J. Appl. Ichthyol.
15 (1999), 67-72
0 1999 Blackwell Wissenschafts-Verlag, Berlin
ISSN 0175-8659
Cardiovascular responses to hypoxia in the Adriatic sturgeon (Acipenser naccario
By C. Agnisola’, D.J. McKenzie’, D. Pellegrino3, P. Bronzi4, B.Tota3 and E.W. Taylo?
’Dipartimento di Fisiologia Generale e Ambientale, Universit&di Napoli “Federico II”, Napoli, Italy; 2SchooI of Biological Sciences,
University of Birmingham, Birmingham, UK; ’Dipartimento di Biologia Cellulare, Universit; della Calabria, Arcavacata di Rende
(Cs). Italy; ‘ENEL, SRI-CRAM Cologno Monzese (Mr), Italy.
Summary
The in vivo cardiovascular responses to hypoxia, and the intrinsic
functional characteristics of the heart in vitro, were determined,
and compared, in the Adriatic sturgeon (Acipenser nnccarii).
During exposure to hypoxia in vivo, blood oxygen content (Cao,)
declined as water 0, partial pressure (Pwo,) was reduced, despite
an increase in haematocrit. The main cardiovascular response was
a reduction in dorsal aortic blood pressure, with a slight
bradycardia, while cardiac output remained constant. Reduced
oxygen content of the perfusate had significant inhibitory effects
on the intrinsic performance of the heart in vitro, causing a
reduction in the heart rate; a reduction in the sensitivity of
responses to increased preload (Frank-Starling response), and a
more rapid decline in power output and stroke volume when
afterload was increased. Overall, the in vitro results suggest that
hypoxia depresses the contractility of the heart (i.e. its inotropic
responses). The reduction in dorsal aortic pressure in vivo may,
therefore, counteract the depressive effects of hypoxia on heart
contractility, and thereby avoid a hypoxic depression of cardiac
output.
Introduction
As sturgeon live in freshwater, marine and estuarine
environments, and often migrate between them, they may
experience a wide range of environmental variation, including
low oxygen levels, or hypoxia. Teleost fish usually exhibit
homeostatic regulation of oxygen uptake during exposure to
hypoxia, maintaining uptake relatively independent of ambient
oxygen levels. This occurs through reflex changes in ventilatory
activity and/or cardiovascular function, which combine to
increase the effectiveness of branchial gaseous exchange (Hughes
and Shelton 1962). Little is known about how hypoxia influences
the cardiovascular and respiratory physiology of sturgeon, which,
as members of the Chondrostei, are an ancient and unique group
of fish.
In teleost fish the typical cardiovascular response to hypoxia
includes a bradycardia and hypertension, with an increase of both
ventral and dorsal aorta pressures, an increase in peripheral
vascular resistance in the systemic circulation and a constancy or
a reduction in resistance in the branchial circulation (Fritsche and
Nilsson 1993). On the other hand, a small hypotension has been
described in elasmobranchs (Piiper et al. 1970; Butler and Taylor
1971). In teleosts, cardiac output often remains constant due to
compensatory changes in stroke volume, according to the Frank-
Starling response (Fritsche and Nilsson 1993). Similar responses
have been described in an agnathan, the hagfish, Myxine
glutinosa (Axelsson et al. 1990) and in an elasmobranch, the
dogfish ScyIiorhinus canicula (Butler and Taylor 1971; Taylor et
al. 1977; Short et al. 1979).
US.Copyright Clearance Center Code Statement: 0175-8659/99/1504-05-0067$14.00/0
Apart from the direct effects of hypoxia on the vasculature
and the heart, this response is mediated in teleosts and
elasmobranchs by reflexes triggered by branchial and extra-
branchial oxygen receptors, and involves both the autonomic
nervous system and catecholamine release from chromaffin tissue
(Randall and Taylor 1989). The reflex responses of the Adriatic
sturgeon, Acipenser naccari, to stimulation of peripheral
chemoreceptors were described by McKenzie et al. (1 995a).
These reflexes were similar to those described in teleosts
(Burleson et al. 1992), and included a bradycardia and
hyperventilation. Acipenser naccarii was apparently unique in
that it possessed no inhibitory cardiac vagal tone in normoxia, a
fact that might influence cardio-circulatory responses and
regulation of blood flow in hypoxia.
A study by Burggren and Randall (1978) reported that the
white sturgeon (Acipenser transrnontanus) exhibited an unusual
metabolic and ventilatory response to hypoxia, behaving as an
“oxygen conformer” and exhibiting a progressive reduction in
rates of oxygen uptake and gill ventilation as water oxygen
partial pressure (Pwo,) was reduced. A similar hypoxic
depression of oxygen uptake has been described in the Adriatic
sturgeon (McKenzie et al. 1995b). The potential role of the
cardiovascular system in these responses to hypoxia is as yet
unclear.
The present paper describes the cardio-circulatory responses
to hypoxia in the Adriatic sturgeon in vivo, and relates these to
the intrinsic characteristics of the heart as determined in vitro.
Materials and Methods
Animals
Immature Adriatic sturgeon (Acipenser naccarii Bonaparte) of
1.5 years of age, either sex, and with a mean live mass of 1.6 f
0.2 kg were maintained at the Experimental Thermal Aquaculture
Plant “La Casella” (Sannato, PC, Italy) in indoor 4 mzfibreglass
tanks with a continuous water supply (23 rt 1”C, pH 7.9, hardness
125 mg I-’ as CaCO,), and fed a diet of pelletted commercial
sturgeon feed (Alma Storioni Prima Fase, Agros, Bolzano, Italy).
An appropriate number of animals (specified below) were
randomly selected as experimental subjects. All in vivo and in
vitro experiments were performed at La Casella.
Surgical procedures for the in vivo study
Sturgeon were anaesthetised in a buffered solution of tricaine
methane sulphonate (MS222, 1:lOOOO w/v in water) and
transferred to a surgical table where they were artificially
ventilated with a MS222 solution at 1:20000 w/v. Two different
surgical procedures were used for the in vivo study, one for
measuring arterial pressures, heart rate, blood 0, status and
haematocrit, the other for measuring in vivo cardiac output. In the
first case, a saline-filled cannula (PE 50 Intramedic) was
68 C. Agnisola et al.

