J Appl Physiol 121: 1374–1378, 2016.
First published September 22, 2016; doi:10.1152/japplphysiol.00428.2016.
Case Studies in Physiology: Ventilation and perfusion in a giraffe– does
size matter?
Görel Nyman,1 Bengt Röken,2 Eva-Maria Hedin,3 and Göran Hedenstierna4
1
Department of Clinical Sciences, Swedish University of Agricultural Sciences, Uppsala, Sweden; 2Kolmården Wild Animal
Park, Kolmården, Sweden; 3Department of Medical Sciences, Cardiology-Arrhythmia, Uppsala University, Uppsala, Sweden;
and 4Hedenstierna Laboratory, Department of Medical Sciences, Uppsala University, Uppsala, Sweden
Submitted 6 May 2016; accepted in final form 19 September 2016
Nyman G, Röken B, Hedin E-M, Hedenstierna G. Case Studies
in Physiology: Ventilation and perfusion in a giraffe – does size
matter? J Appl Physiol 121: 1374 –1378, 2016. First published Sep-
tember 22, 2016; doi:10.1152/japplphysiol.00428.2016.—The trachea
in the giraffe is long but narrow, and dead space ventilation is
considered to be of approximately the same size as in other mammals.
Less is known about the matching between ventilation and lung blood
flow. The lungs in the giraffe are large, up to 1 m high and 0.7 m wide,
and this may cause considerable ventilation/perfusion (VA/Q) mis-
match due to the influence of gravitational forces, which could lead to
hypoxemia. We studied a young giraffe under anesthesia using the
multiple inert gas elimination technique to analyze the VA/Q distri-
bution and arterial oxygenation and compared the results with those
obtained in other species of different sizes, including humans. VA/Q
distribution was broad but unimodal, and the shunt of blood flow
through nonventilated lung regions was essentially absent, suggesting
no lung collapse. The VA/Q match was as good as in the similarly
sized horse and was even comparable to that in smaller sized animals,
including rabbit and rat. The match was also similar to that in
anesthetized humans. Arterial oxygenation was essentially similar in
all studied species. The findings suggest that the efficiency of VA/Q
matching is independent of lung size in the studied mammals that vary
in weight from less than 1 to more than 400 kg.
giraffe; ventilation; gas exchange; ventilation/perfusion
NEW & NOTEWORTHY
Unique measurement of ventilation-perfusion matching in a
giraffe. Surprisingly, the matching was similar to that in the
horse as well as in the rabbit and rat, 1/100 and 1/500 of the
weight of the giraffe, and also similar to that seen in humans.
THE ADULT GIRAFFE REACHES an average height of ~5 m (4, 14)
and heights up to 6.6 m have been reported (16, 35). The
shoulder height is ~3 m (4, 14, 16). The neck is thus as long as
2 m or more. Despite a long trachea, dead space to tidal volume
ratio (VD/VT), as measured by conventional CO2 elimination,
has been found to be similar in size to that of other mammals
and has been explained by a narrow trachea and large tidal
volume (23, 30).
Less is known about pulmonary gas exchange and arterial
oxygenation in the giraffe, although a comment has been made
that arterial oxygen and carbon dioxide tensions (PaO2, PaCO2)
were within normal limits despite the use of anesthetics that
had been used to enable measurements (3). In a tall lung, a
considerable gravitational influence on the distribution of lung
blood flow may exist, with a potential impact on ventilation/
Address for reprint requests and other correspondence: G. Hedenstierna,
Hedenstierna Laboratory, Dept. of Medical Sciences, entr 40:2, 75185 Uppsala
University, Uppsala, Sweden (e-mail: goran.hedenstierna@medsci.uu.se).
1374 8750-7587/16 Copyright © 2016 the American Physiological Society
Downloaded from journals.physiology.org/journal/jappl (046.024.024.224) on October 19, 2022.
perfusion (VA/Q) matching and arterial oxygenation (37).
When a young giraffe was undergoing surgical repair of a
fractured foot, we got an opportunity to conduct gas exchange
measurements during anesthesia, with the giraffe breathing
spontaneously in a partial lateral position. We were also given
time to make a measurement of the VA/Q matching with the
multiple inert gas elimination technique, MIGET (34). We
compared our results with those from previous studies in
different species, including humans.
MATERIALS AND METHODS
A 2-yr-old female giraffe, weight ~400 kg, with a chronic hind leg
lesion (healed epiphyseal fracture) was studied during anesthesia. She
was born and lived at the Kolmarden Wild Animal Park in Sweden.
The ethical board of the park had approved the study, provided that
the clinically indicated anesthesia for correction of the fractured foot
was not prolonged. The study was done in 1990, but the results were
not reported previously.
Anesthesia. Anesthesia was induced with xylazin 100 mg (Rompun
vet.) and 0.2 ml Large Animal Immobilon (etorphine HCl 0.5 mg and
acepromazine 2 mg) injected intramuscularly at the base of neck with
a blow dart. Large Animal Immobilon (0.5 ml) was given intrave-
nously, and the giraffe was stabilized with the trunk in left lateral
position and neck and head in an upright position, the latter adjusted
by careful positioning of snares around the neck (partial lateral
recumbency). The right hind hoof was trimmed, and a supportive cast
of Technovit was applied to adjust the hoof’s angle to the ground.
Anesthesia was reversed by the opioid antagonist Large Animal
Revivon (diprenorphine, 2,4 mg) and alpha-2-adrenoceptor antagonist
atipamezol, 10 mg (Antisedan vet.), both given intravenously. The
giraffe got on its feet without support 150 s after the reversal of the
anesthesia.
Blood gases and oxygen saturation. A Teflon catheter (80 mm
Branüle cannula, external diameter 2.1 mm) was introduced into a
