The Pa Tho Physiology of Hyperventilation Disorders

8/2/2019 The Pa Tho Physiology of Hyperventilation Disorders

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DOI 10.1378/chest.109.2.5161996;109;516-534Chest

William N. Gardner

Disorders

The Pathophysiology of Hyperventilation

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CopyrightPhysicians. It has been published monthly since 1935.CHEST is the official journal of the American College of Chest

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r e u i e w sT h e P a t h o p h y s i o l o g y o f H y p e r v e n t i l a t i o nD i s o r d e r s *W i l l i a m N . G a r d n e r , M B , B S

( C H E S T 1 9 9 6 ; 1 0 9 : 5 1 6 - 3 4 )C S F = c e r e b r o s p i n a l f l u i d ; P a c c > 2 = a r t e r i a l P C O 2 ; P a C 0 2 =a l v e o l a r P C O 2 ; P E T C 0 2 = e n d - t i d a l P C O 2 ; V A = a l v e o l a r vent i l a t i o n ; V c o 2 = C 0 2 p r o d u c t i o n

T T y p e r v e n t i l a t i o n i s b r e a t h i n g i n excess o f m e t a b o l i c. * - * r e q u i r e m e n t s a n d i s a s s o c i a t e d w i t h r e d u c t i o n i na r t e r i a l P C O 2 ( P a C 0 2 ) , r e s p i r a t o r y a l k a l o s i s , a n d a w i d er a n g e o f s y m p t o m s . M a n y p s y c h o s o m a t i c s y n d r o m e sh a v e b e e n d e s c r i b e d i n t h e p a s t i n w h i c h h y p e r v e n t i l a t i o n h a s a v a r i a b l e a n d u n c e r t a i n r o l e , 1 " 6 b u t t h e t e r m' h y p e r v e n t i l a t i o n s y n d r o m e ' was f i r s t u s e d i n 1 9 3 8 t od e s c r i b e p a t i e n t s w i t h t h e s o m a t i c s y m p t o m s o f b o t hh y p o c a p n i a a n d a n x i e t y . 7 T h i s t h e m e h a s b e e n e x t e n d e d b y s u b s e q u e n t a u t h o r s 8 " 1 1 a n d t h e r e h a v e b e e nm a n y r e v i e w s , 1 2 " 1 8 b u t n e v e r t h e l e s s , t h e r e r e m a i n su n c e r t a i n t y a n d l a c k o f consensus a b o u t t h e b o u n d a r i e sa n d even e x i s t e n c e o f t h i s s y n d r o m e . 1 9 ' 2 0M o s t p h y s i c i a n s r e g a r d h y p e r v e n t i l a t i o n a s s y n o n y mous w i t h a n x i e t y a n d t h u s i n v a r i a b l y w i t h i n t h ep r o v i n c e o f t h e p s y c h i a t r i s t s , b u t a n x i e t y m a y b ea b s e n t 2 1 , 2 2 or s e c o n d a r y t o h y p e r v e n t i l a t i o n . 2 3 S o m ed i a g n o s e i t i n t h e p r e s e n c e o f t h e s o m a t i c s y m p t o m s o fh y p o c a p n i a e i t h e r a t r e s t or i n d u c e d b y v o l u n t a r yo v e r b r e a t h i n g w i t h o u t a s s u m p t i o n s a b o u t e t i o l o g y . 2 4S o m e r e g a r d i t a s a h a b i t d i s o r d e r 2 3 or an a b n o r m a l i t yo f r e s p i r a t o r y c o n t r o l . 2 5 M a n y r e f u s e t o r e c o g n i z e i t a sa s e p a r a t e e n t i t y or r e g a r d i t as s e c o n d a r y t o o r g a n i cd i s e a s e . 2 6 H y p e r v e n t i l a t i o n i s r a r e l y d i s c u s s e d i n r e s p i r a t o r y t e x t b o o k s , t h e s y m p t o m s o f h y p o c a p n i a ar er a r e l y r e c o g n i z e d i n c l i n i c a l p r a c t i c e , a n d w h e n t h e ya r e , t h e y ar e r a r e l y m a n a g e d w i t h t h e same c l i n i c a l a n ds c i e n t i f i c o b j e c t i v i t y a s o t h e r o r g a n i c a n d p h y s i o l o g i cd i s t u r b a n c e s .T h e s e u n c e r t a i n t i e s r e f l e c t t h e c o m p l e x i t y o f t h i ss u b j e c t t h a t f a l l s b e t w e e n p s y c h i a t r y , c l i n i c a l m e d i c i n e ,a n d p h y s i o l o g y a n d i s n o t r e c o g n i z e d a s a m a i n s t r e a ms u b j e c t b y a n y o f t h e s e d i s c i p l i n e s . I n o l d e r s t u d i e s ,t e c h n i q u e s t o measure P a C C > 2 a n d d i a g n o s t i c m e t h o d s* F r o m t h e D e p a r t m e n t o f T h o r a c i c M e d i c i n e , K i n g s C o l l e g eS c h o o l o f M e d i c i n e a n d D e n t i s t r y , L o n d o n , U n i t e d K i n g d o m .M a n u s c r i p t r e c e i v e d F e b r u a r y 1 3 , 1 9 9 5 ; r e v i s i o n a c c e p t e d A u g u s tR e p r i n t r e q u e s t s : D r . G a r d n e r , D e p t . o f T h o r a c i c M e d i c i n e , K i n g sC o l l e g e S c h o o l o f M e d i c i n e & D e n t i s t r y , B e s s e m e r R d , L o n d o n S E 59PT