implanted into the dorsal aorta, via the roof of the mouth and records. Flow was determined volumetrically (Houlihan et al.
secured with a cuff and sutures (Randall et al. 1992). The cannula 1988) and cardiac output expressed as ml m i d k g l . Power
was flushed twice a day with heparinised (50 IU 1.') sturgeon output (PO) was expressed as mW g'l of ventricle and was
saline (see below), and was connected to a pressure transducer calculated as (afterload - preload) o cardiac output/60, where
for measurement of heart rate (beats m i d ) and dorsal aortic pressures were expressed as kPa and cardiac output transformed
systolic and diastolic pressures (kPa). For measurement of in ml min-' g-'. Afterload was defined as the mean output
cardiac output (plus associated heart rate), the ventral aorta was pressure and preload as the mean diastolic intra-atrial pressure.
exposed from the side of the isthmus and a flow probe (Crystal Protocols
Biotech VF-I Pulsed Doppler) was secured around the vessel. Hypoxic exposure in vivo.
The probe lead was tunnelled caudally and firmly secured by skin Measurements of the effects of hypoxia on cardioventilatory and
sutures. Pressure and flow signals were displayed and recorded blood-gas variables were made with a protocol involving 30 min.
on a chart recorder (Gould Windograf). stepwise exposures to 3 progressive levels of hypoxia. For the
Following surgery, fish were transferred to individual measurements of the effects on blood pressures, heart rate and
darkened plexiglass chambers (volume 40 1) and allowed to blood variables, normoxic resting cardioventilatory variables
recover for a minimum of 24 h in a continuous flow of water, were recorded and a 1 ml blood sample was collected for
aerated by passage through a gas-exchange column counter- measurement of control, normoxic Pao,, Cao, and haematocrit
current to a stream of air. (blood was replaced with an equal volume of saline). Water Po,
Measurement of blood variables was then reduced through two levels of hypoxia, 6.6 f 0.2 kPa
Under the conditions defined in the experimental protocols (see (moderate hypoxia), and 4.6 f 0.2 kPa (deep hypoxia). Hypoxia
below), blood samples were withdrawn from the dorsal aortic was created by passing water counter-current to a flow of 100%
catheter, and the collected blood was replaced with an equal N, through a gas-exchange column. At the end of the exposure
volume of saline. Arterial blood oxygen partial pressure (Pao,) periods in moderate and deep hypoxia measurements of steady-
was measured with a Radiometer oxygen electrode, thermostatted state heart rates and blood pressures were made and a blood
to the same water temperature as the fish and attached to a sample was taken for measurement of the above mentioned blood
Radiometer PHM73 acid-base analyser. Arterial blood total variables. Measurements of cardiac output (plus heart rate) were
oxygen content (Cao,) was measured with the technique performed on a separate group of animals (N=5, mean weight
described by Tucker (1967) using an Instrumentation 0.99 f 0.13 kg), following exactly the same protocol of stepwise
Laboratories oxygen electrode and IL 1302 blood-gas analyser exposure to hypoxia.
thermostatted to 37OC. In vitro cardiacperformance.
Portions of whole blood were immediately centrifuged upon A group of 5 animals (mean weight 1.18*0.13 kg) was used to
sampling and plasma decanted and weighed. The red cell pellet test the effect of hypoxia on the intrinsic properties of the heart,
was also weighed and haematocrit was calculated from the employing an in vitro isolated working preparation. The
weights of the liquid and cellular portions of the sample, using following criteria were used to set the basal perfusion conditions:
