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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. S c . , W e s t e r n W a s h i n g t o n 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 in THE FACULTY OF GRADUATE (Department  STUDIES  o f Zoology)  We a c c e p t t h i s t h e s i s a s c o n f o r m i n g t a the required  standard  THE UNIVERSITY OF BRITISH COLUMBIA O c t o b e r 1985 © Manabu S h i m i z u , 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  copying of this thesis for scholarly purposes may department  or  by  his or  her  representatives.  be  for extensive  granted by the head of  It is understood  that  copying  publication of this thesis for financial gain shall not be allowed without my permission.  Department of  ZOOLOGY  The University of British Columbia 1956 Main Mall Vancouver, Canada V6T 1Y3 Date  October 1 1 , ,1985  my or  written  ABSTRACT  A c i d - b a s e  balance  submerged  P e k i n  v a r i a b l e s  d e t e r m i n i n g  (PaC02>»  which  plasma)  were  s t e a d i l y -  f o r c e d  ] ,  magnitude  which i n  o f  i n c r e a s e  i n c r e a s e  s i n c e o n l y  plasma be  a c i d i c On  p r e - d i v e decreased p r e - d i v e  P11CO2 a  was a l s o  There  o f  o t h e r  i n  g r e a t  p r o t e i n i n  no  from  cause  d u r i n g  i n  d i v e s ,  r a p i d l y  +  , C a  was about  a f f e c t  plasma a c i d i c ,  t o  -  +  Mg2  ,  a  4  i n  equal i n t h e  a n d  +  t h e f i r s t  C l ~ ) .  two  mequiv/L  T h e o r e t i c a l l y be  t h i s  t h e plasma a s  a t  a l k a l i n e ,  b u t  c o n d i t i o n  c a n  remained  t h e end o f i n c r e a s e d  A r t e r i a l  CO2  e m e r s i o n .  plasma  [ l a c t a t e ]  2  o f  changes  d i v e .  a n d was a l r e a d y  a f t e r  i n c r e a s e  i n c r e a s e  d u r i n g  pH s l o w l y  r e c o v e r y .  a n  PaC02-  emersion  o n e minute  i n c r e a s e  i n  a r t e r i a l  longer  ( K  t h e f o u r - m i n u t e  a f t e r  more  i o n s  b u t t h e r e  t h e i n c r e a s e  hand,  an  a n dp H a  no s i g n i f i c a n t  p r o g r e s s i v e l y  r e c o v e r y  l e v e l  s t r o n g  [SID] s h o u l d  became  much  were  by  d i d n o t change  t h e e n d o f  l e v e l  c o u l d  t e n s i o n  c o n c e n t r a t i o n  There  t i m e .  was accompanied  o n e minute  t h e o t h e r  a c i d  w i t h  +  d u e t o  D u r i n g  weak  CO2  a n d t o t a l  i n c r e a s e d  submergence,  by  ([SID])  t o t a l  important  ( a r t e r i a l  PaC02  i o n d i f f e r e n c e  minutes  balance  f o r c i b l y  A l l  submergence  CNa ].  c o n c e n t r a t i o n s  an  a c i d - b a s e  approximates  i n  p l a t y r h y n c h o s ) .  i o n d i f f e r e n c e  decreased  [ l a c t a t e  (Anas  was s t u d i e d  measured.  D u r i n g  Strong  ducks  s t r o n g  content  i n plasma  p H .  a t  as  t h e  towards  d i v e s . t h e  t e n s i o n o r  below  t h e  T h e r e f o r e , There  i n t h e f i r s t  was,  minute  however, o f  + recovery.  Although  dive values after [lactate t h e r e was acidic  one  change i n t h e t o t a l  [SID].  circulatory  d i v e , the  in recovery  a d j u s t m e n t s and  pH  returned  l a c t a t e produced  increase in  c o u l d o n l y be  be-fore d i v i n g  Since  to pre-dive  tissues  to  levels  acidic  i s a c o n s e q u e n c e of  i n the hypoperfused  due  th  prevented  emersion, confirming that the i n recovery  pre-  plasma p r o t e i n c o n t e n t ,  0^  observed  submergence.  3 were e l e v a t e d , f r o m  Breathing  minute a f t e r  condition  CK  i n a marked r e d u c t i o n i n CSID3.  c o n d i t i o n observed  decreased  1 and  t h e -four—minute  1 resulted no  CNa  +  during  the  iv TABLE OF CONTENTS  Abstract  •  i i  Table o f c o n t e n t s  iv  L i s t of figures  v  L i s t of tables Acknowledgements  vii  Introduction  1  M a t e r i a l s and methods  8  Results  17  Discussion  30  References  >  vi  •••  36  V  L I S T OF FIGURES  Figure  1.  Semilog p l o t o f the dependent a c i d - b a s e v a r i a b l e s a g a i n s t PCO2 i n p l a s m a w h i l e [SID]  4  S e m i l o g p l o t o f the dependent a c i d - b a s e v a r i a b l e s a g a i n s t t h e [SID] i n p l a s m a w h i l e PCO2 and [ A T O T ^ **e c o n s t a n t  6  and  F i g u r e 2.  CATOT^  are  constant  a  F i g u r e 3.  Schematic  diagram of  the  experimental  setup  F i g u r e 4.  Changes i n ( A ) PaC02» <B> C H ] , <C> [ l a c t a t e ] , (D) [ N a ] and <E> [SID] f r o m p r e - d i v e v a l u e s a g a i n s t t h e l e n g t h o f submergence  22  C h a n g e s i n ( A ) PaC02» <B> [ H ] , <C> [ l a c t a t e ] , <D> [ N a ] , <E> [K*] and <F) [SID] f r o m p r e - d i v e v a l u e s a g a i n s t t i m e a f t e r t h e end o f t h e o n e - , t w o - and f o u r - m i n u t e submergence  25  C h a n g e s i n ( A ) P a C 0 2 , <B> pHa and (C) [ l a c t a t e ] a g a i n s t t i m e f r o m t h e end o f t h e t w o - m i n u t e d i v e a f t e r b r e a t h i n g l O O t O2 for f i v e minutes. P r e - d i v e and e n d - d i v e values are included  28  +  13 -  +  F i g u r e 5.  +  -  +  F i g u r e 6.  -  vi L I S T OF TABLES  Table  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 p r e s s u r e and m i n u t e 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 - , t w o - and f o u r - m i n u t e s o f submergence  18  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 ] , C N a 3 , C K ] , CCa2+], [Mg2+], c c i " ] and pHa b e f o r e (A) o n e - , <B> t w o - and <C) f o u r - m i n u t e s o f submergence  20  -  +  +  ACKNOWLEDGEMENTS  I  e x p r e s s my s i n c e r e  Dr.  