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UBC Theses and Dissertations

Acid-base balance in arterial plasma of white Pekin duck (Anas platyrhynchos) during forced submergence… Shimizu, Manabu 1985

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ACID-BASE BALANCE IN ARTERIAL PLASMA OF WHITE PEKIN DUCK <ANAS PLATYRHYNCHOS) DURING FORCED SUBMERGENCE AND RECOVERY By MANABU SHIMIZU B. Sc., Western Washington U n i v e r s i t y , 1983 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE i n THE FACULTY OF GRADUATE STUDIES (Department o f Zoology) We accept t h i s t h e s i s as conforming t a the r e q u i r e d standard THE UNIVERSITY OF BRITISH COLUMBIA October 1985 © Manabu Shimizu, 1985 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department or by his or her representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of ZOOLOGY The University of British Columbia 1956 Main Mall Vancouver, Canada V6T 1Y3 Date October 11, ,1985 A B S T R A C T A c i d - b a s e b a l a n c e i n p l a s m a w a s s t u d i e d i n f o r c i b l y s u b m e r g e d P e k i n d u c k s ( A n a s p l a t y r h y n c h o s ) . A l l i m p o r t a n t v a r i a b l e s d e t e r m i n i n g a c i d - b a s e b a l a n c e ( a r t e r i a l CO2 t e n s i o n (PaC02>» s t r o n g i o n d i f f e r e n c e ( [ S I D ] ) a n d t o t a l p r o t e i n c o n t e n t w h i c h a p p r o x i m a t e s t o t a l w e a k a c i d c o n c e n t r a t i o n i n p l a s m a ) w e r e m e a s u r e d . D u r i n g f o r c e d s u b m e r g e n c e PaC02 i n c r e a s e d a n d p H a s t e a d i l y d e c r e a s e d w i t h t i m e . T h e r e w a s a l s o a n i n c r e a s e i n [ l a c t a t e - ] , w h i c h w a s a c c o m p a n i e d b y a n i n c r e a s e o f e q u a l m a g n i t u d e i n C N a + ] . T h e r e w e r e n o s i g n i f i c a n t c h a n g e s i n t h e c o n c e n t r a t i o n s o f o t h e r s t r o n g i o n s ( K + , C a 2 + , M g 2 + a n d C l ~ ) . S t r o n g i o n d i f f e r e n c e d i d n o t c h a n g e d u r i n g t h e f i r s t t w o m i n u t e s o f s u b m e r g e n c e , b u t t h e r e w a s a b o u t a 4 m e q u i v / L i n c r e a s e b y t h e e n d o f t h e f o u r - m i n u t e d i v e . T h e o r e t i c a l l y a n i n c r e a s e i n [ S I D ] s h o u l d c a u s e p l a s m a t o b e a l k a l i n e , b u t s i n c e p l a s m a b e c a m e p r o g r e s s i v e l y a c i d i c , t h i s c o n d i t i o n c a n o n l y b e d u e t o t h e i n c r e a s e i n PaC02-D u r i n g r e c o v e r y f r o m d i v e s , t h e p l a s m a r e m a i n e d a s a c i d i c o n e m i n u t e a f t e r e m e r s i o n a s a t t h e e n d o f t h e d i v e s . O n t h e o t h e r h a n d , a r t e r i a l p H s l o w l y i n c r e a s e d t o w a r d s t h e p r e - d i v e l e v e l d u r i n g r e c o v e r y . A r t e r i a l CO2 t e n s i o n d e c r e a s e d m u c h m o r e r a p i d l y a n d w a s a l r e a d y a t o r b e l o w t h e p r e - d i v e l e v e l o n e m i n u t e a f t e r e m e r s i o n . T h e r e f o r e , P11CO2 c o u l d n o l o n g e r a f f e c t p l a s m a p H . T h e r e w a s , h o w e v e r , a g r e a t i n c r e a s e i n [ l a c t a t e - ] i n t h e f i r s t m i n u t e o f + + r e c o v e r y . Although CNa 1 and CK 3 were e l e v a t e d , from p r e -d i v e v a l u e s a f t e r the -four—minute d i v e , the i n c r e a s e i n [ l a c t a t e 1 r e s u l t e d i n a marked r e d u c t i o n i n CSID3. Si n c e t h e r e was no change i n the t o t a l plasma p r o t e i n content, th a c i d i c c o n d i t i o n observed i n r e c o v e r y c o u l d o n l y be due t o decreased [SID]. Breathing 0^ be-fore d i v i n g prevented c i r c u l a t o r y adjustments and pH r e t u r n e d to p r e - d i v e l e v e l s one minute a f t e r emersion, c o n f i r m i n g t h a t the a c i d i c c o n d i t i o n observed i n r e c o v e r y i s a consequence of the l a c t a t e produced i n the hypoperfused t i s s u e s d u r i n g submergence. iv TABLE OF CONTENTS Abstract • i i Table of contents i v L i s t of figures v L i s t of tables v i Acknowledgements v i i Introduction 1 Materials and methods 8 Results 17 Discussion 30 References ••• 36 > V LIST OF FIGURES F i g u r e 1 . S e m i l o g p l o t o f t h e dependent a c i d - b a s e 4 v a r i a b l e s a g a i n s t PCO2 i n plasma w h i l e [SID] and CA T O T ^ a r e c o n s t a n t F i g u r e 2. S e m i l o g p l o t o f t h e dependent a c i d - b a s e 6 v a r i a b l e s a g a i n s t t h e [SID] i n plasma w h i l e PCO2 and [AT OT^ a**e c o n s t a n t F i g u r e 3 . Schemat ic diagram o f t h e e x p e r i m e n t a l s e t u p 13 F i g u r e 4 . Changes i n ( A ) PaC02» <B> C H + ] , <C> [ l a c t a t e - ] , 22 (D) [Na + ] and <E> [SID] from p r e - d i v e v a l u e s a g a i n s t the l e n g t h o f submergence F i g u r e 5 . Changes i n ( A ) PaC02» <B> [ H + ] , <C> [ l a c t a t e - ] , 25 <D> [ N a + ] , <E> [K*] and <F) [SID] from p r e - d i v e v a l u e s a g a i n s t t ime a f t e r t h e end of t h e o n e - , two- and f o u r - m i n u t e submergence F i g u r e 6. Changes i n ( A ) P a C 0 2 , <B> pHa and 28 (C) [ l a c t a t e - ] a g a i n s t t ime from t h e end o f t h e two-minute d i v e a f t e r b r e a t h i n g l O O t O2 f o r f i v e m i n u t e s . P r e - d i v e and e n d - d i v e v a l u e s a r e i n c l u d e d v i LIST OF TABLES T a b l e 1 . V a l u e s o f heart, r a t e , mean a r t e r i a l b l o o d 18 p r e s s u r e and minute v e n t i l a t i o n b e f o r e , d u r i n g and a f t e r o n e - , two- and f o u r - m i n u t e s o f submergence T a b l e 2 . P r e - d i v e v a l u e s o f P a C 0 2 » CSID3, [ l a c t a t e - ] , 20 CNa + 3 , C K + ] , CCa2+], [Mg2+], cci"] and pHa b e f o r e (A) o n e - , <B> two- and <C) f o u r - m i n u t e s o f submergence ACKNOWLEDGEMENTS I e x p r e s s my s i n c e r e a p p r e c i a t i o n t o my s u p e r v i s o r . D r . D a v i d R. J o n e s , f o r h i s s u p p o r t which has made i t p o s s i b l e f o r me t o complete t h e r e s e a r c h f o r t h i s d e g r e e . I a l s o thank D r . P . W. Hochachka and D r . D . J . R a n d a l l f o r t h e use o f t h e i r equipment i n t h i s s t u d y . I a p p r e c i a t e t h e h e l p , a d v i c e and s u g g e s t i o n s from D r . W. K. M i l s o m , D r . G . R. J . G a b b o t t , M r . R. A . F u r i l l a , M r . M. Douse, M r . M. H e i e i s , Mr . F . M. Smith and M i s s S . S t e v e n s o n . I thank D r . C . H a r r i s and h i s s t a f f f o r making the s t r o n g i o n measurements on t h e S . M. A . C . system a t S t . P a u l ' s H o s p i t a l , V a n c o u v e r , B . C . . T h i s s t u d y was s u p p o r t e d by g r a n t s from t h e B r i t i s h Columbia Hear t F o u n d a t i o n and N . S . E . R. C . C . awarded t o D r . J o n e s . 