previously determined measurements of density of each fraction the diastolic output pressure was set to 2.0 kPa, while preload
to calculate their relative volumes (McKenzie et al. 1997). was adjusted to obtain a stroke volume of about 0.2 ml kg".
Isolated heart preparation Basal performance was set with reference to stroke volume rather
Each animal was anaesthetised with 0.2 g/l MS-222 and 1 ml/kg then cardiac output, because in vitro heart rate is lower than in
heparin (80 IU/ml) injected into the caudal arteryhein, then vivo (Agnisola et al. 1996). After stabilisation under basal
killed by a sharp blow to the head and pithed, and the heart was conditions, the isolated hearts were challenged with increasing
dissected and cannulated according to Houlihan et al. (1988). A preloads to induce a maximal in vitro Frank-Starling response
double cannula was set in the atrium to measure intra-atrial and power output. Preload was increased through three to four
pressure. As there are valves at the end of the bulbus cordis (the steps to increase cardiac output and then PO, according to the
most rostra1 contractile chamber of the heart, at the entrance to Frank-Starling relationship, up to a plateau for power output with
the ventral aorta), the isolated preparation included a 7-10 mm. maximal volume loading (PO,,). Subsequently, output pressure
length of the ventral aorta, and care was taken to ensure that was increased through the physiological range as far as possible
cannulation of this vessel did not obstruct these valves. Oxygen without affecting cardiac output to induce a homeometric
supply to the myocardium was sustained by perfusing the hearts response (defined as the heart's intrinsic ability to maintain
with oxygenated saline (Davie et a]. 1992). A perfusion chamber resting stroke volume over a range for aortic output pressure,
that allowed subambient extracardiac pressure to be developed Farrell and Jones 1992) under conditions of maximal volume
during the cardiac cycle was used according to Acierno et al. loading. The maximum PO measured under these conditions was
(1990). The perfusion temperature was 20°C. A modification of equivalent to the in vitro power output of the heart under
Cortland's saline was prepared, with the following composition conditions of maximal volume and pressure loading (POvL,pL).
(in mmol/l): 130 NaCI, 5.09 KCI, 0.1 NaH,PO,, 2.5 Na,HPO,, The total perfusion time was 50 min. As the heart rate appeared
1.14 MgCI,, and 2 CaCI,. Glucose was added (1 g 1"). Total to be independent of both preload and afterload (see below), the
osmolarity of this saline was similar to that of sturgeon plasma values of PO,, and PO,,,,, enabled evaluation of in vitro cardiac
(285 osM; Cataldi et al. 1995). pH was adjusted to 7.8 with scope (as given by the ratio between maximal and basal values)
NaHCO, (about 1 g I-') to agree with direct measurements of in sturgeon under these specified conditions. This volume plus
blood pH from chronically cannulated animals under normoxic pressure loading protocol was repeated twice on the same heart,
conditions (Randall et al. 1992). The saline was gassed with using two different levels of oxygen content in the saline. As
99.5% 0,-0.5% CO, (0, tension, Po, = 65.1 kPa) or with 99.5% previously described by Agnisola et al. (1996) control perfusion
air-0.5% CO, (Po, = 19.8 kPa). Intra-atrial pressure and aortic was with oxygenated saline (0, content = 23 ml r', n = 5), to
pressure were measured with Elcomatic pressure transducers compensate for the reduced oxygen content of saline relative to
connected to a chart recorder [Unirecord, UGO BASILE, blood, while aerated saline (0, content = 7 ml r', n = 5 ) was
Comerio, Italy]. Heart rate was determined from the pressure considered to deliver a hypoxic stimulus to the heart.
 Cardiovascular responses to hypoxia 69