jugular vein for inert gas infusion (see below). Arterial blood samples
were taken by puncture of the carotid artery.
Blood was collected in heparinized glass syringes with matched
barrels, capped, and stored on ice for subsequent blood gas analysis
(equipment: ABL 3, Radiometer, Copenhagen, Denmark). Oxygen
saturation and hemoglobin concentration were measured by spectro-
photometry (OSM 3, Radiometer).
Ventilation and cardiac output. The expired minute ventilation was
collected by a specially designed mask that was fitted onto the nose of
the animal. It was made airtight by an inflatable cuff, the surface of
which was covered by aluminum foil to avoid inert gas absorption in
rubber (see below). One-way valves allowed breathing through the
mask and passage of expired gas via metal tubing to a mixing box to
ensure the collection of mixed expired gas. Minute ventilation was
measured by collecting the expired gas in Douglas bags and emptying
them through a gasometer. The giraffe breathed ambient air through-
out the study.
Cardiac output was estimated according to the Fick principle, but
because we were not allowed to collect mixed venous blood, we had
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 VA/Q in a Giraffe • Nyman G et al. 1375
to use jugular venous blood. Arterial and jugular venous O2 concen- uptake was 890 ml/min (STPD), corresponding to ~3 ml·min⫺1·
trations were measured as described above, and a mixed expired gas kg body wt⫺1, similar to human per kilogram body weight. The
sample was used for measurement of O2 and CO2 concentrations expired CO2 amounted to 751 ml/min, resulting in a respiratory
using the polarographic technique (Beckman OM 14) and an infrared
meter (Leybold-Hereus). The measured oxygen uptake was divided by exchange ratio of 0.84. PaO2 was 70 mmHg during air breath-
the arterial-venous oxygen content difference to yield cardiac output. ing, and PaCO2 was 57 mmHg. Thus PaO2 was in the lower
Replacing mixed venous by jugular venous blood may have caused normal range for the supine position in conscious humans and
some overestimation of cardiac output. PaCO2 was slightly increased, indicating a certain hypoventila-
Ventilation/perfusion matching. MIGET, as described by Wagner tion (10). The VD/VT ratio calculated according to the Enghoff
et al. (33) was used, with necessary modifications to make it appli-
cable to the giraffe. In short, six gases (SF6, ethane, cyclopropane, equation was 0.36, corresponding to 0.97 liters, including
enflurane, ether, and acetone), inert in the sense of being chemically apparatus dead space (8). The latter was measured separately
inactive in blood, were dissolved in saline and infused for 25 min at by water displacement with consideration of the volume of the
a rate of 45 ml/min. The infusion time was shorter than the normal 40 nose of the giraffe and amounted to 0.27 liters. The physio-
min because of the limited duration of anesthesia. However, subse- logical dead space thus corresponded to 0.70 liters and the
quent double blood and expired gas samples were taken 3 to 4 min VD/VT ratio to 0.26.
apart, and no gross differences in the inert gas concentrations were Cardiac output, calculated according to the Fick principle
noticed and the calculated retention and excretion values were similar
in the two sets of samples. The blood samples were collected in glass
with peripheral venous blood instead of mixed venous blood,
syringes with matched barrels and stored on ice until analysis. The was 18.1 l/min. The heart rate was 48 beats/min and the stroke
expired gas was collected in special glass syringes with ceramic volume 377 ml. Thus ventilation and circulation data were
barrels. The syringes were kept at 37–38°C by means of a heating much the same as in humans after correction for body size (19).
blanket to avoid water condensation and to keep them gas tight (they Ventilation-perfusion relationships. Two sets of data were
would leak if cooled). With these precautions taken, the blood and gas obtained and both were of high quality, as shown by the very
samples were brought to the laboratory and analyzed, ~3 h later. The low remaining sum of squares, RSS, 0.83 and 1.8 (34). The
blood samples were tonometered against a gas phase and were, former is shown in Fig. 1 and Table 1. Calculations of the
together with the gas samples, injected into a gas chromatograph
(model 5890, Hewlett-Packard) with an electron capture detector to VA/Q distribution revealed a broad single mode, as indicated
measure SF6 and a flame ionization detector for analysis of the other by a rather high logarithmic standard deviation of the perfusion
gases. The inert gas data, together with the measured minute ventila- distribution (log SDQ). A moderate amount of the perfusion
tion and calculated cardiac output (as described above), were submit- was distributed to VA/Q ratios below 0.1 (“low VA/Q”), and
ted to a computer-based analysis for calculation of the distribution of there was essentially no shunt. There was no ventilation to
VA/Q (33). The solubility coefficients of the inert gases were calcu- regions with VA/Q ratios above 10 except dead space (here
lated using blood samples taken before onset of the infusion (33). defined as ventilation of regions with VA/Q ⬎ 100). Dead
space/tidal volume ratio (VD/VT) was 0.33, similar to the dead
RESULTS
space calculated on the basis of CO2 elimination, according to
Ventilation, blood gases, and cardiac output. The expired the Enghoff equation (8) (Table 1). PaO2 predicted from the
ventilation was 18.9 l/min (BTPS) and the respiratory rate 7 VA/Q distribution was 64 mmHg and thus fairly similar to the
breaths/min and thus tidal volume (VT) was 2.7 l. Oxygen measured PaO2.

