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Swimming performance and behaviour of young-

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brevirostrum) under fixed...

Article in Canadian Journal of Zoology · February 2012

DOI: 10.1139/z2012-004

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 345

Swimming performance and behaviour of young-

of-the-year shortnose sturgeon
(Acipenser brevirostrum) under fixed and
increased velocity swimming tests
D. Deslauriers and J.D. Kieffer
Can. J. Zool. Downloaded from www.nrcresearchpress.com by Guangzhou Jinan University on 06/05/13

Abstract: Swimming performance and behaviour in fish has been shown to vary depending on the investigation method. In
this study, an endurance swimming curve was generated for young-of-the-year shortnose sturgeon (Acipenser brevirostrum
LeSueur, 1818) (~7 cm total length, ~2 g) and compared with values determined in a separate incremental swimming (crit-
ical swimming, Ucrit) test. Using video, tail-beat frequency (TBF) was quantified and compared for fish swimming under
both swimming tests. From the endurance-curve analysis, it was found that sturgeon did not display a statistically significant
burst swimming phase. Maximum sustainable swimming speed (calculated to be 18.00 cm·s–1) from the endurance curve oc-
curred at ~80% of Ucrit (22.30 cm·s–1). TBF was similar at all speeds for both swimming tests, except at speeds approaching
Ucrit, where fish displayed TBFs of 4.29 Hz for the endurance protocol and 2.26 Hz for the Ucrit protocol. TBF was more
variable between individuals swimming at the same speed within the Ucrit compared with the endurance protocol. Finally, a
significant negative correlation was found between TBF and Ucrit in individual fish, suggesting that station-holding may be
an important energy saving strategy during swimming in this size class of sturgeon.
Key words: Acipenser brevirostrum, shortnose sturgeon, Ucrit, endurance, tail-beat frequency (TBF), station-holding.
For personal use only.

Résumé : On a démontré que la performance et le comportement de nage chez les poissons varient selon la méthode d’é-
tude retenue. Nous avons produit, dans notre étude, une courbe de nage d’endurance pour les jeunes de l’année de l’estur-
geon à museau court (Acipenser brevirostrum LeSueur, 1818) (~7 cm de longueur totale, ~2 g) et l’avons comparée aux
valeurs obtenues dans un test séparé de nage par incréments (nage critique, Ucrit). Nous avons mesuré par vidéo la fréquence
de battement de la queue (TBF) et l’avons comparée chez les poissons dans les deux tests de nage. L’analyse de la courbe
d’endurance révèle que les esturgeons ne font pas de phase de nage précipitée statistiquement significative. D’après la
courbe d’endurance, la vitesse de nage maximale soutenue (évaluée à 18,00 cm·s–1) se produit à ~80 % de Ucrit (22,30 cm·s–1).
TBF est semblable à toutes les vitesses dans les deux tests de nage, sauf aux vitesses proches de Ucrit où les poissons affi-
chent des TBF de 4,29 Hz dans le protocole d’endurance et de 2,26 Hz dans le protocole de Ucrit. TBF est plus variable en-
tre les individus qui nagent à la même vitesse dans le protocole d’Ucrit que dans le protocole d’endurance. Enfin, il existe
une corrélation négative significative entre TBF et Ucrit chez les poissons individuels, ce qui laisse penser que le maintien
sur place peut être une stratégie pour sauver de l’énergie durant la nage chez cette classe de taille d’esturgeons.
Mots‐clés : Acipenser brevirostrum, esturgeon à museau court, Ucrit, endurance, fréquence de battement de la queue (TBF),
nage sur place.
[Traduit par la Rédaction]

