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Nunn and Lumb’s Applied
Respiratory Physiology
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Nunn and Lumb’s Applied
Respiratory Physiology

NINTH EDITION
Andrew Lumb MB BS FRCA
Consultant Anaesthetist,
St James’s University Hospital,
Leeds, UK
Honorary Clinical Associate Professor,
University of Leeds,
Leeds, UK

Caroline Thomas BSc MBChB MRCS(Ed) FRCA

Consultant Anaesthetist,
St James’s University Hospital,
Leeds, UK
Honorary Senior Clinical Lecturer,
University of Leeds,
Leeds, UK

For additional online content visit ExpertConsult.com

Elsevier

Copyright © 2021, Elsevier Limited. All rights reserved.

First edition 1969
Second edition 1977
Third edition 1987
Fourth edition 1993
Fifth edition 1999
Sixth edition 2005
Seventh edition 2010
Eighth edition 2017

The right of Andrew Lumb and Caroline Thomas to be identified as authors of this work has been asserted by
them in accordance with the Copyright, Designs and Patents Act 1988.

No part of this publication may be reproduced or transmitted in any form or by any means, electronic or
mechanical, including photocopying, recording or any information storage and retrieval system, without
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This book and the individual contributions contained in it are protected under copyright by the Publisher
(other than as may be noted herein).

Notices

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using any information, methods, compounds or experiments described herein. Because of rapid advances
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To Lorraine, Emma and Jenny (AL)
and
Simon, Martha and Ted (CT)
Foreword
It is truly an honor to be asked to write the Foreword to
this newest edition of the landmark text in Respiratory
Physiology. For 50 years this has been the acknowledged
source for education and reference for physicians, scientists
and students whose clinical practice and curiosity involve
the respiratory system. The first edition of the pioneering
text Nunn’s Applied Respiratory Physiology was published
in 1969 with Dr. John Nunn as the Editor. After the
4th edition, the torch was passed to Dr. Andrew Lumb who
maintained the excellent standards and expanded the focus
through the 5th to 8th editions. Now with this 9th edition,
the text has become Nunn and Lumb’s Applied Physiology
and Dr. Lumb has shared the authorship with Dr. Caroline
Thomas.
I have been told that an efficient method of teaching is
to: “Say what you’re going to say, say it, and then repeat
what you said”. The Authors follow this scheme using Key
Points at the beginning of each chapter then following at the
end with an online Summary to reinforce the educational
material. Also, they have maintained the use of a large num-
ber of clear and memorable figures and diagrams. I believe
their frequent use of electrical or hydrostatic models to ex-
plain the underlying physiology is one of the elements that
makes this text so useful to learners.
The Editors have continued the logical progression of
this text, refined in previous editions, beginning with Part 1
on Basic Principles and then building on this foundation to
develop Part 2 on Applied Physiology and finally to extend
to Part 3 on The Physiology of Pulmonary Disease. Part 1
vi
explains respiration starting with the relevant anatomy and
progressing through lung mechanics, pulmonary circulation
and gas exchange to cellular respiration. The final chapter
deals with the non-respiratory functions of the lungs. Part 2
then discusses several specific clinical conditions such as
pregnancy, paediatrics, sleep, extremes of barometric pres-
sure, anaesthesia and air pollution. A very well written and
useful addition to the is 9th edition is Chapter 15 on Obesity.
Obesity has become an epidemic in the developed world and
the ventilatory management of obese patients during mini-
mally invasive surgery is currently a significant clinical issue.
Part 3 on the Physiology of Pulmonary Disease presents
chapters on a range of very important clinical topics from
ventilatory failure to pulmonary surgery. Of particular note
are the up-to-date chapters on acute lung injury and respira-
tory support. This is a rapidly evolving area and recent devel-
opments are presented clearly and concisely.
I find Respiratory Physiology a difficult topic. I tell
students that if the answer is simple, you don’t understand
the question. Yet in the practice of Anaesthesia we are forced
to make important decisions on how to manage the airway,
gas exchange and respiratory mechanics of diverse individu-
al patients on a daily basis. This text helps walk us through
many of these complex clinical decisions. I would like to
thank and congratulate Drs. Lumb and Thomas for their
excellent work.
Peter Slinger
Professor of Anesthesia , University of Toronto
Preface to the Ninth Edition
Over the past five decades Nunn’s Applied Respiratory Physi-
ology has developed into a renowned textbook on respira-
tion, providing both physiologists and clinicians with a
unique fusion of underlying principles and their applica-
tions. After writing four editions, Dr John Nunn retired
in 1991, and a new author was required. As Dr Nunn’s
final research fellow in the Clinical Research Centre in
Harrow, AL was honoured to be chosen as his successor. AL
has now also completed four editions and has chosen a suc-
cessor to lead the project into the future whilst maintaining
the fundamental ethos of the book. As practising clinicians
with a fascination for physiology, the authors of the ninth
edition have again focussed on combining a clear, logical
and comprehensive account of basic respiratory physiology
with a wide range of applications, both physiological and
clinical. This approach acknowledges the popularity of
the book amongst doctors from many medical specialities,
but also provides greater insight into the applications
of respiratory physiology to readers with a scientific back-
ground. The clinical chapters of Part 3 are not intended
to be comprehensive reviews of the pulmonary diseases
considered, but rather to provide a detailed description
of physiological changes, accompanied by a brief account
of the clinical features and treatment of the disease.
In this edition, the number of references provided has
been reduced by around a third in recognition of the ease
with which online searches may now be performed. Refer-
ences retained are either historical or seminal papers, or re-
cent high-quality publications. Key references are identified
by bold type in the reference list following each chapter.
These highlighted references either provide outstanding re-
cent reviews of their subject or describe research that has
made a significant impact on the topic under consideration.
Advances in respiratory physiology since the last edition
are too numerous to mention individually. Appreciation of
the impact of air quality on the lungs continues to develop,
and there is increasing awareness of the global health burden
of pollution. Chapter 20 has been updated in recognition
of this and the publication of new worldwide guidelines
on pollution levels. The harmful effects of hyperoxia are be-
coming more accepted in clinical practice, and the physio-
logical mechanisms of these are described in Chapter 25.
There is much recent literature on this topic, exemplified
by the U-shaped curve of oxygen levels and mortality in
critical care patients (see Fig. 25.6). The optimal strategy for
artificial ventilation in healthy lungs, for example, during
anaesthesia, remains disputed; the section in Chapter 21 has
been updated to reflect the physiological effects of these
strategies, for example, the recent focus on driving pressure
as the potentially damaging component. The role of intra-
operative ventilation strategy in the prevention of postop-
erative pulmonary complications is becoming more clear.
In keeping with its increasing worldwide prevalence,
the effects of severe obesity on the respiratory system are
now brought together into a new chapter for this edition.
Chapter 15 covers the predictable aspects of obesity on res-
piration, such as the effect of the mass of the chest wall and
abdomen on lung mechanics and lung volumes. Less pre-
dictable topics include the effects of obesity hormones on
respiratory control. The chapter also covers the impact of
childhood obesity on lung development, which may lead to
lung dysanapsis in which airway and gas exchange tissues
grow disproportionately.
For this edition the book is printed with a larger page
format to improve the clarity of the figures and tables, and
remains available in both print and electronic format. This
allows readers wishing to dip into the book access to chapter
summaries or individual chapters. For those who own a
print copy, online access is automatically available. This
content includes additional chapters and self-assessment
material, useful for students approaching exams and, new
for this edition, a series of 24 mini-lectures by AL to en-
hance the information provided in print.
We wish to thank the many people who have helped with
the preparation of the book at Elsevier and our colleagues
who have assisted our acquisition of knowledge in subjects
not so close to our own areas of expertise. We are indebted
to Professor Peter Slinger for his kind words in the Foreword
and would like to thank Drs B. Oliver, J. Black, K. McKay
and P. Johnson for permission to use the images in Figure
28.3. Last, but by no means least, we thank our families for
their continuing encouragement and for tolerating preoc-
cupied and reclusive parents/spouses for so long. AL’s
daughter Jenny, when aged 5, often enquired about his ac-
tivities in the study, until one evening she nicely summa-
rized the years of work by confidently stating that ‘if you
don’t breathe, you die’. So what were the other 423 pages
about?
Andrew Lumb and Caroline Thomas
Leeds 2019
vii
Contents