f o r o r g a n i c causes o f h y p e r v e n t i l a t i o n were r u d i m e n t a r y a n d t h e r e i s a d e a r t h o f d a t a b a s e d on m o d e r nq u a n t i t a t i v e c r i t e r i a . F a i l u r e t o d e f i n e p a t i e n t g r o u p sm a k e s i t d i f f i c u l t t o c o m p a r e s t u d i e s . T h e r e i s uncert a i n t y even a t a p h y s i o l o g i c l e v e l i n t h a t c o n t r o l m e c h a n i s m s d u r i n g s t i m u l a t e d b r e a t h i n g h a v e b e e n e x t e n s i v e l y s t u d i e d b u t t h e r e ar e f e w s t u d i e s a t r e s t a n d i nt h e h y p o c a p n i c r a n g e . 2 7I n t h e p r e s e n t c o n t e x t , i t i s r e l e v a n t t o a s k w h e t h e rt h i s t o p i c s h o u l d b e o f i n t e r e s t t o a c h e s t p h y s i c i a n , a n di n d e e d , i t i s r a r e l y f e a t u r e d i n r e s p i r a t o r y c o n f e r e n c e s .A t a p h y s i o l o g i c l e v e l , h y p e r v e n t i l a t i o n i s a p u r e r e s p i r a t o r y d i s t u r b a n c e . I t i s d i f f i c u l t t o g i v e r e l i a b l e f i g u r e sf o r i n c i d e n c e a n d b e c a u s e o f t h e p r o b l e m s o u t l i n e da b o v e , most p u b l i s h e d f i g u r e s s h o u l d b e t r e a t e d w i t hc a u t i o n . Y e t , p h y s i c i a n s i n most s p e c i a l t i e s , e s p e c i a l l yi n r e s p i r a t o r y m e d i c i n e a n d e m e r g e n c y m e d i c i n e , f r e q u e n t l y see p a t i e n t s i n whom h y p e r v e n t i l a t i o n cont r i b u t e s a t l e a s t i n p a r t t o t h e s y m p t o m s , a n d t h e yu s u a l l y f i n d i t d i f f i c u l t t o f o r m u l a t e p o s i t i v e d i a g n o s t i ca n d m a n a g e m e n t s t r a t e g i e s . D i s p r o p o r t i o n a t e b r e a t h l e s s n e s s i s common i n r e s p i r a t o r y p r a c t i c e a n d i s o f t e nr e g a r d e d a s s y n o n y m o u s w i t h h y p e r v e n t i l a t i o n . 2 8I n s e e k i n g a f r a m e o f r e f e r e n c e f o r t h e s e d i f f i c u l td i s o r d e r s , i t m a y b e t o o s i m p l i s t i c t o s e e k a s i n g l e d e f i n i t i o n o f h y p e r v e n t i l a t i o n s y n d r o m e . T h e term i s nowa p p l i e d i n so m a n y d i f f e r e n t c o n t e x t s t h a t i t s u s e f u l n e s sas a s i n g l e e n t i t y must b e q u e s t i o n e d . I t i s c e r t a i n l yi n c o r r e c t t o r e g a r d h y p e r v e n t i l a t i o n a s a s i m p l e mani f e s t a t i o n o f a n x i e t y a n d t h e r e i s o f t e n a c o m p l e xi n t e r a c t i o n a m o n g o r g a n i c r e s p i r a t o r y , p s y c h i a t r i c , a n dp h y s i o l o g i c d i s t u r b a n c e s . A t t h e s i m p l e s t l e v e l , h y p o c a p n i a i s a c l i n i c a l l a b o r a t o r y f i n d i n g i n d i c a t i v e o f ar a n g e o f d i s o r d e r s a n d s h o u l d b e t r e a t e d i n t h e samew a y a s a r a i s e d b l o o d ur ea or a l o w b l o o d g l u c o s e l e v e l .H o w e v e r , i t i s a l s o u s e f u l c l i n i c a l l y t o r e c o g n i z e a r a n g eo f s y n d r o m e s or c l i n i c a l s i t u a t i o n s i n w h i c h h y p e r v e n t i l a t i o n i s o f p r e d o m i n a n t c l i n i c a l i m p o r t a n c e . T h e s eo f t e n o v e r l a p w i t h , b u t ar e n o t n e c e s s a r i l y s y n o n y m o u sw i t h , s y n d r o m e s o f d i s p r o p o r t i o n a t e b r e a t h l e s s n e s s .T h e p h y s i o l o g i c b a s i s a n d c l i n i c a l c o m p l e x i t i e s w i l lnow b e d i s c u s s e d i n more d e t a i l . T h e r e ar e s t i l l m a n yu n k n o w n f a c t o r s . T h i s w i l l b e f o l l o w e d b y a d e s c r i p t i o no f t h e s y m p t o m s o f h y p o c a p n i a , t h e i n d i v i d u a l d i s o r d e r s t h a t c an i n d u c e h y p e r v e n t i l a t i o n , a r a n g e o f s y n -