Introduction and anaerobic performance (Beamish 1978). For these tests,

fish swim against a constant water speed and the time they
For the past ~50 years, researchers have used the incre- swim against the current defines their endurance at that par-
mental (critical swimming, Ucrit) and the fixed (endurance) ticular speed. In theory, three distinct phases of swimming
swimming tests to evaluate swimming performance in fish can be observed: sustained, prolonged, and burst (Beamish
(Brett 1964; Beamish 1978; Kieffer 2010). The Ucrit test is 1978). Sustained swimming includes speeds at which a fish
performed by swimming fish in increasing water velocities can theoretically swim for an infinite period of time. In prac-
(increments) for predetermined periods of time (intervals; tice, sustained swimming is for ≥200 min at a given speed
Brett 1964). In contrast, endurance swimming tests assess (Beamish 1978; Adams et al. 1999). The speeds that fish
performance across the full range of speeds achievable by can maintain between 20 s and 200 min are considered to be
the fish, and thus allow for the quantification of both aerobic within the prolonged phase of swimming. This phase is often
Received 11 August 2011. Accepted 10 January 2012. Published at www.nrcresearchpress.com/cjz on 22 February 2012.
D. Deslauriers. Department of Biology and Canadian Rivers Institute, University of New Brunswick, Saint John, NB E2L 4L5, Canada.
J.D. Kieffer. Department of Biology, University of New Brunswick, Saint John, NB E2L 4L5, Canada.
Corresponding author: James D. Kieffer (e-mail: jkieffer@unb.ca).

