RESEARCH ARTICLE

Swimming Capability and

Swimming Behavior of
Juvenile Acipenser schrenckii
LU CAI1, RACHEL
2
TAUPIER
1
2
, DAVID 1
JOHNSON , ZHIYING TU
1
, GUOYONG LIU ,
AND YINGPING HUANG *
1
Engineering Research Center of Eco‐environment in Three Gorges Reservoir Region, Ministry
of Education, China Three Gorges University, Yichang, PR China
2
School of Natural Sciences and Mathematics, Ferrum College, Ferrum, Virginia

ABSTRACT Acipenser schrenckii, the Amur Sturgeon, was a commercially valuable fish species inhabiting the
Amur (Heilongjiang) River but populations have rapidly declined in recent years. Dams impede A.
schrenckii spawning migration and wild populations were critically endangered. Building fishways
helped maintain fish populations but data on swimming performance and behavior was crucial for
fishway design. To obtain such data on A. schrenckii, a laboratory study of juvenile A. schrenckii
(n ¼ 18, body mass ¼ 32.7  1.2 g, body length ¼ 18.8  0.3 cm) was conducted using a
stepped velocity test carried out in a fish respirometer equipped with a high‐speed video camera at
20°C. Results indicate: (1) The counter‐current swimming capability of A. schrenckii was low with
critical swimming speed of 1.96  0.10 BL/sec. (2) When a linear function was fitted to the data,
oxygen consumption, as a function of swimming speed, was determined to be MO2 ¼ 337.29
þ 128.10U (R2 ¼ 0.971, P < 0.001) and the power value (1.0) of U indicated high swimming
efficiency. (3) Excess post‐exercise oxygen cost was 48.44 mgO2/kg and indicated excellent fatigue
recovery. (4) Cost of transport decreased slowly with increased swimming speed. (5) Increased
swimming speed led to increases in the tail beat frequency and stride length. This investigation
contributed to the basic science of fish swimming behavior and provided data required for the
design of fishways. Innovative methods have allowed cultivation of the species in the Yangtze River
and, if effective fishways could be incorporated into the design of future hydropower projects on
the Amur River, it would contribute to conservation of wild populations of A. schrenckii. The
information provided here contributes to the international effort to save this critically endangered
species. J. Exp. Zool. 319A:149–155, 2013. © 2013 Wiley Periodicals, Inc.
J. Exp. Zool.
319A:149–155, How to cite this article: Cai L, Taupier R, Johnson D, Tu Z, Liu G, Huang Y. 2013. Swimming
2013 capability and swimming behavior of juvenile Acipenser schrenckii J. Exp. Zool. 319A:149–155.

China has the world's largest number of dams, 87,873 by the end
of 2010 (Ministry of Water Resources, People's Republic of
China, 2010), and continues to build dams for flood prevention
and hydropower. Although dams have brought economic
development, they have led to ecological problems such as
impeding or preventing fish migration. Building fishways and
improving their design can effectively mitigate this problem (Yang
et al., 2011), but successful passage of threatened species through
the fishways depends on how well fishway design matches
swimming capability. Knowledge of swimming performance is
therefore important for effective fishway design (Cheong
et al., 2006).
Grant sponsor: National Nature Science Foundation of China;
Grant numbers: 50979049; 51179096.
*Correspondence to: Yingping Huang, Engineering Research Center of
Eco‐environment in Three Gorges Reservoir Region, Ministry of Education,
China Three Gorges University, Hubei, Yichang 443002, PR China. E‐mail:
chem_ctgu@126.com
Received 19 July 2012; Revised 30 November 2012; Accepted 19
December 2012
Published online 28 January 2013 in Wiley Online Library (wiley
onlinelibrary.com).
DOI: 10.1002/jez.1780

© 2013 WILEY PERIODICALS, INC.