1 INTRODUCTION A s e r i e s o f p h y s i o l o g i c a l changes and a d j u s t m e n t s t a k e p l a c e when b i r d s and mammals a r e submerged: apnea , a drop i n the a r t e r i a l oxygen t e n s i o n ( P a 0 2 ) » an i n c r e a s e i n the a r t e r i a l c a r b o n d i o x i d e t e n s i o n (PaC02>, a r e d u c t i o n i n t h e a r t e r i a l pH (pHa) , b r a d y c a r d i a , a r e d i s t r i b u t i o n o f b l o o d f l o w and changes i n metabol ism ( I r v i n g e t a l . . , 1935; I r v i n g , 1939; S c h o l a n d e r , 1940, 1961-1962; A n d e r s e n , 1963a, 1963b, 1963c, 1966; Kooyman e t g l . . , 1981; Jones and B u t l e r , 1982) . These a d j u s t m e n t s a r e m a i n t a i n e d t h r o u g h o u t submergence as l o n g as the l i m i t s o f p h y s i o l o g i c a l t o l e r a n c e a re not exceeded ( I r v i n g e t a l . . , 1935; I r v i n g ejt g l . . , 1941; Kerem and E i s n e r , 1973; Hudson and J o n e s , 1982) . When t h e a n i m a l s u r f a c e s , e f f o r t s a r e made t o r e s t o r e the p h y s i o l o g i c a l v a r i a b l e s to n o r m a l : t h e s e i n v o l v e t a c h y c a r d i a , hyperpnea and i n c r e a s e d b l o o d f l o w t o the p e r i p h e r a l v a s c u l a t u r e , a l l o f which s e r v e t o r e s t o r e t h e l e v e l s o f b l o o d gases and pH t o normal and t o c l e a r m e t a b o l i c end p r o d u c t s which accumulated d u r i n g the d i v e ( B u t l e r and J o n e s , 1968; Murphy et. g l _ . , 1980) . The a r t e r i a l pH f a l l s d u r i n g t h e p e r i o d o f f o r c e d submers ion (Andersen e t a l . . 1965; Andersen and H u s t v e d t , 1967; L i l l o and J o n e s , 1982) and d e s p i t e the p h y s i o l o g i c a l changes which o c c u r a f t e r e m e r s i o n , i t has been r e p o r t e d t h a t pH remains low f o r sometime d u r i n g r e c o v e r y (Andersen e t g l _ . , 1965; Andersen and H u s t v e d t , 1967) . D u r i n g d i v i n g t h e i n c r e a s i n g P 0 C O 2 and l a c t i c a c i d c o n c e n t r a t i o n a r e assumed t o 2 reduce t h e a r t e r i a l pH w h i l e l a c t i c a c i d a l o n e i s c o n s i d e r e d t o keep pH low d u r i n g r e c o v e r y (Andersen e t a l . . , 1965) . One at tempt has been made t o c a l c u l a t e the a c i d - b a s e s t a t u s d u r i n g d i v i n g and r e c o v e r y (Andersen e t a l . . , 1965) , but the r e s u l t was c o n f u s i n g because the method o f a n a l y z i n g a c i d - b a s e s t a t u s was i n a d e q u a t e . The c u r r e n t way o f u n d e r s t a n d i n g a c i d - b a s e s t a t e was i n t r o d u c e d by Davenport (1947) u s i n g t h e H e n d e r s o n - H a s s e l b a l c h e q u a t i o n , [HCO3-3 pH = p K ' + l o g , ( 1 ) . a x PCO2 Where K ' i s t h e d i s s o c i a t i o n c o n s t a n t o f CO2 i n plasma (pK' = 6.10) and CC i s t h e s o l u b i l i t y c o e f f i c i e n t o f CO2 i n plasma (.a = 0 . 0 3 0 1 ) . T h i s e q u a t i o n i s adequate f o r a s o l u t i o n i n which o n l y changes i n PCO2 cause changes i n pH and C H C O 3 - ] . A l t h o u g h the e q u a t i o n i t s e l f i s v a l i d , i t i s o f t e n i n c o r r e c t l y a p p l i e d because i n many p h y s i o l o g i c a l s o l u t i o n s PCO2 i s not t h e o n l y v a r i a b l e a f f e c t i n g t h e a c i d - b a s e b a l a n c e . To a v o i d m i s i n t e r p r e t a t i o n o f the a c i d - b a s e s t a t e , i n f o r m a t i o n on each i o n s p e c i e s i n a s o l u t i o n i s a l s o e s s e n t i a l . In f a c t , t h e r e are two major groups of v a r i a b l e s ; one group c o n t a i n s the s o -c a l l e d independent v a r i a b l e s , whose s t a t e i n a s o l u t i o n i s independent from o t h e r v a r i a b l e s ; t h e o t h e r group c o n t a i n s the s o - c a l l e d dependent v a r i a b l e s , whose s t a t e i s always d e t e r m i n e d by independent v a r i a b l e s ( S t e w a r t , 1978) . There are t h r e e independent v a r i a b l e s a f f e c t i n g a c i d - b a s e s t a t e i n p l a s m a : PaC02» s t r o n g i o n d i f f e r e n c e ([SID] = (sum o f a l l 3 s t r o n g base c a t i o n c o n c e n t r a t i o n ) - (sum o f a l l s t r o n g a c i d a n i o n c o n c e n t r a t i o n ) ) and t o t a l weak a c i d c o n c e n t r a t i o n ( C A T O T ^ >• The l a s t v a r i a b l e i s a lmost e q u a l t o t o t a l plasma p r o t e i n c o n c e n t r a t i o n i n plasma and r a r e l y c h a n g e s . There are s i x dependent v a r i a b l e s : [ H C O 3 - ] , [ C O 3 2 - ] , [HAD ( c o n c e n t r a t i o n o f u n d i s s o c i a t e d weak a c i d s ) , CA~] ( c o n c e n t r a t i o n of d i s s o c i a t e d weak a c i d a n i o n ) , tH +3 and [ O H - ] . E q u a t i o n (1) c o n t a i n s two dependent v a r i a b l e s and o n l y one independent v a r i a b l e , t h e r e f o r e , t h e e q u a t i o n i s not adequate t o d e s c r i b e t h e a c i d - b a s e b a l a n c e o f p l a s m a . In r e a l i t y , i t i s n e c e s s a r y t o s o l v e s i x independent e q u a t i o n s s i m u l t a n e o u s l y t o a c h i e v e an adequate d e s c r i p t i o n ( S t e w a r t , 1981) . Changes i n t h e dependent v a r i a b l e s a re d i f f e r e n t depending on which independent v a r i a b l e ( s ) i s (are) c h a n g e d . F i g u r e s 1 and 2 show how the independent v a r i a b l e s (PCO2 and [ S I D ] , r e s p e c t i v e l y ) a f f e c t t h e a c i d - b a s e s t a t u s . The o b j e c t i v e o f t h e c u r r e n t i n v e s t i g a t i o n was to i d e n t i f y the r e l a t i o n s h i p between t h e a c i d i c c o n d i t i o n o f plasma and the b e h a v i o u r o f t h e t h r e e i n d e p e n d e n t v a r i a b l e s d u r i n g d i v i n g and r e c o v e r y . T h i s was a c c o m p l i s h e d by a s s e s s i n g changes i n PaC02 and c o n c e n t r a t i o n s o f a l l the major s t r o n g i o n s i n t h e plasma which may c o n t r i b u t e t o change i n the [SID] d u r i n g submergence and d u r i n g r e c o v e r y . 