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The Gravity of Giraffe Physiology

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12
Giraffe Cardiovascular Adaptations to Gravity
Alan R. Hargens∗ , Knut Petterson† , and Ronald W. Millard‡
∗
Professor of Orthopaedic Surgery, University of California, San Diego;
†
Kungsladugårdsgatan 110, SE 414 76 Göteborg, Sweden; ‡ Professor of Pharmacology & Cell
Biophysics, University of Cincinnati College of Medicine
GRAVITY AND THE CARDIOVASCULAR SYSTEM
The physiological systems of animals have adapted to Earth’s
gravity over the past hundreds of millions of years. In gen-
eral, gravitational adaptations of the cardiovascular system are
more pronounced in terrestrial species with greater height and
thus greater gravity-dependent gradients of blood pressure
from head to feet. For example, dinosaurs (1), tree-climbing
snakes (2), giraffes (3), and other tall animals have evolved
mechanisms to provide adequate blood flow and nutrition to
their brains while restricting blood flow and tissue swelling
in their legs. Terrestrial animals of short stature and marine
animals probably require much less sophisticated cardiovas-
cular adaptive mechanisms. At the other extreme, aquatic
snakes have little ability to withstand gravity out of water
and rapidly “faint” when placed head above tail (2). More-
over, when gravity is absent even over short periods of time,
astronauts experience orthostatic intolerance upon readapta-
tion to gravity (see Chapter 58). Because humans are rela-
tively tall compared to other species of animals, they too have
developed extensive and sophisticated regulatory mechanisms
to maintain cerebral perfusion and prevent lower extremity
edema while in an upright posture. In fact, most understand-
ing of gravitational mechanisms to date relates to observations
in humans. However, taller terrestrial animals, such as the
giraffe, may allow better understanding of the physiological
adaptations to gravity. For example, blood pressure in giraffes
is high to pump blood to their brain, but high blood pres-
sures in their feet would theoretically cause severe dependent
edema.
Cardiovascular systems generate and regulate blood pres-
sure to provide flow to tissues. This blood flow nourishes tis-
sues by supplying oxygen (O2 ) and other nutrients, and by
removing carbon dioxide (CO2 ) and other waste products.
Transmural pressures (Px : pressure gradients from inside to
outside the circulation) dictate the wall stress (S) and degree
of openness or closure (vessel radius r) experienced by circu-
94
latory structures according to the Law of Laplace: Px = S/r
(4). Normally, five components determine transmural pres-
sure in the circulation: (a) the dynamic pressure resulting
from cardiac pumping against peripheral resistance to blood
flow; (b) dynamic pressures due to inertial forces of blood
during activity (e.g., the footward acceleration of blood in
leg vessels at heel strike during locomotion); (c) pressure due
to the finite compliance (volume/pressure characteristics) of
the systemic vasculature, especially the veins (this pressure
is commonly called the mean circulatory filling pressure); (d)
extravascular pressure of tissues and interstitial fluid (e.g., total
tissue pressure); and (e) an intravascular hydrostatic pressure
due to gravity (gravitational pressure). Although extravascu-
lar tissues and fluid have mass, gravity affects extravascular
hydrostatic pressures to a minor extent because tissue fluids
are bound within cells or are discontinuous in most extravas-
cular spaces (except for cerebrospinal fluid). Structural and
contractile tissues respond dynamically, within physiological
limits and over a long period of time, as adaptive structural
and functional remodeling, to increased (or decreased) levels
of physical stress by increasing (or decreasing) their functional
capability and/or mass (5). Although physiological responses
to stress occur within seconds to minutes, longer-term func-
tional and structural compensatory adjustments to the initia-
tion of stress are detectable within hours (6).
TRANSCAPILLARY FLUID BALANCE
Transcapillary fluid shifts between the microcirculation and
the surrounding tissues (Figure 12.1) are dictated by capillary Fig. 12.1
surface area and the ability of capillary basement membrane
and endothelium to retain plasma proteins. These fluid shifts
are determined by hydrostatic and colloid osmotic pressures
according to the Starling-Landis equation (7):
J c = L p A[(Pc − Pt ) − σp (πc − πt )] (Equation 1)
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G I R A F F E C A R D I O VA S C U L A R A D A P T A T I O N S T O G R AV I T Y 95

Figure 12.1. Starling pressures, which regulate transcapillary fluid balance. Pressure parameters that deter-
mine directions and magnitude of transcapillary exchange include capillary blood pressure Pc , tissue fluid
pressure Pt (directed into capillary when positive or directed into tissue when negative), plasma and cap-
illary colloidal osmotic pressure πc , and tissue fluid colloidal osmotic pressure πt . Precapillary sphincters
(P.S.) regulate Pc and capillary flow. It is generally agreed that a hydrostatic pressure gradient (P t > lymph
pressure Pl ) drains off excess tissue fluid under conditions of net filtration. Relative magnitudes of pressure
in resting tissues are depicted by the size of the arrows. (Reproduced with permission from Hargens AR.