Foreword by Professor Peter Slinger, vi 24. Anaemia, 279

Preface, vii 25. Oxygen Toxicity and Hyperoxia, 285
26. Comparative Respiratory Physiology, 299
Part 1: Basic Principles
1. Functional Anatomy of the Part 3: Physiology of Pulmonary Disease
Respiratory Tract, 2
27. Ventilatory Failure, 316
2. Elastic Forces and Lung Volumes, 14
28. Airways Disease, 324
3. Respiratory System Resistance, 27
29. Pulmonary Vascular Disease, 339
4. Control of Breathing, 42
30. Diseases of the Lung Parenchyma
5. Pulmonary Ventilation, 59 and Pleura, 349
6. The Pulmonary Circulation, 73 31. Acute Lung Injury, 365
7. Distribution of Pulmonary Ventilation and 32. Respiratory Support and Artificial
Perfusion, 88 Ventilation, 375
8. Diffusion of Respiratory Gases, 111 33. Pulmonary Surgery, 398
9. Carbon Dioxide, 122
10. Oxygen, 136 Appendix A Physical Quantities and Units
of Measurement, 412
11. Nonrespiratory Functions of the Lung, 164
Appendix B The Gas Laws, 415
Part 2: Applied Physiology Appendix C Conversion Factors for Gas Volumes,
417
12. Pregnancy, Neonates and Children, 175 Appendix D Symbols and Abbreviations, 418
13. Exercise, 183 Appendix E Mathematical Functions Relevant to
14. Sleep, 191 Respiratory Physiology, 419
15. Obesity, 199
16. High Altitude and Flying, 205 Online Content
17. High Pressure and Diving, 218
34. The Atmosphere
18. Respiration in Closed Environments and
Space, 225 35. The History of Respiratory Physiology
19. Drowning, 233 Test your Knowledge
20. Smoking and Air Pollution, 236 Access via Expert Consult – see inside front cover for instructions.
21. Anaesthesia, 244 Index, 425
22. Changes in the Carbon Dioxide Partial
Pressure, 268
23. Hypoxia, 273