150
Acipenser schrenckii is a rare and commercially valuable fish
species in China. They have mild temperament, strong adaptability
and are benthic feeders. They tolerate a large temperature range
(0–33°C), but grow rapidly and are most active in the range of
13–26°C. Their maximum length is 290 cm and maximum weight
is 160 kg (Zhuang et al., 2002). Wild A. schrenckii inhabit the
Amur (Heilongjiang) River, with small populations in the Wusuli
River and, formerly, in the Nen River. Increased awareness and
innovative methods have led to cultivation of this species in the
Yangtze River (Zhuang et al., 2002) and are crucial to species
conservation. A. schrenckii migrate upstream for spawning in
spring and it takes the larvae about 30 days and begins their
movement downstream (Zhuang et al., 2002).
Research on A. schrenckii has focused on the morphology,
biochemistry and aquaculture of A. schrenckii (Krykhtin and
Svirskii, '97; Zhuang et al., 2002; Shi et al., 2006). The effects of
light on ontogenetic behavior (Zhuang et al., 2003) and of
temperature on oxygen consumption (Song et al., '97) have also
been investigated. Thiem et al. (2011) have study the passage
success and the detailed behaviors within a fishway for sturgeon
(endangered species) migration. In this study, the swimming
performance of A. schrenckii was investigated using a stepped
velocity test in an intermittent‐flow respirometer, equipped with a
high‐speed video camera to record swimming behavior. Numerical
models of A. schrenckii's critical swim speed (Ucrit), oxygen
consumption rate (MO2), cost of transport (COT), tail beat
frequency (TBF), and tail beat amplitude (TBA) were developed
to provide information for fishway design. If effective fishways
were incorporated into the design of future hydropower projects
on the Amur, it would contribute to conservation of wild
populations of A. schrenckii.
MATERIALS AND METHODS
Fish
Juvenile A. schrenckii (n ¼ 18, body mass ¼ 32.7  1.2 g, body
length (BL) ¼ 18.8  0.3 cm) were obtained from an aquaculture
operation in Yichang, Hubei, China (30°560 N; 111°150 E).
The fish were immediately transferred to laboratory aquaria
(1.8 m  1.0 m  0.5 m), where they acclimated for several
weeks. The water was aerated, the temperature was at 10–20°C
approximately, and the photoperiod was 12L:12D approximately.
They were fed daily and any remaining food was removed after
1 hr. Feeding ended 72 hr before testing to prevent an elevated
metabolic rate (Herrmann and Enders, 2000).
Respirometer
Fish were tested in a flume‐type swimming respirometer with a
volume of 11.5 L and a 4.3 L rectangular swim chamber
(39 cm  10 cm  11 cm). The respirometer was submerged in
a 54 L (84 cm  40 cm  16 cm) tank (Fig. 1). The water
temperature was kept at 20  0.5°C by an aquarium heater
J. Exp. Zool.
CAI ET AL.
Figure 1. Schematic diagram of the flume‐type respirometer.
which was under the water. The maximum flow velocity that could
be attained in the chamber was 1.36 m/sec. A flow rectifier
maintained laminar flow in the chamber and a grid at the end of
the chamber prevented fish from being swept away. Water flow
velocity in the respirometer was controlled by a variable‐speed
motor with propeller. Flow velocity was measured by a three‐
dimensional acoustic Doppler velocity meter (Nortek AS, Oslo,
Norway). The respirometer could be flushed with a submersible
pump to maintain the dissolved oxygen (DO) level. The
respirometer was equipped with high‐speed video camera set
over the chamber to record the fish swimming behavior. During
the experiment the respirometer was sealed, and the dissolved
oxygen and temperature in the respirometer were monitored with
a multi‐parameter probe (HACH HQ30d, Loveland, America).
Experimental Design
Constant Velocity Test. The constant velocity tests were conducted
to estimate the routine metabolic rate (RMR or MO2routine) of A.
schrenckii; each test was repeated nine times. Each fish's body
length was measured before the test. The fish were allowed to
adapt to experimental conditions at a flow rate (U) 0.4 BL/sec for
2 hr (Jain et al., '97). The fish swimming speed was assumed to be
the same as the flow rate in the swim chamber. The DO and
temperature of the water were recorded every 10 min for 12 hr.
When the DO fell below 6 mg/L (Tu et al., 2011), the experiment
was interrupted and the water was aerated. When the test was
completed, the A. schrenckii was removed from the swim chamber
and the fork length, total length, and body mass were measured.
The oxygen consumed was calculated based on the decrease in DO
in the sealed respirometer. Oxygen consumed per hour per
kilogram of body mass (MO2) was obtained from Equation (1):
dðDOÞ V
MO2 ¼  ð1Þ
dt m
where V is the volume of the respirometer (L), m the mass of the
selected A. schrenckii, and d(DO)/dt the slope of a linear regression
of DO decrease with time. DO was measured every 10 min and the
slope calculated for each 30 min interval of the test (R2 > 0.99).
SWIMMING CAPABILITY AND BEHAVIOR
With no fish present in the respirometer, the oxygen
consumption rate of the system was <1% of the oxygen
consumption rate of the A. schrenckii, and therefore had a
negligible impact.
Stepped Velocity Test. The stepped velocity tests were conducted to
estimate the critical swimming speed (Ucrit), oxygen consumption
rate (MO2), excess post‐exercise oxygen cost (EPOC), COT, TBF,
and TBA of A. schrenckii; each test was repeated nine times. Fish
body length was measured before the test. As in the constant
velocity tests, fish were allowed to adapt to experimental
conditions at 0.4 BL/sec for 2 hr. During the test, water velocity
was increased by 0.25 BL/sec at 30‐min intervals, starting at
1 BL/sec. The initial DO and temperature in the respirometer were
recorded and measured at 10‐min intervals. When the A.
schrenckii was forced to the back of the swim chamber, the
flow velocity was decreased until it continued swimming. A fish
was regarded as fatigued when it did not swim after the flow
velocity was decreased three times. Once the fish was fatigued, the
velocity was re‐set to 0.4 BL/sec and the DO and temperature in
the respirometer were recorded every 10 min for 1 hr. When the
experiment was completed, the fork length, total length, and body
mass were measured.
The Ucrit was calculated using the flow velocities and step
intervals recorded during the test and the equation below
(Plaut, 2001):
 
