J. Appl. Ichthyol.

18 (2002), 320324 2002 Blackwell Verlag, Berlin ISSN 01758659

Received: September 17, 2001 Accepted: July 10, 2002

Physiology of sturgeon: eects of hypoxia and hypercapnia

By J. J. Cech, Jr. and C. E. Crocker
Department of Wildlife, Fish, and Conservation Biology, University of California at Davis, Davis, California, USA

Summary Physiology is the study of key performance pathways between an organisms phenotypic design (structure) and its patterns of resource use and survival that determine its evolutionary tness. Changes in environmental conditions (e.g. temperature, dissolved gases) may elicit physiological responses, with particular limits and potential costs (e.g. in redirecting energy from growth or reproduction) in shes. The anadromous white sturgeon (WS) Acipenser transmontanus), is native to eastern North Pacic waters where some poorly-circulated inland systems may become hypoxic and some high-density culture systems hypercapnic. In this brief review of our research, we summarize the WS physiological responses to these variables. Juvenile WS (<5 g wet weight) grew more slowly under moderate hypoxic stress (dissolved oxygen ca. 58% of air-saturation), compared with normoxia, at all experimental temperatures (15, 20, and 25 C) apparently due to decreased food consumption rate. Hypoxia also decreased swimming activity and oxygen consumption rate in juvenile WS. Hypercapnic stress (4575 mg CO2 L)1, pH 7.0) also decreased juvenile WS growth, compared with normocapnic (0.52 mg CO2 L)1, pH 8.0) and acidied wellwater (0.52 mg CO2 L)1, pH 7.1) conditions, apparently due to decreased food consumption. Chronic hypercapnia also decreased blood O2 anity (Bohr eect), increased ventilation frequency as well as catecholamines and cortisol concentrations and caused a partially compensated respiratory acidosis. Finally, hypercapnia caused no signicant change in gut blood ow, compared with normocapnia. However, hypercapnia increased stress-induced struggling, which signicantly decreased gut blood ow. Further physiological studies are needed to better understand sturgeon environmental requirements in natural and culture-related settings (including captive breeding programs to restore depleted populations), and their capabilities for dealing with increased negative anthropogenic pressures (including habitat alterations) on their populations. Introduction Physiology is the science that studies of key performance pathways between an organisms phenotypic design (structure) and its patterns of resource use and survival that determine its evolutionary tness (Fig. 1). Sturgeon, as well as other animals, must use their (inherited and developed) structural and design (morphological and genetic) characteristics through biochemical and physiological mechanisms to achieve performance (e.g. tolerances and adjustments to environmental changes, growth, locomotion, and reproduction) levels, to survive (e.g. ghting o pathogens or evading predators) and use needed available resources (e.g. O2, food, mates, living and nesting spaces). Their survival and resource use patterns
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structure sh populations and communities in ecosystems (or in culture systems). The interactions between performance and patterns of resource use in the model (Fig. 1), when using energy as its currency, can be expressed in a bioenergetic relationship: C M G A R SDA E 1 where: C consumed energy (food); M maintenance metabolic energy requirements; G somatic growth energy requirements; A activity energy requirements; R reproreproductive energy requirements; SDA specic dynamic action (energy conversion requirements); E excreted energy in feces and urine. In this relationship, energy uptake by the organism, usually from food, is partitioned into energy conversion categories of body maintenance, growth, activity, reproduction, and specic dynamic action, or excreted (Warren and Davis 1967; Jobling 1993). If an organism is stressed, for example by environmental changes, then energy uptake may be inhibited or energy allocated toward growth or reproduction may be reallocated to maintaining the body and re-establishing homeostasis (Schreck 1981). Changes in environmental conditions (e.g. temperature, dissolved gases, salinity) often elicit physiological responses, which may aect the sh performance levels. Whereas low-level or short-duration changes are dealt with via slight adjustments, stress-inducing water quality changes [e.g. low dissolved O2 (hypoxia), high dissolved CO2 (hypercapnia)] challenge shes, including sturgeons, which may activate a suite of metabolically costly physiological counter-measures. It is important to understand the limits of these responses and their potential costs (e.g. in redirecting energy from growth or reproduction to other energy conversion categories, see equation 1) to meet conservation, management, or commercial goals for protection and utilization of sturgeon resources. The anadromous white sturgeon (WS, Acipenser transmontanus) is native to eastern North Pacic waters (northern California to Gulf of Alaska). In California, there is an enthusiastic coastal and inland WS sportshery and a growing commercial (freshwater) WS culture industry. Because some poorly circulated inland systems may become hypoxic and some high-density culture systems hypercapnic, we measured the WS physiological responses to these variables. The specic objectives of our experiments were to examine growth, acidbase, and cardiorespiratory responses to hypoxia and hypercapnia in juvenile WS. Materials and methods
Experiment series 1: temperature and hypoxia eects on growth