Can. J. Zool. 90: 345–351 (2012) doi:10.1139/Z2012-004 Published by NRC Research Press
 346
associated with high performance variability owing to uneven
levels of activity (Webb 1975; Hammer 1995) and is believed
to be the result of a switch between aerobic and anaerobic
metabolism that might differ among similar-sized fish of the
same species (Nelson et al. 2002). The third or burst swim-
ming phase consists of fast speeds that can be maintained
for <20 s (Jones 1982). During this phase, fish rely solely
on anaerobic energy supplies (i.e., glycogen, PCr, ATP) to
power swimming until their depletion combined with the ac-
cumulation of metabolic wastes (i.e., lactate) leads to exhaus-
tion (Dobson and Hochachka 1987; Kieffer 2000). Endurance
Can. J. Zool. Downloaded from www.nrcresearchpress.com by Guangzhou Jinan University on 06/05/13
swimming tests are not used as commonly as Ucrit swimming
tests, mainly because they are time-consuming to generate
and a large number of fish are required.
Very few studies have attempted to compare behaviour and
performance results from both tests on the same species. By
definition, Ucrit is the maximum speed a fish can swim
aerobically. However, because this test eventually leads to fa-
tigue, it likely represents prolonged activity more accurately
than sustained activity (Webb 1975; Beamish 1978; Kolok
1999; Hasler et al. 2009), and thus, can be an overestimation
of maximum aerobic capacity (Peake and Farrell 2006).
Webb (1971) suggests that during Ucrit swim trials, salmonids
(rainbow trout, Oncorhynchus mykiss (Walbaum, 1792);
sockeye salmon, Oncorhynchus nerka (Walbaum, 1792))
transition from aerobic to anaerobic metabolism at around
For personal use only.
80% of their final Ucrit value. In contrast, Jones (1982) sug-
gests the transition happens near 50% of Ucrit for certain cyp-
rinid species. These conclusions were based on metabolic-
rate measurements, lactate production, and muscle contracti-
bility under various swimming conditions. A different ap-
proach to allow for this comparison is to contrast Ucrit with
the maximum sustainable swimming speed of endurance
swimming tests (i.e., the speed at which fish switch between
the sustained and the prolonged phase). By establishing
where Ucrit values lie within the endurance curve, it is possi-
ble to estimate more accurately the speed at which the onset
of anaerobic metabolism occurs, and thus, help in predicting
the switch between aerobic and anaerobic fuel usages. This
approach might prove useful for certain species, such as the
shortnose sturgeon (Acipenser brevirostrum LeSueur, 1818),
that do not produce large lactate loads during exercise
(Kieffer et al. 2001; Baker et al. 2005).
In a recent study, Deslauriers and Kieffer (2011) found that
there was high variability associated with performance values
obtained from endurance-type tests compared with Ucrit per-
formance values in A. brevirostrum. This variability was
linked in part to different water acceleration rates (i.e., veloc-
ity increments) between tests. In that study, it was also ob-
served, but not quantified, that certain behavioural aspects of
swimming may be different between the two tests. In particu-
lar, sturgeon modify their swimming behaviour by station-
holding (i.e., pressing body and modifying the angle of the
pectoral fins against the bottom of the swim tunnel to gener-
ate negative lift; Arnold et al. 1991) or substrate skimming
(Kieffer et al. 2009) with changes in water speed. It can be
anticipated that tail-beat frequency (TBF) will be negatively
correlated with station-holding at any given velocity. Hence,
station-holding can be assessed by examining the tail-beat
frequency of the fish because when a fish is station-holding,
the caudal fin is not moving. TBF is often used as an index
Can. J. Zool. Vol. 90, 2012
of swimming behaviour and has been shown to correlate pos-
itively with swimming speed (Webb 1986).
The objectives of this study were to (i) generate a com-
plete endurance curve for juvenile A. brevirostrum; (ii) com-
pare Ucrit values to the maximum sustainable swimming
speed values acquired in the endurance test; and (iii) compare
TBF of fish swimming during both Ucrit and endurance pro-
tocols. Meeting these objectives will provide a clearer assess-
ment of whether Ucrit is primarily an aerobic swim test.
Furthermore, the determination of a full endurance curve
will allow us to gain important information on swimming
performance at various speeds in sturgeon, while providing
inflection points that identify the transition between swim
phases. Finally, the discrepancy in data variability generated
from both methods (Deslauriers and Kieffer 2011) might be
explained by assessing behaviour aspects of swimming, such
as TBF.
Materials and methods
Animal husbandry
Fish were the progeny of wild-stock parents originating
from the Saint John River in New Brunswick, Canada. The
fish were provided by Acadian Sturgeon and Caviar Inc.
(Carter’s Point, New Brunswick, Canada). Fish (total length
7.07 ± 0.38 cm (mean ± SD), mass 1.70 ± 0.27 g (mean ±
SD)) were held in 235 L circular tanks at a density of
15.6 g·L–1 in flow-through conditions with a temperature of
15 ± 1 °C and water flow of 10 L·min–1. The fish were fed
daily to satiation (Ewos micro #2 salmon feed) and no food
was given 24 h prior to the swim trial. A total of 71 fish
were used in the experiments.
Flume description
The swimming section was 7.5 cm (width) × 7.5 cm
(height) × 28 cm (length) (Swim-5; Loligo Systems, Tjele,
Denmark). The swim tunnel was fully submerged into an am-
bient tank that allowed for water temperature control (15 °C).
An air stone was placed in the ambient tank to ensure high
oxygen concentrations (≥90% saturation). Ammonia levels
were always low (<0.01 ppm). Because of the relatively
small fish size, a 1.2 mm mesh-sized grid was placed over
the existing downstream grid. The propeller was powered by
an external electro-motor and could accelerate water up to
50 cm·s–1. Flow straighteners were placed up- and down-
stream of the swimming chamber to ensure laminar water
movement. The flume was calibrated using a portable flow-
meter (Loligo Systems, Tjele, Denmark).
Experimental protocol
Critical swimming (Ucrit) protocol
Once placed in the flume tank, individual fish were al-
lowed a habituation period of 30 min at 15 °C at 0 cm·s–1
(n = 6). Following this period, the water velocity was in-
creased by 3 cm·s–1 every 20 min until fatigue occurred.
Fish were considered fatigued when they were impinged on
the downstream grid. Once the fish was impinged on the
rear screen for 5 s, the trial was ended, and the time to fa-
tigue was recorded and the mass and length of the fish were