D a v i d R.  for  me t o  Dr.  P.  Jones,  complete  appreciation  suggestions  the  research  study.  R.  A. F u r i l l a ,  Mr.  F.  M. S m i t h and M i s s S .  and  his staff  M. A . C . s y s t e m  this  made i t  degree.  Randall for  I appreciate  Mr.  S.  for  f r o m D r . W. K . M i l s o m ,  for  my s u p e r v i s o r .  f o r h i s s u p p o r t which has  W. H o c h a c h k a and D r . D . J .  equipment i n t h i s  to  the  the  help,  D r . G . R.  J.  possible  I also  thank  use o f  their  advice  and  Gabbott,  M r . M. D o u s e , M r . M. H e i e i s , Stevenson.  making t h e at  St.  strong  Paul's  I thank D r . C .  i o n measurements  Hospital,  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 f r o m t h e H e a r t F o u n d a t i o n and N . S .  E.  R.  Harris  on  the  Vancouver, B . C . . British  C . C . awarded t o  Columbia  Dr.  Jones.  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 place the  when b i r d s and mammals a r e s u b m e r g e d :  arterial  oxygen  arterial  carbon  arterial  pH ( p H a ) ,  tension  (Pa02)»  dioxide tension  f l o w and changes i n metabolism 1939;  Scholander,  1963c,  1966;  1940,  These adjustments  a drop i n  i n the  a reduction  i n the  a r e d i s t r i b u t i o n of blood (Irving  1961-1962;  Kooyman e t g l . . ,  an i n c r e a s e  (PaC02>,  bradycardia,  apnea,  take  1981;  e t a l . . , 1935;  Andersen,  1963a,  Irving, 1963b,  J o n e s and B u t l e r ,  are maintained throughout  1982).  submergence  long as the l i m i t s of p h y s i o l o g i c a l t o l e r a n c e are not (Irving 1973;  e t a l . . , 1935;  I r v i n g ejt g l . . , 1941;  Hudson and J o n e s ,  1982).  a r e made t o r e s t o r e  normal:  these involve tachycardia,  restore the levels  clear  metabolic  (Butler The  arterial  submersion 1967;  Lillo  1968;  pH r e m a i n s 1965;  a l l o f which s e r v e  which accumulated  1982)  after  low f o r sometime  P 0 C O 2 and l a c t i c  the p h y s i o l o g i c a l  i t h a s been r e p o r t e d  during recovery 1967).  forced  Andersen and H u s t v e d t ,  and d e s p i t e  emersion,  during the dive  1980).  during the period of  A n d e r s e n and H u s t v e d t ,  increasing  vasculature,  (Andersen e t a l . . 1965;  changes which o c c u r  h y p e r p n e a and i n c r e a s e d  Murphy et. g l _ . ,  pH f a l l s  and J o n e s ,  to  o f b l o o d g a s e s and pH t o n o r m a l a n d t o  end p r o d u c t s  and J o n e s ,  surfaces,  the p h y s i o l o g i c a l v a r i a b l e s  blood flow to the peripheral to  exceeded  Kerem and E i s n e r ,  When t h e a n i m a l  efforts  as  that  (Andersen e t g l _ . ,  During d i v i n g  acid concentration  the  a r e assumed  to  2 reduce  the  arterial  pH w h i l e l a c t i c  t o k e e p pH low d u r i n g r e c o v e r y a t t e m p t h a s been made t o diving  and r e c o v e r y  was  (Andersen et  calculate  (Andersen et  was c o n f u s i n g b e c a u s e t h e  acid alone  the  al..,  is  al.. ,  acid-base 1965),  considered 1965).  One  status during  but the  result  method o f a n a l y z i n g a c i d - b a s e  status  inadequate. The c u r r e n t  way o f  understanding acid-base  i n t r o d u c e d by D a v e n p o r t  (1947)  using the  state  was  Henderson-Hasselbalch  equation, [HCO3-3 pH = p K ' + l o g  ,  (1).  a x PCO2 Where K ' (pK'  the  = 6.10)  plasma in  is  dissociation  and CC i s  the  (.a = 0 . 0 3 0 1 ) .  constant of  CO2 i n  solubility coefficient  This equation  solution  w h i c h o n l y c h a n g e s i n PCO2 c a u s e c h a n g e s i n pH and  CHCO3 ].  equation  itself  is  is  valid,  adequate  it  is  only variable  misinterpretation ion species  of  the  groups of  c a l l e d dependent  determined are  three  plasma:  also  variables;  variables,  incorrectly  i n f o r m a t i o n on In f a c t ,  other  ion difference  the  is  is  =  the  always 1978).  acid-base  ([SID]  so-  group c o n t a i n s  (Stewart,  affecting  each  there  in a solution  whose s t a t e  variables  not  To a v o i d  one g r o u p c o n t a i n s  the  -  PCO2 i s  balance.  essential.  whose s t a t e  independent v a r i a b l e s strong  state,  variables;  by i n d e p e n d e n t  PaC02»  acid-base  acid-base  independent v a r i a b l e s ,  independent from o t h e r so-  the  in a solution is  two m a j o r  called  affecting  for  often  a p p l i e d b e c a u s e 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  are  o f CO2 i n a  Although the  the  plasma  There state  (sum o f  all  in  3 strong  base c a t i o n  concentration)  anion concentration)) ( C A T O T ^ >• protein six  The l a s t  variable  concentration  dependent  dissociated  independent v a r i a b l e , to  describe  reality,  it  is  Changes  equal to  achieve  i n the  tH 3  and  +  therefore, acid-base solve  and  The o b j e c t i v e identify  the  during diving  strong the  -  variables  the  balance  of  only  is  not  plasma.  In  description  (Stewart,  variables  the  current  and r e c o v e r y .  are is  (are)  acid-base  changed.  investigation the  acidic  (PCO2  status. was  to  condition  three independent  of  variables  T h i s was a c c o m p l i s h e d by  p l a s m a w h i c h may c o n t r i b u t e  d u r i n g submergence  different  independent v a r i a b l e s  behaviour of the  i n the  and  equation  c h a n g e s i n PaC02 and c o n c e n t r a t i o n s  ions  [SID]  the  of  an a d e q u a t e  affect  are  (concentration  equations  r e l a t i o n s h i p between  p l a s m a and t h e  assessing  of  [HAD  There  independent  dependent  respectively)  changes.  plasma  six  1 and 2 show how t h e  [SID],  acid  [OH ].  