This 34-day experiment examined eects of temperature and hypoxia [produced by dierential aeration of aquaria, featurwww.blackwell.de/synergy

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Eects of hypoxia and hypercapnia

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Fig. 1. Model linking structural (morphologic, genetic) attributes of organisms to population- and community-levels phenomena (biodiversity), via physiological (as well as biochemical and behavioral) pathways of performance (e.g. growth, locomotory abilities, reproductive output), survival (e.g. evading predators, ghting o pathogens), and patterns of resource use (e.g. food, O2, mates, living and nesting space) characteristics. The dashed lines indicate feedbacks between the various components. This model is based on that of Wainwright (1994)

owing wellwater to examine the eects of hypercapnia (Crocker and Cech 1996). Aquarial size and ow rate characteristics matched those in Experiment series 1. These sh were fed a commercial trout chow (Silvercup, Murray, UT) one to three times daily; the wellwater characteristics were the same as those in Experiment series 1. The rst experiment compared normocapnic conditions (aquaria with bubbled air) with hypercapnic conditions (aquaria with bubbled air and with bubbled CO2), which simulated the sturgeon farm production tank CO2 conditions (4575 mg CO2 L)1, pH 7.0). Because much CO2 hydrates with water when dissolved, forming carbonic acid, a signicantly decreased water pH accompanies hypercapnia. To see if the decreased growth resulted from the hypercapnia or from the decreased pH, alone, we conducted two more experiments with dierent size juvenile sh. In these, we decreased aquarial pH via continuous additions of hydrochloric acid to a similar lowered pH (7.1) produced in the hypercapnic aquaria in experiment 1.

Experiment series 4: blood-O2 equilibrium characteristics

ing continuous ows (ca. 0.5 L min)1) of non-chlorinated, wellwater] on growth of 15, young (0.50.6 g mean initial wet weights) juvenile WS in each of six replicated, 38-L (25.4 cm 50.8 cm 30.5 cm) glass aquaria (Cech et al. 1984). Aquaria were separated by 25 mm styrofoam blocks and were housed in a stainless steel cabinet. Temperatures used were 15 (water chiller), 20 (ambient temperature), and 25 C (aquarium heaters in head tank); dissolved oxygen levels were >83% air saturation (normoxia) and 58% air saturation (hypoxia). WS were fed ad libtum rations or adult Artemia and chopped, tubicid worms. Fish were starved for 24 h before weighing (tared beaker of water, Mettler electronic balance) at the start and end of the experiment. Mean (n 6 measurements) wellwater characteristics were: total dissolved solids: 390 mg L)1, total suspended solids: <5.0 mg L)1, total alkalinity: 300 mg L)1, pH 7.8, and hardness: 320 mg L)1.

We also studied blood-O2 equilibrium characteristics to better understand oxygen uptake and delivery characteristics in WS exposed to experimental temperature (15 or 20 C) and dissolved CO2 conditions (Crocker and Cech 1998). Blood was sampled from cannulated WS (n 11 at 15 C, n 6 at 20 C) and loaded into ow-through-type tonometers for equilibration with desired gas mixtures (<1, 30.4, and 76 mm Hg PCO2) and at the temperature set in the water bath. Blood partial pressure of O2 (PO2) and pH data were measured from samples taken and mixed from the various tonometers to construct (duplicate) blood-O2 equilibrium curves and determine the (mean) half-saturation (P50) values.
Experiment series 5: acid-base eects of hypercapnic exposure