4 F i g u r e 1. A s e m i l o g p l o t o f a l l s i x dependent v a r i a b l e s i n plasma a g a i n s t PC02- The s t r o n g i o n d i f f e r e n c e (CSID]) and the t o t a l weak a c i d c o n c e n t r a t i o n < [A-roT^ ' a r e c o n s t a n t a t 40 m e q u i v / L and 20 m e q u i v / L r e s p e c t i v e l y . V a l u e s a r e c a l c u l a t e d w i t h t h e e q u a t i o n s d e s c r i b e d by Stewart (1981) . 5 ~ IO"1 > 0> IO"2 TJ C o " IO"4 < C H C 0 5 3 C A T O T D * 2 0 mequiv/L CSIDD - 4 0 mequiv/L O 1 ° O i io O o 10 - 6 C C 0 5 3 O 10 ,-7 L C H + D • X I0"8 L i i i i i i i i i i i i i i i i • i 20 30 40 50 60 70 80 90 100 Pco2 (mmHg) 6 F i g u r e 2 . A s e m i l o g p l o t o f a l l s i x dependent v a r i a b l e s i n plasma a g a i n s t the s t r o n g i o n d i f f e r e n c e < CSID1). Carbon d i o x i d e t e n s i o n <PC02> and t h e t o t a l weak a c i d c o n c e n t r a t i o n < [ A T O T J > a r e c o n s t a n t a t 30 mm Hg and 20 m e q u i v / L r e s p e c t i v e l y . V a l u e s a r e c a l c u l a t e d w i t h t h e e q u a t i o n s d e s c r i b e d by Stewar t (1981) . C0H"3 J 1 1 1 t i i i 30 4 0 50 60 CSID3 ( m e q u i v / L ) 8 MATERIALS AND METHODS In t h i s s t u d y f i f t e e n a d u l t female White P e k i n ducks (Anas p l a t y r h y n c h o s ) were u s e d . The average body weight of the ducks was 2 . 7 1 0 . 6 <Istandard d e v i a t i o n ) k g . The ducks were k e p t i n an open pen w i t h f r e e a c c e s s t o f o o d and water b e f o r e b e i n g brought i n t o the l a b o r a t o r y f o r e x p e r i m e n t s . Each a n i m a l was u s u a l l y brought t o t h e l a b o r a t o r y one o r two days b e f o r e t h e e x p e r i m e n t s . In the l a b o r a t o r y , t h e a n i m a l was k e p t i n a w i r e cage (610 mm x 560 mm x 560 mm: h e i g h t , width and l e n g t h , r e s p e c t i v e l y ) under an a r t i f i c i a l p h o t o p e r i o d (12 hours day and 12 hours n i g h t ) . B e f o r e s u r g e r y , f o o d and water were g i v e n ad. l i b . . A f t e r the p r e p a r a t o r y p r o c e d u r e s , o n l y water was g i v e n t o t h e a n i m a l . B l o o d v e s s e l c a n n u l a t i o n s were the o n l y s u r g i c a l p r e p a r a t i o n s done on t h e a n i m a l s i n t h i s s t u d y . S u r g e r y was c a r r i e d out under l o c a l a n e s t h e s i a ( X y l o c a i n e 2&, A s t r a , M i s s i s a u g a , O n t a r i o ) . The f l a r e d end o f p o l y e t h y l e n e t u b i n g (PE-90 , C l a y - A d a m s , P a r s i p p a n y , New J e r s e y ) was i n s e r t e d i n t o both t h e l e f t and r i g h t b r a c h i a l a r t e r i e s . The a r t e r i a l c a t h e t e r s were p l a c e d so t h a t the t i p s were near t h e b i f u r c a t i o n w i t h the a o r t a . A l l c a t h e t e r s were f i l l e d w i t h a h e p a r i n i z e d s a l i n e s o l u t i o n <200 UPS u n i t s h e p a r i n per ml o f s a l i n e ) . S u r g e r y was not done under a s e p t i c c o n d i t i o n s but a l l s u r g i c a l i n s t r u m e n t s were s t e r i l i z e d p r i o r t o the o p e r a t i o n . The a n i m a l s were a l l o w e d t w e l v e t o t h i r t y s i x hours t o r e c o v e r from s u r g e r y . 9 H e a r t r a t e (HR) was o b t a i n e d from e l e c t r o c a r d i o g r a m (ECG) t r a c e s . The ECG was r e c o r d e d by p l a c i n g e l e c t r o d e s s u b c u t a n e o u s l y i n t h e l e f t s h o u l d e r and i n t h e r i g h t t h i g h . S i g n a l s were f e d i n t o an i s o l a t e d p r e - a m p l i f i e r (Hodel 11-5407-58, Gould I n s t r u m e n t s , C l e v e l a n d , Ohio) then t o a U n i v e r s a l a m p l i f i e r (Hodel 13-4615-58, G o u l d I n s t r u m e n t s ) . Mean a r t e r i a l b l o o d p r e s s u r e (HABP) was r e c o r d e d t h r o u g h one o f t h e a r t e r i a l c a t h e t e r s . The c a t h e t e r was c o n n e c t e d t o a p r e s s u r e t r a n s d u c e r ( B i o Tech B T - 7 0 , Narco B i o - S y s t e m , H o u s t o n , T e x a s ) . The s i g n a l s were p r o c e s s e d by a p r e s s u r e p r o c e s s o r a m p l i f i e r (Hodel 13-4615-52, Gould I n s t r u m e n t s ) . T i d a l volumes ( V T ) and b r e a t h i n g f r e q u e n c i e s ( f ) were m o n i t o r e d w i t h a pnuemotachograph (No. A547, H e w l e t t - P a c k a r d , Waltham, M a s s a c h u s e t t s ) a t t a c h e d t o a b o d y - p l e t h y s m o g r a p h (chamber s i z e = 150 mm (diameter ) x 500 mm ( l e n g t h ) ) . The p r e s s u r e d i f f e r e n c e a c r o s s t h e pneumotachograph was r e c o r d e d w i t h a p r e s s u r e t r a n s d u c e r (DP103 P r e s s u r e T r a n s d u c e r , V a l i d y n e E n g i n e e r i n g , N o r t h r i d g e , C a l i f o r n i a ) . V e n t i l a t i o n s i g n a l s from t h e t r a n s d u c e r were p r o c e s s e d by an i n t e g r a t o r a m p l i f i e r (Model 13-4615-70, G o u l d Ins t ruments ) t o g i v e i n t e g r a t e d t i d a l volume. Minute v e n t i l a t i o n (v"E) was c a l c u l a t e d from V T and f . Each b l o o d sample c o n s i s t e d o f 2 .6 ml and was c o l l e c t e d a n a e r o b i c a l l y t h r o u g h t h e o t h e r a r t e r i a l c a t h e t e r i n t o a 3 ml p o l y e t h y l e n e s y r i n g e . The s y r i n g e was i m m e d i a t e l y d i s c o n n e c t e d from the c a t h e t e r , s e a l e d t i g h t l y , and then p l a c e d i n an i c e b a t h . A f t e r an exper iment t h e b l o o d sample was a n a l y z e d and p r o c e s s e d . A w e l l mixed 0 .2 ml p o r t i o n o f the b l o o d sample was used t o measure P a C 0 2 » Pa02 and pHa u s i n g a b l o o d gas a n a l y z e r (IL813 p H / B l o o d Gas A n a l y z e r and A c i d Base C a l c u l a t o r , Inst rument L a b o r a t o r i e s , L e x i n g t o n , M a s s a c h u s e t t s ) . The a n a l y z e r was s e t a t a temperature of 3 7 . 0 ° C and t h e b l o o d gas and pHa v a l u e s were a d j u s t e d t o t h e body t e m p e r a t u r e o f the a n i m a l ( 4 1 . 