5 1 6 R e v i e w sCopyright 1996 by American College of Chest Physicians

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3/21

30

2 0VENTILATION

/ m i n 1 0o 4 f ,

1 0 20 30 40E N D - T I D A L P C 0 2 mm H g

F i g u r e 1 . R e l a t i o n s h i p b e t w e e n m i n u t e v e n t i l a t i o n a n d P E T C O 2d u r i n g r e s t i n g b r e a t h i n g i n 8 n o r m a l s u b j e c t s ( c l o s e d s y m b o l s ) a n d1 6 p a t i e n t s w i t h c h r o n i c h y p e r v e n t i l a t i o n ( o p e n s y m b o l s , 2 runs p e rs u b j e c t ) . E a c h s y m b o l i s t h e mean o f 4 0 t o 1 0 0 b r e a t h s ( m o d i f i e df r o m G a r d n e r e t a l 3 1 ) .d r o m e s i n w h i c h , i n our e x p e r i e n c e , t h e s e f a c t o r s c anc o m b i n e t o i n d u c e h y p e r v e n t i l a t i o n i n c l i n i c a l p r a c t i c e ,a n d f i n a l l y a p r a c t i c a l s c h e m e f o r t r e a t m e n t o f t h e s ep a t i e n t s i n a c h e s t c l i n i c .

P h y s i o l o g yH y p e r v e n t i l a t i o n i s an a b n o r m a l i t y o f r e s p i r a t o r yc o n t r o l a n d i s d e f i n e d a s b r e a t h i n g i n excess o f m e t a b o l i c r e q u i r e m e n t s , i e , C O 2 p r o d u c t i o n . I n s u b j e c t sw i t h n o r m a l l u n g s , P a C 0 2 i s v e r y c l o s e t o a l v e o l a r P C O 2( P A C O 2 ) , w h i c h i n t u r n c an b e a p p r o x i m a t e d b y e n d -t i d a l P c o 2 ( P e t C 0 2 ) . A l v e o l a r v e n t i l a t i o n ( V A ) , C 0 2