Interstitial Fluid Pressure and Lymph Flow. In: Skalak R, Chien S, eds. Handbook of Bioengineering. New
York: McGraw-Hill; 1986:1–35.)

where Jc = net transcapillary fluid shift, Lp = hydraulic con-
ductivity of capillary wall, A = capillary membrane filtration
area, Pc = capillary blood pressure, Pt = tissue fluid pres-
sure, σp = reflection coefficient for plasma proteins, πc =
capillary blood colloid osmotic pressure, and πt = tissue fluid
colloid osmotic pressure.
Again, because blood has continuous columns within the
body, local Pc is greatly affected by gravity. On the other hand,
because tissue fluids are discontinuous in the body, Pt is rela-
tively unaffected by gravity.
In most tissues, a net flux of fluid and plasma proteins
occurs across the capillary membrane (endothelial cells [ECs]
and basement membrane), through the interstitial spaces, and
into the lymphatic system (see Figure 12.1). The hydraulic
conductivity of capillary wall Lp is determined by endothelial
function as well as capillary basement membrane thickness.
Generally, the reflection coefficient for plasma proteins, σ p ,
is determined by the permeability properties of the capillary
wall. Some tissues, such as muscle and skin, have relatively high
σp (≈0.9) such that protein transport rates are low. In other
tissues that have lower σ p , such as lung and liver, πt is almost
equivalent to πc . In these tissues, Lp and filtration rates are rel-
atively high and lymphatic drainage is critical for tissue fluid
balance. Recent evidence also indicates that lymphatic vessels
have a two-valve system, with primary valves at the level of the
initial lymphatic endothelium and a secondary valve system in
the lumen of the transporting lymphatics that facilitates unidi-
rectional transport towards the lymphatic nodes and thoracic
duct (8).
MICROCIRCULATORY ADAPTATION
TO GRAVITY
Transcapillary ultrafiltration, Jc , in the human calf is signifi-
cantly lower than that in the forearm in supine subjects (9).
Although postural changes are associated with increases in
precapillary resistance, altering pressure and flow characteris-
tics within exchange vessels of the dependent limb, no changes
occur in the capillary surface area available for exchange
(10). The autonomic regulation of capillary hemodynamics
in dependent limbs appears to involve a veno-arteriolar reflex
(11). Exploration of dependent limb capillary filtration has
revealed that the α-adrenergic receptor-mediated vasocon-
strictor response in the circulatory system of the human leg is
enhanced upon standing and may represent an adaptive mech-
anism that limits blood pooling and transcapillary ultrafiltra-
tion in the leg during upright position (12). However, should
autonomic nervous system dysfunction (i.e., dysautonomia)
exist, profound hypotension will occur without the typical
reflex tachycardia when standing abruptly. In such individ-
uals, dependent limb vasoconstriction control is decreased
significantly (13). In this regard, orthostatic hypotension is a
common consequence of such dysfunction in elderly patients.
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96 ENDOTHELIAL BIOMEDICINE
Because the human feet and head experience the great-
est change in blood pressure with altered vectors of gravity,
it is worthwhile to understand their structure and functional
responses to gravitational stimuli. Previously, we found that
elevation of blood pressure to the head produced by a transi-
tion from 60 degrees head-up tilt (HUT) to 6 degrees head-
down tilt (HDT) increases microcirculatory blood flow in
skin of the head (14). This pressure and flow increase may
be expected in a tissue not commonly exposed to increased
blood pressure and therefore, not able to regulate blood flow
closely to prevent edema. On the other hand, moving the
body from 6-degree HDT to 60-degree HUT actually reduces
microcirculatory flow in the foot. This response suggests that
the microcirculation of the lower body, normally adapted to
upright posture on Earth, is able to regulate capillary pres-
sure and flow adequately to prevent dependent edema. Com-
bined with vasoconstriction and a thicker capillary basement
membrane in dependent tissues to reduce hydraulic conduc-
tivity (15), other mechanisms for preventing dependent edema
include lymphatic drainage (7) and venous pumps in the leg
(16).