d e p e n d i n g on w h i c h i n d e p e n d e n t v a r i a b l e ( s ) Figures  total  CA~] ( c o n c e n t r a t i o n  two d e p e n d e n t  necessary to  simultaneously to 1981).  the  strong  concentration  [CO32-],  -  contains  adequate  almost  [HCO3 ],  weak a c i d s ) ,  (1)  weak a c i d  is  weak a c i d a n i o n ) ,  Equation  (sum o f a l l  i n p l a s m a and r a r e l y  variables:  of u n d i s s o c i a t e d  one  and t o t a l  -  and d u r i n g  of to  recovery.  all  the  change  major in  4  F i g u r e 1. A s e m i l o g p l o t of a l l s i x dependent v a r i a b l e s i n p l a s m a a g a i n s t PC02The s t r o n g i o n d i f f e r e n c e ( C S I D ] ) 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-roT^ ' constant at 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 . Values are 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 S t e w a r t ( 1 9 8 1 ) . a  r  e  5  ~ IO"  1  CHC053  > 0>  IO"  2  mequiv/L CSIDD - 4 0 mequiv/L  CATOTD * 2 0  TJ C  o  " IO" <  4  CC053  O O i io O  1°  -6  o 10  O  ,-7 L  10  CH D +  •  X  I0" L 8  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 Pco (mmHg) 2  6  Figure 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 t h e 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> a n d t h e t o t a l weak a c i d concentration <[ATOTJ > c o n s t a n t a t 30 mm Hg and 20 m e q u i v / L respectively. Values are c a l c u l a t e d with the equations d e s c r i b e d by S t e w a r t ( 1 9 8 1 ) . a  r  e  C0H"3  J  1  30 CSID3  1  1  t  i  40 50 (mequiv/L)  i  i  60  8 MATERIALS AND METHODS  In t h i s  study  fifteen  (Anas p l a t y r h y n c h o s ) the  before  <Istandard d e v i a t i o n )  into  the  the  experiments.  i n a wire cage  w i d t h and l e n g t h , photoperiod surgery,  (12  out  Missisauga,  cannulations  under l o c a l Ontario).  mm x 560  were t h e  animals  heparinized  saline  saline).  Surgery  surgical  operation.  so t h a t aorta.  solution was n o t  instruments  the  tips  from  the  the  animal.  surgical Surgery  2&,  tubing  was i n s e r t e d The  was  Astra,  polyethylene  arteries.  into  arterial  were n e a r  the  A l l c a t h e t e r s were f i l l e d  with a  <200 UPS u n i t s h e p a r i n p e r ml o f  done u n d e r a s e p t i c c o n d i t i o n s were s t e r i l i z e d  The a n i m a l s were a l l o w e d  recover  Before  study.  New J e r s e y )  brachial  animal height,  After  (Xylocaine  The f l a r e d end o f  the  two  artificial  only  in this  one o r  mm:  hours n i g h t ) .  anesthesia  and r i g h t  with the  experiments.  laboratory  u n d e r an  Clay-Adams, Parsippany,  bifurcation  hours to  for  o n l y w a t e r was g i v e n t o  done on t h e  left  The d u c k s  laboratory,  mm x 560  h o u r s day and 12  c a t h e t e r s were p l a c e d  all  (610  procedures,  preparations  both the  In t h e  respectively)  Blood v e s s e l  (PE-90,  the  of  f o o d and w a t e r  f o o d and w a t e r were g i v e n ad. l i b . .  preparatory  carried  to  kg.  access to  laboratory  E a c h a n i m a l was u s u a l l y b r o u g h t days b e f o r e  White P e k i n ducks  The a v e r a g e body w e i g h t  i n an open pen w i t h f r e e  being brought  was k e p t  female  were u s e d .  d u c k s was 2 . 7 1 0 . 6  were k e p t  adult  surgery.  p r i o r to  twelve  to  but  the  thirty  six  9 Heart traces.  rate  (HR) was o b t a i n e d f r o m e l e c t r o c a r d i o g r a m (ECG)  The ECG was r e c o r d e d  subcutaneously  i n the l e f t  by p l a c i n g  s h o u l d e r and i n t h e r i g h t  S i g n a l s were f e d i n t o an i s o l a t e d 5407-58,  Gould  Universal  Mean a r t e r i a l  catheters.  pressure  transducer  Houston,  Texas).  Tidal  Gould  The c a t h e t e r  Instruments).  was c o n n e c t e d  The s i g n a l s were p r o c e s s e d  T  attached  by a  Gould  ( V ) and b r e a t h i n g f r e q u e n c i e s  m o n i t o r e d w i t h a pnuemotachograph  (No. A547,  (f)  (chamber  size  pressure  d i f f e r e n c e a c r o s s t h e pneumotachograph  = 150 mm ( d i a m e t e r )  x 500 mm ( l e n g t h ) ) .  (DP103 P r e s s u r e  from t h e t r a n s d u c e r  amplifier  (Model 1 3 - 4 6 1 5 - 7 0 ,  integrated  t i d a l volume.  calculated  from V  T  were p r o c e s s e d Gould  placed  was r e c o r d e d  Ventilation  by an  Instruments)  integrator  to  give  (v" ) was E  and f .  through the other  polyethylene syringe. disconnected  The  Transducer,  Minute v e n t i l a t i o n  Each b l o o d sample c o n s i s t e d anaerobically  were  to a body-plethysmograph  Validyne Engineering, Northridge, C a l i f o r n i a ) . signals  pressure  Hewlett-Packard,  Massachusetts)  transducer  to a  Instruments).  Waltham,  with a pressure  t h r o u g h one  Narco B i o - S y s t e m ,  a m p l i f i e r (Hodel 13-4615-52,  volumes  (Hodel 11-  (HABP) was r e c o r d e d  ( B i o Tech B T - 7 0 ,  thigh.  Ohio) then t o a  (Hodel 13-4615-58,  blood pressure  of the a r t e r i a l  processor  pre-amplifier  Instruments,Cleveland,  amplifier  electrodes  ml and was c o l l e c t e d  arterial  catheter  i n t o a 3 ml  The s y r i n g e was i m m e d i a t e l y  from t h e c a t h e t e r ,  i n an i c e b a t h .  of 2.6  sealed  tightly,  A f t e r an e x p e r i m e n t  and t h e n  the blood  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  the  measure  b l o o d s a m p l e was u s e d t o  a b l o o d gas  analyzer  Instrument  Massachusetts).  