tf
U crit ¼ Up þ  Ut ð2Þ
ti
where Up (BL/sec) is the final velocity recorded before the fish
became fatigued, Ut (BL/sec) the velocity increment, tf (min) the
time elapsed at fatigue velocity, and ti (min) the prescribed interval
time. Drag was considered to be negligible as the volume of the
fish was <10% of the swim chamber (Webb, '71).
Oxygen consumption was calculated both before and after the
fish became fatigued, with data fitting for each period. The
correlation, before fatigue, between the oxygen consumption rate
and the swimming velocity was fitted to a linear function:
MO2 ¼ a þ bU ð3Þ
where a and b were constants obtained from the linear function
and U (BL/sec) was the swimming velocity. The oxygen
consumption rate after fatigue was obtained by fitting the data
to an exponential function (Lee et al., 2003):
MO2 ¼ a þ bect ð4Þ
where a, b, and c were the constants obtained from the function, e
was the natural logarithm and t (hr) the time. The excess post‐
exercise oxygen cost (EPOC, mgO2/kg) was the area under the
151
curve of the exponential function remaining above the RMR
baseline.
The COT (J/(kg  m)) was the energy expended in swimming. In
order to derived the correlation between COT and U, MO2 (mgO2/
(kg  hr)) was multiplied by the oxycaloric value of 14.1 J/mgO2
and then divided by U (m/sec). The function was as follows
(Hepher, '88; Videler, '93):
COT ¼ aU 1 þ bU c1 ð5Þ
where a and b were constants and U was the swimming velocity.
The video recordings of fish swimming during the tests were
manipulated with a computer program to obtain TBF and TBA.
Stride length (LS, cm) was calculated using the following function
(Videler and Wardle, '91):
U
LS ¼ ð6Þ
TBF
where U (BL/sec) was the swimming velocity and TBF (beat/sec)
was the TBF.
All experimental data was analyzed with Origin 8.1. The data
were expressed as mean  SEM. The significant level of F‐test
was set as P < 0.001.
RESULTS
The physical attributes of A. schrenckii, along with calculated
results were shown in Table 1. Ucrit was 1.96  0.10 BL/sec. The
correlation (Fig. 2) between the oxygen consumption rate and
swim velocity was calculated using Equation (3),
MO2 ¼ 337:29 þ 128:10U (R2 ¼ 0.971, P < 0.001).
The excess post‐exercise oxygen consumption rate during
recovery from fatigue was fitted to an exponential function (4),
MO2 ¼ 291:28 þ 332:52e6:81t (R2 ¼ 0.997, P < 0.001). Figure 3
indicates that oxygen consumption rate rapidly decreased after
fatigue and 40–50 min was required for recovery. The calculated
EPOC was 48.44 mgO2/kg.
COT versus U was displayed in Figure 4, indicating an inverse
relationship and a good fit was obtained using Equation (5),
COT ¼ 4:68U 1 þ 5:26U 0:54 (R2 ¼ 0.986, P < 0.001).
Correlation between U and TBF was shown in Figure 5 and the
data were fitted to the equation, TBF ¼ 1.73 þ 0.96U (R2 ¼
0.983, P < 0.001). The highest flow velocity was above Ucrit and
did not correlate as well as the other velocities. The relationship
between TBA and U was erratic; the TBA increased dramatically,
decreased and then increased slowly (TBA were 4.8, 7.4, 6.8, 7.1,
7.6, 7.3 cm, respectively, from 1 to 2.25 BL/sec). The highest flow
rate was above Ucrit and the fish was struggling to maintain
position. The data were fitted to several functions but none gave
an acceptable fit. A good fit between LS and U was obtained using
the function, LS ¼ 3.24 þ 1.75U (R2 ¼ 0.998, P < 0.001). All of
the standard errors for the coefficients of functions were shown in
Table 2.
J. Exp. Zool.
152 CAI ET AL.

