UNIVERSITI PUTRA MALAYSIA
QUANTITATIVE OBSERVATIONS ON THE PULMONARY
ANATOMY OF THE DOMESTIC FOWL AND OTHER
GROUND-DWELLING BIRDS
M. K. VIDYADARAN
FPV 1987 2
It
entitled
the
is
hereby
Domestic
Vidyadaran,
scope,
certified
that we
have
read
this
thesis
'Quantitative Observations on the Pulmonary Anatomy
Fowl
and
quality,
and
Other Ground-Dwelling
Birds'
by
M.K.
in our opinion it is satisfactory in terms
and
presentation as partial fulfilment of
requirements for the degree of Doctor of Philosophy
AL
AINUDDIN, Ph.D.
Assoc. Professor/Dean of Graduate Studies
Universiti Pertanian Malaysia
(Chairman Board of Examiners)
A.1!l::.
iJ::.1f:?
HUG�!!��
D.t
G.M.
Sc.
D.
Research Unit for Comparative Animal
Respiration
University of Bristol, England
(External Examiner)
... .f.� ...
VYVYAN HOWARD, Ph.D.
Department of Anatomy
University of Liverpool, Fngland
(External Examiner)
of
of
the
A.S.
KING,
Ph.D.
Professor of Veterinary Anatomy
University of Liverpool
England
( Internal
�'��
..,... -
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•
•
•
6'
>r".r...-....t
. .. . ...
I
-....__ ,.
__'�
Examiner )
-�/
..... . ....
"
.........
TENGKU AZt·tI TENGKU IBRAHIH,
�t e
Associ
/
"
Ph.D.
Professor Head,
Department of Animal Science,
Faculty of Veterinary Medicine and Animal Science
Universiti Pertanian Malaysia
( Internal
Examiner )
�·KASSIM·BIN·HAMiD:·Ph:D:·
Associate Professor
Faculty of Veterinary Medicine and Animal Science
Universiti Pertanian Halaysia
( Internal
/
Examiner Supervisor )
This
thesis
was
submitted
to the
Senate
of
Universiti
Pertanian l'-alaysia and was accepted as partial fulfilment of
requirements for the degree of Doctor of Philosophy
\ � MAY 1987
Il3.te:
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ZAINUDIDN, Ph.D.
Associate Professor / Dean,
Graduate Studies.
the
QUANTITATIVE OBSERVATIONS 00 THE PUIMONARY ANATOMY OF THE
lXMESTIC FCWL AND orHER GROUND-CWELLING BIRDS
by
M. K. Vidyadaran
A
thes i s subm itted in pa rt ial fulf ilment
of
the
requi rements for t he deg ree of Doctor of P hi losophy in
the Faculty of Veterinary Medicine and Anima l Scien ce ,
University Pertan ian Malays ia .
February 1987
This thesis is dedicated to my parents
Mr. M. P . K. Menon
and
Mrs. Lakshmi Menon
ii
Many
people
have
me
helped
by
a
Professor A.
to
encouragement
from
and
King for advice,
unfailing support at
I am
efforts
all
of
stages,
and
right
introducing me to quantitative biology to supervision of
this thesis.
his
S.
spectrum
In particular,
assistance in carrying out this project.
indebted
wide
I also wish to thank Dr.
Kassim bin Hamid
for
advice and encouragement throughout the period of my work
at Universiti Pertanian Malaysia.
I
would
Ganesamurthi
Universiti
like
who
Lai
Choo May,
transmission
my
express
provided
Pertanian
valuable assistance,
Dr.
to
able
Mr.
P.
assistance
at
appreciative of
the
technical
I am
Malaysia.
gratitute
carrnents and advice by Mrs.
Mrs.
J.
Henry and Mr.
electron microscopy.
to
D.
Ho Ooi
Z.
King,
Kuan
I am grateful to Dr.
J.N.
Maina for his kindness and for the stimulating discussions
many aspects of quantitative
Sheffield,
Malaysia,
and
Dr.
Mak
on
biology.
I also wish to thank Mr.
of University of Liverpoo l,
on
G.
Dr.