p r o d u c t i o n ( V C O 2 ) , a n d P A C O 2 are l i n k e d t h r o u g h ane q u a t i o n w i t h t h e g e n e r a l f o r m o f V C O 2 a V A * P a C C > 2 /( b a r o m e t r i c p r e s s u r e - H 2 0 v a p o r p r e s s u r e ) , t h e g r a p h i cm a n i f e s t a t i o n o f w h i c h i s t h e " m e t a b o l i c h y p e r b o l a . " 2 9An e x a m p l e o f t h i s i s s h o w n i n F i g u r e 1 i n w h i c h meanm i n u t e v e n t i l a t i o n i s p l o t t e d a g a i n s t mean P E T C O 2 f o ra g r o u p o f p a t i e n t s w i t h c h r o n i c h y p e r v e n t i l a t i o n a n da m a t c h e d c o n t r o l g r o u p . T h i s r e l a t i o n s h i p i s o n l y t r u l ya p p l i c a b l e t o t h e s t e a d y - s t a t e a n d e q u i v a l e n t r e l a t i o n s h i p s h a v e n o t b e e n d e r i v e d f o r t h e n o n s t e a d y - s t a t e i nw h i c h C O 2 p r o d u c t i o n b y t h e t i s s u e s m a y d i f f e r f r o me x c r e t i o n a t t h e l u n g . T h i s r e l a t i o n s h i p i m p l i e s t h a t a ta c o n s t a n t V C O 2 an i n c r e a s e i n V A must b e a s s o c i a t e dw i t h a r e c i p r o c a l f a l l o f P a C C > 2 . S t a t e d a n o t h e r w a y ,P a C 0 2 b e l o w t h e n o r m a l r a n g e ( h y p o c a p n i a ) i s i n v a r i a b l y a s s o c i a t e d w i t h a r a i s e d V A a n d e x c e s s i v e r e s p i r a t o r y " d r i v e . " F o l l o w i n g 1 5 t o 2 0 min o f v o l u n t a r yh y p e r v e n t i l a t i o n t o b e l o w 2 0 mmH g , h y p o c a p n i a c anb e m a i n t a i n e d w i t h o n l y an o c c a s i o n a l l a r g e b r e a t h , 3 0b u t even c h r o n i c p e r s i s t e n t h y p e r v e n t i l a t i o n i s a s s o c i a t e d w i t h a t l e a s t 10% i n c r e a s e i n v e n t i l a t i o n . 3 1 I t i s af u n d a m e n t a l p r i n c i p l e i n d e a l i n g w i t h a n y h y p e r v e n t i -l a t i o n - r e l a t e d d i s o r d e r s t h a t t h e cause or causes o f t h i si n c r e a s e d d r i v e s h o u l d a l w a y s b e s o u g h t a n d d o c u m e n t e d .