GRAVITATIONAL PHYSIOLOGY
IN THE GIRAFFE
The giraffe, Giraffa camelopardalis, represents a unique mam-
malian model for developmental studies of tissue adaptation
to increasing load bearing. Whereas fetal giraffes develop in
a quasi-weightless milieu, postpartum giraffes must contend
with increasing load bearing in their dependent tissues as they
grow to heights of over 5 meters. Previous studies of the adult
giraffe cardiovascular system (17–19) indicate that this system
is certainly a unique model for investigating the physiology of
being tall and adaptations to large and variable gradients of
blood pressure. These early studies of blood pressures doc-
ument that arterial pressure near the giraffe heart is about
twice that in humans to provide more normal blood pressure
and perfusion to the brain. In the context of higher systemic
blood pressure and longer hydrostatic columns of blood from
heart to feet, the giraffe circulatory system exhibits significant
increases in vascular wall muscle thickness, capillary basement
membrane thickness, and other vascular control systems in
dependent blood vessels that, in concert, participate in capil-
lary fluid dynamics and edema prevention.
During the 1985 Giraffe Physiology Expedition to Africa,
our studies focused on hemodynamic adaptations and edema
prevention in the legs of adult giraffes (3,20). Mean values for
arterial and venous pressures for giraffes and humans qual-
itatively match the expected gravitation pressure gradients,
using the heart as a reference for fluid discontinuity between
Fig. 12.2 the head and foot (Figure 12.2). Although Pc was not mea-
sured directly in the giraffe, it is probably near local venous
pressure (Pv ) in the feet (150 mm Hg) and nearly 10 to 20
mm Hg at the top of the giraffe’s neck (3). Importantly, πc was
identical in the giraffe and human and, therefore, contrary to
an early, unpublished theory by August Krogh. Thus, πc in
the giraffe foot offers no unusual resorptive pressure for pre-
venting dependent edema. Because of rapid circulation and
mixing of blood within the cardiovascular system, it is diffi-
cult to imagine how πc could be significantly higher in the
dependent tissues of the giraffe. Although some Pt values in
the neck were negative, the average bodily Pt ranged between
1 and 6 mm Hg, except under the tight skin and fascia of
the extremities, where mean Pt was 44 mm Hg. Interestingly,
πt in legs was very low (1 mm Hg), except in foot samples
that were contaminated by blood. This finding provided evi-
dence that giraffe capillaries are highly impermeable to plasma
proteins and that σ p approximates unity in lower limbs. This
conclusion was supported by studies of peripheral lymph that
indicated only trace amounts of protein were present and πl
equaled zero.
Of importance, we found that the blood and tissue
fluid pressures that determine transcapillary ultrafiltration
are highly variable with exercise (Figure 12.3). These pres- Fig. 12.3
sures, combined with a tight skin and fascial “antigravity suit,”
move venous blood and tissue fluid upward against grav-
ity, thus preventing pooling of blood and edema in depen-
dent tissues. More proximally, an active skeletal-muscle pump
aids venous return. The nonhydrostatic pressure gradient
down the giraffe’s jugular vein indicates that blood cascades
down from the head, and that circulation above heart level
does not depend on a siphon-like principle, as earlier pro-
posed (21). Other edema-preventing mechanisms include
dependent precapillary vasoconstriction, abundant lympha-
tic drainage, and very low capillary permeability to plasma
proteins.
Developmental alterations are present in the load-bearing
tissues of newborn compared with adult giraffes (22). For
example, vascular wall thickness closely relates to local blood
pressure in the developing giraffe. Tissue samples were col-
lected from four 5- to 6-year-old, 3.5- to 4-meter, male and
female giraffes during the 1985 Giraffe Physiology Expedition;
and from three newborns and two 25- and 35-year-old, 5-
meter adult giraffes from the Cincinnati and Cheyenne Moun-
tain Zoos. Arterioles and other microcirculatory vessels were
sampled from skin and muscle of the head, neck, thorax, legs,
and feet. Although we never had the opportunity to mea-
sure blood pressures in newborn giraffes, based on compa-
rable studies of other species, the arterial pressures of baby
giraffes are probably significantly lower at heart level and in
dependent tissues than are those in adult giraffes. It is appar-
ent that dependent arteries in adults had much thicker walls
than did those in newborn giraffes (Figure 12.4). In adult Fig. 12.4
animals, arterial wall hypertrophy correlates directly with the
degree of tissue dependency (Table 12-1). Thus, the wall Table 12.1
thickness-to-lumen ratios of large arteries increase from head
to foot (except for one “ankle” artery that didn’t follow this
pattern).