Settle and Dr. R. J. Bansali
N. T.
Tian Kwan of
James of University of
Universiti
Pertanian
for valuable help on statistical analysis.
I am especially appreciative of the help provided by the
Department
of Wildlife and National Parks for
J ungle Fo.vls for the experirrent.
iii
supplying
Red
The
financial
support
for
also
grateful to CICHE and Association
Universities
came
work
from
to which I am much grateful.
Universiti Pertanian Malaysia,
am
much of the
of
I
Commonwealth
for providing grants which enabled me
to
visit
the Departments of Veterinary Anatomy in Liverpool and Nairobi
for
training
and
discussions
on
electron
microscopy
and
stereology.
To
Sharmili,
my
I
wife,
am
Shamala
and my
indebted for their
daughters,
love,
support throughout the period of studies.
iv
Shamini
and
forebearance
and
"I
often
say that when you can rreasure
speaking
about
and express it in numbers you kno.v
about
it;
kno.vledge
the
but
when you cannot express it
you
are
sanething
numbers
is of a rreagre and unsatisfactory kind;
beginning of kno.vledge,
thoughts,
in
what
your
it rray
be
in
your
(cited by Dunnill,
1968 )
but you have scarcely,
advanced to the stage of science".
Lord Kelvin,
v
1883
TABLE CF CONI'ENI'S
Page
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TABLE CF CONTENI'S. . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . .
iii
vi
LIST CF TABLES. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xi
LIST CF FIGURES
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x���
LIST OF ABBREVIATIONS. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
xv
ABSTRAcr
xvii
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INTROD�ION.
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Lung-Air Sac System of Birds:
Morphogenesis
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General Aspects
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Structure of the Adult Lung
Function of the Adult Lung
LITERATURE
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REVIE:W . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
• •
Domestic fowl (Gallus gallus var damesticus)
(Gallus gallus)
Jungle Fowl
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Stereological Studies of the Adult Avian Lung
Red
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14
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White-breasted Water-Hen (Amaurornis
phoenicurus )
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(Cairina rnoschata)
Muscovy Duck
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Stereological Studies of the Juvenile
a vian lung
OBJECTIVES.
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21
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MATERIALS AND MEl'HODS .
Animals . .
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Fixation of Lungs
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Estimation of Lung Volume
Sampling P rocedure
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Ti ssue P rocessi ng for Microscopy
Lig ht Microscopy
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Stereological Analysi s of Histological
Sections and Electron Microg raphs
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Hi stological Sections
Electron Micrographs
Stereology
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Volume Density
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Transmi ssion Electron Microscopy .
Scanning Electron Microscopy
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Surface Area and Surface Densi ty
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Harmonic Mean Thickness . . . . . . . . . . . . . . . . . . . . . . . . . . . .
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Arithmetic Mean Thickness
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Pubnonary D iffusing capacity
Carcass Evaluation P rocedure
Stati stical Analysi s
RESULT S .
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G ross morphology
(i ) Adult lung
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Quali tative Observations
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(ii ) Juveni le lung .
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vii
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H i stological Character i stics . . . . . . . . . . . . . . . . . . . . . . . . .
59
( i ) Adult lung . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
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( i i ) Juvenile lung . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
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Transm i ssion Electron Microscopy . . . . . . . . . . . . . . . . . . . . . .
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( i ) Adult lung . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
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( i i ) Juvenile lung . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
64
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Quantitative Observations . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
76
Bod y weight , lung volume , and lung
volume per unit body weight . . . . . . . . . . . . . . . . . . . . . . . . .
77
Analy s i s of the relat ive volume and
the absolute volume of the main components
of the lung . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
80
Volume per unit body weight of the main
canponent of the lung . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
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Volume densities and the absolute volumes
of the main components of the exchange area . . . . . . . .
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Volume per unit body weight of the main
canpo nents of the exchange ti ssue . . . . . . . . . . . . . . .
..
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Analys i s of the di str ibution of ai r and blood i n
the lung . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
89
Analy s i s of the surface areas and surface
area per unit body we ight of the res i stance
barr iers of the ai r-haemoglobin pathway
of the lung . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
94
Analysis of the surface area of the blood-gas
barrier per unit volume of the exchange tissue
( St/Vx ) , and volume of capillary blood per unit
surface area of the blood-gas ba rr ier (Vc/St )
96
Scanning Electron Microscopy . . . . . . . . . .
