n s which a r e t o be e x p e c t e d a t s e a , a t 38 kHz, a p p l y i n g t h e d i s t r i b u t i o n s o f tilt a n g l e (Fig.. 4 ) t o a l l l e n g t h
g r o u p s a r e shown i n F i g . 6 f o r cod and h e r r i n g . F i g . 6A shows t h a t t h e e x p e c t e d mean v a l u e o f a t a r g e t s t r e n g t h d i s t r i b u t i o n of cod
w i l l b e 8-9 dB lower t h a n t h e c o r r e s p o n d i n g maximum v a l u e s . The r e s u l t s a r e compared w i t h t h e f i e l d 0bs.ervation.s made by M I D T T U N and NAKKEN (1971). Assuming a l l f i s h o b s e r v e d t o be h o r i z o n t a l , t h e e x p e c t e d TS-length r e l a t i o n f o r o b s e r v a t i o n s w i t h a 6O t r a n s d u c e r beamwidth w i l l 'be a s i n d i c a t e d by l i n e I1
( F i g . 6A). For h e r r i n g t h e e x p e c t e d t a r g e t s t r e n g t h s a t s e a w i l l b e 6 dB lower t h a n t h e c o r r e s p o n d i n g maximum v a l u e s ( F i g . 6 B ) . The d i f f e r e n c e s between day and n i g h t v a l u e s a r e i n s i g n i - f i c a n t . The r e l a t i v e l y s m a l l d i f f e r e n c e between t h e e x p e c t e d and t h e maximum o b s e r v e d v a l u e s o f TS a t s m a l l f i s h l e n g t h s
( F i g . 6A), i s c a u s e d by t h e l e s ; d i r e c t i v i t y o f s m a l l f i s h e s .
,
The e f f e c t of swimming on t a r g e t s t r e n g t h i s shown i n F i g . 7 and F i g . 8 . The movements o f t h e f i s h i n t r o d u c e d a v a r i a t i o n i n t a r g e t s t r e n g t h and t h i s v a r i a t i o n i n c r e a s e s w i t h i n c r e a s i n g swimming a c t i v i t y . NO s i g n i f i c a n t change i n mean v a l u e s of t a r g e t s t r e n g t h c a n be s e e n ( T a b l e 4 ) . F i g . 8 i n d i c a t e s a p e r i o d i c r e l a t i o n between t a r g e t s t r e q t h a n d t a i l b e a t .I n T a b l e 5 i s shown a comparison between t h e maximum d o r s a l and maximum s i d e a s p e c t t a r g e t s t r e n g t h . None of t h e s p e c i e s o b s e r v e d
shows a l a r g e r mean d i f f e r e n c e t h a n 4 dB and s i g n i g i c a n t d i f f e r e n c e s a r e o b t a i n e d o n l y f o r c o d , h e r r i n g and s p r a t . F i g . 9 which
p r e s e n t s t a r g e t s t r e n g t h a s a f u n c t i o n of r o l l a n g l e i n d i c a t e s , however, t h a t t h e cod may have c o n s i d e r a b l y lower t a r g e t s t r e n g t h s i a t r o l l a n g l e s l a r g e r t h a n a p p r o x i m a t e l y 30°.
The r e l a t i o n between mean v a l u e s o f maximum d o r s a l a s p e c t t a r g e t s t r e n g t h i n e a c h l e n g t h g r o u p and t h e a n g l e between 6 dB p o i n t s i n t h e d i r e c t i v i t y p a t t e r n ( F i g . 3 ) i s shown i n F i g . 1 0 . The t h r e e c u r v e s a r e s i g n i f i c a n t l y s e p a r a t e d and t h e o b t a i n e d v a l u e s c o r - r e s p o n d s t o t h e f i e l d o b s e r v a t i o n s made by M I D T T U N and NAKKEN
( 1 9 7 1 ) .