A B
Fractional Retention (R) and Excretion (E)

1.0 R
Ventilation and Perfusion

0.8

E
0.6

0.4

0.2

VA/Q
0.0
0.0 0.001 0.1 10.0 1000

Blood:Gas partition coefficient, log scale

Fig. 1. MIGET data in the spontaneously breathing, anesthetized giraffe in partial lateral position. A: retention (R) and excretion (E) of the 6 gases plotted against
their blood/gas partition coefficients (indicated by the small circles from low to high coefficient: SF6, ethane, cyclopropane, enflurane, ether, acetone). Solid lines,
curve calculated for an assumed homogeneous lung with similar dead space and shunt as in the studied giraffe. Dashed lines, curve fitted to the measured data.
B: ventilation/perfusion distribution (VA/Q). Compare with rabbit, horse, and humans in Fig. 2.

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1376 VA/Q in a Giraffe • Nyman G et al.

Table 1. PaO2, PaCO2, and VA/Q data in different species. All but conscious humans have been studied during intravenous
anesthesia (in humans, low concentration of anesthetic gas was added to the intravenous drugs)
Species Pos./O2 n PaO2, mmHg PaCO2, mmHg logSDQ logSDV QS/QT, % LowVA/Q, %

Giraffe (present study) lat/0.21 1 70 57 1.07 1.02 1 8

Horse (21, 22) lat/0.21 12 65 ⫾ 10 49 ⫾ 5 0.81 ⫾ 0.44 5⫾3 5⫾7
Sheep (6) lat/0.21 8 76 ⫾ 5 46 ⫾ 6 0.53 ⫾ 0.06 1.29 ⫾ 0.80 2⫾1 1⫾1
Rabbit (17, 18) sup/0.34 23 81 ⫾ 22* 38 ⫾ 12 0.84 ⫾ 0.23 0.77 ⫾ 0.30 2⫾3 1⫹2
Rat (2) lat/0.21 10 87 ⫾ 5 30 ⫾ 3 1.06 ⫾ 0.18 1.06 ⫾ 0.11 4⫾1
Man (15) lat/0.30 15 70 ⫾ 8* 35 ⫾ 1 1.02 ⫾ 0.21 1.46 ⫾ 0.11 4⫾1 7⫾3
Conscious man (27) sup/0.21 7 99 ⫾ 2 39 ⫾ 1 0.38 ⫾ 0.03 0.37 ⫾ 0.04 0⫾0
Values are means ⫾ SD. Giraffe, horse, sheep, and wake humans have been studied during spontaneous breathing. Rabbit, rat, and anesthetized humans have
been studied during mechanical ventilation. Pos., position, lateral or supine; O2, inspired oxygen fraction, 0.21 (air) or higher. *PaO2 recalculated to the expected
value during air breathing.