We believe the developing giraffe provides an excellent
model for investigations of adaptive mechanisms to natural
hypertension found in the adult giraffe. Arteries of the feet
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G I R A F F E C A R D I O VA S C U L A R A D A P T A T I O N S T O G R AV I T Y 97

Figure 12.2. Mean arterial Pa , venous Pv , and Starling pressures (Pc , πc , Pt , and πt ) in giraffe (right) as
compared to human (left) at hydrostatic levels between the head and feet. In the giraffe, lymph samples
obtained from the leg had only trace amounts of protein and π1 = 0. Foot samples for π t often were
contaminated by blood and therefore were less reliable. Pt beneath the tight skin and fascia of the legs ranged
between 40 and 50 mm Hg and indicated the presence of a “natural anti-G suit” in the giraffe. Data for
pressures in humans are obtained from several sources in addition to our previous studies. (Reproduced
with permission from Hargens AR, Millard RW, Pettersson K, van Hoven W, Gershuni DH, Johansen K.
Transcapillary Fluid Balance in the Giraffe. In: Staub ND, Hogg JC, Hargens AR eds. Interstitial-Lymphatic
Liquid and Solute Movement, Advances in Microcirculation. Basel, Switzerland: S. Karger; 1987:195–202.)

are exposed to blood pressures greater than 400 mm Hg in
adult giraffes (3). These vessels have developed pronounced
smooth-muscle hypertrophy and narrow lumens to accom-
modate these extraordinarily high blood pressures. Our pre-
vious report of a less-than-normal arterial pressure gradient
from heart to foot in the upright, stationary giraffe (3) sug-
gests that the reduction in lumen cross-sectional area plays
some role in blood pressure reduction to dependent tissues.
It is interesting that the arterial wall hypertrophy apparently
is confined to dependent vessels with diameters over 400 μm
microns in adults and is not observed in newborn giraffes or
in vessels near the head of adult giraffes.
CAPILLARY BASEMENT MEMBRANE OF THE
GIRAFFE VASCULAR SYSTEM
In 1971, Williamson and co-workers (15) measured the capil-
lary basement membrane width (CBMW) in muscle obtained
from human infants (n = 19, 9F/1M, 1-day to 6-month-olds),
children (n = 4, 1F/3M, 3- to 5-year-olds), and adults (n =
17, 6F/11M, 36- to 75-year-olds) obtained within 6 hours post-
mortem. They also evaluated muscle from one adult giraffe for
capillary basement membrane thickness. The measurements
in human samples illustrate a significant age-dependent effect,
with the thickest basement membrane measured in the adult
capillaries. The results also illustrate increased CBMW in the
lower legs of adults. CBMW in human infants ranges from
538 to 563 Å and does not vary significantly with anatomic
location. In children, CBMW is significantly thicker (806 to
1,076 Å) in all anatomical sites sampled (pectoral, abdomi-
nal, thigh, calf) than that in the same sample sites of infants.
As in infants, no difference in the CBMW is noted among
anatomical locations in children. In contrast, the CBMW of
adults is significantly greater than that in children and infants,
and shows progressive thickening in dependent tissues mov-
ing down toward the feet (pectoral = 964 Å, calf = 1,894 Å).
In giraffe, the CBMW increases from 780 Å at the neck to
1,437 Å in the front and hind legs. These data are con-
sistent with the hypothesis that CBMW is strongly influ-
enced by intravascular hydrostatic pressure, which is most
extreme in the long hydrostatic column of the giraffe circu-
latory system. This increase in CBMW in the leg of giraffes
may be an important contributing factor to microcirculatory
fluid homeostasis in dependent tissues with high intravascular
pressure.
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98 ENDOTHELIAL BIOMEDICINE

Figure 12.3. Range of averaged mean blood and tissue fluid pressures in neck and foot of walking giraffes.