Experiment series 2: hypoxia eects on oxygen consumption and activity

We investigated eects of hypoxia on oxygen consumption rates (OCR) and activity rates of dierent size juvenile WS at life-stage-appropriate temperatures (1020 C, Crocker and Cech 1997). Wellwater characteristics were the same as those in Experiment series 1. Using glass closed respirometers, a dissolved oxygen meter, and counterows of water and N2 to produce the hypoxia, OCR (lg O2 g)1 h)1) were calculated using standard techniques (Cech 1990). An overhead video camera and recorder were used to estimate sh swimming (cm h)1) by counting the lines (painted on each respirometer) crossed as the sh swam along the long axis of the respirometer (Crocker and Cech 1997).

Experiment series 3: hypercapnia eects on growth

We conducted another growth study (three experiments) with juvenile white sturgeon in four replicated aquaria of

We also investigated acid-base eects of hypercapnic exposure in cannulated, juvenile WS (Crocker and Cech 1998). Cannulae were implanted into anesthetized (MS-222, 0.1 g L)1) sturgeon (n 46, 0.81.4 kg wet weight) by threading PE-50 polyethylene tubing through a 17 ga. hypodermic needle which had been inserted through the skin and ventral musculature to the hemal canal. Upon assuring that the cannula was in the dorsal aorta (by observing the bright red color and the relatively high pressure of the blood advancing in the cannula), the needle was removed and the cannula was lled with heparinized saline and sutured to the skin at the point of entry. Sturgeon were placed in 30-L, cylindrical ow-through-type chambers, which were situated in temperature controlled water baths. Wellwater characteristics were the same as those in Experiment series 1. Acrylic gas stripping columns and an electronic gas ow meter regulated dissolved CO2 partial pressures (PCO2) in the water entering the chambers. Experimental sh were exposed to either normocapnic (controls) or hypercapnic (30 mm Hg PCO2) conditions over a 96-h experimental period. Blood samples were withdrawn at intervals for measurements of PCO2, PO2, pH, and plasma (norepinephrine), (epinephrine), and (cortisol) (Crocker and Cech 1998). In addition, ventilatory frequency was visually counted, and blood bicarbonate ion concentration was calculated using the HendersonHasselbalch relationship (Davenport 1974). These data were used to compare the relative performance responses of normocapnic and hypercapnic WS.

322 Experiment series 6: cardiovascular responses to hypercapnia

J. J. Cech, Jr. and C. E. Crocker

To measure WS cardiovascular responses to hypercapnia, we surgically implanted ultrasonic blood ow probes on its major arteries (Crocker et al. 2000). We set up the sh (n 28, 1.52.8 kg wet weight) in the same chamber/water bath/gas mixer as in Experiment series 5, to regulate dissolved CO2 in the chambers inowing water. Because we were interested in whether hypercapnia may be limiting gut blood ow, and thereby limiting nutrient uptake and growth of these cultured sh, we measured gut blood ow with one of the blood ow probes (Transonic, ultrasonic) on the celiomesenteric artery and compared it with the total cardiac output (measured using a ow probe on the ventral aorta, Crocker et al. 2000). Results and discussion
Experiment series 1

As temperature increased from 15 to 20 C, under normoxic (or, near-air-saturated) conditions, juvenile WS specic growth rate signicantly increased from 1.6 to 2.6% body weight day)1. In contrast, a further temperature increase to 25 C resulted in only a slight, non-signicant increase (to 2.9% body weight day)1). Further, mildly hypoxic conditions (at 58% of air-saturated conditions) signicantly decreased growth at all temperatures (Cech et al. 1984). This hypoxiainduced decrease in growth appeared to result from decreased food consumption rates and/or decreased energy storage (although these were not quantied). Further, plasma cortisol and catecholamine levels were not quantied in these small sh. Thomas et al. (1992) measured signicant increases in both epinephrine and norepinephrine with exposure to lower dissolved oxygen levels (<20% of air saturation) in larger (7501 200 g wet weight) trout (Salmo fario). A comparative study on the sympatric striped bass (Morone saxatilis) of the same size showed some interesting dierences. Striped bass growth rate increased with each increase in temperature, under normoxia, including to 25 C, where the fastest rates of growth were shown (3.2 % body weight day)1). This species was introduced to California waters in the late 1870s, and their signicant growth rate increase with the 5 C increase to 25 C probably reects the warmer conditions of Chesapeake Bay on the USA Atlantic Coast, where striped bass evolved. Growth of the young striped bass was also sensitive to hypoxia, although not signicantly so at the lowest temperature (15 C), where its growth rate is the lowest (Cech et al. 1984).