0 ° C> by t h e i n t e r n a l computer . A l l samples were p r o c e s s e d w i t h i n t h i r t y minutes a f t e r t h e e x p e r i m e n t . Plasma l a c t i c a c i d c o n c e n t r a t i o n was de termined w i t h an e n z y m a t i c a s s a y k i t ( L a c t i c A c i d D e t e r m i n a t i o n K i t No. 826-UV, SIGMA C h e m i c a l s , S t . L o u i s , M i s s o u r i ) . A w e l l mixed 0 .3 ml p o r t i o n o f t h e b l o o d sample was c e n t r i f u g e d i m m e d i a t e l y a f t e r an exper iment t o s e p a r a t e the plasma from t h e c e l l s . An a l i q u o t o f 100 ftl o f s e p a r a t e d plasma was added t o c h i l l e d 200 ftl o f 8Je ( v o l u m e t r i c ) p e r c h l o r i c a c i d and a g i t a t e d v i g o r o u s l y f o r 30 seconds w i t h a mixer (Mixer 5432, E p p e n d o r f , Westbury , New Y o r k ) . The sample was p l a c e d i n an i c e b a t h f o r f i v e minutes and then c e n t r i f u g e d f o r 10 minutes t o s e p a r a t e t h e c l e a r s u p e r n a t a n t from t h e c o a g u l a t e d p r o t e i n s . The s u p e r n a t a n t was s t o r e d i n a r e f r i g e r a t o r a t 0 - 5 ° C . The c o n c e n t r a t i o n o f l a c t i c a c i d was measured w i t h i n f o u r days o f s a m p l i n g . One p a r t o f g l y c i n b u f f e r (No. 826-3 , SIGMA C h e m i c a l s ) was mixed w i t h two p a r t s o f d i s t i l l e d w a t e r . N i c o t i n a m i d e a d e n i n e d i n u c l e o t i d e (NAD) (No. 260-110, SIGMA C h e m i c a l s ) was added t o the b u f f e r s o l u t i o n i n a r a t i o o f 10 mg o f NAD t o 6 .0 11 ml o f t h e b u f f e r s o l u t i o n . Both 980 jil o f t h e b u f f e r m i x t u r e and 10 ftl o f t h e p r o c e s s e d b l o o d sample were p l a c e d and then mixed t h r o u g h l y i n a s p e c t r o p h o t o m e t r i c c u v e t t e which had a l i g h t path l e n g t h o f 10 mm. With a s p e c t r o p h o t o m e t e r (SP8-400 UV/VIS S p e c t r o p h o t o m e t e r , UNICAM, Cambridge , E n g l a n d ) , t h e i n i t i a l a b s o r p t i o n was measured b e f o r e 10 ftl o f enzyme ( l a c t a t e dehydrogenase (LDH), No. 826-10, SIGMA C h e m i c a l s ) was a d d e d . The f i n a l a b s o r p t i o n was measured 45 minutes a f t e r LDH was a d d e d . The c o n c e n t r a t i o n o f l a c t i c a c i d was c a l c u l a t e d from t h e d i f f e r e n c e between t h e two r e a d i n g s . C o n c e n t r a t i o n s o f s t r o n g i n o r g a n i c i o n s ( [ N a + 3 , IK+l and [ C l ~ ] ) , t o t a l c a l c i u m and t o t a l p r o t e i n c o n t e n t s i n t h e serum were d e t e r m i n e d w i t h a S . M . A . C . system ( S e q u e n t i a l M u l t i p l e A n a l y z e r Computer s y s t e m , T e c h n i c o n I n s t r u m e n t s , T a r r y t o n , New Y o r k ) . A volume o f 2 .0 ml o f t h e b l o o d sample was p l a c e d i n a s i l i c o n e - c o a t e d e v a c u a t e d serum tube ( V a c u t a i n e r , B e c t o n -D i c k i n s o n , R u t h e r f o r d , New J e r s e y ) i m m e d i a t e d l y f o l l o w i n g an e x p e r i m e n t . The sample s t o o d a t room t e m p e r a t u r e f o r 20 minutes t o a l l o w i t t o c l o t . The serum was s e p a r a t e d by c e n t r i f u g a t i o n f o r f i v e m i n u t e s . An a l i q u o t o f 800 /tl o f t h e s e p a r a t e d serum was t r a n s f e r e d t o a n o t h e r tube and kept f r o z e n a t - 2 0 ° C u n t i l i t was a n a l y z e d by t h e S . M . A . C . s y s t e m . The samples were a n a l y z e d w i t h i n t h r e e days o f t h e e x p e r i m e n t s . The magnesium i o n c o n c e n t r a t i o n was d e t e r m i n e d by an a tomic a b s o r p t i o n s p e c t r o p h o t o m e t e r (Model 2380 Atomic A b s o r p t i o n S p e c t r o p h o t o m e t e r , P a r k i n - E l m e r , N o r w a l k , C o n n e c t i c u t ) . An a l i q u o t o f 10 ftl o f t h e serum, which was 12 s e p a r a t e d i n the above p r o c e d u r e was d i l u t e d t o 1000 pi w i t h d i s t i l l e d w a t e r . The d i l u t e d sample was put t h r o u g h the s p e c t r o p h o t o m e t e r w i t h an a u t o m a t i c s a m p l i n g system (Model A S -3 A u t o s a m p l i n g System, P a r k i n - E l m e r ) . A m i x t u r e o f a c e t y l e n e ( C 2 H 2 ) and a i r was used w i t h a f l o w s p o i l e r i n t h e s p e c t r o p h o t o m e t e r . A b s o r p t i o n measurements were t a k e n a t a wavelength o f 422.7 nm. The c o n c e n t r a t i o n was c a l c u l a t e d by an i n t e r n a l computer . The [SID3 was c a l c u l a t e d from [ N a + ] , C K + ] , [ C a 2 + ] , [ M g 2 + ] , [ C I - ] and [ l a c t a t e - ] . The c a l c i u m i o n c o n c e n t r a t i o n was c a l c u l a t e d from t h e t o t a l serum c a l c i u m c o n t e n t . The i o n i z e d c a l c i u m c o n c e n t r a t i o n was d e t e r m i n e d on t h e b a s i s o f t h e assumption t h a t i o n i c c a l c i u m c o m p r i s e d 47 .5 per c e n t o f t o t a l plasma c a l c i u m ( B i a n c h i , 1968) . P r o t o c o l The a n i m a l was p l a c e d i n a t e m p e r a t u r e - c o n t r o l l e d b o d y -ple thysmograph (see F i g u r e 3 ) . Wings and l e g s were l i g h t l y r e s t r a i n e d w i t h f i l a m e n t t a p e . The head o f t h e a n i m a l extended out o f the ple thysmograph t h r o u g h a d e n t a l dam c o l l a r so t h a t t h e a n i m a l c o u l d b r e a t h room a i r . The c a t h e t e r s and the ECG l e a d s were f e d out t h r o u g h an a i r t i g h t h o l e . The pneumotachograph was a t t a c h e d t o a p o r t i n t h e ple thysmograph t o measure t h e a i r f l o w c r e a t e d by b r e a t h i n g movements o f the a n i m a l . The head o f t h e a n i m a l was p l a c e d i n a r e s t r a i n e r so t h a t t h e head c o u l d be lowered under t h e w a t e r . Once the a n i m a l was p l a c e d i n t h e ple thysmograph and a l l o f t h e c o n n e c t i o n s ( c a t h e t e r s , ECG, e t c . ) were made, t h e a n i m a l was 13 F i g u r e 3 . A s c h e m a t i c diagram o f the e x p e r i m e n t a l s e t u p . Dimensions o f t h e b o d y - p l e t h y s m o g r a p h a r e 150 mm (diamter ) x 500 mm ( l e n g t h ) i n the chamber d i m e n s i o n and t h e volume o f the chamber i s a p p r o x i m a t e l y 8 .8 L . blood sam catheter blood pressure catheter - i f pneumotachograph 15 l e f t u n d i s t u r b e d f o r t h i r t y minutes t o one hour b e f o r e t h e exper iment began, so t h a t i t c o u l d r e a c h a s t e a d y s t a t e . B e f o r e t h e f i r s t exper iment and a f t e r each e x p e r i m e n t , 0 .3 ml o f a r t e r i a l b l o o d was withdrawn f o r h e m a t o c r i t measurements. N i n e ducks were f o r c i b l y submerged once o r t w i c e f o r p e r i o d s o f one , two o r f o u r m i n u t e s . A l l o f t h e measurements were made between one minute b e f o r e submersion and t e n minutes a f t e r e m e r s i o n . S i x b l o o d samples were c o l l e c t e d a t t h e