Negative venous and subcutaneous pressures during exercise help prevent dependent edema. With head
upright, mean arterial pressure below the jaw ranged between 36 and 155 mm Hg (e.g., 145/55 mm Hg for
systolic/diastolic pressures). Drinking water in head-down posture raised carotid pressures to 330/240 mm
Hg. (Reproduced with permission from Hargens AR, Millard RW, Pettersson K, van Hoven W, Gershuni DH,
Johansen K. Transcapillary Fluid Balance in the Giraffe. In: Staub ND, Hogg JC, Hargens AR eds. Interstitial-
Lymphatic Liquid and Solute Movement, Advances in Microcirculation. Basel, Switzerland: S. Karger; 1987:195–
202.)

Figure 12.4. Four-hundred-micron diameter artery from foot skin of adult giraffe has thick wall (left)
compared with a similar artery from foot skin of newborn giraffe (right).
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G I R A F F E C A R D I O VA S C U L A R A D A P T A T I O N S T O G R AV I T Y 99

Table 12-1: Ratios of Smooth-Muscle Wall Thickness/ Table 12-2: Innervation of Giraffe Vasculature
Lumen Radius (w/r) for Arteries from the Neck to the Feet
of Adult Giraffes Adventitial Media
Innervation Innervation
Artery w/r Outer Diameter (mm)
Load bearing Tibial artery + −
Carotid 0.15–0.20 9.5–1.0 vessels Popliteal artery ++ −
Brachial 0.33–0.43 5.5–7.5 Metatarsal artery + −
Femoral 0.65–0.68 4.0–4.2 Tarsal artery ++ +
Ulnar 0.70 7.2 Saphenous vein − −
Radial 0.70 5.4 Metatarsal vein − −
“Ankle” 0.33 4.6 Central vessels Ascending aorta ++ ++
Metatarsal 0.51–0.81 3.1–4.8 Pulmonary artery + −
Digital 0.56 4.0 Superior caval vein − −
Inferior caval vein − −
(Reproduced with permission from Hargens AR, Gershuni DH, Danzig LA,
Millard RW, Pettersson K. Tissue adaptations to gravitational stress: newborn Head and neck Carotid artery +++ +++
versus adult giraffes. Physiologist. 1988;31:S110–S113.) Jugular vein + −

INNERVATION OF THE GIRAFFE
VASCULAR SYSTEM
Giraffes are tall animals, and changing posture will there-
fore cause major circulatory changes, including an increase
of hydrostatic pressure in the head and neck during head-
down drinking. However, the structure of the cardiovascular
system in the giraffe is adapted gravitationally to the normal
upright posture of the animal. Because short-term regulation
of blood pressure and flow is determined by activity in the
sympathetic nervous system, we examined and compared the
innervation of different segments of the cardiovascular system
(23). In general, efferent, sympathetic nerves control cardiac
function, arterial resistance, and venous capacitance.
Tissue specimens were obtained from two young male
giraffes killed as a part of a conservation-culling program
in Africa. We used antisera against dopamine-β-hydroxylase
(DβH, marker of adrenergic nerves), neuropeptide Y (NPY, a
peptide transmitter in sympathetic nerves), as well as gen-
eral markers for nerve terminals (synapsin 1) and nerve
fibers/axon bundles (neurofilament, NF). With these anti-
bodies, a network of nerve terminals and fibers were demon-
strated. The distribution of nerve elements were similar using
different antisera. However, although all antibodies visual-
ized varicose nerve terminals indicative of adrenergic fibers,
Table 12.2 the NF antibody also marked nonautonomic axon bundles
(Table 12-2).
In all arterial blood vessels, a network of nerve terminals
with DβH-, NPY-, and Syn1-positive fibers is found. NF-
positive fibers are sparsely located at the medioadventitial bor-
der, and thick paravascular bundles are found frequently. In
the medial layer of the popliteal artery, NF-positive nerve fibers
also are observed; but the other antibodies used give no label-
ing, indicating that the nerves are not adrenergic. In general,
arteries from the extremities have a relatively sparse innerva-
tion and are usually restricted to the medioadventitial border,
although some evidence for a sparse sympathetic innervation
The density of nerve terminal (axons) near smooth muscle cells ranges from −
(no nerves observed) to +++ (a dense network present).
(Reproduced with permission from Nilsson O, Booj S, Dahlstrom A, Hargens
AR, Millard RW, Pettersson KS. Sympathetic innervation of the cardiovascular
system in the giraffe. Blood Vessels. 1988;25:299–307.)
of the outermost layer of the media is found. The ascending
aorta has a denser innervation, with sympathetic fibers fre-
quently penetrating into the outer medial layers. The carotid
arteries have the densest innervation of all arteries. The nerves
penetrate deep into the media, with fibers reaching almost to
the intima layer.