MO2 (mgO2/(kg  hr))

Kf, condition factor ¼ 100M/LF3; N, number of A. schrenckii; M, body mass; LB body length; LF, fork length; LT, total length; RMR value of every A. schrenckii found by regressive RMR via statistics
416.71–884.31
The range of

—
The range of
U (BL/sec)

1.00–2.50
0.40
RMR (mgO2/(kg  hr))
295.38  10.42

Figure 2. Oxygen consumption rate (MO2, mean  SEM) as a

function of swimming speed (linear function: MO2 ¼ 337:29þ
—

128:10U , R2 ¼ 0.971, P < 0.001).

0.37  0.03
0.37  0.02

DISCUSSION
The work of fishways was poor in China. Because most fishways in
Kf
Table 1. The morphological parameters of the A. schrenckii and the oxygen consumption rate.

China were built by counterfeited other countries' fishways, but

the migration fish were different so that the design of fishways
should be different (Wang and Guo, 2005). The suitable parameter
of swimming performance for fishways should be derived. The
24.1  0.5
23.4  0.7
LT (cm)

swimming performance of A. schrenckii was investigated and

20.8  0.4
20.0  0.5
LF (cm)

(P < 0.05), U, flow velocity; MO2, oxygen consumption rate.

19.0  0.4
18.6  0.5
LB (cm)
33.2  1.2
32.2  2.2
M (g)
N
9
9
characteristic

Figure 3. Excess post‐exercise oxygen consumption related to

time. Area under the curve was used to determine EPOC, with RMR
Constant
Velocity

Stepped

indicated as a baseline (exponential function: MO2 ¼ 291:28þ

332:52e6:81t , R2 ¼ 0.997, P < 0.001).