DISCUSSION
The major findings in this study of an anesthetized, sponta-
neously breathing giraffe were a small dead space, a broad but
uniform VA/Q distribution, and essentially no shunt.
The small dead space in a giraffe with a VD/VT ratio of 0.26
after subtraction of apparatus dead space (determined using the
inert gas elimination technique) is in agreement with previous
studies that have used CO2 elimination technique (23, 30). The
narrow trachea reduces dead space ventilation and contributes
to “normal” VD/VT but should impose a high respiratory
resistance during physical exercise. The horse has a shorter and
wider trachea, giving the same dead space volume as in the
giraffe, but with less respiratory resistance (29).
The fair match of VA/Q ratios and the minor shunt may
appear surprising in view of the size of the animal (36). The
lung in an adult giraffe male, body weight 1 200 kg, is ~80 cm
dorsoventrally (isogravitational direction) and 65 cm from one
side of the diaphragmatic lobes to the other (gravitational
direction in the lateral recumbency; from measurements on a
giraffe in the Wild Animal Park). These dimensions were
smaller in the young giraffe studied, which weighed 400 kg
and had a lung height of ~55– 60 cm and lung width of 45–50
cm. This is, however, comparable to horses of similar weight
that have been studied with MIGET (see below).
That shunt was almost absent may suggest that no atelectasis
had developed. Large atelectasis is frequently seen in anesthe-
tized, supine horses of similar size. However, horses in the
lateral position have less atelectasis than they do in the supine
position (25), and the giraffe was studied in the “partial” lateral
position. Moreover, the giraffe was breathing ambient air,
which delays formation of atelectasis. This is because atelec-
tasis is caused by absorption of gas behind closed airways and
this absorption takes longer during the breathing of air than
during the breathing of oxygen (5, 7). A short duration of
anesthesia (here around 25 min) may not have been enough to
cause atelectasis when air was breathed. The fair VA/Q match
in view of the large lung may also possibly be related to a more
even distribution of the perfusion of the lung, with no consis-
tent gravitational gradient, as shown by Hlastala et al. (12) in
the conscious upright horse. Alternatively, ventilation is dis-
tributed mainly to more dependent lung regions where perfu-
sion should be expected to be the largest. Active control
mechanisms of ventilation and perfusion distribution to opti-
mize VA/Q matching do exist, e.g., hypoxic pulmonary vaso-
constriction, but the anesthesia that was provided during the
studies may have interfered with these mechanisms (20). Thus
the conscious horse and humans have better VA/Q matching
than during anesthesia (26, 32). The concept of fractal distri-
bution of ventilation and lung blood flow, independent of
gravity and, possibly, of body size, is another potential mech-
anism of more homogeneous VA/Q distribution (9).
Another consequence of large lung dimensions, as in the
giraffe, could be stratified heterogeneity of the air in alveoli, as
seen in the Varanid lizard with its relatively uncomplicated
lung and large airspaces (13). Stratified heterogeneity will
more likely be seen for a heavy gas molecule and cause
increased retention of the gas enflurane, which was used in the
MIGET in the Varanid lizard study (13). However, we found
no increased retention of enflurane in our giraffe, as can be
inferred from the retention data in Fig. 1A. Moreover, PaO2 can
be predicted from the VA/Q match, assuming complete equil-
ibration of gases across the alveoli-capillary membranes and no
stratified heterogeneity. Because we found no difference be-
tween the measured and the predicted PaO2, this is a further
sign that there was no diffusion impairment in our giraffe,
either over alveolar-capillary membranes [as in pulmonary
fibrosis (1) or by stratified heterogeneity in alveolar gas (13)].
The match of VA/Q in the anesthetized giraffe was as good
as in a horse of similar weight, which has a vertical lung height
of less than 65 cm, and the giraffe has, if anything, a smaller
shunt (21, 22) (Table 1). Anesthetized sheep in the lateral
position, weight 38 –97 kg, also have a similar degree of VA/Q
mismatch, as inferred from logSDQ and logSDV (6). The
giraffe compares favorably with an anesthetized human, who,
in lateral position, has a vertical lung height of 20 –30 cm and
still has a broad VA/Q dispersion and a clear shunt (15) (Table
1). Even more interesting is the fact that anesthetized rabbits
(studied supine) and rats (studied in the lateral position), with
1/100 and 1/500 of the weight of the horse and giraffe and a
vertical lung height of 1–5 cm, have similar degrees of VA/Q
mismatch (2, 17, 18) (Table 1). Most of these animals have
been selected for comparison because they had been investi-
gated during anesthesia in the lateral position, similar to the
giraffe in our study. However, anesthetized, supine, and me-
chanically ventilated pigs and dogs show essentially similar
results (28, 31). Thus the sizes of the studied mammals do not
seem to matter. It should be mentioned that the giraffe has a
bronchial ramification and lobular division similar to other
mammals (24), so gross lung anatomy does not differ between
the studied species.
Examples of VA/Q distributions in anesthetized horse, rab-
bit, and conscious and anesthetized humans are shown in Fig.