In veins, the sympathetic innervation is sparse. In fact,
veins from the lower part of the body and caval veins appear
devoid of adrenergic innervation. However, NF-positive, par-
avascular axon bundles are found, and the expected adrenergic
fibers at the medioadventitial border are absent. In the jugular
veins, however, we observed such adrenergic fibers, although
their density still appears sparse.
These observations show that a sympathetic innervation of
the vascular system exists in giraffes, but the innervation pat-
tern is nonuniformly distributed. It is surprising that the caval
and load-bearing veins apparently lack sympathetic innerva-
tion. This is in sharp contrast to man, wherein veins in the
extremities, and especially capacitance veins, have a rich sup-
ply of adrenergic fibers, the activity of which is important for
venous return from the legs (24). We can only speculate on
why giraffes lack this type of innervation, or alternatively, use
other transmitters than those for which we stained. However,
the larger veins in the legs of giraffe run beneath both the tight
skin and a stiff fascia. Thus, the capacitance of these veins may
be limited despite their high venous pressures reported (see
above), so little capacitance is present to regulate.
Sympathetic nerves were, however, found in the jugular
vein. During upright posture, these veins do not need active
regulation, but this could be important during head-down
drinking, when the pressure gradient is reversed and jugu-
lar pressure increases. Valves also are present in the jugular
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100 ENDOTHELIAL BIOMEDICINE
veins (25), and these could—together with sympathetic
nerves—contribute to prevent blood flow in the retrograde
direction and pooling of blood in the head during those short
periods when giraffes lower their head below heart level to
drink.
The innervation of limb arteries is similar to that found in
most mammalian species, and innervation is restricted largely
to the medioadventitial border. However, as the thickness of
the medial layer can be extreme in these vessels, the efficiency
in the spreading of excitation from the nerves must be very
efficient if these nerves are to control the contractions of the
entire medial sheet. The density of the innervation of large
arteries increases from foot to head. In the aorta, we found
fibers in the outer medial layer. In the carotid arteries, sympa-
thetic fibres are found penetrating deep into the media, almost
reaching the intima layer. The physiological significance of
this dense innervation is not known. However, the pattern we
observed is consistent with an adaptive pattern, with dense
innervation in vessels subjected to the largest transient fluctu-
ations during head-down drinking, and a less dense sympa-
thetic innervation occurring in the vasculature of load-bearing
tissues.
In summary, our results concerning tissue adaptations to
increased load bearing in the developing giraffe, along with
those of other investigators, provide intriguing findings that
deserve further investigation, especially from an EC stand-
point. More studies of the developmental biology of giraffes
are needed to elucidate those mechanisms by which tissues
adapt to increased blood pressure in the feet. Such knowledge
may provide useful information for understanding structural
and functional adaptations of blood vessels in human health
and disease.
KEY POINTS
r Despite very high local blood pressures, dependent
edema in the giraffe is prevented by a tight “anti-
gravity suit,” muscle pumping of blood and lymph
toward the heart, and vascular adaptations.
r Vascular walls and capillary basement membranes
increase thickness in dependent tissues.
r Some vascular adaptations in the giraffe are devel-
opmental in nature.
r Surprisingly, arteries and veins of the lower body
have sparse sympathetic innervation.
Future Goals
r To understand better EC structure and function and
their role in the local regulation of giraffe circulations
of blood and lymph
r To determine which adaptations are developmental
in nature.
r To investigate cerebral circulations, in order to un-
derstand adaptive mechanisms during head-down
drinking behavior in giraffes.
ACKNOWLEDGMENTS
These studies were supported by grants from NSF (DCB-8409253),
NIH (HL-32703), National Geographic Society (3072–85), Depart-
ment of Veterans Affairs, and NASA. We thank Professor Wouter
van Hoven and the Department of Zoology, University of Pretoria,
South Africa, for providing giraffes and facilities for our 1985 Giraffe
Physiology Expedition. We thank Dr. D.G.A. Meltzer and Roodeplaat
Research Labs, Dr. Richard Burroughs and the National Zoological
Gardens in Pretoria, and Dr. Frik J. Stegmann and the University of
Pretoria Veterinary School at Onderstepoort. We thank Dr. Paul Calle,
the Cheyenne Mountain Zoo, and the Cincinnati Zoo for providing
specimens of baby and aged giraffes.
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