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Analys i s of the absolute volume s of
erythrocytes , erythrocyte nucleus ,
erythrocyte cytoplasm, and the
percentage volume of pulmonary capil lary
haematocrit . . .
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99
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Analysi s of the thickness of the blood-gas
( ti ssue ) barr ier and the plasma layer
Analysis of the pulmonary diffusing
capacit ies of the resi stance barr iers
of the air-haemoglobin pathways
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Carcass Evaluat ion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
DISCUSSION . .
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Critique of the Methodology
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Error s due to sample size and interval
Error s due to perfusion , fixation ,
osmolarity and tissue processing
Error s due to estimat ion of lung
volume by Scherle' s method
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Error s due to stereological method s
Error s to possible genetic impurity
of the Red Jungle Fo.vl . . . . .
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10 1
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III
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12 1
Pulmonary Stereo1ogy of the Adult
Female Dctne st ic Fo.vl . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
12 3
Pulmonary Stereo1ogy of the Adult
Female Red Jungle Fo.vl . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
138
Pulmonary Stereo1ogy of the Adult
Female White-breasted Water-Hen
148
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Pulmonary Stereology of the Adult
Female Muscovy Duck . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
158
Pulmonary Stereology of the
Developing Lung . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
168
SUr.t>1ARy
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COOCLUSIONS
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BI BLI()SRAPHY
APPENDICES
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18 1
19 7
211
LI sr OF TABLES
Page
Table
I
Number of aniffi3.1s used for var ious
investigations . .
•• ••
24
II
Body weight and lung volume . . . . . . . . . . . . . . . . . . . . . . . .
78
III
Relative and Absolute volumes of
the ffi3. in components of the lung . . . . . . . . . . . . . . . . . .
81
Absolute volume per uni t body weight
of the main components of the lung . . . . . . . . . . . . . . . . .
83
Relative and Absolute volumes of
the ffi3. i n components of the exchange
ti ssue of the lung . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
85
Absolute volume per uni t body weight of
the ffi3. i n components of the exchange tissue . . . . . . . . .
87
Analysis of the volume di str ibution
of the air in the lung . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
90
Analys i s of the volume di str ibution
of the blood in the lung . . . . . . . . . . . . . . . . . . . . . . . . . . .
92
The relative proportions of the lung
formed by air and blood . . . . . . . . . . . . . . . . . . . . . . . . . . . .
93
The sur face areas of the resi stance
barr ier s of the air-haemoglobin
pathway of the lung
95
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IV
Va
Vb
VI a
VIb
VIc
VII
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VIllb
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.
Specific surface areas and surface
to volume ratios of the resi stance
barr ier s of the gas exchange tissue . . . . . . . . . . . . . . . .
98
Absolute volume of the erythrocyte components:
Pulmonary and Venous haerna tocr its . . . . . . . . . . . . . . . . . .
100
Thickness of the blood-gas ( ti ssue )
barr ier and the plasma layer . . . . . . . . . . . . . . . . . . . . . . .
10 2
Anatomical
pulmonary
dif fusing
capaci ties of the resi stance barr ier s
of the air-haemoglob i n pathway for oxygen . . . . . . . . . .
10 5
xi
Xb
� natomical pulmonary diffusing capacities
of the resistance barriers of the ai rhaemoglobi n pathway for oxygen
.
XI
XII
.
.
.
.
.
.
.
.
.
.
Compa risons of pulmonary parameters of
the domestic fowl and the Red Jungle Fowl
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
Allomet ric functions of the volume of
lung with body weight , dressing pe rcentage
and carcass components
.
.
xii
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
10 7
110
112
LIST Of FIGURES
Page
F igure
1.
2.
3.
4.
Sem idiagrarnmatic medial v i ew of the
r ight lung of Gallus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
4
Medial view of the right lung of
Gallus . . . . . . . . . ., . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
6
Systematic stratif ied sampling
procedure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
29
An electron micrograph superimpo sed
w i th a quadratic lattice grid
35
•
5.