f o l l o w i n g t i m e s : one minute b e f o r e submersion ( " p r e - d i v e " s a m p l e ) , f i f t e e n seconds b e f o r e emers ion ( " e n d - d i v e " s a m p l e ) , and one , t h r e e , f i v e and t e n minutes a f t e r e m e r s i o n . I f the h e m a t o c r i t measurement i n d i c a t e d more than a 5% d r o p , t h e second d i y e was c a n c e l l e d . The a n i m a l was a l l o w e d a t l e a s t two h o u r s between d i v e s t o r e c o v e r . S i x ducks were f o r c i b l y submerged t w i c e , each t ime f o r a p e r i o d o f two m i n u t e s , a f t e r b r e a t h i n g 100& 02 f o r f i v e m i n u t e s . A p l a s t i c bag w i t h an i n l e t f o r oxygen was p l a c e d o v e r t h e h e a d . The head was lowered i n t o t h e empty f u n n e l (see F i g u r e 3 ) . Oxygen was a d m i n i s t e r e d a t a r a t e o f 5 L per m i n u t e . A f t e r f i v e minutes t h e p l a s t i c bag was removed and water was poured i n t o the f u n n e l t o b e g i n a d i v e . The a n i m a l was s u r f a c e d i n t o room a i r . The r e s t o f t h e p r o c e d u r e was t h e same as t h e above exper iment except t h a t o n l y 0 .6 ml b l o o d was c o l l e c t e d f o r each b l o o d s a m p l e . The s t r o n g i n o r g a n i c i o n s were not measured i n t h i s e x p e r i m e n t . Where s t a t i s t i c a l s i g n i f i c a n c e s a r e i n d i c a t e d i n t h e t e x t , t h e d e c i s i o n s were made from r e s u l t s o f a n a l y s i s of 16 v a r i a n c e o r S t u d e n t ' s t - t e s t . The c r i t i c a l l i m i t f o r s i g n i f i c a n c e was s e t a t a p r o b a b i l i t y o f 5 per c e n t <P = 0 . 0 5 ) . S t a n d a r d e r r o r s a r e a s s o c i a t e d w i t h s t a t i s t i c a l mean v a l u e s i n the t e x t u n l e s s o t h e r w i s e s t a t e d . 17 RESULTS Heart, r a t e s t a r t e d t o f a l l n o t i c e a b l y between twenty t o f o r t y seconds a f t e r submersion i n most o f the d i v e s . H e a r t r a t e c o n t i n u e d t o f a l l even one minute a f t e r t h e b e g i n n i n g of the d i v e s . The deepes t b r a d y c a r d i a was o b s e r v e d i n two- and f o u r - m i n u t e d i v e s a f t e r about 80 seconds from t h e b e g i n n i n g o f the d i v e . H e a r t r a t e was around 20 b e a t s per minute (bpm) f o r the r e s t o f d i v e . Once t h e a n i m a l s s u r f a c e d , t h e h e a r t r a t e s i n c r e a s e d above p r e - d i v e v a l u e s . Mean a r t e r i a l b l o o d p r e s s u r e d i d not show a g r e a t change d u r i n g t h e e x p e r i m e n t s e x c e p t i n two- and f o u r - m i n u t e d i v e s when a s l i g h t h y p o t e n s i o n o c c u r r e d a t t h e end o f t h e d i v e and i n a l l d i v e s s h o r t p e r i o d s o f h y p e r t e n s i o n o c c u r r e d i m m e d i a t e l y a f t e r e m e r s i o n . B r e a t h i n g i n c r e a s e d a f t e r a l l d i v e s . H y p e r v e n t i l a t i o n l a s t e d l o n g e r a f t e r the l o n g e r d i v e s . A l l o f the above d a t a a r e p r e s e n t e d i n T a b l e 1. These r e s u l t s a r e i n agreement w i t h t h o s e i n p r e v i o u s r e p o r t s ( A n d e r s e n , 1963a; Jones and P u r v e s , 1970; B u t l e r and J o n e s , 1971; J o n e s , 1973; L i l l o and J o n e s , 1982; Mangalam, 19S4) . P r e - d i v e v a l u e s o f P a C 0 2 » c o n c e n t r a t i o n s o f s t r o n g i o n s , [SID] and pHa a r e p r e s e n t e d i n T a b l e 2 . In a l l d i v e s , PaC02 i n c r e a s e d s i g n i f i c a n t l y . The p a r t i a l p r e s s u r e o f carbon d i o x i d e i n t h e a r t e r i a l b l o o d i n c r e a s e d more o r l e s s l i n e a r l y as d i v e d u r a t i o n i n c r e a s e d ( F i g u r e 4A>. Hydrogen i o n c o n c e n t r a t i o n i n c r e a s e d , i n a s i m i l a r f a s h i o n , w i t h d i v e d u r a t i o n ( F i g u r e 4 B ) . Plasma l a c t a t e i o n c o n c e n t r a t i o n IS T a b l e 1. V a l u e s o f h e a r t r a t e , mean a r t e r i a l b l o o d p r e s s u r e and minute v e n t i l a t i o n b e f o r e , d u r i n g and a f t e r t h e o n e - , two-and f o u r - m i n u t e submers ion are i n d i c a t e d a t the same t ime as the b l o o d v a r i a b l e s were measured. * V a l u e i s s i g n i f i c a n t l y -d i f f e r e n t <P < 0.05) from p r e - d i v e v a l u e . TIME AFTER EMERSXON C minute> n P.D. E.D. 1 3 5 10 Heart Rate (bps) 4-*in dive 5 178 20.6" 247* ' 239" 245" 238* ±S.E. ±17 ±1.3 ±50 ±7 ±15 ±16 2-sin dive 5 239 21.4' 315" 275" 289" 386* ±S.E. ±24 ±1 .6 ±51 ±11 ±21 ±70 1-sin dive 5 170 52.4* 158 185 189 183 ±5.E. 5 ±12 ±1.7 ±7 ±8 ±8 ±6 M.A.B.P. On Hg) 4-»in dive 5 194 149* 201 167 175 175 ±S.E. ±16 ±10 ±11 ±14 ±10 •12 2-sin dive 5 180 142* 185 169 166 171 ±S.E. ±11 ±9 ±9 ±11 ±14 ±12 1-sin dive 5 166 174 168 156 157 162 ±S.E. ±6 ±15 ±13 ±7 ±6 ±8 (L/nln) 4-«in dive 5 0.921 3.17* 2.66" 2.07" 1.50* ±S.E. ±0.061 ±0.47 ±0.34 ±0.33 ±0.19 2-sin dive 5 1.23 2.31" 1.64" 1.56 1 .40 ±S.E. ±0.21 ±0.49 ±0.24 ±0.22 ±0.15 1-sin dive 5 0.805 1.37" 0.924 0.950 0.745 ±S.E. ±0.034 ±0.14 ±0.094 ±0.163 ±0.076 * - significantly different C P < 0 .05) fro* a pre-dive neon H.A.B.P. - Mean Arterial Blood Pressure 20 T a b l e 2. P r e - d i v e v a l u e s o f PaC02» [ S I D ] , [ l a c t a t e " ] , [ N a + ] , [ k + ] , [ C a 2 + ] , [ M g 2 + ] , [ C I - ] and pHa b e f o r e (A) o n e - , <B) two-and C O f o u r - m i n u t e submergence a r e i n d i c a t e d . Numbers are p r e s e n t e d as m e a n s i s t a n d a r d e r r o r s <n = 5 f o r each v a l u e ) . 21 PaC02 <mm Hg) ISID3 ( * e q u i v / L ) [ L a c t a t e " ] (mequiv/L) CNa*3 (mequiv/L) [K*] (mequiv/L) [Ca 2*3 (mequiv/L) [Mg2*3 (mequiv/L) CC1"] (mequiv/L) pHo 34.5*1.9 38.9+0.4 1.88+0.32 14411 3.210.1 3.810.4 1.610.1 11210 7.44110.014 30.2+1.6 39.311.5 1.90+0.24 14213 3.9+0.5 3.910.5 1.910.1 l l O l l 7.435+0.031 31.811.9 39.210.e 1.8110.36 140+1 3.310.2 4.7+0.3 1.710.1 106 + 1 7.43210.015 22 F i g u r e 4 . Changes i n <A) PaC02# <B) [H + ] , <C) [ l a c t a t e - ] , <D) [Na*3 and <E) [SID] from p r e - d i v e v a l u e s are p l o t t e d a g a i n s t the l e n g t h o f submergence. Each p o i n t was o b t a i n e d from measurements made a t t h e end o f t h e d i v e whose d u r a t i o n was as i n d i c a t e d i n the f i g u r e . Symbols and b a r s on the graph i n d i c a t e means and i a t a n d a r d e r r o r s <n = 5 f o r each mean). •Value i s s i g n i f i c a n t l y d i f f e r e n t (P < 0.05) from p r e - d i v e v a l u e . #Values are s i g n i f i c a n t l y d i f f e r e n t <P < 0.05) from each o t h e r (# i s a number) . 