A comparison o f a l l t h e o b s e r v e d t a r g e t s t r e n g t h s f o r t h e two f r e q u e n c i e s a p p l i e d i s madein F i g . 11, where a l s o a f r e q u e n c y d i f f e r e n c e o f 2 . 4 dB ( d e r i v e d from e q u . TS = 24.5 l o g l 0 L
-
4 . 5l o g n - 26.4, McCARTNEY and STUBBS 1971) i s i n d i c a t e d . F i g . 11
i n d i c a t e s t h a t t h e d i f f e r e n c e i n t a r g e t s t r e n g t h between 38 kHz and 1 2 0 kHz v a r i e s w i t h t h e magnitude of t a r g e t s t r e n g t h
( f i s h l e n g t h ) . DISCUSSION
The s l o p e s of t h e r e g r e s s i o n l i n e s a t 38 kHz f o r c o d , s a i t h e and p o l l a c k a r e i n a c c o r d a n c e w i t h t h e r e s u l t s r e p o r t e d by McCARTNEY and S T U B B S . ( ~ ~ ~ ~ ) . So i s a l s o t h e s l o p e a t 1 2 0 kHz f o r c o d , w h i l e t h e l i n e s f o r s a i t h e and p o l l a c k a t t h i s f r e - quency show s m a l l e r s l o p e s , comparable t o t h e f i n d i n g of LOVE
( 1 9 7 1 ) . The d i f f e r e n c e i n t h e s l o p e s between 38 kHz and 120 kHz f o r p o l l a c k may, however, n o t be s i g n i f i c a n t s i n c e t h e
l e n g t h r a n g e of t h e o b s e r v e d f i s h e s a r e narrow and t h e v a r i a t i o n from specimen t o specimen i s l a r g e . The s l o p e s f o r h e r r i n g and s p r a t a r e b o t h s m a l l e r t h a n t h o s e found f o r t h e g a d o i d s p e c i e s . The a p p e a r e n t d i f f e r e n c e between h e r r i n g and s p r a t a r e n o t
s i g n i f i c a n t and t h e d a t a c o u l d p r o b a b l y have been t r e a t e d a s from one s p e c i e s , r e s u l t i n g i n s l o p e s o f a p p r o x i m a t e l y 16.0 and 20.5 dB/decade a t 38 and 1 2 0 kHz r e s p e c t i v e l y . F o r f i s h e s o f l e n g t h s 6 - 1 2 cm t h e d o r s a l a s p e c t t a r g e t s t r e n g t h s o f g a d o i d s a n d c l u p e o i d s a r e a p p r o x i m a t e l y e q u a l . F o r b i g g e r f i s h t h e d o r s a l a s p e c t t a r g e t s t r e n g t h of t h e c l u p e o i d s w i l l b e lower a s compared t o t h e g a d o i d s , t h e d i f f e r e n c e between a 35 cm cod and a 35 cm h e r r i n g b e i n g 7-8 dB. T a b l e 5 shows t h a t t h e s i d e a s p e c t t a r g e t s t r e n g t h o f cod i s 4 dB lower t h a n t h e d o r s a l a s p e c t t a r g e t
s t r e n g t h w h i l e h e r r i n g seem t o have a 3.5 dB d i f f e r e n c e t h e
o p p o s i t e way ( 3 8 kHz). T h i s i n d i c a t e s t h a t h e r r i n g and cod have a p p r o x i m a t e l y e q u a l s i d e a s p e c t t a r g e t s t r e n g t h s and c o n s e q u e n t l y a r e e q u a l a s t a r g e t s f o r h o r i z o n t a l working s o n a r s .
The s m a l l d i f f e r e n c e between t h e c a l c u l a t e d v a l u e s which a r e t o be e x p e c t e d a t s e a and t h e f i e l d o b s e r v a t i o n of t a r g e t s t r e n g t h s
( F i g . 6A) a r e w e l l w i t h i n t h e l i m i t s o f c a l i b r a t i o n a c c u r a c y . However, a s b o t h t h e f i e l d o b s e r v a t i o n s o f t a r g e t s t r e n g t h and t h e d a t a on t i l t a n g l e d i s t r i b u t i o n a r e o b t a i n e d on spawning cod good agreement s h o u l d be e x p e c t e d . L i n e I11 i n F i g . 6 i s b a s e d
on t h e a s s u m p t i o n t h a t a l l k n g t h g r o u p s have e q u a l t i l t a n g l e d i s t r i b u t i o n s . To what e x t e n t t h i s h o l d s good i s n o t known a s d a t a on t i l t a n g l e d i s t r i b u t i o n a c c o r d i n g t o l e n g t h , s p e c i e s and s e a s o n i s l a c k i n g .