C O 2 S t o r e sH y p e r v e n t i l a t i o n w a s h e s o u t C O 2 s t o r e s . T h e s e aree x t e n s i v e , a p p r o x i m a t i n g 1 2 0 L . O n l y a b o u t 1 0 0 mL o fC O 2 i s s t o r e d i n g a s p o c k e t s , t h e amount d e p e n d i n g ont h e p a r t i a l p r e s s u r e o f C O 2 a n d i t s p h y s i c a l s o l u t i o n .T h e r e m a i n d e r i s s t o r e d i n o t h e r b o d y c o m p a r t m e n t sb u t e s p e c i a l l y i n b o n e , s k e l e t a l m u s c l e , a n d v i s c e r a 3 2 ori t i s c h e m i c a l l y b o u n d a s b i c a r b o n a t e . A l l t i s s u e s h a v et h e i r own C O 2 d i s s o c i a t i o n curves. C O 2 i s e a s i l yt r a n s f e r r e d t o a n d f r o m most b o d y c o m p a r t m e n t s , b u tb o n e s t o r e s ar e n o t e a s i l y a v a i l a b l e .A t t h e b e g i n n i n g o f v o l u n t a r y h y p e r v e n t i l a t i o n ,t h e r e i s a d r a m a t i c r e d u c t i o n o f P A C O 2 . C O 2 e l i m i n a t i o n a t t h e l u n g s i n c r e a s e s i n i t i a l l y b u t d r o p s w i t h i n af e w m i n u t e s a n d e v e n t u a l l y r e t u r n s t o n o r m a l a s b a l ance i s r e s t o r e d b e t w e e n w a s h o u t f r o m t h e l u n g s a n dt i s s u e s . 3 2 " 3 5 T h e r e i s a 5 0 % c h a n g e i n C O 2 c o n t e n t o ft h e b o d y w i t h i n 5 m i n o f a c h a n g e i n P A C O 2 . T h e r e i si n i t i a l l y o n l y m o b i l i z a t i o n o f C O 2 f r o m t h e a l v e o l a r g a sa n d b l o o d i n t h e p u l m o n a r y v e i n s , l e f t s i d e o f t h e h e a r t ,a n d t h e f i r s t p a r t o f t h e s y s t e m i c c i r c u l a t i o n . T h e p e r i p h e r a l s t o r e s ar e more d e p e n d e n t on t h e venous P C O 2a n d c a r d i a c o u t p u t . T h e C O 2 c o n t e n t o f b l o o d c an b ec a l c u l a t e d v i a t h e C O 2 d i s s o c i a t i o n curve t h a t i s l i n e a rb e t w e e n a P C O 2 o f 3 0 a n d 8 0 mmH g , b u t i t i s mored i f f i c u l t t o e s t i m a t e c h a n g e s i n o t h e r b o d y s t o r e s b e cause v a r i o u s o r g a n s d o n o t a l l e q u i l i b r a t e a t t h e samer a t e .A c i d - B a s e C h a n g e s

P a C 0 2 i s l i n k e d t o a r t e r i a l h y d r o g e n a n d b i c a r b o n a t e i o n s v i a t h e H e n d e r s o n - H a s s e l b a c h e q u a t i o n . A f t e rt h e s t a r t o f f o r c e d b r e a t h i n g , t h e r e i s a r a p i d o n s e t o fr e s p i r a t o r y a l k a l o s i s w i t h r i s e o f a r t e r i a l b l o o d p H , f a l lo f H + , a n d f a l l o f b i c a r b o n a t e . p H b e g i n s t o r i s e w i t h i n5 t o 2 0 s , r e a c h e s a maximum a t 1 0 t o 1 5 m i n , a n d f a l l st o n o r m a l w i t h i n 5 min o f t h e e n d o f f o r c e d b r e a t h i n g . 3 3 " 3 5 A r b u s e t a l 3 6 d o u b l e d v e n t i l a t i o n i n a n e s t h e -t i s e d p a t i e n t s f o r u p t o 9 0 m i n . A c i d - b a s e a d j u s t m e n t swere c o m p l e t e w i t h i n a f e w m i n u t e s , t h e r e l a t i o n b e tween H+ a n d P C O 2 b e i n g d e s c r i b e d a s H + = 0 . 7 4P C O 2 + 1 0 . 4 w i t h H+ i n n m o l / L a n d P a C C > 2 i n mm H g .A t a P a C C > 2 o f 4 5 mm H g , s i g n i f i c a n c e b a n d s were 4 0 . 4t o 4 6 . 7 n m o l / L f o r H + , 7 . 3 9 t o 7 . 3 3 f o r p H , 2 4 . 2 t o 2 7 . 9f o r H C O 3 " , a n d a t a P a C 0 2 o f 1 5 mm H g , t h e y were1 8 . 3 t o 2 4 . 6 n m o l / L f o r H + , 7 . 7 4 t o 7 . 6 1 f o r p H , a n d1 5 . 3 t o 2 0 . 5 f o r H C O 3 - .I n t h e b r a i n , a n d p r o b a b l y i n o t h e r o r g a n s , r e s p i r a t o r y a l k a l o s i s i s a s s o c i a t e d w i t h g r e a t l y i n c r e a s e d g l y -c o l y s i s w i t h p r o d u c t i o n o f l a c t i c a n d o t h e r a c i d s . 3 7 R e n a l c o m p e n s a t i o n i s p r o b a b l y u n i m p o r t a n t i n h u m a n sa n d a c i d - b a s e a d a p t a t i o n i s u n l i k e l y t o m a k e a m a j o rc o n t r i b u t i o n t o t h e m a i n t e n a n c e o f h y p o c a p n i a i n p a t i e n t s w i t h c h r o n i c h y p e r v e n t i l a t i o n . I n t h e s t u d y o fA r b u s e t a l , 3 6 mean r e n a l e x c r e t i o n o f H C C > 3 ~ d u r i n g