The a n a l y z e r  C and t h e  b l o o d gas  body t e m p e r a t u r e o f computer. after  A l l samples  the  enzymatic  lactic  of the  an e x p e r i m e n t  ftl for  of  of  8Je  30  a temperature  values  (41.0°  were p r o c e s s e d  of  were a d j u s t e d  C> by t h e  to  the  internal  within thirty  minutes  100  St.  Louis, Missouri).  ftl  separate the of separated  supernatant  from the  was s t o r e d  concentration  of  lactic  cells.  a c i d and a g i t a t e d  (Mixer 5432,  centrifuged  for  in a refrigerator  at  a c i d was measured  An 200  Westbury, five  separate  proteins.  after  vigorously  bath f o r  10 m i n u t e s t o  ml  chilled  Eppendorf,  i n an i c e  coagulated  826-UV,  immediately  p l a s m a was added t o  perchloric  with a mixer  and t h e n  w i t h an  A w e l l mixed 0 . 3  plasma from the  The s a m p l e was p l a c e d  supernatant  was d e t e r m i n e d  ( 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.  (volumetric)  New Y o r k ) .  clear  animal  at  Lexington,  b l o o d s a m p l e was c e n t r i f u g e d to  seconds  minutes  and pHa  acid concentration  assay k i t  SIGMA C h e m i c a l s ,  aliquot  was s e t  Acid  experiment.  Plasma  portion  the  Laboratories,  of  Pa02 and pHa u s i n g  ( I L 8 1 3 p H / B l o o d Gas A n a l y z e r and  Base C a l c u l a t o r ,  37.0°  PaC02»  ml p o r t i o n  the  The  0 - 5 ° C.  The  w i t h i n f o u r days  of  sampling. One p a r t o f g l y c i n b u f f e r was mixed w i t h two p a r t s o f adenine d i n u c l e o t i d e added t o  the  (No. 8 2 6 - 3 ,  SIGMA  d i s t i l l e d water.  (NAD) ( N o . 2 6 0 - 1 1 0 ,  buffer solution in a ratio  Chemicals)  Nicotinamide  SIGMA C h e m i c a l s ) of  10  mg o f  NAD t o  was 6.0  11 ml o f t h e b u f f e r s o l u t i o n . and  B o t h 980 jil o f t h e b u f f e r  10 ftl o f t h e p r o c e s s e d  b l o o d s a m p l e were p l a c e d and t h e n  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 light  path  l e n g t h o f 10 mm.  UV/VIS Spectrophotometer, initial  added.  The f i n a l  was a d d e d .  UNICAM,  (LDH),  The c o n c e n t r a t i o n  Concentrations total  A volume o f 2 . 0  of l a c t i c  inorganic  system  serum t u b e  New J e r s e y )  The s a m p l e s t o o d  serum was t r a n s f e r e d  - 2 0 ° C until  ([Na 3,  atomic  Multiple  T a r r y t o n , New  (Vacutainer,  a t room t e m p e r a t u r e  Becton-  f o r 20  The serum was s e p a r a t e d  by  An a l i q u o t o f 800 / t l o f t h e  to another  t u b e and k e p t  i t was a n a l y z e d by t h e S . M . A . C .  absorption spectrophotometer  Absorption Spectrophotometer,  in a  i m m e d i a t e d l y f o l l o w i n g an  magnesium i o n c o n c e n t r a t i o n  Connecticut) .  IK+l and  i n t h e serum  (Sequential  s a m p l e s were a n a l y z e d w i t h i n t h r e e d a y s o f t h e The  +  protein contents  c e n t r i f ugation f o r f i v e minutes.  at  ions  Technicon Instruments,  minutes t o a l l o w i t t o c l o t .  separated  a c i d was c a l c u l a t e d  ml o f t h e b l o o d s a m p l e was p l a c e d  evacuated  Rutherford,  experiment.  SIGMA C h e m i c a l s ) was  t h e two r e a d i n g s .  of strong  A n a l y z e r Computer s y s t e m ,  Dickinson,  10 ftl o f enzyme  No. 826-10,  c a l c i u m and t o t a l  silicone-coated  Cambridge, England), the  before  were d e t e r m i n e d w i t h a S . M . A . C .  York).  (SP8-400  a b s o r p t i o n was measured 45 m i n u t e s a f t e r LDH  f r o m t h e d i f f e r e n c e between  [Cl~]),  c u v e t t e w h i c h had a  With a 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 was measured  ( l a c t a t e dehydrogenase  mixture  system.  frozen The  experiments.  was d e t e r m i n e d by an (Model 2380 A t o m i c  P a r k i n - E l m e r , Norwalk,  An a l i q u o t o f 10 ftl o f t h e s e r u m ,  w h i c h was  12 separated  i n t h e above p r o c e d u r e  distilled  water.  The d i l u t e d s a m p l e was p u t t h r o u g h t h e  spectrophotometer  w i t h an a u t o m a t i c  3 Autosampling System,  spectrophotometer.  Parkin-Elmer).  an i n t e r n a l The ],  [SID3  was c a l c u l a t e d  from  [ C I ] and [ l a c t a t e ] . -  ionized  of  acetylene  i n the were t a k e n a t a was c a l c u l a t e d by  from t h e t o t a l  calcium concentration  plasma c a l c i u m  [Na ], +  CK ], [Ca +  2 +  ],  The c a l c i u m i o n c o n c e n t r a t i o n  -  assumption t h a t  total  The c o n c e n t r a t i o n  (Model A S -  computer.  was c a l c u l a t e d  the  A mixture  A b s o r p t i o n measurements  w a v e l e n g t h o f 4 2 2 . 7 nm.  2 +  sampling system  and a i r was u s e d w i t h a f l o w s p o i l e r  (C2H2)  [Mg  was d i l u t e d t o 1000 pi w i t h  serum c a l c i u m c o n t e n t .  The  was d e t e r m i n e d on t h e b a s i s  i o n i c calcium comprised 47.5 (Bianchi,  of  per cent of  1968).  Protocol 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 plethysmograph restrained extended so t h a t the  (see F i g u r e 3 ) .  with filament tape.  measure  that  room a i r .  to a port  The  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 t h e  The head o f t h e a n i m a l was p l a c e d  was p l a c e d  connections  hole.  i n the plethysmograph  in a restrainer  t h e head c o u l d be l o w e r e d u n d e r t h e w a t e r .  animal  dam c o l l a r  The c a t h e t e r s a n d  were f e d o u t t h r o u g h an a i r t i g h t was a t t a c h e d  lightly  T h e head o f t h e a n i m a l  the animal could breath  ECG l e a d s  animal.  