F i g . 6 B shows t h a t t h e change i n t h e e x p e c t e d t a r g e t s t r e n g t h s f o r h e r r i n g from day t o n i g h t was i n s i g n i f i c a n t , a l t h o u g h b o t h t h e mean and t h e s p r e a d o f t h e tilt a n g l e d i s t r i b u t i o n s changed from d a y t o n i g h t . AS t h e mean v a l u e of t h e day o b s e r v a t i o n s o f tilt a n g l e i s much c l o s e r t o t h e a n g l e o f maximum d o r s a l a s p e c t t a r g e t s t r e n g t h t h a n t h e mean of t h e n i g h t o b s e r v a t i o n s
( F i g . 4 ) , t h i s w i l l cornpensatethe i n c r e m e n t i n s p r e a d from n i g h t t o d a y .
C o n s i d e r i n g F i g . 6 i t i s s e e n t h a t c h a n g e s o f tilt a n g l e d i s t r i - b u t i o n s b o t h f o r h e r r i n g andcod may have c o n s i d e r a b l e e f f e c t s on mean v a l u e s o f t a r g e t s t r e n g t h . T h i s i s a m a t t e r which c a n l e a d
t o c e r i o u c e r r o r s b o t h i n s i z i n g and abundance e s t i m a t i o n . R e l i a b l e e s t i m a t e s of t a r g e t s t r e n g t h s of i n d i v i d u a l f i s h a t s e a c a n o n l y be o b t a i n e d when t h e f i s h e s a r e s c a t t e r e d
.
Whens u c h e s t i m a t e s a r e u s e d t o c a l c u l a t e d e n s i t i e s of s c h o o l i n g f i s h t h e d e n s i t y e s t i m a t e s w i l l be c o r r e c t i f t h e t i l t a n g l e d i s t r i - b u t i o n a r e e q u a l f o r s c a t t e r e d and s c h o o l i n g f i s h . I f n o t , l a r g e e r r o r s m i g h t be i n t r o d u c e d . More i n f o r m a t i o n on tilt a n g l e
d i s t r i b u t i o n s r e l a t e d t o t h e d e n s i t y of f i s h c o n s e n t r a t i o n s w i l l t h e r e f o r e improve t h e abundance e s t i m a t i o n w i t h a c o u s t i c e q u i p m e n t . I t i s i m p o r t a n t t o know i f t h e t a r g e t s t r e n g t h o b s e r v a t i o n s made on s t u n n e d o r dead f i s h a r e v a l i d f o r f r e e swimming i n d i v i d u a l s . I n t h e e x p e r i m e n t s done w i t h l i v e f i s h , t h e body movements of t h e f i s h were o b s e r v e d t o be s i m i l a r t o f r e e swimming f i s h . Most o f t h e r e c o r d i n g s were o b t a i n e d when t h e f i s h had a swimming a c t i v i t y comparable t o a " f a s t c r u i s i n g " , s i t u a t i o n . F o r p u r p o s e s o f
s i z i n g , i d e n t i f i c a t i o n and abundance e s t i m a t i o n , t h e a v e r a g e v a l u e of t a r g e t s t r e n g t h i s t h e i m p o r t a n t p a r a m e t e r . Although
t h e f i s h o b s e r v e d ( T a b l e 4 ) a r e t o o few f o r s a f e c o n c l u s i o n s , t h e r e were no i n d i c a t i o n s t h a t t h e o b s e r v e d p e r i o d i c t a r g e t s t r e n g t h
v a r i a t i o n i n f l u e n c e d t h e mean v a l u e s i g n i f i c a n t l y . What seems
p r o b a b l e , however, i s an i n c r e a s e d v a r i a n c e on t a r g e t s t r e n g t h d u e t o swimrning. T h i s i s p a r t i q u l a r l y c l e a r f o r t h e o b s e r v e d s a i t h e . The q u e s t i o n o f why t h e t a r g e t s t r e n g t h v a r i a t i o n s