Data analysis The effects of hypoxia on cardiovascular function in sturgeon

Two-way ANOVA or repeated measure ANOVA, with Student- are described in Figure 1.Moderate hypoxia (6.6 kPa) induced a
Newman-Keuls as post hoc test, or paired Student’s t-test were slight but significant bradycardia, associated with a significant
used to compare data as appropriate. Apparent differences reduction in dorsal aortic pressure, which was accounted for by a
between mean values were assigned significance at a confidence reduction of both unchanged by exposure to deep hypoxia.
level of 95% (P< 0.05). Measurement of blood flow revealed that the relatively slight
hypoxic bradycardia was not accompanied by a reduction in
3.5 1 cardiac output (Figure 2), systolic and diastolic pressures (Figure
1). These effects were indicating that a compensatory increase in
T stroke volume had occurred. Thus, the main circulatory response
3.0 - 100
to hypoxia in sturgeon seems to be a reduction in blood pressure
and vascular resistance, accompanied by a slight bradycardia.
2.5 - 75 ‘

75

2.0 -
Q
0-
0
r
4 z
1.5 - 50
v)

3
5’
50 .
I * 4

25

0.5

’.OI
0 ’ I I I 0 25 -

Fig. 1: Effect of hypoxia on in vivo cardiovascular functional 0 -

parameters in sturgeon. fH (beats min -I) = heart rate; PDA(kPa) = dorsal
aorta pressure (squares = systolic; circles = diastolic). fi, values:
normoxia, 18.5 kPa; moderate hypoxia, 6.6 kPa; deep hypoxia, 4.6 kPa.
Data are means (iSE) of 6 determinations. *significantdifference from
the normoxic value (repeated measures ANOVA, P<0.05).

Results
In vivo cardiovascular responses to hypoxia
Hypoxia elicited a reduction in pao,, which was accompanied by Fig. 2: Effect Of hypoxia O n in V i V O heart rate (f”) and cardiac Output
a significant reduction in total oxygen content of arterial blood, (co) in sturgeon. Data are means (* S.E.) of 5 determinations. pwo,
despite an increased haematocrit (Table 1). values as in Fig.1. *significant difference from the normoxic value
(repeated measures ANOVA, W0.05).
Table 1. In vitro cardiac performance in relation to oxygen supply
Effects of moderate (F’wo, 6.6 kPa) and deep (F’wo, 4.6 kPa) hypoxia on
Intrinsic heart rate (fH), as measured in the isolated heart
arterial blood oxygen levels and haematocrit in A. nuccurii.
preparation under basal perfusion conditions was relatively low
F’wo, Pao, Cao, Haematocrit (oxygenated saline: 23.7 f 2.8 beats m i d ; aerated saline: 19.7 f
(kPa) (Ma) (vol Yo) (%I 2.9 beats m i d ) . Under both perfusion conditions, heart rate was
independent of both preload and afterload (repeated measures
18.5 9.93*1.20 10.78&1.I9 22.13i0.87 ANOVA). The mean rate calculated from all measurements,
irrespective of loading conditions, was significantly higher under
6.6 2.4&0.3* 6.4&1.11* 26.64*0.90* oxygenated perfusion as compared to aerated perfusion
conditions (27.2 f 0.6 vs 22.0 f 0.8 beats m i d , respectively).
4.6 1.8&0.34* 4.93i0.98** 25.48*1.22* A reduction in the oxygen content of the perfusate
significantly reduced the extent to which increased preload
*Significantly different from the normoxic value (P<0.05);** stimulated positive inotropic responses by the heart (Frank-
significantly different from both normoxic and moderately hypoxic Starling response). That is, when perfusate Po, was reduced from
values (Repeated measure ANOVA) 65 to 20 kPa, a much larger increase in preload was required to
stimulate the same increase in cardiac output (Figure 3, left
panel). When the Frank-Starling response was expressed as the
ratio between the change in power output (PO,, minus basal) and
the corresponding change in preload, a 4-fold greater value was
obtained under oxygenated as compared to aerated perfusion
conditions (2.64 vs 0.67, respectively).
The ability of the heart to respond to increases in afterload
(Figure 3, right panel) was also significantly reduced in hypoxia.
The maximum in vitro power output, where the heart was
operating under volume plus pressure loading conditions
(POvL,pL)(see Agnisola et al. 1996), was significantly lower in
aerated versus oxygenated saline (0.73 f 0.19 vs 1.05 f 0.22 mW
g-’, respectively), and was attained at a lower output pressure
(4.22iZ0.01 kPa in aerated vs 5.02iZ0.02 kPa in oxygenated
saline). As a consequence, cardiac scope was significantly
reduced by hypoxia (2.9f0.5 in aerated vs 4.6k0.9 in
oxygenated).
Discussion
There are limits to the validity of comparisons which can be
made between the in vifro and in vivo data from sturgeon
(Agnisola et al. 1996). This is in part because of the low oxygen
content of saline relative to blood, which contains haemoglobin
as an oxygen transporter, and the absence of the coronary
circulation in the in vitro preparation. However, both of these