measured. Fish were swum only once to avoid training
Published by NRC Research Press
 Deslauriers and Kieffer
effects. Critical swimming (Ucrit) was calculated as Ucrit
(cm·s–1) = Vf + [(T1/t) × dv], where Vf is the speed (cm·s–1)
of the last completed swimming period, dv is the velocity in-
crement (cm·s–1), t is the time interval (min), and T1 (min) is
the time swum at the final velocity before fatigue (Brett
1964).
Endurance swimming protocol
Once placed in the flume tank, individual fish were al-
lowed a habituation period of 30 min at 15 °C at 0 cm·s–1.
Following this period, the water velocity was increased to
Can. J. Zool. Downloaded from www.nrcresearchpress.com by Guangzhou Jinan University on 06/05/13
15 cm·s–1 for 1 min, allowing the fish to orient against the
current, before water velocity was adjusted within 5 s to the
test speed. Once the appropriate water speed was reached, the
water velocity was kept constant throughout the swimming
trial. Individual fish were swum at speeds ranging from 6 to
42 cm·s–1 with 3 cm·s–1 increments.The trial was terminated
when the fish had swum for 270 min or if the fish had fa-
tigued and was impinged on the downstream screen for 5 s.
Time to fatigue, mass, and length for each fish were recorded
following the swim trial. Fish were swum only once to avoid
training effects. Five fish were tested individually at each
speed. A total of 65 fish were used for this experiment.
The experimental protocols were approved by the Univer-
sity of New Brunswick Animal Care Committee, following
guidelines established by the Canadian Council on Animal
For personal use only.
Care.
Video analysis
A video camera (Sony DCR-SR40; Sony Inc., Tokyo, Ja-
pan) was installed on a tripod 1 m above the swim flume
with a full view of the entire swimming area. Video footage
allowed for the quantification of TBF at various speeds dur-
ing both Ucrit and endurance swimming tests. Station-holding
behaviour could not be quantified using this approach, as this
would require filming swimming fish from the side view of
the flume (Kieffer et al. 2009). One tail beat was considered
complete when the upper lobe of the caudal fin had com-
pleted the full amplitude of its range and returned to its ini-
tial position. Videos were imported into Windows Movie
Maker software where frame by frame analysis (30 frames·s–1)
was performed. TBF (Hz) was counted by observation of the
upper lobe of the caudal fin. TBF was calculated on 6 and 5
(per speed) fish for Ucrit and endurance tests, respectively.
For the Ucrit swim trials, video was taken for the 1st min
and the 19th min for each time interval. Both TBF values
for each time interval were averaged. If a fish did not com-
plete an interval, only the TBF value calculated during the
first min was used.
Once the test velocity was reached for the endurance swim
trials, video was taken during the 1st min and the 19th min
for each fish at every velocity and TBF was averaged. TBF
(Hz) was calculated only for the 1st min if the fish did not
reach the 20 min mark.
Statistical analysis
Prior to any statistical analyses, data were tested for nor-
mality (Anderson–Darling test) and homogeneity of variance
(Levene’s test). Data for Ucrit and endurance were tested us-
ing linear regression analysis to determine if body length
had any influence on swimming performance. Endurance
347
swimming data were not normally distributed and variance
was not homogeneous; therefore these data were log-
transformed prior to statistical analysis. A piecewise regres-
sion analysis on the data for the endurance curve was per-
formed. Piecewise regression uses established model-fitting
techniques to test whether the relationship between two vari-
ables is the same over the entire range of the sampled inde-
pendent variable (Crawley 2007). Breakpoints, where the
relationship between the dependent and independent varia-
bles changed, were used to estimate the transition points be-
tween swim phases. An ANOVA on the residuals was run
sequentially to determine the best-fit model while comparing
it with the nonsegmented regression. TBF was compared be-
tween swimming tests for the range of speeds leading up to
Ucrit (22.30 cm·s–1) using an ANCOVA with velocity and
swimming test as factors and fish length as covariate. Simple
regression analysis was used to determine the relationship be-
tween TBF and velocity for both endurance and Ucrit tests.
Results were considered significant at a level of p < 0.05.
All statistical tests on TBF were performed using Minitab
version 16 software (Minitab Inc., State College, Pennsylva-
nia, USA). The endurance curve was analyzed using R soft-
ware (R Foundation for Statistical Computing, Vienna,
Austria).
Results
Fish length was not correlated with swimming perform-
ance for either Ucrit (p = 0.294) or endurance (p = 0.117)
swimming tests. The Ucrit values of juvenile sturgeon were
22.30 ± 5.17 cm·s–1 (mean ± SD). Endurance data were
non-normal (p < 0.005) following log-transformation, but
variances were homogeneous (p = 0.426). As a result, the
log-transformed data were analyzed using a piecewise regres-
sion; this analysis showed that the data were best fit as two
separate lines with a break at 18 cm·s–1 (Fig. 1): left-hand
slope equals loge(endurance) = 5.99 – 0.07V and right-hand
slope equals loge(endurance) = 4.84 – 0.15V), where endur-
ance is measured in min and velocity (V) is measured in
cm·s–1 (multiple r2 = 82.69%).
The left-hand slope represents the sustained phase of
swimming, whereas the right-hand slope represents the pro-
longed phase of swimming. The slope breaks at 18.00 cm·s–1,
suggesting that speeds <18 cm·s–1 fall within the sustained
phase, whereas speeds ≥18 cm·s–1 are within the prolonged
phase of swimming. The model with two “pieces” fits the
data significantly better than the one-piece model (one-way
ANOVA, p < 0.005). However, the three-piece model (with
an additional break point at 33.00 cm·s–1) was found to be
nonsignificant (one-way ANOVA, p = 0.107).
Out of the 20 fish swimming within the sustained phase,
16 fish reached the 200 min mark (i.e., the standard criterion)
and were able to swim for the additional 70 min. Fish swim-
ming at 6 and 9 cm·s–1 (10 fish) all reached the 270 min
mark, whereas 6 out of 10 fish were able to complete the en-
tirety of the swim trials at 12 and 15 cm·s–1.
Fish length did not significantly influence TBF (p =
0.131). Both velocity and type of swimming test (i.e., Ucrit
and endurance) significantly affected the TBF (p < 0.005)
(Fig. 2). Mean TBFs were 1.15 and 1.61 Hz at 6 and
9 cm·s–1, respectively, following the Ucrit protocol; these
Published by NRC Research Press
 348 Can. J. Zool. Vol. 90, 2012