CHEST / 1 0 9 / 2 / F E B R U A R Y , 1 9 9 6 5 1 7Copyright 1996 by American College of Chest Physicians

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b o t h t h e c o n t r o l a n d h y p o c a p n i c p e r i o d wa s l e s s t h a n1 m E q / h w i t h t h e c o n c l u s i o n t h a t t h e c o n t r i b u t i o n o fr e n a l H C 0 3 - l o s s t o t h e r e d u c t i o n i n e x t r a c e l l u l a rH C O 3 " s t o r e s was n e g l i g i b l e , one t h i r d o f t h i s r e d u c t i o n b e i n g t h e r e s u l t o f b u f f e r i n g b y h e m o g l o b i n a n dt h e r e m a i n d e r b u f f e r i n g b y t h e t i s s u e s . R e n a l e x c r e t i o nwas l i t d e i n f l u e n c e d b y a v a r i e t y o f p r e m e d i c a t i o n s ,o x y g e n m i x t u r e s , a n d a n e s t h e t i c s . I n more c h r o n i ch y p e r v e n t i l a t i o n , 3 8 h y p o c a p n i c - i n d u c e d s u p p r e s s i o no f r e n a l a c i d e x c r e t i o n was i n d e p e n d e n t o f p l a s m a H+a n d H C 0 3 - a n d wa s t o o s m a l l t o c o m p e n s a t e f o r t h ed i r e c t a l k a l e m i c e f f e c t o f h y p o c a p n i a . T h i s was i nc o n t r a s t t o s t u d i e s i n d o g s 3 9 i n w h i c h r e n a l a c i d secret i o n b y d i s t a l t u b u l a r e x c h a n g e w i t h s o d i u m a l m o s tc o m p l e t e l y c o r r e c t e d t h e a r t e r i a l p H w h e n P a C 0 2 wasr e d u c e d t o 1 5 mm H g b y s t e p w i s e r e d u c t i o n o fi n s p i r e d o x y g e n over 4 t o 5 d a y s i n an e n v i r o n m e n t a lc h a m b e r .T h e u n i m p o r t a n c e o f r e n a l c o m p e n s a t i o n i n h u m a n sh a s b e e n c o n f i r m e d b y s t u d i e s a t h i g h a l t i t u d e . D u r i n g8 d a y s s t a y a t a l t i t u d e b y a g r o u p o f l o w l a n d e r s , cere-b r o s p i n a l f l u i d ( C S F ) H C 0 3 ~ r e m a i n e d 4 t o 5 m E ql o w e r t h a n c o n t r o l v a l u e s a f t e r 1 t o 2 d a y s a t a l t i t u d ea n d r e n a l b u f f e r b a s e e x c r e t i o n r e d u c e d b l o o d s t a n d a r d b i c a r b o n a t e o n l y b y 1 m E q / L a f t e r 1 w e e k . 4 0 As i m i l a r f a i l u r e o f r e v e r s a l o f r e s p i r a t o r y a l k a l o s i s h a sb e e n r e p o r t e d i n h u m a n s over a 5 - d a y s t a y a t 1 2 , 0 0 0f e e t 4 1 a n d over a m o n t h a t a s i m u l a t e d a l t i t u d e o f u pt o 2 2 , 0 0 0 f e e t . 4 2 CSF c o m p e n s a t i o n i s a l s o i n c o m p l e t e ,CSF p H i n h u m a n s s h o w i n g o n l y 50% c o m p e n s a t i o na t t h e e n d o f 2 6 h o f v o l u n t a r y o v e r b r e a t h i n g t o aP a C 0 2 o f 3 1 mm H g . 4 3 M o r e c o m p l e t e c o m p e n s a t i o no f p H o c c u r r e d i n l o n g - t e r m r e s i d e n t s a t an a l t i t u d e o f3 , 9 0 0 t o 4 , 5 0 0 m 4 4C o n t r o l o f B r e a t h i n gO b v i o u s i n c r e a s e i n e i t h e r r a t e or d e p t h o f r e s p i r a t o r y movement i s n o t n e c e s s a r i l y a s s o c i a t e d w i t hh y p o c a p n i a . F o r e x a m p l e , a l a r g e t i d a l v o l u m e c an b ea s s o c i a t e d w i t h a s l o w r e s p i r a t o r y f r e q u e n c y w i t h noc h a n g e i n V A or P a C 0 2 - I n p a n t i n g , a l o w t i d a l v o l u m em a y b e a s s o c i a t e d w i t h f a i l u r e t o c l e a r t h e d e a d - s p a c e 4 5l e a d i n g t o C O 2 r e t e n t i o n w h i c h m a i n t a i n s P a C 0 2 a tn o r m a l l e v e l s d e s p i t e a c l i n i c a l l y d r a m a t i c i n c r e a s e i ne x p i r e d v e n t i l a t i o n . S u c h r e s p o n s e s h a v e b e e n d o c u m e n t e d i n a p a t i e n t w i t h a c e n t r a l n e u r o n a l l e s i o n , 4 6a n d we h a v e seen a n u m b e r o f d e m e n t e d p a t i e n t s w i t hr a p i d s h a l l o w r e s p i r a t i o n b u t a n o r m a l P a C 0 2 -T h e c h e m o r e c e p t o r s w i l l p r e v e n t P a C 0 2 f r o m r i s i n gm u c h a b o v e t h e r e s t i n g v a l u e b u t v a g a l r e f l e x e s arep r o b a b l y i n a c t i v e i n h y p o c a p n i a 4 1 a n d t h e r e a p p e a rt o b e f e w f e e d b a c k c o n t r o l m e c h a n i s m s t o p r e v e n tP a C 0 2 f r o m f a l l i n g d u r i n g h y p e r v e n t i l a t i o n . H o w e v e r ,t h e r e r e m a i n m a n y u n c e r t a i n t i e s a b o u t c o n t r o l o fb r e a t h i n g i n t h e h y p o c a p n i a r a n g e . 2 7 A t t h e l o w e r e n d