displayed signicant decreases in OCR with exposure to mild (51% of air-saturation levels) hypoxia (Table 1). Because we could estimate swimming activity from the young sturgeons movements in the respirometers (crossing lines painted on the glass), analysis of the videotapes showed that hypoxia also signicantly decreased activity (Table 1). This phenomenon occurred regardless of the order to which the sturgeon were exposed to hypoxia: last or rst. Therefore, the signicantly decreased metabolic rate of juvenile WS was at least partially due to decreased activity. The juvenile WS did not show a compensatory increase in OCR when returned to normoxic conditions, after exposure to hypoxia. This response is consistent with the hypometabolic (decreased overall metabolic) response to hypoxia, in part due to decreased activity levels in our juvenile WS, seen also in adult WS (Burggren and Randall 1978). This response may serve WS well in natural habitats where decreased activity, at least in the juveniles, would decrease oxygen demand, thereby conserving oxygen resources in hypoxic habitats until conditions improved. This response diers from that seen in Siberian sturgeon (A. baerii), which do not decrease OCR during hypoxia and show a compensatory OCR increase after return to normoxia (Nonnotte et al. 1993; Maxime et al. 1995). This compensatory increase in OCR repays the oxygen debt associated with anaerobic contributions to overall metabolism (measurable by plasma lactic acid increases), during hypoxia (Nonnotte et al. 1993; Maxime et al. 1995). This distinctly dierent pattern, compared with WS, may stem from habitat-related dierences in the Siberian sturgeons life history (Ruban and Sokolov 1987; McDowall 1988).

Experiment series 3

Experiment series 2

Expressed as mg O2 consumed per gram body weight per hour, the small (0.2 g body weight at 10 C), medium (1.9 g at 16 C), and large (63 g at 20 C) body weight WS all

WS growth performance was signicantly decreased in the hypercapnic conditions (1.17% body weight day)1), compared with normocapnic conditions (2.86% body weight day)1) in experiment 1 of this series (Crocker and Cech 1996). As in the growth experiment with hypoxia (Experiment series 1), the decreased growth was apparently due to a decreased food consumption rate. In neither of the second two experiments did the decreased pH signicantly aect growth, leading us to conclude that the decreased appetite and decreased growth rate seen in the rst experiment were due to the high CO2, not due to the resulting pH decrease (Crocker and Cech 1996). Hypercapnia (5060 mg L)1, pH 6.156.30) also decreased growth of rainbow trout (Oncorhynchus mykiss) after a 330-day chronic exposure (Smart et al. 1979). Sophisticated, high-density WS culture systems (e.g. Stolt Sea Farm Corporation in Elverta, California) may become hypercapnic. Such high densities of sh are possible with the carefully metered addition of pure O2 into the water inows, increasing the quantity of dissolved O2 for the sh to respire.

Table 1 Mean (SE, n 159) juvenile white sturgeon oxygen consumption rate (OCR, mg O2 g)1 h)1) and estimated swimming activity (ESA, cm h)1) at 10 C (0.2 0.0 g live weight), 16 C (1.9 0.1 g), and 20 C (63.1 4.0 g) during normoxia and hypoxia. All hypoxia means are signicantly lower (P < 0.05) than normoxic controls at the same temperature (Crocker and Cech 1997) Normoxia 10 C OCR ESA 266 30 44 3 16 C 210 30 646 22 20 C 178 10 235 10 Hypoxia 10 C 59 13 61 16 C 131 20 225 11 20 C 139 14 28 4