40

c
2m! --
L E
+.r
L m
20
*:
a -

*
0 0
1.o 2.0
U Y
J T

-I-

6. e
0 0
20 30
T

0 0 I I I I
1 2 3 4 5 6

Preload, kPa Afterload, kPa

Fig. 3: Performance of isolated sturgeon hearts perfused under oxygenated (squares) or aerated (circles) conditions with variable preload (left-
hand column) and with variable afterload at maximum preload (right-hand column). Data are means (*S.E.)of 5 determinations.
Cardiovascularresponses to hypoxia 71

problems are somewhat offset by the fact that the fish heart Apparently, some sturgeon species display a different
pumps venous blood in vivo, so that the Po, gradient between behaviour, behaving as oxygen conformers even in moderate
perfusate and myocardium in vitro is, in fact, higher with respect hypoxia (Burggren and Randall 1978; McKenzie et al. 1995b).
to in vivo. In addition, it may be significant that in vitro McKenzie et al. (1995b) found that A. naccarii exhibits a
preparations of the sturgeon heart are characterised by relatively significant reduction in oxygen uptake at the same level of
low heart rates, presumably because of the absence of the moderate hypoxia at which cardiac output was measured in the
excitatory adrenergic control that is exerted in vivo (McKenzie et present study. Thus, the present results indicate that the hypoxic
al. 1995a; Agnisola et al. 1996), such that the oxygen demand of depression of oxygen uptake is not linked to reduced perfusion of
the heart is likely to be lower than in vivo. This may, in part, systemic tissues, and raise the question of how this unusual
explain the ability of the isolated hearts to simulate the in vivo metabolic response occurs. It should be reported, by contrast, that
resting stroke volume (but not the cardiac output) under close to studies on the Siberian sturgeon (A. baeri) indicated that this
in vivo loading conditions, and under both oxygenated and species showed a typical piscine hyperventilation and regulation
aerated perfusion conditions. Thus, despite the lack of an exact of oxygen uptake in hypoxia, down to a critical Pwo, below
matching between in vitro and in vivo cardiac performance, the which aerobic metabolism was depressed (Nonnotte et al. 1993;
effects of changes in the oxygen content of the perfusate on in Maxime et al. 1995).
vitro function may nonetheless give some useful insights into the In conclusion, a reduction in blood pressure and vascular
direct consequences of hypoxia and reduced blood oxygen resistance in vivo may represent an adaptive mechanism in
content on the heart in vivo. sturgeon, which helps to offset the depressive effects of
The in vivo data indicate that the circulatory response of hypoxaemia on the heart, and maintain an adequate blood flow.
sturgeon to hypoxia involves a slight bradycardia, with a Hypoxic depression of oxygen uptake in A. naccarii (McKenzie
compensatory increase in stroke volume, accompanied by a et al. 1995b) does not appear to be linked to reduced perfusion of
reduction in systemic blood pressure and peripheral vascular peripheral tissues.
resistance, so that cardiac output remains unchanged. It is worth
nothing that, while hypertension is the usual response of blood Acknowledgements
pressure to hypoxia in teleosts, a small hypotension has been The study was supported by the EC grant AIR1 CT92- 186.
described in elasmobranchs (Piiper et al. 1970; Butler and Taylor
1971). This suggests that, although the cardioventilatory control
systems in sturgeon are similar in many respects to those of other
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Acipenser naccarii
A splendid hatchery specimen, this 160 cm individual is
now part of the expanding brood stock of A. naccarii
residing at Azienda Agricola V.I.P. di Giovannini
Giacinto, Orzinuovi (BS). Italy.