Fig. 1. Endurance curve (log-transformed) for juvenile shortnose sturgeon (Acipenser brevirostrum) (n = 5 per speed). The first segment of
the curve represents sustained swimming (loge(endurance) = 5.99 – 0.07V, where V is velocity), whereas the second segment represents pro-
longed swimming (loge(endurance) = 4.84 – 0.15V). The break in the line occurs at 18 cm·s–1.
1000

100
Log e (endurance) (min)

10
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0.1

0.01
0 5 10 15 20 25 30 35 40 45
Velocity (cm·s –1)

Fig. 2. Tail-beat frequency (TBF) data (mean ± SE) for both endurance (n = 5 per speed) and critical (Ucrit) (n = 6 per speed until 18 cm·s–1;
n = 5 at 21 cm·s–1; n = 3 at 24 cm·s–1; n = 1 at 27, 30, and 33 cm·s–1) swimming tests for juvenile shortnose sturgeon (Acipenser breviros-
For personal use only.

trum). The vertical broken line represents the critical swimming speed. The asterisk represents a significant difference at speeds below Ucrit.
8
Ucrit = 22.30 cm·s–1

6
Tail-Beat Frequency (Hz)

5
*
4

3
Endurance
Ucrit
2

0
0 5 10 15 20 25 30 35 40 45
Velocity (cm·s –1)

were not significantly different than those determined for the
endurance test at the same speeds (1.79 and 2.16 Hz). TBF
values were 4.52 Hz at 21 cm·s–1 for the endurance protocol
and were significantly different than those for the Ucrit proto-
col (2.26 Hz) at the same speed. TBF values between 12 and
18 cm·s–1 for both swimming tests were statistically similar
(Fig. 2). Regression analysis for TBF for both swimming
tests can be described as follows: TBFendurance = 1.360 +
0.123V (r2 = 56.5%) and TBFUcrit = 1.22 – 0.002V3 +
0.046V2 – 0.248V (r2 = 37.9%).
The regression calculated from the Ucrit swimming test
represents TBF values for speeds up to 21 cm·s–1. At higher
velocities, the data are skewed by a few fish (i.e., three fish at
24 cm·s–1 and one fish at 27, 30, and 33 cm·s–1) and there-
fore do not give a representative value for the entire group
(Fig. 2). The variability was the highest for the endurance
swimming trial (i.e., in swimming time values) with coeffi-
cients of variation (COV) ranging from 37.6% at 39 cm·s–1
to 186.6% at 27 cm·s–1 (n = 5 for each speed). The critical
swimming test showed less variation (i.e., in Ucrit values)
with a COV of 23.2% (n = 6). In comparison, TBF data
showed less variability within the endurance swimming test
(mean COV: 36.0%) than for the Ucrit test (mean COV:
51.5%) for speeds ranging from 6 to 21 cm·s–1 (Table 1).