J. Exp. Zool.
SWIMMING CAPABILITY AND BEHAVIOR
Figure 4. Cost of transport (COT, mean  SEM) as a function
of swimming speed (function: COT ¼ 4:68U 1 þ 5:26U 0:54 ,
R2 ¼ 0.986, P < 0.001).
mathematical models obtained for Ucrit, MO2, EPOC, COT, TBF, and
TBA. When A. schrenckii migrated at spring and autumn, the
water temperature was 10–25°C. The experimental temperature
(20°C) was in the range so that our experimental data were helpful
to the design of fishways.
Critical swimming speed (1.96  0.10 BL/sec at 20°C) was
positively correlated with sustained maximum swimming speed
Figure 5. Tail beat frequency (TBF, mean  SEM) as a function of
swimming speed (linear regression: TBF ¼ 1.73 þ 0.96U, R2 ¼
0.983, P < 0.001).
153
Table 2. Standard error for the coefficients of functions.
Parameters a b c
MO2 ¼ a þ bU 14.93 9.78 —
MO2 ¼ a þ bect 3.77 7.91 0.40
COT ¼ aU1 þ bUc1 12.91 12.76 1.05
TBF ¼ a þ bU 0.08 0.06 —
LS ¼ a þ bU 0.50 0.39 —
MO2, oxygen consumption rate; COT, cost of transport; TBF, tail beat
frequency; LS, stride length; U, swimming speed.
(Zheng et al., 2010). The critical swimming speed was relatively
low, which was similar to A. fulvescens in Peake et al.'s ('97) study,
and to be expected, given A. schrenckii's preferred habitat in the
benthic zone of slow flowing rivers (Zhuang et al., 2002). The
velocity of fishway slot and entrance must be below Ucrit for
successful A. schrenckii migration.
Reported power values for U in the MO2 power function
ðMO2 ¼ a þ bU c Þ ranged from 1.1 to 3.0 and were inversely
related to swimming efficiency (Wieser, '91). If we fitted the data
to a power function, the curve would be similar to straight
line and the power value would be 1.31, within the reported
range. However, the linear function was more concise than the
power function. The power value determined in a linear function
was 1. Peake (2005) had studied swimming and respiration of
white sturgeon and lake sturgeon, and then reported the
correlation between MO2 and U, but function was not derived.
However, we saw this correlation figure and found that if the data
were fitted to function, the curve would be close to a linear so that
the power value would be close to 1 approximately. The low
power value in our study, therefore, was close to other sturgeon.
This value was on the low end of the range, indicating that
A. schrenckii swimming efficiency was high and cost‐of‐
transport was low.
The excess post‐exercise oxygen cost (EPOC) was used to
determine recovery time after fatigue and resting pools were
usually required in fishways. The lower the EPOC, the faster the
fish recovers (Lee et al., 2003) and the EPOC was a critical
parameter for the design of fishway resting pools. Successful
sturgeon fishways should incorporate rapid‐velocity sections
between somewhat slower sections (resting pools) for rest and
recovery (Webber et al., 2007). The EPOC for A. schrenckii was
48.44 mgO2/kg, lower than for salmon (Lee et al., 2003), so resting
pools can be less extensive.
Figure 4 indicates that COT decreases with increasing U and was
consistent with other studies (Tu et al., 2011). This was because
RMR was constant and accounted for a larger proportion of MO2 at
low swimming speeds. However, after a rapid initial decrease, COT
J. Exp. Zool.
154
for A. schrenckii decreased slowly with increasing U, an indication
of efficient swimming.
Other investigators have reported a linear correlation between
TBF and U (Videler and Wardle, '91; Ohlberger et al., 2007b) with a
slope inversely related to swimming efficiency (Videler and
Wardle, '91; Videler, '93; Steinhausen et al., 2005; Tu et al., 2011).
The slope obtained for A. schrenckii was 0.96 (Fig. 5) and, for
comparison, the slopes for Schizothorax chongi, Oncorhynchus
mykiss, Cyprinus carpio, and Rutilus rutilus were, respectively,
0.33, 0.56, 1.31, 1.30 (Videler, '93; Ohlberger et al., 2007b; Tu
et al., 2011).
Except for the large increase from a swimming speed of
1–1.25 BL/sec, the change of TBA with U was small and increased
swimming speed depended more on changes in TBF and LS than
TBA. The width of the fishway minimal slot should be wider than
the maximum TBA of the A. schrenckii.
Water temperature and fish size also have a significant effect on
swimming performance (Ohlberger et al., 2007a; Lyon et al., 2008)
and should be further investigated in the A. schrenckii. However,
the data provided by this investigation on the swimming
performance of A. schrenckii were crucial for the design of an
effective fishway and contributed to the international effort
to maintain wild populations of this critically endangered
species.
ACKNOWLEDGMENTS
We thank Miss Zhang Long‐long who also contributed to the work
providing assistance with the writing.
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