Eects of hypoxia and hypercapnia Table 2 Juvenile white sturgeon half-saturation (P50) values (mm Hg) for normocapnic (<1 mm Hg PCO2), low hypercapnic (30.4 mm Hg PCO2) and high hypercapnic (76 mm Hg PCO2) conditions at two temperatures (Crocker and Cech 1998). Data are means from duplicate measurements from pooled samples (n 11 sh at 15 C, n 6 sh at 20 C) Temperature 15 C 20 C Arterial P50 13 16 Low hypercapnic P50 31 39 High hypercapnic P50 34 44

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increase in the hypercapnic sh, presumably due to the hyperventilation. As expected, arterial pH signicantly decreased in the hypercapnic sh. However, after about 2 days, it started to show a compensation toward that of the normocapnic sturgeon, due to the signicant accumulation of bicarbonate ion. This response parallels that in most teleosts studied so far, but the WS response (ca. 35% of pH restoration) is more limited and slower. For example, Eddy et al. (1977) described a ca. 91% arterial pH restoration after a 7-day hypercapnic exposure in rainbow trout; the marine teleost, Conger, recovered 92% of its initial pH depression after only 30 h hypercapnic exposure (Toews et al. 1983).

However, the high respiring biomass of the sturgeon produces much CO2. Because CO2 is 2530 times as soluble in water than is O2, depending on the temperature, the CO2 tends to stays dissolved, producing hypercapnic conditions.

Experiment series 6

Experiment series 4

Increasing CO2 decreased WS blood O2 anity, from the arterial-type CO2 levels through low- and high-hypercapnic levels we would associate with the high density WS culture (Crocker and Cech 1998). This Bohr eect-type shift results in higher P50 values at increased PCO2 (Table 2). Note that although low hypercapnic conditions (30.4 mm Hg PCO2) dramatically decreased the oxygen anity (increased P50), compared with normocapnic conditions at each temperature, further PCO2 increases (to 76 mm Hg PCO2) do not substantially further decrease blood O2 anity (P50 only increases by 3 and 5 mm Hg at 15 C and 20 C, respectively). Thus, WS in high hypercapnic conditions at 15 or 20 C are able to adequately saturate their blood with oxygen for respiratory purposes under high hypercapnia, almost as much as under low hypercapnia. Studies by Nikinmaa and Soivio (1979) do not show this pattern in rainbow trout blood, albeit at lower PCO2 levels (up to 11 mm Hg PCO2).
Experiment series 5

Although hypercapnic exposure did not aect gut blood ow, the struggling event signicantly decreased the ow (Table 4). Further, we measured an increased number of struggle events when the sturgeon were exposed to hypercapnia, compared to when it was exposed to normocapnia. Thus, the stress associated with hypercapnia could increase struggling, with consequent decreases in gut blood ow, under hypercapnic culture conditions (Crocker et al. 2000). Interestingly, the relative gut blood ow (% of cardiac output) we measured in WS was ca. half that measured in Atlantic cod (Gadus morhua, Axelsson and Fritsche 1991), indicating a possible reduced splanchnic perfusion scope in WS, as compared with Atlantic cod.

Overall

As expected, the arterial CO2 conditions directly reected those in the water (Table 3). Hypercapnic exposure induced stress, as shown in signicantly increased plasma catecholamines (norepinephrine and epinephrine) and cortisol concentrations. Ventilatory frequency consistently increased (hyperventilation) under hypercapnia, compared with the normocapnic controls. Arterial oxygen tension showed either no dierence or a slight

Although WS can complete their life cycle in freshwater environments, the Sacramento River is an important spawning site for at least one anadromous population on the California coast. Their migratory path may extend from either marine conditions in the Pacic Ocean or brackish conditions in the San Francisco Bay Estuary, through the interconnected conuence of the Sacramento and San Joaquin rivers to spawning sites further upstream in the Sacramento system. Localized, poor circulation of water in some of these slough habitats can lead to environmental hypoxia, environmental hypercapnia, or increased temperatures. We were interested in how white sturgeon respond to these environmental factors, especially regarding the performance characteristics to survive and use needed available resources (Fig. 1). Generally, juvenile WS growth (G, in equation 1) decreases with exposure to hypoxia or hypercapnia (Experiment series 1 and 3) can be at least partially explained by decreased food consumption rates