( F i g . 8 ) seems r e l a t e d t o e a c h t a i l b e a t c y c l e and n o t t o e a c h h a l f c y c l e , c a n n o t b e answered from t h e s e i n v e s t i g a t i o n s . The r e l a t i o n between maximurn d o r s a l a s p e c t t a r g e t s t r e n g t h and t h e a n g l e between t h e 6 dB p o i n t s i n t h e d i r e c t i v i t y p a t t e r n
( F i g . 1 0 ) show s i g n i f i c a n t d i f f e r e n c e s between t h e 3 s p e c i e s ( c o d , s a i t h e and h e r r i n g ) when t h e mean v a l u e s a r e c o n s i d e r e d . The v a l u e s f o r l a r g e cod and c o a l f i s h a r e i n c l o s e agreement w i t h t h e f i e l d o b s e r v a t i o n s made by M I D T T U N and NAKKEN ( 1 9 7 1 ) . The v a r i a t i o n s from specimen t o specimen w i t h i n t h e same s p e c i e s a r e , however, l a r g e and a s i m i l a r p l o t t o t h a t o f F i g . 1 0 o f i n d i v i d u a l f i s h would show a l a r g e d e g r e e o f o v e r l a p . M I D T T U N and NAKKEN ( 1 9 7 1 ) s u g g e s t t h a t s u c h p l o t s m i g h t be used f o r
i d e n t i f i c a t i o n a c c o r d i n g t o s p e c i e s . F i g . 10 i n d i c a t e s t h a t t h i s s h o u l d be w i t h i n r e a c h f o r t h e 3 s p e c i e s u n d e r c o n s i d e r a t i o n , when t h e y a r e unmixed. When mixed r e c o r d i n g s o c c u r , i t . w i l l p r o b a b l y be e x t r e m e l y d i f f i c u l t o r i m p o s s i b l e t o d i s c r i m i n a t e between s p e c i e s by t h i s method.
F i g . 10 shows a l s o t h a t t h e d o r s a l a s p e c t t a r g e t s t r e n g t h o f i n d i v i d u a l cod a t 3 8 kHz d e c r e a s e l e s s w i t h t i l t a n g l e t h a n f o r s a i t h e and h e r r i n g . T h i s means t h a t v a r i a t i o n i n t i l t a n g l e
d i s t r i b u t i o n s m i g h t l e a d t o l a r g e r e r r o r s i n s i z i n g and abundance
1
e s t i m a t i o n f o r t h e two l a t t e r s p e c i e s t h a n f o r co&. F o r s m a l l f i s h ( l o w L / A ) , c h a n g e s i n t i l t a n g l e a r e o f l e s s i m p o r t a n c e f o r a l l 3 s p e ' c i e s , d u e t o t h e r e l a t i v e l y low d i r e c t i v i t y of s m a l l f i s h . The d a t a p l o t i n F i g . 11 w i l l f i t a s t r a i g h t l i n e r e l a t i o n s h i p
( k l o g h l where k i s a c o n s t a n t ) a t t a r g e t s t r e n g t h s below - 3 0 dB I
I
i f t h e m a c k e r e l i s e x c l u d e d . The c u r v e d s h a p e of t h e p l o t I
i
c o n s i d e r i n g a l l o b s e r v a t i o n s , a r e p r o b a b l y c a u s e d by t h e f a c t t h a t m e r e l y a l l o u r d a t a a r e ~ ~ t h i n t h e r e g i o n o f i n t e r f e r e n c e e f f e c t s .
ACKNOWLEDGEMENTS
--
The authors wish to thank I. HOFF who was responcible for the
l
electronics and together with W. LØTVEDT and J. VESTNES tookI
part in the data collecting; G. VESTNES and A. STORLER whol
wer.e of invaluable help during the planning and preparation of the work; P , EIDE and G. HELLE who did the programming;B. BRIGTSEN, B. BRYNHILDSEN, H. GILL, S. MYKLEVOLL and A. RAKNES
I
who all contributed during the analyses and preparation of the manuscript.REFERENCES
BARHAM, E.G. 19.?0. Deep-Sea Fishes: lethargy and vertical orienta- tion. In Proceedings of an international symposium
-
onbiological sound scattering in the ocean, Warrenton, Virginia: 100-118 (Ed. G. Brooke Farquhar)
.
BELTESTAD, A.K. 1973. Beiteadferd og vertikalvandring hos O-gruppe sild (Clupea harengus L.) i relasjon til lysintensitet.