Published by NRC Research Press

 Deslauriers and Kieffer 349

Fig. 3. Mean tail-beat frequency (TBF) per velocity increment for individual shortnose sturgeon (Acipenser brevirostrum) (n = 6) under the
critical (Ucrit) swimming test. Values noted above the last data point represent the Ucrit of each fish. Individual fish are represented by different
symbols.
5.0

4.5 15.30
19.65

4.0 20.85
Tail-Beat Frequency (Hz)
3.5
23.10

3.0
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2.5 24.15

2.0

1.5

1.0

0.5
30.75

0
0 5 10 15 20 25 30 35

Velocity (cm·s –1)

For personal use only.

Table 1. Coefficient of variation (%) for both endurance
(n = 5) and critical (Ucrit) (n = 6) swimming tests at
velocities up to critical swimming speed for shortnose
sturgeon (Acipenser brevirostrum).
Velocity (cm·s–1) Endurance (%)
6 48.72
9 53.87
12 28.61
15 21.25
18 34.12
21 29.34
Lastly, TBF for each fish was plotted against test velocity
(Fig. 3) to allow for the visualization of behavioural perform-
ance (i.e., TBF) and its relationship with Ucrit. The results in-
dicate that a negative relationship exists between TBF and
Ucrit values (Fig. 3).
Discussion
Under the endurance protocol, juvenile A. brevirostrum are
able to swim within the sustained and prolonged phases over
the speeds tested. These results are similar to those obtained
for juvenile lake sturgeon (Acipenser fulvescens Rafinesque,
1817) (Peake et al. 1997; Hoover et al. 2005) and white stur-
geon (Acipenser transmontanus Richardson, 1836) (Boysen
and Hoover 2009). In contrast, Adams et al. (1999) showed
that a third (burst) phase also existed for the pallid sturgeon
(Scaphirhynchus albus (Forbes and Richardson, 1905)) over
the speeds tested. This switch from prolonged to burst swim-
ming occurred at ~2.68 BL·s–1 (where BL is body length) for
~15 cm S. albus, whereas burst swimming in A. brevirostrum
was not evident statistically (i.e., no change in slope), even at
speeds approaching 42 cm·s–1 (~6 BL·s–1; Fig. 1).
It has commonly been assumed that if a fish swims for
200 min or longer at a particular speed during the endurance
swimming test, it is swimming in the sustained range of
speeds (Brett 1967). However, this criterion has not been
consistently followed across studies, and experiment termina-
Ucrit (%) tion points include 240 min (Beamish 1978), 480 min
55.83 (Adams et al. 1999), and several days (Walker and Pull
61.43 1973; Beamish 1978). In the current study, a period of
32.74 270 min was chosen to reflect sustainable swimming speeds,
35.57 mostly to verify if A. brevirostrum were able to withstand
51.07 sustainable speeds longer than the prescribed 200 min. At
72.40 slower speeds, all 16 fish that reached the 200 min point
were able to swim at the same speed for the additional
70 min. Although all fish were able to swim for 270 min at
speeds of 6 and 9 cm·s–1, 40% of fish were unable to swim
for the full 270 min at speeds of 12 and 15 cm·s–1, thus sug-
gesting that they may be swimming within the prolonged
phase. However, for fish swimming at speeds ≥18 cm·s–1,
performance was significantly reduced because all fish except
one (which swam for 193 min) were unable to maintain
swimming for longer than 35 min regardless of water veloc-
ity (Fig. 1). These results suggest that the slope of the rela-
tionship between swim speed and endurance changes at
18 cm·s–1 and implies that this change in performance identi-
fies the transition from sustained to prolonged swimming in
A. brevirostrum (Fig. 1).
The second objective of this study was to relate fish per-
formance determined using the critical swimming test with
those achieved using the endurance test. As defined, Ucrit is
often considered to be closely related to the maximum sus-
tainable swimming speed for similar-sized fish of the same
species (Hammer 1995). In practice, Ucrit is likely only an es-
timator of aerobic swimming capacity in fish (i.e., where the
highest oxygen consumption rates are achieved), because as
fish are approaching Ucrit, they are using bouts of rapid (pre-