Table 3 Mean (SE) water (W) and arterial blood (A) gas/acid-base, stress hormone (norepinephrine, NE; epinephrine, E; cortisol) levels, and gill ventilatory frequency (Vf) during normocapnia and 96 h hypercapnia in juvenile white sturgeon (n 46, Crocker and Cech 1998) Normocapnia control PW CO2 (mm Hg) PA CO2 (mm Hg) NE (nmol L)1) E (nmol L)1) Cortisol (ng ml)1) Vf (breaths min)1) PO2 (mm Hg) pHA ) (mmol L)1) HCO3 0.3 2.7 64.2 23.9 35.8 59 112.4 7.76 7.1 0.0 0.0 16.4 5.4 9.5 2 4.7 0.02 0.6 0h 0.3 2.7 23.8 15.1 16.3 64 118.5 7.79 7.4 0.0 0.0 4.7 0.8 2.1 2 1.5 0.02 0.9 96 h 0.3 2.5 51.0 19.8 13.6 58 122.7 7.74 6.6 0.0 0.0 13.4 5.6 3.4 2 2.6 0.02 0.6 Hypercapnia control 0.3 2.4 44.4 22.0 49.1 61.0 115.7 7.73 7.2 0.4 0.2 9.6 3.4 9.3 2.1 3.1 0.02 0.5 0h 28.3 24.8 366.7 36.9 44.6 105.2 130.2 7.16 13.9 8.3* 5.2* 90** 10.5** 7.7** 4.7* 3.6* 0.08* 0.6* 96 h 29.3 25.1 68.6 37.7 71.5 108.2 116.0 7.32 21.1 4.4* 1.3* 24.2 3.6** 21.0** 5.7* 4.2 0.03* 1.7*

*indicate signicant dierences (P < 0.05) from control values and **from normocapnic values at the same sampling time.

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J. J. Cech, Jr. and C. E. Crocker

Table 4 Mean (SE, n 28) values for gut blood ow (GBF) as a percent of cardiac output in juvenile white sturgeon during normocapnic and hypercanic exposure, before and after a struggle event (Crocker et al. 2000) Normocapnia GBF (%) 16.3 1.2 Normocap-struggle 3.3 1.1* Hypercapnia 15.8 2.6 Hypercap-struggle 9.3 2.6*