Thesis, .Univ. of Bergen. L1n
CUSHING, D.H. 1968. Direct estimation of a fish population acousti- cally. J.Fish.Res.Bd.Canada, 25(11): 2349-2364.
/ \ CRAIG, R.E. and FORBES, S. 1969. A sonar for fish counting. FiskDir.
Skr.Ser.HavUnders., 15: 210-219.
HASLETT, R.W.G. 1969. The target strengths of fish. J.Sound.Vib.
9: 181-191.
-
HOLLIDAY, D.V. 1972. Resonance structure in echoes from schooled pelagic fish. J.Acoust.Soc.Am.,51: 1322-1332.
LOVE, R.H. 1969. Maximum side-aspect target strength of an indi- vidual fish. J.Acoust.Soc.Am., 46: 746-752.
LOVE, R.H. 1971. Dorsal aspect target strength of an individual fish. J.Acoust.Soc,Am., 49: 816-823.
McCARTNEY, B.S. and STUBBS, A.R. 1971. Measurements of the acoustic target strengths of fish in dorsal aspect,
including swimbladder resonance. J.Sound Vib. 15(3):397-420.
MIDTTUN, L. 1966. Note on measurement of target strength of fish at sea. Coun;Meet.int.Coun.Explor.Sea, 1966 (F9): 1-3.
[ ~ i m e ~ g
MIDTTUN, L. and HOFF. I. 1962. Measurements of the reflection of sound by fish. FiskDir.Skr.Ser.HavUnders., 13(3): 1-18.
1
MIDTTUN, L. and NAKKEN, 0. 1971. On acoustic identification, sizing and abundance estimation of fish. FiskDir.Skr.Ser.
HavUnders., 16: 36-48.
OLSEN, K. 1971. Orientation measurements of cod in Lofoten obtained from underwater photographs and their relation to target strength. Coun.Meet.int.Coun.Explor.Sea, 1971 (B17): 1-8.
[~imeoJ
SHIBATA, K. 1970. Study on details of ultrasonic reflection from individual fish. Bull.Fac.Fish.Nagasaky Univ. 29: 1-82.
Table 1. Length distribution of observed fish.
Species
Length groups, cm
5- 8- 11- 14- 17- 20- 25- 30- 40- 50- 60- 70- 80- 90- 100- Total 7 10 13 16 19 24 29 39 49 59 69 79 89 99 130
God Saithe Pollack Mackerel
Herring 9 11 1 1 1 7 2 41
Sprat 3 7 1 0 7 2 29
Haddock 2 12 1 15
Blue whiting 10 10
Whiting 6 2 8
Spiny dogfish Wrasse
Ballan Wrasse Trout
Horse mackerel 1
~umpsucker 1 1
Poor cod 1 1
Prawn 1 1 2
Total 8 26 37 15 6 35 39 108 15 28 19 2 2 1 2 . 343 ,
F-l W
1 4
T a b l e 2 . C a l c u l a t e d s l o p e (m) a n d c o n s t a n t ( b ) of r e q r e s s i o n l i n e s . TS = m Logl0 L
+
b , t h e c o r r e l a t i o n c o e f f i c i e n t r , t h e s t a n d a r d e r r o r s a n d t h e n u m b e r o f f i s hY X
m e a s u r e d N .
S p e c i e s F r e q . N c m b r s
Y X
kHz d B d B d B
C o d 3 8 7 3 2 4 . 5 - 6 6 . 6 O . 9 7 2 2 . 0 2
1 2 0 7 2 2 4 . 6 - 6 7 . 6 0 . 9 5 5 2 . 2 8
S a i t h e 3 8 6 8 2 3 . 3 - 6 4 . 9 0 . 9 7 5 1 . 4 4
1 2 0 6 8 2 0 . 1 - 6 0 . 1 O . 9 4 8 1 . 8 5
P o l l a c k 3 8 4 6 2 2 . 7 - 6 5 . 5 0 . 8 7 9 1 . 5 0
1 2 0 4 6 1 7 . 5 - 5 6 . 4 0 . 7 5 4 1 . 8 6
H e r r i n g 3 8 3 8 1 3 . 6 - 5 6 . 8 0 . 8 5 1 1 . 5 1 1 2 0 4 1 1 8 . 8 - 6 2 . 4 O . 8 9 0 1 . 7 9
S p r a t 3 8 2 9 1 7 . 2 - 6 0 . 8 0 . 7 8 4 1 . 6 6
1 2 0 2 9 2 1 . 4 - 6 6 . O 0 . 8 1 9 1 . 8 3
T a b l e 3 . Mean v a l u e s
(E)
a n d s t a n d a r d d e v i a t i o n s ( S t . d e v . ) of t a r g e t s t r e n g t h a c c o r d i n g t o l e n g h t ( L ) . N i s t h e n u m b e r of f i s h m e a s u r e d .S w e c i e s F r e q . N L TS S t . d e v .