viii
Videos Table of Contents

1. Elastic resistance of the respiratory system

2. Elastic resistance from surface forces
3. Compliance and static lung volumes
4. Respiratory system resistance
5. Active control of airway calibre
6. Ventilation
7. Lung perfusion
8. Ventilation/perfusion relationships
9. Dead space and shunt
10. Carbon dioxide carriage in blood
11. Oxygen cascade
12. Oxygen carriage in blood
13. Airway lining fluid
14. Defence against inhaled substances
15. Breathing during sleep
16. Breathing at altitude
17. Diving
18. Respiration and microgravity
19. Breathing in closed environments
20. Long term space travel
21. Physiology of general anaesthesia
22. Ventilation in theatre
23. Oxygen toxicity
24. Physiology of hyperoxia

ix
This page intentionally left blank
 PART I

Basic Principles

1
1
Functional Anatomy
of the Respiratory Tract
K E Y P O I N TS
• In addition to conducting air to and from the lungs, the
nose, mouth and pharynx have other important functions
including speech, swallowing and airway protection.
• Starting at the trachea, the airway divides about 23 times,
terminating in an estimated 30 000 pulmonary acini, each
containing more than 10 000 alveoli.
T
his chapter is not a comprehensive account of respi-
ratory anatomy but concentrates on those aspects
that are most relevant to an understanding of func-
tion. The respiratory muscles are covered in Chapter 5.
Mouth, Nose and Pharynx
Breathing is normally possible through either the nose or
the mouth, the two alternative air passages converging in
the oropharynx. Nasal breathing is the norm and has two
major advantages over mouth breathing: filtration of par-
ticulate matter by the vibrissae hairs and better humidifica-
tion of inspired gas. Humidification by the nose is highly
efficient because the nasal septum and turbinates increase
the surface area of mucosa available for evaporation and
produce turbulent flow, increasing contact between the
mucosa and air. However, the nose may offer more resis-
tance to airflow than the mouth, particularly when ob-
structed by polyps, adenoids or congestion of the nasal
mucosa. Nasal resistance may make oral breathing obliga-
tory, and many children and adults breathe only or partly
through their mouths at rest. With increasing levels of exer-
cise in normal adults, the respiratory minute volume in-
creases, and at a level of around 35 L.min21 the oral airway
comes into play. Deflection of gas into either the nasal or
the oral route is under voluntary control and accomplished
with the soft palate, tongue and lips. These functions
are best considered in relation to a midline sagittal section
(Fig. 1.1).
Figure 1.1, A, shows the normal position for nose breath-
ing: the mouth is closed by occlusion of the lips, and the
2
• The alveolar wall is ideally designed to provide the minimal
physical barrier to gas transfer, whilst also being structurally
strong enough to resist the large mechanical forces applied
to the lung.
tongue is lying against the hard palate. The soft palate is
clear of the posterior pharyngeal wall. Figure 1.1, B, shows
forced mouth breathing, for instance when blowing through
the mouth without pinching the nose. The soft palate be-
comes rigid and is arched upwards and backwards by con-
traction of tensor and levator palati to lie against a band of
the superior constrictor of the pharynx known as Passavant’s
ridge, which, together with the soft palate, forms the
palatopharyngeal sphincter. Note also that the orifices of
the pharyngotympanic (Eustachian) tubes lie above the pal-
atopharyngeal sphincter and can be inflated by the subject
only when the nose is pinched. As the mouth pressure is
raised, this tends to force the soft palate against the posterior
pharyngeal wall to act as a valve. The combined palatopha-
ryngeal sphincter and valvular action of the soft palate is
very strong and can easily withstand mouth pressures in
excess of 10 kPa (100 cmH2O).
Figure 1.1, C, shows the occlusion of the respiratory tract
during a Valsalva manoeuvre. The airway is occluded at
many sites: the lips are closed, the tongue is in contact with
the hard palate anteriorly, the palatopharyngeal sphincter is
tightly closed, the epiglottis is in contact with the posterior
pharyngeal wall, and the vocal folds are closed, becoming
visible in the midline in the figure.
During swallowing the nasopharynx is occluded by con-
traction of both tensor and levator palati. The larynx is ele-
vated 2 to 3 cm by contraction of the infrahyoid muscles,
stylopharyngeus and palatopharyngeus, coming to lie under
the epiglottis. In addition, the aryepiglottic folds are ap-
proximated, causing total occlusion of the entrance to the
larynx. This extremely effective protection of the larynx is

