Rev. Physiol. Biochem. Pharmacol., Vol.

86
© by Springer-Verlag 1979

Mechanisms of Gas Exchange in Bird Lungs

PETER SCHEID *

Contents
I. I n t r o d u c t i o n . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
II. A n a t o m y o f t h e R e s p i r a t o r y S y s t e m . . . . . . . . . . . . . . . . . . . . . . . . 138
A. L o c a t i o n o f Lungs and Air Sacs . . . . . . . . . . . . . . . . . . . . . . . . . 140
B. Bronchial S y s t e m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140
C. Microscopic S t r u c t u r e o f t h e P a r a b r o n c h i . . . . . . . . . . . . . . . . . . . 142
III. V e n t i l a t i o n o f R e s p i r a t o r y Tract . . . . . . . . . . . . . . . . . . . . . . . . . . 143
A. V e n t i l a t i o n o f Air Sacs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
B. Air F l o w P a t t e r n in R e s p i r a t o r y Tract . . . . . . . . . . . . . . . . . . . . . 144
C. M e c h a n i s m s Involved in R e c t i f y i n g Lung Air F l o w . . . . . . . . . . . . . 147
D. Significance o f U n i d i r e c t i o n a l F l o w . . . . . . . . . . . . . . . . . . . . . . 148
IV. Gas T r a n s p o r t P r o p e r t i e s o f Blood . . . . . . . . . . . . . . . . . . . . . . . . . 149
A. O x y g e n Dissociation Curve . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
B. C a r b o n D i o x i d e Dissociation Curve . . . . . . . . . . . . . . . . . . . . . . . 152
C. Effective Dissociation Curves . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
V. Basic C o n c e p t s for P a r a b r o n c h i a l Gas E x c h a n g e . . . . . . . . . . . . . . . . 153
A. Basic P a r a m e t e r s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
B. Structural A r r a n g e m e n t o f t h e P a r a b r o n c h i a l Lung . . . . . . . . . . . . 155
VI. Cross-Current M o d e l for Gas E x c h a n g e in t h e Ideal P a r a b r o n c h u s . . . . . 156
A. A s s u m p t i o n s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 156
B. Q u a n t i t a t i v e R e l a t i o n s h i p s . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158
C. S o m e Peculiarities o f t h e Cross-Current S y s t e m . . . . . . . . . . . . . . . 159
VII. A p p l i c a b i l i t y o f Cross-Current M o d e l t o t h e P a r a b r o n c h i a l Lung . . . . . . 164
A. D i f f u s i o n in Air Capillaries . . . . . . . . . . . . . . . . . . . . . . . . . . . . t 64
B. S t e a d y S t a t e C o n d i t i o n s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167
C. Linear Dissociation Curves . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168
D. I n h o m o g e n e i t i e s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169
E. N e o p u l m o and Gas C o m p o s i t i o n in Caudal Air Sacs . . . . . . . . . . . . 172
VIII. M e a s u r e m e n t o f Gas E x c h a n g e U n d e r Various C o n d i t i o n s . . . . . . . . . . 173
A. At Rest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . t73
B. Resting C o n d i t i o n s with Elevated Gas E x c h a n g e R a t e s . . . . . . . . . . 174
C. In R u n n i n g Birds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175
D. During Flight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175
E. During Diving . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176
F. Gas E x c h a n g e and Heat E x c h a n g e . . . . . . . . . . . . . . . . . . . . . . . . 177
References ............................................. 178

* A b t e i l u n g Physiologie, M a x - P l a n c k - I n s t i t u t ffir e x p e r i m e n t e l l e M e d i z i n , H e r m a n n -
Rein-StrafSe 3, D - 3400 G 6 t t i n g e n , F R G .
138 P. Scheid:

I. Introduction

Scientific descriptions of the anatomy and physiology of avian respiration

date back to the sixteenth century (cf. Duncker, 1971). While most of this
work remained qualitative and many topics controversial, application in
recent years of modern techniques in a number of laboratories has helped
to obtain a much clearer picture of the structure and function of the respi-
ratory system in birds. It is not the intention of this article to provide a
comprehensive review of the physiology of avian respiration; some excel-
lent textbook articles have recently appeared (e.g., Fedde, 1976). Rather
it will highlight mechanisms of gas exchange in the avian parabronchial
lung for two reasons. First, considerable progress has been achieved in our
understanding in this area, yet many recent reviews have omitted its de-
scription; secondly, the physiology of gas exchange in birds has occupied
the author's own interest for some time.
In thus restricting the scope of the report, many topics which are close-
ly related to avian respiration and which have progressed considerably over
the past years are omitted. The interested reader will find, however, a num-
ber of reviews on several aspects of avian respiration (Romanoff and Ro-
manoff, 1949; Rornanoff, 1967; King and Molony, 1971; Jones and Jo-
hansen, 1972; Jones, 1972; Lasiewski, 1972; Calder and King, 1974; Free-
man and Vince, 1974; Dawson, 1975; De]ours, 1975; Schmidt-Nielsen,
1975 ; Fedde, 1976;Jones, 1976;Piiper and Scheid, 1977;Bouverot, 1978;
White, 1978) and may consult the proceedings of recent conferences
(Piiper, 1972, 1978a;Scheid, 1974). References to the many papers on gas
exchange during development of birds may be obtained from Piiper
(1978a).
This review is mainly written to address physiologists who are familiar
with the general principles of respiratory physiology in mammals, and will
therefore concentrate main'ly on those aspects in respect of which birds
differ from mammals. Emphasis is, moreover, laid on those factors which
appear to be common for all birds; species differences will generally not
be addressed. Values for physiologic parameters will only be quoted as far
as they help the understanding o f basic principles.
The anatomical terminology used is adopted from King (1979); for
physiologic terminology, the proposals of Piiper et al. (1971) are followed.

II. Anatomoy of the Respiratory System

Application of modern techniques has led to a considerable advancement

in our understanding of the structure of the avian respiratory system. Of