were  out o f the plethysmograph through a d e n t a l  pneumotachograph to  Wings and l e g s  body-  so  Once t h e  i n t h e p l e t h y s m o g r a p h and a l l o f t h e  (catheters,  ECG,  etc.)  were made,  t h e a n i m a l was  13  Figure 3. A schematic diagram of the experimental s e t u p . D i m e n s i o n s 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 ( d i a m t e r ) 500 mm ( l e n g t h ) i n t h e chamber d i m e n s i o n and t h e volume o f chamber i s a p p r o x i m a t e l y 8 . 8 L .  x the  blood s a m catheter  blood p r e s s u r e catheter  -if pneumotachograph  15 left  undisturbed for t h i r t y  experiment  began,  Before the  first  of a r t e r i a l  so t h a t  minutes to  it  experiment  one h o u r b e f o r e  could reach and a f t e r  a steady  state.  each experiment,  b l o o d was w i t h d r a w n f o r h e m a t o c r i t  two o r f o u r m i n u t e s .  were made between one m i n u t e b e f o r e after  emersion.  following  f i f t e e n seconds  and o n e ,  three,  before emersion  measurement  ("pre-dive"  over the (see  head.  minute.  after  the  collected  Oxygen was a d m i n i s t e r e d a t  above experiment  were n o t measured  in this  the  plastic to  the least  time f o r  for  a  five  funnel  a r a t e of 5 L per  bag was removed and  begin a d i v e .  The r e s t  of the  The s t r o n g  The a n i m a l  p r o c e d u r e was  except t h a t o n l y 0.6  blood sample.  Where s t a t i s t i c a l text,  funnel  i n t o room a i r .  f o r each  the  bag w i t h an i n l e t f o r o x y g e n was p l a c e d  w a t e r was p o u r e d i n t o t h e  same a s  each  b r e a t h i n g 100& 02  A f t e r f i v e minutes the  was s u r f a c e d  If  recover.  The head was l o w e r e d i n t o t h e empty  Figure 3).  sample),  emersion.  S i x d u c k s were f o r c i b l y submerged t w i c e ,  A plastic  minutes  the  The a n i m a l was a l l o w e d a t  two h o u r s between d i v e s t o  p e r i o d o f two m i n u t e s ,  at  i n d i c a t e d more t h a n a 5% d r o p ,  s e c o n d d i y e was c a n c e l l e d .  minutes.  measurements  ("end-dive"  f i v e and t e n m i n u t e s a f t e r  ml  for  s u b m e r s i o n and t e n  one m i n u t e b e f o r e s u b m e r s i o n  sample),  hematocrit  A l l of the  S i x b l o o d s a m p l e s were c o l l e c t e d  times:  0.3  measurements.  N i n e d u c k s were f o r c i b l y submerged o n c e o r t w i c e periods of one,  the  ml b l o o d  inorganic  ions  experiment.  s i g n i f i c a n c e s are  indicated in  d e c i s i o n s were made f r o m r e s u l t s  of a n a l y s i s  the of  the was  16 variance or Student's  t-test.  The c r i t i c a l  limit  for  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 p e r c e n t <P = 0 . 0 5 ) . mean v a l u e s  Standard e r r o r s i n the t e x t  are associated  unless otherwise  with  stated.  statistical  17  RESULTS  Heart, r a t e s t a r t e d forty  seconds  after  rate continued to the  dives.  to  submersion  fall  dive.  the  rest  d i d not twoat  Heart  above  Once t h e  80  seconds  after  after  longer  the  i n T a b l e 1.  all  Butler  and J o n e s ,  Mangalam,  are  the  minute the  (bpm)  heart  blood  pressure  hypotension  of  Breathing longer presented  i n agreement w i t h t h o s e J o n e s and P u r v e s ,  1973;  Lillo  PaC02»  concentrations  in  occurred  periods  emersion.  for  rates  except  above d a t a a r e  1963a;  and  beginning of  experiments  Jones,  of  presented  significantly.  i n the  arterial  dive duration  concentration duration  A l l of  1971;  i n two-  Hyperventilation lasted  (Andersen,  values  and pHa a r e  dioxide  dives.  Heart  in 1970;  and J o n e s ,  1982;  19S4).  Pre-dive  increased  dives.  to  beginning of  from the  Mean a r t e r i a l  immediately a f t e r  These r e s u l t s  previous reports  the  beats per  during the  twenty  dives.  d i v e and i n a l l d i v e s s h o r t  occurred  increased  the  animals s u r f a c e d ,  pre-dive values.  show a g r e a t c h a n g e  hypertension  as  about  r a t e was a r o u n d 20  end o f t h e  between  was o b s e r v e d  and f o u r - m i n u t e d i v e s when a s l i g h t  the  [SID]  i n most o f  The d e e p e s t b r a d y c a r d i a  of d i v e .  increased  noticeably  e v e n one m i n u t e a f t e r  four-minute dives after the  fall  The p a r t i a l  increased  4B).  (Figure  In a l l  pressure  blood increased  increased,  (Figure  i n Table 2.  more o r  4A>.  lactate  ion  strong dives,  of  ions, PaC02  carbon  less  linearly  Hydrogen i o n  in a similar fashion,  Plasma  of  with dive  concentration  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 m i n u t e 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 - , t w o and f o u r - m i n u t e s u b m e r s i o n a r e i n d i c a t e d a t t h e same t i m e as t h e b l o o d v a r i a b l e s were m e a s u r e d . *Value i s significantlyd 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 .  TIME n  P.D.  Heart Rate (bps) 178 4-*in dive 5 ±S.E. ±17 239 2-sin dive 5 ±S.E. ±24 170 1-sin dive 5 ±5.E. 5 ±12 M.A.B.P. O nHg) 4-»in dive 5 194 ±S.E. ±16 180 2-sin dive 5 ±S.E. ±11 166 1-sin dive 5 ±S.E. ±6 (L/nln) 0.921 4-«in dive 5 ±S.E. ±0.061 1.23 2-sin dive 5 ±S.E. ±0.21 0.805 1-sin dive 5 ±S.E. ±0.034  E.D.  