NC
T 6
L
E 4
SP
A
B
VF
C
• Fig. 1.1 ​Magnetic resonance imaging scans showing median sagittal
sections of the pharynx in a normal subject. (A) Normal nasal breathing
with the oral airway occluded by lips and tongue. (B) Deliberate oral
breathing with the nasal airway occluded by elevation and backwards
movement of the soft palate. (C) A Valsalva manoeuvre in which the
subject deliberately tries to exhale against a closed airway. Data acqui-
sition for scans (A) and (B) took 45 s, so anatomical differences
between inspiration and expiration will not be visible. I am indebted​
to Professor M. Bellamy for being the subject. E, Epiglottis; L, larynx;
NC, nasal cavity; SP, soft palate; T, tongue; VF, vocal fold.
CHAPTER 1 Functional Anatomy of the Respiratory Tract 3
10
Oral cavity
Cross-sectional area (cm2)
8
Hypopharynx
Oropharynx
Bronchi
Incisors
Glottis
Trachea
0
0 5 10 15 20 25 30 35
Distance along airway (cm)
• Fig. 1.2 ​Normal acoustic reflectometry pattern of airway cross-
sectional area during mouth breathing.
capable of withstanding pharyngeal pressures as high as
80 kPa (800 cmH2O), which may be generated during
swallowing.
Upper airway cross-sectional areas can be estimated from
conventional radiographs, magnetic resonance imaging
(MRI) as in Figure 1.1 or acoustic pharyngometry. In
the latter technique, a single sound pulse with a duration of
100 ms is generated within the apparatus and passes along
the airway of the subject. Recording of the timing and
frequency of sound waves reflected back from the airway
allows calculation of the cross-sectional area, which is then
presented as a function of the distance travelled along the
airway (Fig. 1.2). Acoustic pharyngometry measurements
correlate well with MRI scans of the airway, and the
technique is now sufficiently developed for use in clinical
situations such as estimating airway size in patients with
sleep-disordered breathing (Chapter 14).1
The Larynx
The larynx evolved in the lungfish for the protection of the
airway during such activities as feeding and perfusion of the
gills with water. Although protection of the airway remains
important, the larynx now has many other functions, all
involving some degree of laryngeal occlusion.
Speech
Phonation, the laryngeal component of speech, requires a
combination of changes in position, tension and mass of
the vocal folds (cords). Rotation of the arytenoid cartilages
by the posterior cricoarytenoid muscles opens the vocal
folds, while contraction of the lateral cricoarytenoid and
oblique arytenoid muscles opposes this. With the vocal
folds almost closed, the respiratory muscles generate a posi-
tive pressure of 5 to 35 cmH2O, which may then be re-
leased by slight opening of the vocal folds to produce sound
waves. The cricothyroid muscle tilts the cricoid and aryte-
noid cartilages backwards and also moves them posteriorly
in relation to the thyroid cartilage. This produces up to
50% elongation and therefore tensioning of the vocal
4 PA RT I Basic Principles
folds, an action opposed by the thyroarytenoid muscles,
which draw the arytenoid cartilages forwards towards the
thyroid, shortening and relaxing the vocal folds. Tension-
ing of the folds results in both transverse and longitudinal
resonance of the vocal fold, allowing the formation of com-
plex sound waves. The deeper fibres of the thyroarytenoids
comprise the vocales muscles, which exert fine control
over the pitch of the voice by creating slight variations in
both the tension and mass of the vocal folds. A more dra-
matic example of the effect of vocal fold mass on voice
production occurs with inflammation of the laryngeal
mucosa and the resulting hoarse voice or complete inability
to phonate.
Effort Closure
Tighter occlusion of the larynx, known as effort closure, is
required for making expulsive efforts. It is also needed to
lock the thoracic cage, securing the origin of the muscles of
the upper arm arising from the rib cage, thus increasing the
power which can be transmitted to the arm. In addition to
simple apposition of the vocal folds described previously,
the aryepiglottic muscles and their continuation, the
oblique and transverse arytenoids, act as a powerful sphinc-
ter capable of closing the inlet of the larynx by bringing the
aryepiglottic folds tightly together. The full process enables
the larynx to withstand the highest pressures which can
be generated in the thorax, usually at least 12 kPa
(120 cmH2O) and often more. Sudden release of the
obstruction is essential for effective coughing (page 46),
when the linear velocity of air through the larynx is said
to approach the speed of sound.
Laryngeal muscles are involved in controlling airway re-
sistance, particularly during expiration, and this aspect of
vocal fold function is described in Chapter 5.
The Tracheobronchial Tree
An accurate and complete model of the branching pattern
of the human bronchial tree remains elusive, although sev-
eral different models have been described. The most useful
and widely accepted approach remains that of Weibel,2 who
numbered successive generations of air passages from the
trachea (generation 0) down to alveolar sacs (generation 23).
This ‘regular dichotomy’ model assumes that each bronchus
regularly divides into two approximately equal size daugh-
ter bronchi. As a rough approximation it may therefore be
assumed that the number of passages in each generation is
double that in the previous generation, and the number of
air passages in each generation is approximately indicated
by the number 2 raised to the power of the generation num-
ber. This formula indicates one trachea, two main bronchi,
four lobar bronchi, 16 segmental bronchi and so on.
However, this mathematical relationship is unlikely to be
true in practice, where bronchus length is variable, pairs
of daughter bronchi are often unequal in size, and trifurca-
tions may occur.
Work using computed tomography to reconstruct, in
three dimensions, the branching pattern of the airways has
shown that a regular dichotomy system does occur for at
least the first six generations.3 Beyond this point, the same
study demonstrated trifurcation of some bronchi and air-
ways that terminated at generation 8. Table 1.1 traces the
characteristics of progressive generations of airways in the
respiratory tract.
Trachea (Generation 0)
The adult trachea has a mean diameter of 1.8 cm and length
of 11 cm. Anteriorly it comprises a row of U-shaped carti-
lages which are joined posteriorly by a fibrous membrane
incorporating the trachealis muscle (Fig. 1.3). The part of
the trachea in the neck is not subjected to intrathoracic
pressure changes, but it is very vulnerable to pressures aris-
ing in the neck due, for example, to tumours or haematoma
formation. An external pressure of the order of 4 kPa
(40 cmH2O) is sufficient to occlude the trachea. Within the
chest, the trachea can be compressed by raised intrathoracic
pressure during, for example, a cough, when the decreased
diameter increases the linear velocity of gas flow, and there-
fore the efficiency of removal of secretions (page 47).
Main, Lobar and Segmental Bronchi
(Generations 1–4)
The trachea bifurcates asymmetrically, and the right bron-
chus is wider and makes a smaller angle with the long axis
of the trachea. Foreign bodies therefore tend to enter the
right bronchus in preference to the left. Main, lobar and
segmental bronchi have firm cartilaginous support in their
walls that is U-shaped in the main bronchi, but in the form
of irregularly shaped and helical plates lower down, with
bronchial muscle between. Bronchi in this group (down to
generation 4) are sufficiently regular to be individually
named (Fig. 1.4). The total cross-sectional area of the respi-
ratory tract is minimal at the third generation (Fig. 1.5).
These bronchi are subjected to the full effect of changes
in intrathoracic pressure and will collapse when the intra-
thoracic pressure exceeds the intraluminar pressure by
around 5 kPa (50 cmH2O). This occurs in the larger bron-
chi during a forced expiration, limiting peak expiratory flow
rate (see Fig. 3.7).
Small Bronchi (Generations 5–11)
The small bronchi extend through about seven generations,
with their diameter progressively falling from 3.5 to 1 mm.
Down to the level of the smallest true bronchi, air passages
lie in close proximity to branches of the pulmonary artery in
a sheath containing pulmonary lymphatics, which can be
distended with oedema fluid, giving rise to the characteristic
‘cuffing’ responsible for the earliest radiographic changes in
pulmonary oedema. Because these air passages are not di-
rectly attached to the lung parenchyma, they are not subject
TABLE
1.1 Structural Characteristics of the Air Passages