•
•
•
•
•
An electron micrograph superimpo sed
w ith a random short line test grid
•
6.
8a .
8b.
.
.
.
•
•
•
•
•
•
•
lOa .
lOb.
lla .
•
•
•
•
•
•
•
•
•
•
•
•
.
•
•
•
•
•
•
•
•
•
•
•
•
•
.
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
37
39
Scanning electron micrograph of the
lung of the White-breasted Water-Hen
( Arnaurorni s phoenicurus ) . . . . . . . . . . . . . . . . . . . . . . . . . .
69
Scanning electron micrographs of the
parabronchi of the lung . a . �dult
domestic fowl , b. Adult Red Jungle Fowl . . . . . . . . . . .
70
Scanni ng electron micrographs of the
pa rabronchi of the lung . a . Muscovy Duck ,
b . Adult White-breasted Water-Hen
71
•
9.
.
P rinciples used in estimating surface
area and harmonic mean thickness of
the resi stance barr iers
.
7.
•
•
•
•
•
•
•
•
•
•
Scanning electron micrograph of the
exchange tissue of the lung of the Whitebreasted Water-Hen (Arnaurornis phoenicurus )
•
•
•
•
•
•
•
•
•
•
.
•
•
•
73
Scanning electron micrograph ( Latex Cast ) of
the air cap illaries of the exchange tissue of
domestic fowl (Gallus gallus var domesticus ) . . . . . .
74
Scanning electron microg raph (Latex Cast ) of
the blood cap illar ies of the exchange tissue
of the danes tic fowl. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
75
T ransmiss ion electron microg raphs of the
exchange tissue of the Adult lung . a . Dome stic
fowl ,
b. Red Jungle Fowl . . . . . . . . . . . . . . . . . . . . . . . . .
61
xiii
llb.
Transmi ssion electron micrographs of the
exchange ti ssue of the adult lung . c .Muscovy
Duck , d. White-breasted water-Hen
•
12a .
12b.
12c.
•
•
•
•
•
•
.
•
•
•
•
•
•
.
•
62
Transmi ssion electron micrograph of the
exchange tissue of the lung of the
danestic fa.vl at day 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . .
65
Transmiss ion electron micrograph of the
exchange ti ssue of the lung of the
danestic fa.vl at day 7 . . . . . . . . . . . . . . . . . . . . . . . . . . . .
66
Transmi ss ion electron micrograph o f the
exchange ti ssue of the lung of the
danestic fa.vl at day 30 . . . . . . . . . . . . . . . . . . . . . . . . . . .
67
xiv
LIsr CF ABBREVIATIONS
Deo
2
erythrocytes
DLo
total
Drno
oxygen
diffusing
anatomical
2
for oxygen
2
oxygen
oxygen
Dpo
oxygen
Dto
oxygen
2
2
diffusing
diffusing
capacity
(conductance)
pulmonary
capacity
diffusing
of
capacity
the
of
capacity
membrane
for
plasma
for
the
oxygen
diffusing
capacity
of
the
blood-gas
(tissue) barrier for oxygen
Hc
pulmonary capillary haematocrit
He
venous haematocrit
r
correlation coefficient
Sa
surface area of the air capillary epithelium
Sc
surface area of the blood capillary epithelium
s. d.
standard deviation
Se
surface area of the capillary erythrocytes
Sp
surface area of the plasma layer
St
surface area of the blood-gas (tissue) barrier
Va
volume of the lumen of the air capillaries
Vb
volume
of the wall and lumen
vessels
of
the
blood
larger than capillaries
Vc
volume of the lumen of the blood capillaries
Ve
volume of the pulmonary capillary erythrocytes
Vee
volume
of
the
cytoplasm
of
the
pulmonary
capillary erythrocytes
Ven
of
volume
of
capillary
the
nucleus
erythrocytes
AV
of
the
pulmonary
VL
volume of the fixed lung (left lung x 2 )
Vlb
volume of the lumina of parabronchi
secondary bronchi ( including atria )
Vp
volume of the wall and lumen of
bronchus
vt
volume of the blood-gas ( tissue) barrier
Vtn
volume of the tissue not involved
exchange
Vx
volume of the exchange tissue of the lung
W
body weight
Ehp
harmonic mean thickness of the plasma
Eht
harmonic
( ti ssue )
Et
mean thickness
of
the
the
in
and
prlinary
gaseous
blood-gas
barrier
arithmetic mean thickness
( ti ssue ) barrier
of
the
blood-gas
Specific values are those standardized against body
for example, St/W means the specific surface area
tissue barrier .