AFTER EMERSXON C minute>  1  20.6" ±1.3 21.4' ±1 .6 52.4* ±1.7  247* ±50 315" ±51 158 ±7  149* ±10 142* ±9 174 ±15  201 ±11 185 ±9 168 ±13 3.17* ±0.47 2.31" ±0.49 1.37" ±0.14  3  '  5  10  239" ±7 275" ±11 185 ±8  245" ±15 289" ±21 189 ±8  238* ±16 386* ±70 183 ±6  167 ±14 169 ±11 156 ±7  175 ±10 166 ±14 157 ±6  175 •12 171 ±12 162 ±8  2.66" ±0.34 1.64" ±0.24 0.924 ±0.094  .05) f r o * a pre-dive neon * - significantly different C P < 0 H.A.B.P. - Mean A r t e r i a l Blood Pressure  2.07" ±0.33 1.56 ±0.22 0.950 ±0.163  1.50* ±0.19 1 .40 ±0.15 0.745 ±0.076  20  T a b l e 2. P r e - d i v e v a l u e s of PaC02» [SID], [ l a c t a t e " ] , [Na ], [k ], [Ca ], [ M g ] , [ C I ] and pHa b e f o r e (A) o n e - , <B) t w o 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 a r e p r e s e n t e d a s 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 e a c h v a l u e ) . +  +  2 +  2 +  -  21  PaC02  <mm  ISID3  (*equiv/L)  [Lactate"] CNa*3 [K*]  Hg)  (mequiv/L)  (mequiv/L) (mequiv/L)  34.5*1.9  30.2+1.6  31.811.9  38.9+0.4  39.311.5  39.210.e  1.88+0.32  1.90+0.24  1.8110.36  14411  14213  140+1  3.210.1  3.9+0.5  3.310.2  [Ca *3  (mequiv/L)  3.810.4  3.910.5  4.7+0.3  [Mg2*3  (mequiv/L)  1.610.1  1.910.1  1.710.1  11210  l l O l l  106 + 1  7.44110.014  7.435+0.031  7.43210.015  2  CC1"] pHo  (mequiv/L)  22  F i g u r e 4. Changes i n < A ) PaC02# <B) [H ] , <C) [lactate ], <D) [Na*3 and <E) [SID] f r o m 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 the l e n g t h o f submergence. E a c h p o i n t was o b t a i n e d f r o m 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 t h e f i g u r e . S y m b o l s and b a r s on t h e g r a p h 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 e a c h 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 . 0 5 ) from p r e - d i v e value. # V a l u e s a r e 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 e a c h o t h e r (# i s a n u m b e r ) . +  -  30 A PoC0 (mm Hg) 2  0  B  25 A C H  4  :  1.2  (nequiv/L) 0 5 A Clactate"3 (mequiv/L)  A C Na*J (mequiv/L)  0 4 A C SIDD (mequiv/L)  TIME  (min)  increased dive.  s i g n i f i c a n t l y during d i v e s except  At the  end o f  the  four-minute dive, the  end o f  i n the  [lactate ]  higher than at  (Figure  4C).  Serum s o d i u m i o n c o n c e n t r a t i o n  (Figure  4D).  The c o n c e n t r a t i o n s  the  two-minute d i v e also  other  increased  i o n s and  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 c h a n g e .  strong  increased  ion difference  was  -  significantly  of the  one-minute  s l i g h t l y and t h e  total  The  value at  end o f t h e  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 t h a n  dive  (Figure  [SID]  p o s t - d i v e measurements  lower than the  length of the d i v e . five  minutes a f t e r  concentration after  the  the d i v e .  towards  dive  i n the were  first  larger  a l l dives,  was l e s s Lactate  Large  than a f t e r  minutes recovery  minutes  [lactate ] -  the  and r e t u r n e d t o 5D).  the  the or  values  in  the  longer  gradually  end o f  dives  decreased  [lactate ] -  were  the one-minute d i v e  other  two  dives  slowly returned  to  the  remained s i g n i f i c a n t l y  except f o l l o w i n g  Serum s o d i u m i o n c o n c e n t r a t i o n s  (Figure  increased  after  ion concentration  higher after minute d i v e s .  the  of  three  minute a f t e r  although a f t e r  however,  ten  PaC02  Hydrogen i o n  increases  pre-dive values,  during diving  5A).  Hydrogen i o n c o n c e n t r a t i o n  pre-dive values.  5C>.  (Figure  decrease u n t i l  s i g n i f i c a n t l y from p r e - d i v e  The i n c r e a s e s  increase  (Figure  the  were made,  pre-dive values regardless  continued to  animal surfaced  observed a f t e r the  It  increased  ( F i g u r e 5B>.  pre-  4E).  When t h e f i r s t was a l r e a d y  the  were  the  one-  lower  than  p r e - d i v e l e v e l i n a few  Serum p o t a s s i u m i o n  a b o u t 2 m e q u i v / L one m i n u t e a f t e r  concentration the  four-minute  25  F i g u r e 5. Changes i n <A> P O C 0 2 P <B) [ H ] , <C) [lactate ], <D) C N a 3 , <E) Ck*3 and (F) [SID] f r o m 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 i m e a f t e r t h e end o f t h e o n e - ( A ) , t w o - < • ) and f o u r - m i n u t e < # ) s u b m e r g e n c e . S y m b o l s and b a r s on t h e g r a p h 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 e a c h 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  ( m m Hg)  -10 20 A L H * D  (nequi v / L )  0 10 A  Cloctote"]  (mequiv/ L)  0 3 (mequiv/L)  A  CK  +  J  (mequiv/L)  0  -2 A C SIDD (mequiv/L)  -10  3  5 TIME  (min)  27 dive,  however,  measurement protein  (Figure  the  returned 5E).  the decrease  other  two d i v e s  Pa02 i n c r e a s e d decreased  surfaced.  after  (Figure  of breathing  t o 35517 mm H g .  100& o x y g e n ,  The a r t e r i a l  than  after  surfacing.  i n the two-minute (Figure  returned  emersion.  the pre-dive  minute a f t e r comparable  after  significantly  CO2 tension  and t h e i n c r e a s e  was  level  The a r t e r i a l C O 2 one m i n u t e  after  d r o p i n pHa d u r i n g t h e  to the pre-dive  (Figure  level  dive i n the previous  to the pre-dive  surfacing  the animal  In t h e d i v e s  6 A and F i g u r e 4 A ) .  pHa r e t u r n e d  dive.  to the p r e - d i v e  The a r t e r i a l  T h e r e was a s i g n i f i c a n t  however,  after  O2 tension  t o 12012 mm Hg one m i n u t e a f t e r  s i g n i f i c a n t l y during diving,  experiments  6B).  level  The r e d u c t i o n  by one i n pHa was  to that of the four-minute dive of the previous  s e r i e s of experiments.  A l t h o u g h t h e r e was a  increase  the increase  of  period.  5F).  t h e d i v e and r e c o v e r y .  than  total  the one-minute  100* 0 2 » HR a n d MABP d i d n o t change  increased  dive,  however,  Minute v e n t i l a t i o n returned  breathing  tension  during the recovery  i n [SID] was s i g n i f i c a n t l y l e s s  w i t h i n t h r e e minutes  greater  i o n s and t h e  t o 32719 mm Hg by t h e end o f t h e t w o - m i n u t e  was r e d u c e d  during  strong  by t h e n e x t  was s i g n i f i c a n t l y l o w e r t h a n t h e  i n a l l cases,  A f t e r f o u r minutes  It  to the pre-dive value  The o t h e r  ion difference  pre-dive value ,  +  c o n t e n t d i d n o t change  The s t r o n g  dive  [K ]  in [lactate ], -  t h e i n c r e a s e s seen  significant  was much s m a l l e r  i n the previous experiments  t h a n any  (Figure 6 C > .  