Mean
Diameter Area
Generation Number (mm) Supplied Cartilage Muscle Nutrition Emplacement Epithelium

Trachea 0 1 18 Both lungs

Links open end
Individual U-shaped
Main bronchi 1 2 12 of cartilage
lungs
2 4 8
Lobar bronchi g g g Lobes Columnar ​
Within connective ​
3 8 5 tissue sheath ciliated ​
alongside arterial epithelium
Segmental ​ Conducting Irregular
4 16 4 Segments Helical bands From the bron- vessels
bronchi airways shape
chial circulation
5 32 3

CHAPTER 1 Functional Anatomy of the Respiratory Tract

g g g Secondary
Small bronchi
lobules
11 2000 1
Bronchioles 12 4000 1
g g g Strong helical
Terminal ​ Cuboidal
muscle bands Embedded directly in
bronchioles 14 16 000 0.7
the lung paren-
15 32 000 Muscle bands ​ chyma Cuboidal to
Respiratory ​ g g 0.4
Pulmonary between ​ flat between
bronchioles Absent
Acinar ​ 18 260 000 acinus alveoli alveoli
airways From the pulmo-
19 520 000
Alveolar g g nary circulation
0.3 Thin bands in ​ Form the lung ​ Alveolar ​
ducts
22 4 000 000 alveolar septa parenchyma epithelium
Alveolar sacs 23 8 000 000 0.2