30 A PoC0 2 (mm Hg) 0 25 A C H 4 : (nequiv/L) 0 5 A Clactate"3 (mequiv/L) A C Na*J (mequiv/L) 0 4 A C SIDD (mequiv/L) 1.2 B TIME (min) i n c r e a s e d s i g n i f i c a n t l y d u r i n g d i v e s except i n the one-minute d i v e . At the end o f the f o u r - m i n u t e d i v e , [ l a c t a t e - ] was s i g n i f i c a n t l y h i g h e r t h a n a t the end o f the two-minute d i v e ( F i g u r e 4 C ) . Serum sodium i o n c o n c e n t r a t i o n a l s o i n c r e a s e d ( F i g u r e 4 D ) . The c o n c e n t r a t i o n s o f t h e o t h e r i o n s and t o t a l serum p r o t e i n c o n t e n t showed no s i g n i f i c a n t change . The s t r o n g i o n d i f f e r e n c e i n c r e a s e d s l i g h t l y and t h e v a l u e a t t h e end o f the f o u r - m i n u t e d i v e was s i g n i f i c a n t l y h i g h e r than p r e -d i v e [SID] ( F i g u r e 4 E ) . When t h e f i r s t p o s t - d i v e measurements were made, PaC02 was a l r e a d y lower than t h e p r e - d i v e v a l u e s r e g a r d l e s s o f the l e n g t h o f t h e d i v e . I t c o n t i n u e d t o d e c r e a s e u n t i l t h r e e o r f i v e minutes a f t e r t h e d i v e ( F i g u r e 5 A ) . Hydrogen i o n c o n c e n t r a t i o n i n c r e a s e d s i g n i f i c a n t l y from p r e - d i v e v a l u e s a f t e r t h e a n i m a l s u r f a c e d i n the f i r s t minute a f t e r t h e end o f t h e d i v e . The i n c r e a s e s were l a r g e r a f t e r t h e l o n g e r d i v e s ( F i g u r e 5B>. Hydrogen i o n c o n c e n t r a t i o n g r a d u a l l y d e c r e a s e d towards p r e - d i v e v a l u e s . L a r g e i n c r e a s e s i n [ l a c t a t e - ] were o b s e r v e d a f t e r a l l d i v e s , a l t h o u g h a f t e r t h e o n e - m i n u t e d i v e t h e i n c r e a s e was l e s s than a f t e r the o t h e r two d i v e s ( F i g u r e 5C>. L a c t a t e i o n c o n c e n t r a t i o n s l o w l y r e t u r n e d t o the p r e - d i v e v a l u e s , however, [ l a c t a t e - ] remained s i g n i f i c a n t l y h i g h e r a f t e r t e n minutes r e c o v e r y except f o l l o w i n g t h e one-minute d i v e s . Serum sodium i o n c o n c e n t r a t i o n s were lower than d u r i n g d i v i n g and r e t u r n e d t o the p r e - d i v e l e v e l i n a few minutes ( F i g u r e 5 D ) . Serum p o t a s s i u m i o n c o n c e n t r a t i o n i n c r e a s e d about 2 m e q u i v / L one minute a f t e r t h e f o u r - m i n u t e 25 F i g u r e 5 . Changes i n <A> POC02P <B) [H + ] , <C) [ l a c t a t e - ] , <D) CNa + 3 , <E) Ck*3 and (F) [SID] from p r e - d i v e v a l u e s a r e p l o t t e d a g a i n s t t ime a f t e r the end o f the one- ( A ) , two- < • ) and f o u r - m i n u t e < # ) submergence. Symbols and b a r s on the graph i n d i c a t e means and i s t a n d a r d e r r o r s <n = 5 f o r each mean). " V a l u e i s s i g n i f i c a n t l y d i f f e r e n t <P < 0.05) from p r e - d i v e v a l u e . 26 (mm Hg) -10 20 A L H * D (nequi v / L ) 0 10 A C l o c t o t e " ] ( m e q u i v / L ) 0 3 ( m e q u i v / L ) A CK + J ( m e q u i v / L ) 0 -2 A C S I D D ( m e q u i v / L ) -10 3 5 TIME (min) 27 d i v e , however, [K + ] r e t u r n e d t o t h e p r e - d i v e v a l u e by the next measurement ( F i g u r e 5 E ) . The o t h e r s t r o n g i o n s and t h e t o t a l p r o t e i n c o n t e n t d i d not change d u r i n g t h e r e c o v e r y p e r i o d . The s t r o n g i o n d i f f e r e n c e was s i g n i f i c a n t l y lower than the p r e - d i v e v a l u e i n a l l c a s e s , however, a f t e r t h e one-minute d i v e , t h e d e c r e a s e i n [SID] was s i g n i f i c a n t l y l e s s than a f t e r t h e o t h e r two d i v e s ( F i g u r e 5 F ) . A f t e r f o u r minutes o f b r e a t h i n g 100& oxygen, t h e p r e - d i v e Pa02 i n c r e a s e d t o 35517 mm H g . The a r t e r i a l O 2 t e n s i o n d e c r e a s e d t o 32719 mm Hg by the end o f t h e two-minute d i v e . I t was r e d u c e d t o 12012 mm Hg one minute a f t e r t h e a n i m a l s u r f a c e d . M i n u t e v e n t i l a t i o n r e t u r n e d t o the p r e - d i v e l e v e l w i t h i n t h r e e minutes a f t e r s u r f a c i n g . In the d i v e s a f t e r b r e a t h i n g 100* 0 2 » HR and MABP d i d not change s i g n i f i c a n t l y d u r i n g the d i v e and r e c o v e r y . The a r t e r i a l C O 2 t e n s i o n i n c r e a s e d s i g n i f i c a n t l y d u r i n g d i v i n g , and the i n c r e a s e was g r e a t e r t h a n i n t h e two-minute d i v e i n the p r e v i o u s e x p e r i m e n t s ( F i g u r e 6 A and F i g u r e 4 A ) . The a r t e r i a l C O 2 t e n s i o n r e t u r n e d t o t h e p r e - d i v e l e v e l one minute a f t e r e m e r s i o n . There was a s i g n i f i c a n t drop i n pHa d u r i n g t h e d i v e , however, pHa r e t u r n e d t o the p r e - d i v e l e v e l by one minute a f t e r s u r f a c i n g ( F i g u r e 6 B ) . The r e d u c t i o n i n pHa was comparable t o t h a t o f t h e f o u r - m i n u t e d i v e o f t h e p r e v i o u s s e r i e s o f e x p e r i m e n t s . A l t h o u g h t h e r e was a s i g n i f i c a n t i n c r e a s e i n [ l a c t a t e - ] , t h e i n c r e a s e was much s m a l l e r than any o f the i n c r e a s e s seen i n the p r e v i o u s e x p e r i m e n t s ( F i g u r e 6 C > . 28 F i g u r e 6 . Changes i n ( A ) P a C 0 2 , <B) pHa and <C> [ l a c t a t e - ] a r e p l o t t e d a g a i n s t t ime from the end o f t h e two-minute d i v e s a f t e r b r e a t h i n g 1 0 0 H O 2 f o r f i v e m i n u t e s . P r e - d i v e (PD) and e n d - d i v e (ED) v a l u e s are i n c l u d e d . " V a l u e i s s i g n i f i c a n t l y d i f f e r e n t (P < 0 . 