(C). These hypercapnia-related growth decreases may also be mediated by a signicantly decreased blood circulation to digestive organs, from stress-induced struggling events (Experiment series 6). Further, hypoxic exposure decreased energy allocated to activity (A) and, possibly, maintenance metabolism (M) in these sh (Experiment series 2). Regarding metabolism, hypercapnia decreased blood-O2 anity (Experiment series 4) and provoked a stress response in WS with slow acid-base compensation (Experiment series 5). Physiological studies help bridge the gap between earlier, more descriptive studies and the population and community dynamics models that will be needed to predict distributional and abundance patterns. Further physiological studies are needed to better understand environmental requirements of sturgeons, in natural and culture-related settings (including captive breeding programs to restore depleted populations), and their capabilities for dealing with increased humaninduced pressures (e.g. habitat alteration) on their populations. Acknowledgments We thank S. I. Doroshov and his students and colleagues, A. P. Farrell, K. A. Gamperl, S. J. Mitchell, T. E. Wragg, M. J. Massingill, E. B. Kim, L. E. Sharp, D. Stier, S. S. Hung, H. Zhou, C. A. Myrick, D. Lauritzen, M. Dege, P. B. Moyle, P. S. Young, C. Swanson, M. Sheely, M. Gonzalez, and J. Heublein for useful discussions and technical assistance; J. E. Colt and D. Stevens for water quality data, J. T. Michaels and A. Bunter of Stolt Sea Farm (Elverta, CA) for WS; and P. Lutes and W. Bentley (UC Davis Center for Aquatic Biology and Aquaculture) for logistical and facilities support. We appreciate support from a UC Davis Patricia Roberts Harris Fellowship (CEC), a SFSU Research Infrastructure in Minority Institutions award (National Center for Research Resources, Oce of Research on Minority Health, National Institutes of Health #5-P20 RR11805 to CEC), and a UC Agricultural Experiment Station grant (# 3455-H to JJC). References
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Crocker, C. E.; Cech, J. J. Jr., 1997: Eects of environmental hypoxia on oxygen consumption rate and swimming activity in juvenile white sturgeon, Acipenser transmontanus, in relation to temperature and life intervals. Env. Biol. Fish. 50, 383389. Crocker, C. E.; Cech, J. J. Jr., 1998: Eects of hypercapnia on bloodgas and acid-base status in the white sturgeon, Acipenser transmontanus. J. Comp. Physiol. B168, 5060. Crocker, C. E.; Farrell, A. P.; Gamperl, A. K.; Cech, J. J. Jr., 2000: Cardiorespiratory responses of white sturgeon to environmental hypercapnia. Am. J. Physiol.: Reg., Integr. and Comp. Physiol. 279, R617R628. Davenport, H. W., 1974: The ABC of Acid-base Chemistry, 6th edn. Chicago University Press, Chicago, 124 pp. Eddy, F. B.; Lomholt, J. P.; Weber, R. E.; Johansen, K., 1977: Blood respiratory properties of rainbow trout (Salmo gairdneri) kept in water of high CO2 tension. J. Exp. Biol. 67, 3747. Jobling, M., 1993: Bioenergetics: feed intake and energy partitioning. In: Fish ecophysiology (Fish and sheries series 9). J. C. Rankin and F. B. Jensen (Eds). Chapman and Hall, London, 421 pp. Maxime, V.; Nonnotte, G.; Peyraud, C.; Williot, P.; Truchot, J. P., 1995: Circulatory and respiratory eects of an hypoxic stress in the Siberian sturgeon. Resp. Physiol. 100, 203212. McDowall, R. M., 1988: Diadromy in Fishes. Croom Helm, London, 308 pp. Nonnotte, G.; Maxime, V.; Truchot, J. P.; Williot, P.; Peyraud, C., 1993: Respiratory responses to progressive hypoxia in the sturgeon, Acipenser baeri. Resp. Physiol. 91, 7182. Nikinmaa, M.; Soivio, A., 1979: Oxygen dissociation curves and oxygen capacities of blood of a freshwater sh, Salmo gairdneri. Ann. Sool. Fennici 16, 217221. Ruban, G. I.; Sokolov, L. I., 1987: Morphological variability of Siberian sturgeon, Acipenser baeri, in the Lena River in relation with its culture in warm waters. J. Ichthyol. 26, 8894. Schreck, C. B., 1981: Stress and compensation in teleostean shes: response to social and physical factors. In: Stress and sh. A. D. Pickering (Ed.). Academic Press, London, pp. 295321. Smart, G. R.; Know, D.; Harrison, J. G.; Ralph, J. A.; Richards, R. H.; Cowey, C. B., 1979: Nephrocalciriosis in rainbow trout Salmo gairdneri Richardson: the eect of exposure to elevated CO2 concentrations. J. Fish. Dis. 2, 279289. Thomas, S.; Perry, S. F.; Pennec, Y.; Maxime, V., 1992: Metabolic acidosis and the response of the trout, Salmo fario, to acute severe hypoxia. J. Exp. Biol. 87, 91104. Toews, D. P.; Holeton, G. F.; Heisler, N., 1983: Regulation of the acid-base status during environmental hypercapnia in the marine teleost sh Conger conger. J. Exp. Biol. 107, 920. Wainwright, P. C., 1994: Functional morphology as a tool in ecological research. In: Ecological morphology: integrative organismal biology. P. C. Wainwright and S. M. Reilly (Eds). Chicago University Press, Chicago, pp. 4259. Warren, C. E.; Davis, G. E., 1967: Laboratory studies on the feeding, bioenergetics, and growth of sh. In: The Biological Basis of Freshwater Fish Production. S. D. Gerking (Ed.). Blackwell Science Publisher, Oxford. pp. 175214. Authors addresses: Joseph J. Cech, Jr. (for correspondence), Department of Wildlife, Fish, and Conservation Biology, University of California, Davis, 1 Shields Avenue, Davis, CA 95616, USA. E-mail: jjcech@ucdavis.edu C. E. Crocker (present address) Department of Biology, San Francisco State University, 1600 Holloway Ave., San Francisco, CA 94132 USA.