28  Figure 6 . C h a n g e s i n ( A ) P a C 0 2 , <B) pHa and <C> [lactate ] a r e p l o t t e d a g a i n s t t i m e f r o m t h e end o f t h e t w o - m i n u t e d i v e s a f t e r breathing 1 0 0 H O2 f o r f i v e minutes. P r e - d i v e (PD) and e n d - d i v e (ED) v a l u e s a r e i n c l u d e d . "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 . 0 5 ) f r o m p r e - d i v e v a l u e . -  rO  30 DISCUSSION  The d i v i n g investigation (Andersen,  responses  i n the  1963a; J o n e s and P u r v e s ,  Butler  1978;  Mangalam, 1 9 8 4 ) .  respiratory  and T a y l o r ,  responses  are also  1973;  1970;  Jones,  observed  i n agreement w i t h p r e v i o u s r e p o r t s  P a 0 2 and pHa a r e c o m p a r a b l e  Scheid,  Lillo  and J o n e s ,  ( J o n e s and H o l e t o n ,  1974; 1975;  Bouverot, Bouverot  The p a r t i a l  o f carbon  In t h i s  Bouverot,  1965).  In t h e s e experiments  minutes  and P a C 0 2 i n c r e a s e d  A similar  previously  about  The i n c r e a s e  during i n the less  increase  in  fourteen  i n PaC02 was  (1965) s t u d y .  i n P11CO2 i n t h e c u r r e n t  The d i f f e r e n c e  arterial  60 mm Hg f r o m t h e p r e - d i v e  4 . 3 mm H g / m i n i n A n d e r s e n e t a l . . ' s  (1965)  1983).  ( A n d e r s e n e t al_. ,  d u c k s were d i v e d f o r  about  t h a n A n d e r s e n ejt al_. ' s  K a w a s h i r o and  P o C 0 2 r o s e more o r  (Andersen e t a l . . , 1965).  3 mm H g / m i n .  1974;  dioxide i n the  value  rate of the increase  H i l d w e i n , and  M i l s o m et. a l . ,  dive duration.  PaC02 w i t h d i v e t i m e was r e p o r t e d  values  reported  metabolism accumulates  investigation,  with increasing  (Bamford  with values  S i n c e ducks a r e apneic  CO2 p r o d u c e d by a e r o b i c  circulation. linearly  1979;  recovery  The p r e - d i v e  and Jammes,  et al.. ,  pressure  1982).  1972;  Hill,  blood rose during d i v i n g . diving,  J o n e s and West,  during the p o s t - d i v e  for  Le G o f f ,  and J o n e s ,  B o t h t h e c a r d i o v a s c u l a r and  1976;  previously  reports  Butler  1973;  and J o n e s , PaC02,  current  a r e i n agreement w i t h p r e v i o u s  1971;  period  observed  The  s t u d y was h i g h e r  v a l u e by a p p r o x i m a t e l y i n the rate of the increase  could  be a r e s u l t , o f Because  the  current  study,  the  s t a t e of  the  a n i m a l s were o n l y they could  be an i n c r e a s e possible that  in their the  lightly  struggle  aerobic  differences  maximum c a r d i o v a s c u l a r increases  i n PaC02>  the  two-minute d i v e a f t e r b r a d y c a r d i a and t h e  in  the  is  two-minute d i v e a f t e r  aerobic  m e t a b o l i s m c a u s e d by t h e  and  CH D +  Figure In  increase  in  the  the thus  the  reduced  [CO3  achieving  by t h e  2 -  ],  [A ]  affect  results  was much l a r g e r However,  result  and  -  the  i n which t h e r e  of  dependent v a r i a b l e s ;  blood supply to (Jones et  peripheral tissues As a r e s u l t  a c i d accumulates  lactate  of  an  of was than  the  increased Increased  [HCO3 ] , -  [OH"3 d e c r e a s e  -  increases  acid anion,  therefore,  changes  t h e change  i n the  Among s t r o n g b a s e  the  al_. ,  diving  1979) .  falls, the  As t h e  [HA]  (see  slowly  [lactate ]  total  -  strong  cations,  only  fully  is content  anaerobic metabolism,  i n t o the during  C l ~ , does n o t change in  oxygen  are  anaerobic metabolism  increased  leaks  responses  peripheral tissues  i n these t i s s u e s .  ion apparently  [lactate }  strong  i n the  also  h i g h oxygen l e v e l .  W h i t e P e k i n d u c k s , when t h e  increases. lactic  suggested  be t h e  is  would  1).  expressed, greatly  and  could  the  submersion  breathing a i r .  increase  changes  during  in PaC02  greater  PaCC<2 c a u s e s  in PoC02  It  rate of  b r e a t h i n g oxygen  increase  in  more and h e n c e t h e r e  i n the  This  submergence.  restrained  metabolism.  adjustments  the  no  animal during  are  central  i n the  of  circulation,  submersion.  The  other  dive,  solely responsible for  acid anion the  A s m a l l amount  sodium  concentration. ion increases  its  concentration  during submersion.  increase  in  +  diving.  However,  current  [K ]  study.  during CK 3  p r o l o n g e d <ie. d i d not  +  A n d e r s e n <1965) r e p o r t e d  change  8 minutes)  results.  The maximum d i v e d u r a t i o n i n t h e  An i n c r e a s e  in  [Na  been r e p o r t e d .  The s o u r c e  are  It  not  known.  which g e n e r a l l y 1983).  increased The diving  the  changes  in  of  this  content  air,  lactate the  as  study  the  never in  [Na ]  interstitial  fluid  < E c k e r t and  +  Randall,  a little  a result,  plasma a c i d - b a s e  i n the  diving.  higher  the  [SID]  Strong  the  in PaC02-  that  seen  in  and,  [SID]  forced  becomes ion  differences  In t h e  and t h e i n the  ion concentration breathing  is  small.  consequently,  ion balance.  