0 5 ) from p r e - d i v e v a l u e . rO 30 DISCUSSION The d i v i n g r e s p o n s e s o b s e r v e d i n the c u r r e n t i n v e s t i g a t i o n a r e i n agreement w i t h p r e v i o u s r e p o r t s ( A n d e r s e n , 1963a; Jones and P u r v e s , 1970; B u t l e r and J o n e s , 1971; B u t l e r and T a y l o r , 1973; J o n e s , 1973; Jones and West, 1978; Mangalam, 1984) . Both the c a r d i o v a s c u l a r and r e s p i r a t o r y r e s p o n s e s o b s e r v e d d u r i n g the p o s t - d i v e r e c o v e r y p e r i o d are a l s o i n agreement w i t h p r e v i o u s r e p o r t s (Bamford and J o n e s , 1976; L i l l o and J o n e s , 1982) . The p r e - d i v e v a l u e s f o r PaC02, Pa02 and pHa a r e comparable w i t h v a l u e s r e p o r t e d p r e v i o u s l y (Jones and H o l e t o n , 1972; B o u v e r o t , H i l d w e i n , and Le G o f f , 1974; B o u v e r o t , H i l l , and Jammes, 1974; Kawashiro and S c h e i d , 1975; Bouverot e t a l . . , 1979; Milsom et. a l . , 1983) . The p a r t i a l p r e s s u r e o f carbon d i o x i d e i n t h e a r t e r i a l b l o o d r o s e d u r i n g d i v i n g . S i n c e ducks a r e a p n e i c d u r i n g d i v i n g , CO2 produced by a e r o b i c metabolism accumula tes i n t h e c i r c u l a t i o n . In t h i s i n v e s t i g a t i o n , PoC02 r o s e more or l e s s l i n e a r l y w i t h i n c r e a s i n g d i v e d u r a t i o n . A s i m i l a r i n c r e a s e i n PaC02 w i t h d i v e t ime was r e p o r t e d p r e v i o u s l y (Andersen e t al_. , 1965) . In t h e s e e x p e r i m e n t s ducks were d i v e d f o r f o u r t e e n minutes and PaC02 i n c r e a s e d about 60 mm Hg from t h e p r e - d i v e v a l u e (Andersen e t a l . . , 1965) . The i n c r e a s e i n PaC02 was about 4 . 3 mm Hg/min i n Andersen e t a l . . ' s (1965) s t u d y . The r a t e o f t h e i n c r e a s e i n P11CO2 i n the c u r r e n t s t u d y was h i g h e r than Andersen ejt al_. ' s (1965) v a l u e by a p p r o x i m a t e l y 3 mm H g / m i n . The d i f f e r e n c e i n t h e r a t e o f t h e i n c r e a s e c o u l d be a r e s u l t , o f t h e s t a t e o f t h e a n i m a l d u r i n g submergence. Because t h e a n i m a l s were o n l y l i g h t l y r e s t r a i n e d i n t h e c u r r e n t s t u d y , they c o u l d s t r u g g l e more and hence t h e r e would be an i n c r e a s e i n t h e i r a e r o b i c m e t a b o l i s m . I t i s a l s o p o s s i b l e t h a t the d i f f e r e n c e s i n t h e r a t e o f a c h i e v i n g the maximum c a r d i o v a s c u l a r a d j u s t m e n t s d u r i n g submers ion a f f e c t the i n c r e a s e s i n P a C 0 2 > T h i s i s sugges ted by t h e r e s u l t s o f the two-minute d i v e a f t e r b r e a t h i n g oxygen i n which t h e r e was no b r a d y c a r d i a and the i n c r e a s e i n P a C 0 2 was much l a r g e r than i n t h e two-minute d i v e a f t e r b r e a t h i n g a i r . However, the g r e a t e r i n c r e a s e i n P o C 0 2 c o u l d be the r e s u l t o f an i n c r e a s e d a e r o b i c metabol ism caused by the h i g h oxygen l e v e l . I n c r e a s e d PaCC<2 c a u s e s changes i n the dependent v a r i a b l e s ; [ H C O 3 - ] , [HA] and CH+D i n c r e a s e and [ C O 3 2 - ] , [ A - ] and [OH"3 d e c r e a s e (see F i g u r e 1 ) . In White P e k i n d u c k s , when the d i v i n g r e s p o n s e s a r e f u l l y e x p r e s s e d , t h e b l o o d s u p p l y t o the p e r i p h e r a l t i s s u e s i s g r e a t l y r e d u c e d (Jones e t al_. , 1979) . As t h e oxygen c o n t e n t i n the p e r i p h e r a l t i s s u e s f a l l s , a n a e r o b i c metabol ism i n c r e a s e s . As a r e s u l t o f t h e i n c r e a s e d a n a e r o b i c m e t a b o l i s m , l a c t i c a c i d accumula tes i n t h e s e t i s s u e s . A s m a l l amount o f t h e l a c t a t e i o n a p p a r e n t l y l e a k s i n t o t h e c e n t r a l c i r c u l a t i o n , t h u s [ l a c t a t e - } i n c r e a s e s s l o w l y d u r i n g s u b m e r s i o n . The o t h e r s t r o n g a c i d a n i o n , C l ~ , does not change i n t h e d i v e , t h e r e f o r e , changes i n [ l a c t a t e - ] a r e s o l e l y r e s p o n s i b l e f o r t h e change i n t h e t o t a l s t r o n g a c i d a n i o n c o n c e n t r a t i o n . Among s t r o n g base c a t i o n s , o n l y t h e sodium i o n i n c r e a s e s i t s c o n c e n t r a t i o n d u r i n g s u b m e r s i o n . Andersen <1965) r e p o r t e d an i n c r e a s e i n [K + ] d u r i n g p r o l o n g e d <ie. 8 minutes) f o r c e d d i v i n g . However, CK+3 d i d not change d u r i n g d i v i n g i n t h e c u r r e n t s t u d y . The maximum d i v e d u r a t i o n i n t h e c u r r e n t s t u d y was f o u r minutes which might e x p l a i n the d i f f e r e n c e i n t h e r e s u l t s . An i n c r e a s e i n [Na + ] d u r i n g submers ion has never been r e p o r t e d . The s o u r c e and cause o f t h e i n c r e a s e i n [Na + ] a re not known. I t p r o b a b l y comes from t h e i n t e r s t i t i a l f l u i d which g e n e r a l l y has the h i g h e s t [Na + ] <Eckert and R a n d a l l , 1983) . The i n c r e a s e i n CNa +] was g e n e r a l l y a l i t t l e h i g h e r than t h e i n c r e a s e i n [ l a c t a t e - ] , a n d , as a r e s u l t , t h e [SID] i n c r e a s e d o n l y s l i g h t l y . The changes i n t h e plasma a c i d - b a s e b a l a n c e d u r i n g f o r c e d d i v i n g came m a i n l y from PaCC-2* The plasma becomes i n c r e a s i n g l y a c i d i c d u r i n g d i v i n g . S t r o n g i o n d i f f e r e n c e s a f f e c t i o n b a l a n c e i n t h e d i r e c t i o n o f a l k a l i n i t y , a l t h o u g h the e f f e c t o f t h i s s l i g h t change i n [SID] i s s m a l l . The serum p r o t e i n c o n t e n t d i d not change a n d , c o n s e q u e n t l y , t h e t o t a l weak a c i d d i d not a f f e c t t h e i o n b a l a n c e . The a c i d i c c o n d i t i o n d e v e l o p i n g d u r i n g d i v i n g , t h e r e f o r e , s o l e l y depends on t h e i n c r e a s e i n P a C 0 2 - In t h e two-minute d i v e a f t e r b r e a t h i n g lOOJs oxygen P 0 C O 2 i n c r e a s e d more t h a n t h e two-minute d i v e a f t e r b r e a t h i n g a i r and t h e d e c r e a s e i n pHa was comparable t o t h a t seen i n t h e f o u r - m i n u t e d i v e a f t e r b r e a t h i n g a i r , i n which P a C 0 2 i n c r e a s e d by a s i m i l a r amount. The l a c t a t e i o n c o n c e n t r a t i o n d i d not i n c r e a s e s i g n i f i c a n t l y i n t h e d i v e a f t e r b r e a t h i n g 1 0 0 & oxygen. 