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 VA/Q in a Giraffe • Nyman G et al. 1377

A B
l/min l/min
0.05 Rabbit 3.0 Horse

0.04 QS/QT
2.0 8.4%
VD/VT VD/VT
0.03 53%
36%*
0.02
1.0
VA/Q
0.01
QS/QT 1.5%
VA/Q
0.00 0.0
0 0.01 0.1 1 10 100 10 100
0 0.01 0.1 1

Ventilation ™ and Perfusion ˜

C l/min
Man awake
D
7.0 Man anesthetized
l/min
0.4
5.0 VD/VT VD/VT
31% QS/QT 4.6%
37%
0.3

3.0 0.2

QS/QT 0.1
1.0 0.8% VA/Q
0.0 VA/Q 0.0
0 0.01 0.1 1 10 100 0 0.01 0.1 1 10 100
Fig. 2. Ventilation/perfusion distributions (VA/Q) in the anesthetized rabbit and horse (top) and in conscious and anesthetized humans (bottom). QS/QT, shunt
in percent of cardiac output; VD/VT, dead space in percent of tidal volume, including apparatus dead space except in the rabbit where apparatus dead space was
subtracted because of proportionally larger connecting tubes (*). [Top right republished with permission of John Wiley and Sons, Inc. (21); permission conveyed
through Copyright Clearance Center, Inc. (21). Top left reproduced with permission of the European Respiratory Society © (17). Bottom left republished with
permission of Wolters Kluwer Health, Inc. (32), and bottom right republished with permission of John Wiley and Sons, Inc. (15); permission conveyed through
Copyright Clearance Center, Inc.]

2. It should be mentioned that the “low VA/Q” seen in the
giraffe as well as in anesthetized horses and humans may be
converted to absorption atelectasis and cause shunt if higher
concentrations of oxygen concentration are breathed (5).
There are limitations in our study. First, our findings are
limited to one animal only, and it was not a full grown giraffe.
Second, the giraffe was in partial lateral recumbency with its
body in the lateral position and its neck and head in the upright
position, whereas the horse was in the fully lateral position.
However, the abdomen and chest were in a similar lateral
position in both species, so the gravitational influence on the
lung should have been similar. The rabbit had been studied in
a supine position. Finally, anesthesia in humans was induced
during breathing 100% oxygen and continued with ventilation
with 40% oxygen. If ventilation had been with air, as in the
giraffe and horse, the shunt might have been smaller, but the
dispersion of ventilation because of more of “low” VA/Q
regions would have been even larger, as mentioned above.
Also, ventilation was not spontaneous in anesthetized humans
or the rabbit but provided by a mechanical ventilator. However,
no significant differences in shunt and VA/Q distribution have
been seen between spontaneous and mechanical ventilation in
anesthetized humans (11).
In summary, the anesthetized giraffe had a small dead space
ventilation, comparable to that of the horse. The ventilation/
perfusion matching was surprisingly efficient and shunt was
small. These findings compared well with anesthetized horse
and man and even with rat and rabbit. Thus the size of the
studied mammals, from 0.7 to 3 kg to more than 400 kg did not
matter; the gas exchanging capacity of the lung was more or
less the same.
GRANTS
The study was supported by the Swedish Research Council (5315) and the
Swedish Heart-Lung Fund.
DISCLOSURES
No conflicts of interest, financial or otherwise, are declared by the author(s).
AUTHOR CONTRIBUTIONS
G.N., B.R., and G.H. conceived and designed research; G.N. and E.-M.H.
performed experiments; G.N., E.-M.H., and G.H. interpreted results of exper-
iments; G.N., B.R., and G.H. edited and revised manuscript; G.N., B.R.,

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1378 VA/Q in a Giraffe • Nyman G et al.
E.-M.H., and G.H. approved final version of manuscript; E.-M.H. analyzed
data; E.-M.H. prepared figures; G.H. drafted manuscript.
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