i n which P a C 0 2  dive after  during  d i r e c t i o n of a l k a l i n i t y , although  d i d not change affect  balance  The p l a s m a  s l i g h t change  breathing a i r  comparable t o breathing  and,  -  The serum  the  total  The a c i d i c  therefore,  solely  two-minute d i v e  lOOJs o x y g e n P 0 C O 2 i n c r e a s e d  dive after  in  [lactate ],  developing during d i v i n g ,  increase  breathing  The  +  in  increase  was g e n e r a l l y  +  a c i d i c during  ion balance  condition the  CNa ]  [Na ]  current  s u b m e r s i o n has  and c a u s e o f t h e  highest  in  i n the  weak a c i d d i d n o t  on  during  came m a i n l y f r o m PaCC-2*  effect  protein  ]  the  only s l i g h t l y .  increasingly affect  has the  increase  +  in  difference  p r o b a b l y comes f r o m t h e  The i n c r e a s e  than the  forced  during diving  was f o u r m i n u t e s w h i c h m i g h t e x p l a i n t h e  an  depends  after  more t h a n t h e  two-minute  d e c r e a s e i n pHa was  four-minute dive increased d i d not  by a s i m i l a r  increase  1 0 0 & oxygen.  after amount.  significantly  33 When d u c k s s u r f a c e , readjust  they  the cardiovascular measurements  i m m e d i a t e l y resume  system  The  first  the  d i v e and PaCC<2 was a l r e a d y  The  large  lower  than  decrease  lower  d r o p f r o m e n d - d i v e PaC02  and J o n e s , after  the other  of sustained  dive.  Neverthless,  levels  d r i v e t h e plasma a c i d - b a s e from t h a t  i n the  dives.  than  i n the -  value  on t h e plasma a c i d - b a s e  the  PaC02  opposite +  (see F i g u r e  was s m a l l .  balance  after  [HA] and [ H ]  [A~] and [ 0 H ~ ] i n c r e a s e  from t h e p r e - d i v e  P o C 0 2 was  breathing  CHC03 ],  first  The l a r g e r  pre-dive  balance  value.  elevated  dives,  t h e d r o p f r o m t h e e n d - d i v e PaCC<2 v a l u e  difference PaC02  the lower  t h e end o f  i s greatly  elevated  during diving;  d e c r e a s e and [ C O 3 2 - ] , Although  takes place  two s h o r t e r  1982).  the p r e - d i v e  After four-minute  longer  direction  than  when b r e a t h i n g  1982).  is a result  and J o n e s ,  were made one m i n u t e a f t e r  minute o f t h e r e c o v e r y (Lillo  (Butler  b r e a t h i n g and  1).  was l a r g e ,  the  The e f f e c t  during recovery  of  is  small. Unlike during submersion, period  [SID] d e c r e a s e s f r o m p r e - d i v e  were l a r g e [SID] and  results  after  -  apparently released  levels.  the one-minute  i n decreased  [HCO3-] ,  +  Lactate retained  dive.  The d e c r e a s e d [A~] and [ O H ] ,  2  -  2).  were due t o a l a r g e  Major changes i n  increase in  i o n s produced d u r i n g f o r c e d mostly  diving  are  i n t h e h y p o p e r f u s e d t i s s u e s and  once t h e c i r c u l a t i o n  sodium i o n d e c r e a s e d  recovery  The d e c r e a s e s  [CC<3 ~] ,  [ H ] and [HA] ( s e e F i g u r e  during recovery  [lactate ].  of  except  increased  [SID]  i n the post-dive  i s restored.  to the pre-dive  The c o n c e n t r a t i o n  value  i n the  first  34 minute of  the  recovery  except a f t e r  the  which  CNa ] was a few m e q u i v / L h i g h e r  first  f i v e minutes of  after  the  four-minute dive.  after  the  dive.  values  for  the  minute d i v e , not in  [lactate ]  of  decreased  the  longer  controlling acidic  the  d i v e and  the  dependent  pressure  of carbon  Because the relatively the  small,  change  in  increase  in  The in  PaC02  [lactate ] -  1 0 0 * oxygen,  in  variables  in P0CO2  any e f f e c t  depends m o s t l y  rapidly  as  is  [SID].  different.  arterial  [SID].  from the  The change  The  blood,  on c h a n g e s i n is  small,  to  is  alkaline.  is  [SID] -  in dives  the  the  partial  which  [lactate ].  such as  pHo r e t u r n s  state in  The p l a s m a  however,  pre-dive  on a c i d - b a s e  higher  variable  during diving, is  lactate  lower.  p r e - d i v e v a l u e s h o u l d make p l a s m a  recovery  breathing  was t h e r e f o r e  ions,  of  was  -  was  increase  strong  quantity  fourit  large  [lactate ]  balance  minute  pre-dive  although  other  independent  +  A f t e r the  CC1~],  i n the  d i o x i d e i n the  difference  large  just  in  i n CK ]  one  were a t  +  period.  dives,  [SID] major  CK 3  A larger  plasma a c i d - b a s e  s t a t e of  by  the  during recovery  lower than the  change  longer  was s e e n  Because of the  during recovery.  was p r o d u c e d i n t h e  of  recovery  significant.  During recovery  is  the  and no l a r g e  -  after  The i n c r e a s e  t h e r e was an i n c r e a s e  statistically  [SID] ion  rest  the  T h e r e was an i n c r e a s e  The c o n c e n t r a t i o n s  in  than p r e - d i v e f o r  +  recovery.  four-minute dive  pre-dive  value  was  neutralized during If  the  after value  recovery.  plasma o f  the  d u c k s becomes  during submersion.  acidic  During the  due t o  post-dive  an  increase  recovery  35 period the of the another all  plasma  increased  is  breathing,  acidic,  [lactate ]. d u r i n g the  Carbon d i o x i d e i s w h i l e the  however  lactate  first  ion i s  released  the  blood c i r c u l a t i o n  suggest that  the  prolonged post-dive  set in and  twenty i n the  minutes a f t e r plasma  [lactate ], -  [lactate ] -  ion balance  however, has  long dives  yet  the to  be  resumes.  is  due t o  is  acidic  because  f r o m one c a u s e  minute of  removed by t h e  p e r i p h e r y as  to  it  The t r a n s i t i o n  -  takes place  dives.  also  the  recovery  from It  correlated  correlation  the  is  tempting  with the  maintained between  established.  in  elevated  hyperpnea which l a s t s  the  to  to ten  off-  increase  hyperpnea  36 REFERENCES  Andersen,  H . 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