5
6 PA RT I Basic Principles

ceases to be a factor in maintaining patency. However,

• Fig. 1.3 ​The normal trachea as viewed during a rigid bronchoscopy
(page 399). The ridges of the cartilage rings are seen anteriorly, and the
longitudinal fibres of the trachealis muscle are seen posteriorly, dividing
at the carina and continuing down both right and left main bronchi. The
less acute angle of the right main bronchus from the trachea can be
seen, with its lumen clearly visible, illustrating why inhaled objects
preferentially enter the right lung.
to direct traction and rely for their patency on cartilage
within their walls and on the transmural pressure gradient,
which is normally positive from lumen to intrathoracic
space. In the normal subject this pressure gradient is seldom
reversed and, even during a forced expiration, the intralumi-
nar pressure in the small bronchi rapidly rises to more than
80% of the alveolar pressure, which is more than the extra-
mural (intrathoracic) pressure.
Bronchioles (Generations 12–14)
An important change occurs at about the 11th genera-
tion, where the internal diameter is around 1 mm. Carti-
lage disappears from the airway wall below this level and
beyond this level the air passages are directly embedded
in the lung parenchyma, the elastic recoil of which holds
the air passages open like the guy ropes of a tent. There-
fore the calibre of the airways beyond the 11th generation
is mainly influenced by lung volume because the forces
holding their lumina open are stronger at higher lung
volumes. The converse of this factor causes airway closure
at reduced lung volume (see Chapter 3). In succeeding
generations, the number of bronchioles increases far
more rapidly than the calibre diminishes (Table 1.1).
Therefore the total cross-sectional area increases until, in
the terminal bronchioles, it is about 100 times the area at
the level of the large bronchi (Fig. 1.5). Thus the flow
resistance of these smaller air passages (,2 mm diameter)
is negligible under normal conditions. However, the resis-
tance of the bronchioles can increase to very high values
when their strong helical muscular bands are contracted
by the mechanisms described in Chapters 3 and 28.
Down to the terminal bronchiole the air passages are
referred to as conducting airways, which derive their
nutrition from the bronchial circulation and are thus
influenced by systemic arterial blood gas levels. Beyond
this point the smaller air passages are referred to as acinar
airways and rely upon the pulmonary circulation for their
metabolic needs.
Respiratory Bronchioles (Generations 15–18)
Down to the smallest bronchioles, the functions of the air
passages are solely conduction and humidification. Beyond
this point there is a gradual transition from conduction to
gas exchange. In the four generations of respiratory bron-
chioles there is a gradual increase in the number of alveoli
in their walls. Like the bronchioles, the respiratory bronchi-
oles are embedded in lung parenchyma; however, they
have a well-defined muscle layer with bands which loop
over the opening of the alveolar ducts and the mouths
of the mural alveoli. There is no significant change in the

Right Left

UPPER UPPER
Apical Apical
Posterior Posterior
Anterior Anterior
MIDDLE Lingula
Lateral Superior
Medial Inferior
LOWER
Lateral basal
Anterior basal LOWER
Posterior basal Lateral basal
Medial basal Posterior basal
Apical Anterior basal
Apical

• Fig. 1.4 Lobes and bronchopulmonary segments of the lungs. Red, upper lobes; blue, lower lobes;
green, right middle lobe. The 19 major lung segments are labelled.
 CHAPTER 1 Functional Anatomy of the Respiratory Tract 7

Terminal bronchiole

Total cross-sectional area of airway (cm2)

10 000 A

1000
Respiratory bronchioles
100

10

Alveolar ducts
0
0 5 10 15 20 23
Airway generation

• Fig. 1.5 ​The total cross-sectional area of the air passages at different
generations of the airways. Note that the minimum cross-sectional
area is at generation 3 (lobar to segmental bronchi). The total cross-
sectional area becomes very large in the smaller air passages, ap- Alveolar sacs
proaching a square metre in the alveolar ducts.

calibre of advancing generations of respiratory bronchioles

(,0.4 mm diameter).

Alveolar Ducts (Generations 19–22)

Alveolar ducts arise from the terminal respiratory bronchi-
ole, from which they differ by having no walls other than
the mouths of mural alveoli (,20 in number). The alveolar
septa comprise a series of rings forming the walls of the al-
veolar ducts and containing smooth muscle. Approximately
35% of the alveolar gas resides in the alveolar ducts and the
alveoli that arise directly from them.

Alveolar Sacs (Generation 23)

The last generation of the air passages differs from alveolar
ducts solely because they are blind. It is estimated that
about 17 alveoli arise from each alveolar sac and account for
about half of the total number of alveoli.