o f t h e u b i q u i t o u s v e n t i l a t i o n vs P E T C O 2 r e s p o n s e curvet o i n s p i r e d C O 2 , 4 8 t h e r e i s a d i s c o n t i n u i t y or " d o g - l e g , "s u g g e s t i n g r e d u c t i o n or c e s s a t i o n o f c h e m o r e c e p t o ra c t i v i t y . 4 9 ' 5 0 I n n o r m a l q u i e t u n s t i m u l a t e d b r e a t h i n g ,v e n t i l a t i o n i s l o c a t e d j u s t on t h e d o g - l e g near t h ej u n c t i o n o f t h e d i s c o n t i n u i t y w i t h t h e s t e e p l y s l o p i n gp a r t o f t h e c u r v e , 5 1 c o n s i s t e n t w i t h a c h e m o r e c e p t o rt h r e s h o l d j u s t b e l o w r e s t i n g . T h e p e r i p h e r a l c h e m o r e c e p t o r t h r e s h o l d i s p r o b a b l y l o w e r t h a n t h a t f o r t h ec e n t r a l c h e m o r e c e p t o r s . 5 2 , 5 3A p n e a u s u a l l y occurs d u r i n g p a s s i v e h y p e r v e n t i l a t i o n b o t h i n a n i m a l s 5 4 a n d i n h u m a n s d u r i n g s t a t e s o fa l t e r e d c o n s c i o u s n e s s a n d s l e e p . 5 5 , 5 6 T h e r e i s moreu n c e r t a i n t y i n t h e a w a k e s t a t e i n h u m a n s w i t h d e s c r i p t i o n s o f a p n e a , 5 7 " 5 9 h y p e r p n e a , 6 0 " 6 2 a n d n o r m a l b r e a t h i n g 5 5 ' 6 3 ' 6 ^ f o l l o w i n g v o l u n t a r y o v e r b r e a t h i n g .We a n a l y z e d a p n e i c p a u s e s o f g r e a t e r t h a n 6 s i nn o r m a l s u b j e c t s d u r i n g v a r i o u s r a t e s o f r e c o v e r y o fP E T C O 2 f o l l o w i n g v o l u n t a r y o v e r b r e a t h i n g i n m i l dh y p e r o x i a . 6 5 I m m e d i a t e l y a f t e r h y p e r v e n t i l a t i o n , t h e r ewas h e i g h t e n e d b r e a t h i n g l a s t i n g a b o u t a m i n u t e cons i s t e n t w i t h a f t e r d i s c h a r g e 6 6 A p n e i c p a u s e s t h e n occ u r r e d i n t e r m i t t e n t l y f o r u p t o 1 0 min u n t i l P E T C O 2rose a b o v e a t h r e s h o l d , on a v e r a g e 3 . 