weight;
of the
In the text and Tables , all lung values are totals for
left and right lungs together .
the
The anatomical terminology adopted in thi s thesis is that used
by
the
International
Cammi ttee
on
Avian
Anatanical
the
Nanina
Nanenclature
( I . e . A . A . N. ) and publi shed in
Anatamica Avium (King, 19 79 ) .
The taxonomic nomenclature follows Gruson , ( 1976 ) .
xvi
An abstract of the thesis presented to the Senate of
Universiti Pertanian Malaysia in partial fulf ilment of
the requirements for the Degree of Doctor of Philosophy
QUANI'ITATIVE OBSERVATIONS ON THE PULMONARY ANATOOY
OF THE IXMEsrIC F'O'lL AND arHER GROUND-r:wELLI� BIRDS
by
M . K . Vidyadaran
June , 1986
Supervisor
Associate Professor Dr . Kassim bin Hamid
Faculty
Veterinary Medicine and Animal Science ,
Universiti Pertanian Malaysia, Malaysia
Co-supervisor
Professor Dr . A . S . King
Department
Veterinary Anatomy ,
University of Liverpool , England
The
pulmonary
(Gallus
gallus
(Gallus
gallus) ,
Whi te-breasted
juveniles
stereology
variant
of the adult
damesticus) ,
the Muscovy Duck
Water-H�
the
domestic
Red
( Cairina
Jungle
of
the
domestic fowl and
investigated.
The
lung of the dcxnestic fowl was
wi th that of the Red Jungle Fowl at day 1 ,
Fowl
rroschata) ,
(Arraurornis phoenicurus),
Red
fowl
Jungle
day 7,
the
and
Fowl
the
were
canpared
and day
30 .
A
multistage sampling technique was used for
stereology .
counting,
Standard
stereological
procedures
pulmonary
of
I;X)int
intersection counting , and measurement of intercept
xvii
length,
area,
surface
were employed for estbnating volume density,
and harmonic mean thickness respectively. The anatomical
diffusing capacity was estbnated from Weibel's IIDdel.
The
pulmonary stereological characteristics of the
Red
Jungle Favl are broadly similar to those of the danestic favl,
but
the Red Jungle Favl has a higher specific volume
lung
and a thinner blood-gas (tissue) barrier,
of
the
with a higher
than the
specific diffusing capacity of the barrier for oxygen
danestic favl.
The lung of the White-breasted Water-Hen,
a ground-
to
dwelling bird, has stereological characteristics similar
sane
flying birds
(non-passerine),
all
ground-dwelling
birds
thus showing that
have
inferior
not
pulmonary
stereological characteristics.
The
which
danestic Muscovy Duck has pulmonary characteristics
are
broadly
the
to
those
reported
for
other
except for a greater thickness of the blood-gas
Anseriformes,
(tissue)
slinilar
�
T e specific oxygen diffusing capacity
barrier.
blood-gas
(tissue) barrier was well below that of
of
other
anseriforms and birds in general.
The
juvenile
of
lJoth the danestic fowl
Jungle Favl has a thicker blood-gas
adult.
barrier
In
was
each
the
Red
(tissue) barrier than
the
age group examined,
thicker
the blood-gas
in the danestic fowl than
Jungle Favl.
x-viii
and
in
(tissue)
the
Red
�t
is
day 7 the pulmonary stereology of the domestic
vastly
suggested
inferior to that of the Red
Jungle
Fowl.
fowl
It
is
to meet
that lung develo�nt has not been adequate
the rapid increase in body weight.
The
pathophysiological
effects
values
of same of the parameters are
volume
and volume
greater
total
of
the
stereological
discussed.