M a c k e r e l 3 8 1 6 2 9
-
3 4 - 4 0 . 3 2 . 7Il 2 3 3 5
-
4 1 - 3 8 . 6 3 . 01 2 0 1 6 2 9
-
3 4 - 4 1 . 9 4 . 0I! . 2 2 3 5
-
4 1 - 4 0 . 6 3 . 6Horse 3 8 1 3 3 - 3 4 . 0
m a c k e r e l 1 2 0 1 3 3 - 3 0 . 9
-
H a d d o c k 3 8 1 3 2 8
-
3 8 - 3 2 . 1 1 . 8Il 1 4 8 - 2 8 . O
-
i 1 2 0 1 4 2 8
-
3 8 - 3 0 . 7 1 . 5Il 1 4 8 - 2 7 . 6
-
B l u e 3 8 1 0 3 1
-
3 5 - 3 2 . 0 1 . 8w h i t i n g 1 2 0 9 3 1
-
3 5 - 3 3 . 3 2 . 7W h i t i n g 3 8 4 2 1
-
2 2 - 3 5 . 4 0 . 4Il 1 2 8 - 3 2 . 2
-
Il 2 3 8 , 3 8 - 3 2 . 3 1 . 9
1 2 0 5 2 1
-
2 2 - 3 2 . O 1 . 911 1 2 8 - 3 0 . 8
-
Il 2 3 8 / 3 8 - 2 9 . 5 0 . 7
S p i n y 3 8 3 8 1 , 1 2 0 , 1 2 0 - 2 2 . 8 O . 4
d o g f i s h 1 2 8 3 8 1 , 1 2 0 , 1 2 0 - 2 2 . 1 4 . 2
P r a w n
B a l l a n 3 8 2 1 9 , 2 0 - 3 6 . 8 0 . 1
Wrasse 1 2 0 2 1 9 , 2 0 - 3 5 . 5 O . 5
Wrasse 3 8 2 1 7 , 2 4 - 3 6 . O 2 . 0
1 2 0 2 1 7 , 2 4 - 3 5 . O 2 . 5
T r o u t
L u m p s u c k e r P o o r cod
TabPe 4 . C o r r e s p o n d i n g d o r s a l a s p e c t t a r g e t s t r e n g t h (TS, dB) of swimming and dead f i s h . The swimming f i s h a r e t measured a t t i l t a n g l e s o f maximum o b t a i n a b l e T S .
S p e c i e s , Mean TS Max T s o f t h e C o r r e s p . TS of l e n g t h i n d u r i n g s w i m . a r e s p . f i s h o b s . t h e TS-length r e l .
cm (95% c o n f . l i m . ) a s dead ( 9 5 % c o n f . l i m . )
Cod, 59 -24.3 ( + 5 . 0 ) -24.6
( - 3 . 2 )
Cod, 69 -23.5 ( + 3 . 5 ) -23.9
( - 2 . 5 )
S a i t h e , 53 - 2 7 . 0 ( + 8 . 0 ) Not o b s . - 2 4 . 5 ( I 2 . 8 ) ( - 2 . 5 )
Table 5. Mean values (ATS) and standard deviations (st.dev.) of the di£ference between maximum dorsal and maximum side aspect target strength (ATS). N is the number of fish measured.
Species Freq. N
0 3
St.dev.kHz dB dB
Cod 3 8 7 4 . 0 3 . 0
1 2 0 7 2 . 5 2 . 6
Saithe 3 8 2 9 0.1 1 . 6
Herr ing 3 8 6 - 3 . 5 1 . 6
1 2 0 6 -1.2 2.2
Sprat 3 8 4 - 3 . 0 1 . 0
1 2 0 4 -2.8 2.5
Mackerel 3 8 6 - 1 . 5 2 . 9
1 2 0 5 -3.0 1 . 6
Fig. 1. Experimental set up. 1) Fish suspension,
2) hoisting system, 3 ) tilting system, 4) trans- ducers, 38 KHz and 120 KHz, 5) transducer base and 6 ) raft.