4 mm H g b e l o wr e s t i n g P E T C O 2 , w h e n n o r m a l b r e a t h i n g s u d d e n l y r e s u m e d . T h e occurrence a n d l e n g t h o f t h e p a u s e s werei n d e p e n d e n t o f t h e r a t e o f r e c o v e r y o f P E T C O 2 . T h e s ef i n d i n g s s u g g e s t t h a t i n a w a k e h u m a n s , t h e r e i s nof e e d b a c k c h e m i c a l c o n t r o l f r o m t h e i n t r a c r a n i a lc h e m o r e c e p t o r s w h e n P E T C O 2 i s more t h a n a b o u t 4mm H g b e l o w r e s t i n g , b r e a t h i n g p r o b a b l y b e i n gd e t e r m i n e d b y t h e r e s p i r a t o r y c e n t e r s m o d i f i e d b y" f e e d f o r e w a r d " d r i v e s f r o m t h e c o r t e x a n d p e r i p h e r y . 2 7S i m i l a r c o n c l u s i o n s were a r r i v e d a t b y d e B a c k e r e t a l 5 9w h o s t u d i e d t h e e f f e c t on b r e a t h i n g p a t t e r n o f venousC O 2 u n l o a d i n g b y h e m o d i a l y s i s , a n d b y Cummin e t a l 6 2w h o f o u n d r e d u c t i o n o f v e n t i l a t o r y r e s p o n s e s t o t r a n s i e n t p u l s e s o f C O 2 w h e n P E T C O 2 was l o w f o l l o w i n gv o l u n t a r y h y p e r v e n t i l a t i o n i n n o r m a l h u m a n s . Wec o u l d n o t c o n f i r m p r e v i o u s r e p o r t s t h a t a p n e a s oc c u r r e d o n l y i n s u b j e c t s f a m i l i a r w i t h t h e e q u i p m e n tu s e d t o measure b r e a t h i n g , 5 8 some o f our s u b j e c t sc o n s i s t e n d y s h o w i n g f e w a n d o t h e r s m a n y a p n e a s ,s u g g e s t i n g t h a t t h e t e n d e n c y t o a p n e i c p a u s e s i s ac h a r a c t e r i s t i c o f t h e s u b j e c t .A f t e r d i s c h a r g e or " p o s t s t i m u l u s p o t e n t i a t i o n " i m m e d i a t e l y f o l l o w i n g v o l u n t a r y o v e r b r e a t h i n g h a s b e e nd e s c r i b e d f o l l o w i n g a r a n g e o f s t i m u l i 6 6 a n d a f t e r v o l u n t a r y o v e r b r e a t h i n g a t a c o n s t a n t P E T C O 2 6 1 T h e r o l eo f a f t e r d i s c h a r g e i n p e r p e t u a t i o n o f h y p o c a p n i a i n p a t i e n t s w i t h h y p e r v e n t i l a t i o n r e m a i n s u n c e r t a i n .

S y m p t o m sH y p e r v e n t i l a t i o n i s u n l i k e l y t o b e a p r o b l e m t o ap a t i e n t u n l e s s i t r e s u l t s i n s y m p t o m s , a n d t h e w a y i nw h i c h t h e s e s y m p t o m s are i n d u c e d m a y h o l d t h e k e y

5 1 8 R e v i e w s

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