33 When ducks s u r f a c e , they i m m e d i a t e l y resume b r e a t h i n g and r e a d j u s t the c a r d i o v a s c u l a r system ( B u t l e r and J o n e s , 1982) . The f i r s t measurements were made one minute a f t e r the end of t h e d i v e and PaCC<2 was a l r e a d y lower than the p r e - d i v e v a l u e . The l a r g e drop from e n d - d i v e PaC02 t a k e s p l a c e i n the f i r s t minute o f t h e r e c o v e r y when b r e a t h i n g i s g r e a t l y e l e v a t e d ( L i l l o and J o n e s , 1982) . A f t e r f o u r - m i n u t e d i v e s , P o C 0 2 was lower than a f t e r t h e o t h e r two s h o r t e r d i v e s . The l a r g e r d e c r e a s e i s a r e s u l t o f s u s t a i n e d e l e v a t e d b r e a t h i n g a f t e r t h e l o n g e r d i v e . N e v e r t h l e s s , t h e lower than p r e - d i v e P a C 0 2 l e v e l s d r i v e t h e plasma a c i d - b a s e b a l a n c e i n t h e o p p o s i t e d i r e c t i o n from t h a t d u r i n g d i v i n g ; C H C 0 3 - ] , [HA] and [H + ] d e c r e a s e and [ C O 3 2 - ] , [A~] and [ 0 H ~ ] i n c r e a s e (see F i g u r e 1 ) . A l t h o u g h the drop from t h e e n d - d i v e PaCC<2 v a l u e was l a r g e , the d i f f e r e n c e from t h e p r e - d i v e v a l u e was s m a l l . The e f f e c t of P a C 0 2 on the plasma a c i d - b a s e b a l a n c e d u r i n g r e c o v e r y i s s m a l l . U n l i k e d u r i n g s u b m e r s i o n , i n the p o s t - d i v e r e c o v e r y p e r i o d [SID] d e c r e a s e s from p r e - d i v e l e v e l s . The d e c r e a s e s were l a r g e e x c e p t a f t e r t h e o n e - m i n u t e d i v e . The d e c r e a s e d [SID] r e s u l t s i n d e c r e a s e d [ H C O 3 - ] , [CC<3 2~] , [A~] and [OH - ] , and i n c r e a s e d [H + ] and [HA] (see F i g u r e 2 ) . Major changes i n [SID] d u r i n g r e c o v e r y were due t o a l a r g e i n c r e a s e i n [ l a c t a t e - ] . L a c t a t e i o n s produced d u r i n g f o r c e d d i v i n g a r e a p p a r e n t l y r e t a i n e d m o s t l y i n the h y p o p e r f u s e d t i s s u e s and r e l e a s e d once t h e c i r c u l a t i o n i s r e s t o r e d . The c o n c e n t r a t i o n o f sodium i o n d e c r e a s e d t o t h e p r e - d i v e v a l u e i n the f i r s t 34 minute of t h e r e c o v e r y except a f t e r the f o u r - m i n u t e d i v e i n which CNa +] was a few m e q u i v / L h i g h e r t h a n p r e - d i v e f o r t h e f i r s t f i v e minutes o f r e c o v e r y . There was an i n c r e a s e i n CK + ] a f t e r t h e f o u r - m i n u t e d i v e . The i n c r e a s e was seen one minute a f t e r the d i v e . The c o n c e n t r a t i o n s o f CK+3 were a t p r e - d i v e v a l u e s f o r t h e r e s t o f t h e r e c o v e r y p e r i o d . A f t e r t h e f o u r -minute d i v e , t h e r e was an i n c r e a s e i n C C 1 ~ ] , a l t h o u g h i t was not s t a t i s t i c a l l y s i g n i f i c a n t . Because o f t h e l a r g e i n c r e a s e i n [ l a c t a t e - ] and no l a r g e change i n t h e o t h e r s t r o n g i o n s , [SID] d e c r e a s e d d u r i n g r e c o v e r y . A l a r g e r q u a n t i t y o f l a c t a t e i o n was produced i n the l o n g e r d i v e s , [ l a c t a t e - ] was h i g h e r a f t e r t h e l o n g e r d i v e and [SID] was t h e r e f o r e l o w e r . D u r i n g r e c o v e r y the major independent v a r i a b l e c o n t r o l l i n g t h e plasma a c i d - b a s e b a l a n c e i s [ S I D ] . The plasma i s a c i d i c d u r i n g r e c o v e r y j u s t as d u r i n g d i v i n g , however, the s t a t e o f t h e dependent v a r i a b l e s i s d i f f e r e n t . The p a r t i a l p r e s s u r e o f c a r b o n d i o x i d e i n t h e a r t e r i a l b l o o d , which i s lower than t h e p r e - d i v e v a l u e s h o u l d make plasma a l k a l i n e . Because t h e d i f f e r e n c e i n P 0 C O 2 from t h e p r e - d i v e v a l u e was r e l a t i v e l y s m a l l , any e f f e c t on a c i d - b a s e s t a t e i s n e u t r a l i z e d by the l a r g e change i n [ S I D ] . The change i n [SID] d u r i n g r e c o v e r y depends m o s t l y on changes i n [ l a c t a t e - ] . I f the i n c r e a s e i n [ l a c t a t e - ] i s s m a l l , such as i n d i v e s a f t e r b r e a t h i n g 1 0 0 * oxygen, pHo r e t u r n s t o the p r e - d i v e v a l u e r a p i d l y i n r e c o v e r y . The plasma o f t h e ducks becomes a c i d i c due t o an i n c r e a s e i n P a C 0 2 d u r i n g s u b m e r s i o n . D u r i n g t h e p o s t - d i v e r e c o v e r y 35 p e r i o d t h e plasma i s a l s o a c i d i c , however i t i s a c i d i c because o f t h e i n c r e a s e d [ l a c t a t e - ] . The t r a n s i t i o n from one cause t o a n o t h e r t a k e s p l a c e d u r i n g the f i r s t minute o f t h e r e c o v e r y i n a l l d i v e s . Carbon d i o x i d e i s removed by the e l e v a t e d b r e a t h i n g , w h i l e the l a c t a t e i o n i s r e l e a s e d from the p e r i p h e r y as t h e b l o o d c i r c u l a t i o n resumes . I t i s t e m p t i n g t o s u g g e s t t h a t the p r o l o n g e d p o s t - d i v e hyperpnea which l a s t s t e n t o twenty minutes a f t e r l o n g d i v e s i s c o r r e l a t e d w i t h t h e o f f -s e t i n the plasma i o n b a l a n c e due t o t h e m a i n t a i n e d i n c r e a s e i n [ l a c t a t e - ] , however, t h e c o r r e l a t i o n between hyperpnea and [ l a c t a t e - ] has y e t t o be e s t a b l i s h e d . 36 REFERENCES A n d e r s e n , H . T . , 1963a. F a c t o r s d e t e r m i n i n g t h e c i r c u l a t o r y a d j u s t m e n t s t o d i v i n g . I . Water i m m e r s i o n . A c t a P h y s i o l . S c a n d . 58 :173-185 . A n d e r s e n , H . T . , 1963b. F a c t o r s d e t e r m i n i n g t h e c i r c u l a t o r y a d j u s t m e n t s t o d i v i n g . I I . A s p h y x i a . A c t a P h y s i o l . S c a n d . 58 :186-200 . A n d e r s e n , H . T . , 1963c. 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J . and E . W. T a y l o r , 1973. The e f f e c t of h y p e r c a p n i c h y p o x i a , accompanied by d i f f e r e n t l e v e l s o f l u n g v e n t i l a t i o n , on h e a r t r a t e i n t h e d u c k . R e s p i r . P h y s i o l . 19 :176-187 . D a v e n p o r t , H . W. , 1947. The ABC o f a c i d - b a s e c h e m i s t r y . The U n i v e r s i t y o f C h i c a g o P r e s s , C h i c a g o . E c k e r t , R . , and D . J . R a n d a l l , 1983. Animal p h y s i o l o g y , second e d i t i o n . W. H . Freeman and C o . , San F r a n c i s c o . Hudson, D. M. , and D . R. J o n e s , 1982. Remarkable b l o o d c a t e c h o l a m i n e l e v e l s i n f o r c e d d i v e d d u c k s . J . E x p . Z o o l . 224 :451-456. I r v i n g , L . , 1939. R e s p i r a t i o n i n d i v i n g mammals. P h y s i o l . Rev . 19 :112-134. I r v i n g , L . , 0 . M. S o l a n d t , D . Y., S o l a n d t and K. C . F i s h e r m , 1935. 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