- - - w - - - -
)
T r a n s d u c e r s
Integrator
i-l
R e c o r d e r
I
C a m e ra
Hydrophone
Freq.
I L 1
At tenuator
I
meter
ta
Fig. 2. Block diagram of instrumentatbon.
ni (D
O rt
Y ul
Pi c+
Q Y (D O 3
a s ri-
-
,b rTARGET STRENGTH, dB
I I I I I l I I I
* W W W W W N N N
m
TARGET STWENGSW. d B
HEAD DOWN
T I L T ANGLE, DEGREES HEAD UP
Fig. 4. Distribution of field observations of tilt angle. 1) Cod (mean length 80 c m ) , day and night (OLSEN 1971), 2 ) herring (mean length 13 cm), night and 3) herring, day (BELTESTAD 1973).
I I 1 I I I I
- 5 0
-
L5 - L O - 3 5 - 3 0 - 2 5 - 2 0TARGET CTRENGTH, dB
L
l I l I I I I
- 5 0 - L5 - L O - 3 5 - 3 0 - 2 5 - 2 0
TARGET STRENGTH, d B
Fig. 54. Observations of maximum dorsal aspect target strength on cod and the regression line.
,
I l I I I I
- 35 - 30 - 25 - 20
-50 -L5 -40
TARGET STRENGTH, dB
I l I I I I I
-LO
-
35 - 30 - 25 - 20- 50 - L5
TARGET STRENGTH, d B
Fig. 5B. Observations of maximum dorsal aspect target strength on saithe and the regression line.
l I l l I l 1
- 4 5 - 4 0 - 35 - 30 - 2 5 - 2 0
- 50
TARGET STRENGTH, dB
I I I I I l 1
-
50 - 45 - 4 0-
35 - 30 - 2 5 - 20TARGET STRENGTH, dB
Fig. 5C. Observations of maximum dorsal aspect target strength o n pollack and the regression line.
TS = 13.6 l o g L
-
5 6 . 81 o
TARGET STRENGTH, dB
TS : 18.8 l o g L - 6 2 . 4 1 o
1
l I l I l I I
-
50 - 4 5 - 4 0 -35-
30 - 2 5-
2 0TARGET STRENGTH, d 8
F i g . 5D. O b s e r v a t i o n s o f maximum d o r s a l a s p e c t t a r g e t s t r e n g t h on h e r r i n g and t h e r e g r e s s i o n l i n e .
TS i 17.2 l o g L
-
6 0 . 810 L -
TS = 21.4 Log L - 6 6 . 0 10
I I I I I I
4
-
Fig. 5E. Observations of maximum dorsal aspect target strength on sprat and the regression line.
- 4 0 - 3 5
-
30 - 2 5 - 2 0-
5 0-
45TARGET STRENGTH , dB
l I I I 1 I
-
L5 -LO - 35-
30-
2 5 - 2 0-
50TARGET STRENGTH, d 8
J
I 1 l I 1 I l I- 5 5 - 5 0
-
L 5-
LO - 3 5-
3 0-
2 5-
2 0TARGET STRENGTH, d B
/w -
,g:
x iFig. 5F. Mean values of observations of maximum dorsal aspect target strength of 1) mackerel, 2) horse mackerel
,
3) haddock,
4) blue whi ting, 5 ) whit-ing, 6 ) spiny dogfich, 7 prawn and 8) the cod regression line.