Pulmonary Acinus
This is defined as the region of lung supplied by a first-order
respiratory bronchiole, and includes the respiratory bron-
chioles, alveolar ducts and alveolar sacs distal to a single
terminal bronchiole (Fig. 1.6). This represents the afore-
mentioned generations 15 to 23, but in practice the num-
ber of generations within a single acinus is quite variable,
being between six and 12 divisions beyond the terminal
bronchiole. A human lung contains about 30 000 acini, B
each with a diameter of around 3.5 mm and containing in
excess of 10 000 alveoli. A single pulmonary acinus is prob- • Fig. 1.6 ​(A) Schematic diagram of a single pulmonary acinus show-
ably the equivalent of the alveolus when it is considered ing four generations between the terminal bronchiole and the alveolar
from a functional standpoint, as gas movement within the sacs. The average number of generations in the human lung is eight,
but may be as many as 12. (B) Section of rabbit lung showing respira-
acinus when breathing at rest is by diffusion rather than tory bronchioles leading to alveolar ducts and sacs. Human alveoli
tidal ventilation. Acinar morphometry therefore becomes would be considerably larger. Scale bar 5 0.5 mm. (Photograph kindly
crucial,4 in particular the path length between the start of supplied by Professor E. R. Weibel.)
8 PA RT I Basic Principles
the acinus and the most distal alveolus, which in humans is
between 5 and 12 mm.
Respiratory Epithelium
Before inspired air reaches the alveoli it must be ‘conditioned’,
that is, warmed and humidified, and airborne particles, patho-
gens and irritant chemicals removed. These tasks are under-
taken by the respiratory epithelium and its overlying layer
of airway lining fluid, and are described in Chapter 11. To
facilitate these functions the respiratory epithelium contains
numerous cell types.
Ciliated Epithelial Cells5
These are the most abundant cell type in the respiratory
epithelium. In the nose, pharynx and larger airways the
epithelial cells are pseudostratified, gradually changing to a
single layer of columnar cells in bronchi, cuboidal cells in
bronchioles and finally thinning further to merge with the
type I alveolar epithelial cells (see later). They are differenti-
ated from either basal or secretory cells (see later) and
are characterized by the presence of around 300 cilia per
cell (page 165). The ratio of secretory to ciliated cells in
the airway decreases in more distal airways from about
equal in the trachea to almost three-quarters ciliated in the
bronchioles.
Goblet Cells
These are present at a density of approximately 6000 per
mm2 (in the trachea) and are responsible for producing
the thick layer of mucus that lines all but the smallest
conducting airways (page 165).
Airway Glands6
Submucosal glands occur predominantly in the trachea and
larger bronchi, diminishing in both size and numbers in
more distal airways. The glands consist of a series of branch-
ing ducts, ending with a single terminal duct opening into
the airway and contain both serous cells and mucous cells,
with serous cells occurring in the gland acinus, whereas
mucous cells are found closer to the collecting duct. The
serous cells have the highest levels of membrane-bound
cystic fibrosis transmembrane conductance regulator in the
lung (Chapter 28).
Basal Cells
These cells lie underneath the columnar cells, giving rise to
the pseudostratified appearance, and are absent in the bron-
chioles and beyond. They are the stem cells responsible for
producing new epithelial and goblet cells.
Mast Cells
The lungs contain numerous mast cells which are located
underneath the epithelial cells of the airways and in the al-
veolar septa. Some also lie free in the lumen of the airways
and may be recovered by bronchial lavage. Their important
role in bronchoconstriction is described in Chapter 28.
Club Cells (Formerly Clara Cells)
These nonciliated bronchiolar epithelial cells are found in
the mucosa of the terminal bronchioles, where they may be
the precursor of epithelial cells in the absence of basal cells.
They are metabolically active, secreting a club cell secretory
protein which has antioxidant and immune-modulatory
functions.7
Neuroepithelial Cells
These cells are found throughout the bronchial tree but oc-
cur in larger numbers in the terminal bronchioles. They
may be found individually or in clusters as neuroepithelial
bodies, and are of uncertain function in the adult lung.
Present in foetal lung tissue in a greater number, they may
have a role in controlling lung development. Similar cells
elsewhere in the body secrete a variety of amines and pep-
tides such as calcitonin, gastrin-releasing peptide, calcitonin
gene-related peptide and serotonin.
The Alveoli
The mean total number of alveoli has been estimated as
400 million, but ranges from about 270 to 790 million,
correlating with the height of the subject and total lung
volume.8 The size of the alveoli is dependent on lung
volume, but at functional residual capacity (FRC) they
are larger in the upper part of the lung because of gravity.
At total lung capacity this situation reverses, and there
are estimated to be 32 alveoli per mm3 at the lung apices
compared with 21 at the lung bases.9 At FRC the mean
diameter of a single alveolus is 0.2 mm, and the total
surface area of the alveoli is around 130 m2.
The Alveolar Septa
The septa are under tension generated partly by collagen
and elastin fibres, but more by surface tension at the
air–fluid interface (page 14). They are therefore generally
flat, making the alveoli polyhedral rather than spherical.
The septa are perforated by small fenestrations known as the
pores of Kohn (Fig. 1.7), which provide collateral ventila-
tion between alveoli. Collateral ventilation also occurs be-
tween small bronchioles and neighbouring alveoli (Lambert
channels) and through interbronchiolar pathways of
Martin, and is more pronounced in patients with emphy-
sema (page 332) and in some other species of mammal
(page 310).
On one side of the alveolar wall the capillary endotheli-
um and the alveolar epithelium are closely apposed, with
almost no interstitial space, such that the total thickness
from gas to blood is around 0.3 mm (Figs 1.8 and 1.9).10
This may be considered the ‘active’ side of the capillary, and
gas exchange must be more efficient on this side. The other
side of the capillary, which may be considered the ‘service’
side, is usually more than 1- to 2-mm thick, and contains
a recognizable interstitial space containing elastin and
collagen fibres, nerve endings and occasional migrant
Another random document with
no related content on Scribd:
The religionist expects God to wash all his sins off the slate merely
because, when he is about to die, he says he is sorry. If there be a
God, cannot He look into such a shrunken little soul and see that
there is nothing in it but a fear of death and a horror of the unknown?
I am not an atheist, because for all I know to the contrary there may
be a God, or any number of Gods, but to me the God worshiped by
my forefathers and by the religionists of to-day is a cruel,
preposterous creation conceived by a people who felt the need of
chastisement. He is a celestial traffic cop, hounded by whimpering
weaklings who beseech Him to tell them they are on the right road,
and yet keep trying to show Him which way the traffic should go. In
the Christian and Jewish conceptions of the Heavenly Father I can
see nothing that is fit for a civilized man to worship; indeed, the
nearer a man approaches civilization and intelligence, the less need
there is for him to worship anything. Conversely it is the stupid,
illiterate man, knowing neither how to read nor how to think, who is
most often the religious fanatic. He understands nothing and is afraid
of everything; he goes through life as a small boy goes past a
graveyard at night, whistling to keep up his courage. He requires
religion and its twin, superstition, to give him strength to contemplate
the wonders of the sunset and the falling rain.
For my part, I simply refuse to worry about God. If there is a God, I
hope that I may in time find favor in His sight and obtain my share of
the spiritual loot; there is nothing that I can do about it. And if there is
no God, there is nothing I can do about that, either. I profess neither
knowledge nor theory about the Supreme Being and the heavenly
wonders; knowing nothing, I believe nothing, and believing nothing, I
am prepared to believe anything, asking only reasonably correct
information and authentic signs. These I fail to find in selfish prayers,
constant squabbling over the wishes of the Lord and the building of
magnificent temples within sight and hearing of the ramshackle
tenements of the poor. I do not believe that I shall ever find them, for
“Wherefore and whence we are ye cannot know,
Nor where life springs, nor whither life doth go.”
Without religion I thoroughly enjoy the business of living. I am
oppressed by no dreadful taboos, and I am without fear; I set myself
no standards save those of ordinary self-respect, decent
consideration of the rights and privileges of others, and the
observance of the laws of the land except Prohibition. To my own
satisfaction, at least, I have proved that religion and the Church are
not at all necessary to a full and happy life. And if I am thus a sinner
and my chance of ultimate salvation forfeit, then the fault lies at the
door of those fanatics whose method of teaching religion to a child
was, and is, to hammer it into his head by constant threats of terrible
punishment, by drawing torturing word pictures of Hell, by describing
God as a vicious, vindictive old man, by scolding and tormenting and
laying down taboos until the poor child’s brain whirls in an agony of
fright and misery. I know of no better way to salute them than to refer
them to certain words of their own Savior, to be found in the thirty-
fourth verse of the twenty-third chapter of the Book of St. Luke.