Surface
to volume relationships suggest a
length
of blood capillaries in
the
to
possible
exchange
tissue of the domestic fowl than in the Red Jungle Fowl,
with
the possibility of greater resistance to pulmonary blood flow.
Scanning electron microscopy showed that both the
and
air
capillaries
were
tortuous.
The
air
resembled irregular chambers connected by small
the
blood
capillaries
tubes,
capillaries formed a network of tubes
blood
of
while
uniform
diameter.
Carcass evaluation indicated that although muscle, bone,
etc.
were highly correlated with volume of lung,
is still the best carnparator.�
xu
body weight
Abstrak tesis yang dikemukakan kepada Senat
Universiti Pertanian Malaysia sebagai sebahagian
daripada keper1uan untuk Ijazah Doktor Falsafah
QUANI'ITATIVE OBSERVATIONS ON THE PUlMONARY ANATCMY
OF THE IXMEsrIC FCWL AND arHER GROUND-rwELLING BIRDS
by
M . K . Vidyadaran
June , 1986
Dr . Kassirn bin Hamid
Ketua penyelia
Profesor Madya
Fakulti
Kedoktoran Veterinar dan Sains Perternakan,
Universiti Pertanian Malaysia , Malaysia
Penyelia
Profesor Dr . A. S . King
Department
Veterinary Anatany ,
University of Liverpool , England
Stereologi
pulmonari
variant danesticus) ,
Muskovi
( Cairina
ayam
peliharaan
ayam hutan merah (Gallus
rroschata )
burung
itik
( Amaurornis
dan anak-anak ayam peliharaan dan
dewasa ,
hutan
merah
telah diselidiki.
ayam
peliharaan dengan ayam hutan merah pada
Perbandingan
antara
urnur
ayam
paru-paru
7 dan 30
1,
dilakukan.
Teknik
persampelan
pelbagai lapisan
telah
stereologi pulmonari.
untuk
pengiraan-pengiraan titik , antara bahagian, dan
jarak
intersep ,
telah
xx
Prosedur stereo1ogi
digunakan
untuk
bagi
gallus
gallus ) ,
Uak-uak
phoenicurus )
had
<Gallus
digunakan
untuk
yang
biasa
ukuran
anggaran
ketwnpatan volum, luas
pe rmukaan dan
min harmonik ketebalan .
Keupayaan difusi anatom i adalah dianggarkan
oontoh
mengikut
Weibel .
Sifat-sifat
stereologi pu lmonari ayam hutan merah
sarna
keseluruhannya adalah
hutan
merah
tinggi
dan
keupayaan
mempunyai
rintangan
difusi
pada
dengan ayam peliharaan , tetapi ayam
volum paru-paru spe s i f ik
darah gas
yang
lebih
yang
lebih
nipi s ,
dengan
lebih
tinggi
spes ifik oleh rintangan yang
untuk oksigen daripada ayam pe l i haraan .
Paru-paru
bu rung Uak-uak,
yang amnya menetap
sarna
tanah ,
mempunyai
burung
yang terbang (bukan paserine) , rrenunjukkan bukan
burung
jenis
sifat stereologi yang
di
di
atas
tanah
mempunyai
dengan
sifat
atas
setengah
sernua
stereologi
pulmonari yang bennutu rendah ( inferior).
I tik Muskovi pe l i haraan mempunyai sifat-sifat
pulmonar i
yang banyak persamaan dengan yang dilaporkan bagi Anseriforrre s
yang
lai n ,
Keupayaan
kecuali
difusi
ketebalan
rintangan
oksigen spes if i k bag i
darah-gas
rintangan
( ti su) .
darah-gas
( ti su) adalah rendah daripada jeni s anseriforrres yang lain dan
burung-burung amnya .
Kedua-dua
mempunyai
yang
rintangan
dewa sa .
r intangan
anak
Di
darah-gas
ayam pe l i haraan dan
darah-gas ( tisu)
dalam
seln1l11pulan
(ti su)
adalah
peli haraan daripada ayam hutan merah .
xxi
ayam
lebih
umur
lebih
hutan
tebal
yang
tebal
merah
dar ipada
diper iksa ,
bag i
ayam