E
2 0 -x
z
Z W -r 10
-
8
-
6
- -
L
-
% /
/ , l
, "
0, +
2/ L - / / O 3
i A L
0
,/'i
Q) Ø
v
50 0
O 0 0
\ o
60 0
0
l
@J 70 0
0
- - -
l 8
i
I I I r I I I I
-
5 5 - 5 0 - L 5 - L O - 3 5 - 30 - 25 - 2 0TARGET STRENGTH , d 9
4 l
I I l - I I I I
-50 - 45 - 4 0 - 35 - 3 0 - 2 5 - 2 0
TARGET STRENGTH, d B
F i g . $6. D o r s a l a s p e c t t a r g e t s t r e n g t h - l e n g t h r e l a t i o n s a t 38 KHz f o r cod and h e r r i n g . 1) Observed maximum v a l u e s , 11) mean v a l u e s f o r tilt a n g l e s w i t h i n
- +
3O of a n g l e o f maximum v a l u e ( c o r r e s - ponds t o 6O t r a n s d u c e r beam w i d t h ),
111) expec- t e d v a l u e s i n t h e f i e l d ( d e r i v e d from F i g . 4 ) . 1) F i e l d o b s e r v a t i o n o f mean v a l u e ( M I D T T U N and NAKKEN 1 9 7 1 ) , 2 ) day and 3 ) n i g h t o b s e r v a t i o n s of tilt a n g l e .8 - -
6 - -
4 -
l I l l I I
- 5 0 - L5 - 4 0 - 35 - 3 0 - 2 5 - 2 O
TARGET STRENGTH , d B
- 2 1
1
38 K H zi -
21B
m u
- -
23I
C (3 z
W - 2 5 a
C cn - 2 7
-
2 9W g - 3 1 a - 3 3
C - 3 5
- 41 &
I
C o 3 o 2 0 1 O O 1 o 2 o 3 o C O
- 2 1
l
38 K H zc
;
m u-
- 23I
l-
2 -
2 5W a
l-
-
2 7cn
l- - 2 9
W
*
a - 3 1a - 3 3
l- - 3 5
-
L1H E A D D O W N H E A D U P
T I L 1 A N G L E , D E G R E E S
Fig. 7. Observations of dorsal aspect target strength
1
of a swimming saithe. A ) Low, B ) moderate and C ) high swimming activity.~
- 2 1 1 38 KHzul
- 2 1 38 KHz
W
-
2 5- 2 7
T A I L BEAT C Y C L E
Fig. 8. Observations of dorsal aspect target strength of a swimming cod (69 cm). A) At zero tilt angle and B) at tilt angle of maximum target strength (5O).
DORSAL ASPECT 1
- - -
2F i g . 9 . T a r g e t s t r e n g t h a s a f u n c t i o n of r o l l a n g l e f o r 1) c o d , 2 ) s a i t h e and 3 ) h e r r i n g .
F i g . 1 0 . orres spond ing v a l u e s o f maximum d o r s a l a s p e c t t a r g e t s t r e n g t h and a n g l e between p o i n t s o f h a l f maximum a m p l i t u d e (6dB points). 1) Cod,
2 ) s a i t h e , 3 ) h e r r i n g , 4 ) and 5 ) f i e l d o b s e r - v a t i o n s o f cod and s a i t h e ( M I D T T U N and NAKKEN 1 9 7 1 ) . The v a l u e s a r e a v e r a g e d o v e r t h e i n d i - c a t e d number of f i s h .
-50 , c m T l , : ~ ~ ~ ~ ~ ~ ~ i ~ ~ r ~ ~ ~ ~ ~ ~ ~ r - ~ t ~ - ~ ~ ~ ~
5 1 O 15 20 2 5 3 0 3 5 40
ANGLE BETWEEN 6 dB POINTS, DEGREES
- 2 0 -
- 2 2 - - 2 4 -
- 2 6 -
M - 2 8 -
I x -
0 -30 -
(L1 -o - 3 2 -
I
5 - 3 L -
z w - U:
; n - 3 6 -
t- -
W
C
O 1 O 2
O 3
+ L
A 5
, I I I l I I ~ I
- 2 2 - 2 0 -18 -16 TARGET STRENGTH d B , 120 K H Z
F i g . 11 Maximum d o r s a l a s p e c t t a r g e t s t r e n g t h o f i n d i v i d u a l f i s h a t two f r e q u e c i e s , 3 8 kHz a n d 120 kHz. 1) Cod, 2 ) m a c k e r e l , 3 ) s a i t h e , 4 ) p o l l a c k and 5 ) h e r r i n g a n d s p r a t . F u l l l i n e : McCARTNEY and STUBBS 1970 ( 4 . 5 loT>)
,
b r o k e n l i n e : c u r v e f i t t e d t o t h e d a t a .