If I ever have a son, which now seems unlikely, his boyhood will be
quite different from my own. For him Sunday shall be a day of rest
and pleasure; there shall be no taboos, and no attendance upon
church and Sunday school unless their performances are more
interesting than other available entertainment. They now rank just
below the moving pictures, and are therefore last. I shall bring my
son in contact with the sacred books of the Christians, the Jews, the
Buddhists and of all the other religions as rapidly as he is able to
comprehend them, and he shall be permitted to choose his own
religion if he decides that a religion is necessary to his happiness
and peace of mind. But if he shows any signs of becoming a
preacher, priest or rabbi, or even a Brother, I shall whale hell out of
him. I am that intolerant.
THE END
 A NOTE ON THE TYPE IN
WHICH THIS BOOK IS SET
This book is composed on the Linotype in Bodoni, so-
called after its designer, Giambattista Bodoni (1740-1813)
a celebrated Italian scholar and printer. Bodoni planned
his type especially for use on the more smoothly finished
papers that came into vogue late in the eighteenth century
and drew his letters with a mechanical regularity that is
readily apparent on comparison with the less formal old
style. Other characteristics that will be noted are the
square serifs without fillet and the marked contrast
between the light and heavy strokes.

SET UP, ELECTROTYPED AND PRINTED

BY THE VAIL-BALLOU PRESS, INC.,
BINGHAMTON, N. Y. · ESPARTO
PAPER MANUFACTURED IN SCOTLAND,
AND FURNISHED BY W.
F. ETHERINGTON & CO., NEW
YORK · BOUND BY H.
WOLFF ESTATE,
NEW YORK.
 TRANSCRIBER’S NOTES:
Obvious typographical errors have been corrected.
Inconsistencies in hyphenation have been
standardized.
Archaic or variant spelling has been retained.
On page 39, based upon context, “We could have no
indoor games” in the original should be “We could have
no outdoor games”; this eBook reflects this change.
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