Published by NRC Research Press

 350
sumably anaerobic) swimming (Peake 2008). Thus, critical
swimming speeds likely overestimate the maximum sustain-
able swimming speed found using endurance swimming tests
(Webb 1971; Johnston and Moon 1980; Peake et al. 1995).
Consequently, Ucrit is often found to occur at speeds within
the prolonged swimming phase rather than at the inflection
point (transition point; 18 cm·s–1 in this study) between the
sustained and prolonged phases of swimming. For A. fulves-
cens, Peake et al. (1995) showed that the maximum sustain-
able swimming speed (endurance protocol) was equal to
15.5% of Ucrit for 15 cm fish. This ratio is low when com-
Can. J. Zool. Downloaded from www.nrcresearchpress.com by Guangzhou Jinan University on 06/05/13
pared with that for the S. albus, which showed a maximum
sustainable swimming speed equal to 77.6% of Ucrit (Adams
et al. 1999, 2003). For A. brevirostrum, maximum sustain-
able swimming speed is equal to 80.7% Ucrit. This result in-
dicates that the fish are potentially swimming using anaerobic
metabolism for the remaining ~20% of speeds in the Ucrit
tests. This information might be useful when investigating
the effects of biotic or abiotic factors solely on aerobic proc-
esses using Ucrit methodology. The transition between sus-
tained and prolonged swimming should be confirmed using
physiological measurements (e.g., respirometry and haema-
tology); however, this may be difficult to illustrate because it
has been shown that juvenile A. brevirostrum have a low fac-
torial scope and do not produce large levels of muscle
(~6 mmol/L) and blood (~2 mmol/L) lactate during manual
For personal use only.
chasing (Kieffer et al. 2001). Thus, our results may serve as
a better predictor to estimate the onset of anaerobic metabo-
lism in juvenile sturgeon.
While swimming at different speeds, sturgeon have been
shown to modify their swimming behaviour by using station-
holding, substrate skimming, and burst-and-glide behaviours
(Peake 2004; Hoover et al. 2005; Kieffer et al. 2009). This
behaviour was also observed in the present study. However,
because the bottom of the flume was made of plastic (as op-
posed to more adherent substrates such as sand), station-
holding appeared more difficult as water velocity increased.
At these velocities, sturgeons would hold ground once reach-
ing the upstream section of the flume, slide downstream
without swimming actively, and then swim upstream again
once their caudal fin would touch the downstream screen.
For the present study, swimming behaviour under the two
swimming conditions was assessed by determining TBF at
the various swimming speeds. TBF has been shown to in-
crease as a function of swimming speed in many sturgeon
species (Webb 1986; McKenzie et al. 2001; Parsons et al.
2003; Allen et al. 2006), and it has been used to estimate
swimming speed and oxygen consumption (Geist et al.
2005). Two important differences between protocols, how-
ever, could influence the TBF values: (1) under the Ucrit pro-
tocol, the same fish were filmed at all velocities, whereas in
the endurance protocol, different fish were swum and video-
taped at a single velocity; (2) acceleration rates differed be-
tween the two protocols. Within the Ucrit swimming test, the
water speed increased constantly at a rate of 0.42 BL·s–1
every 20 min. During the endurance swimming test, the ac-
celeration rates ranged from 2.55 to 5.94 BL·s–1. Since fish
in the Ucrit swimming test fatigued at slower speeds presum-
ably because of accumulated fatigue, only speeds leading up
to Ucrit were available for direct comparison across both
swimming tests. TBF under the Ucrit swimming protocol was
Can. J. Zool. Vol. 90, 2012
highly variable among individuals within a single speed
when compared with fish that swum under the endurance
protocol at the same speeds (Table 1). This variability might
reflect differences in swimming ability among individuals of
a same population. The use of station-holding during swim-
ming has been noted for several sturgeon species (Adams et
al. 2003; Hoover et al. 2005) and is believed to be a means
to conserve energy during swimming (Kieffer et al. 2009).
Low acceleration rates within Ucrit protocols might enable
weaker performers to use station-holding behaviour, particu-
larly if they are not strong swimmers at faster speeds, and
thus improve their overall swimming performance. This be-
havioural compensation is not observed to the same degree
in the endurance protocol, as is confirmed by lower TBF var-
iability (Table 1) and higher (although not significantly)
mean TBF at speeds ranging from 6 to 18 cm·s–1. Because
the initial acceleration rates are much higher in the endurance
test, station-holding behaviour may not be performed to the
same extent as it is for the Ucrit swimming test. Interestingly,
Farrell (2008) demonstrated that higher acceleration rates in-
creased swimming performance for the non-station-holding
adult rainbow trout. Beamish (1978) suggested initiating
swimming trials at higher speeds to specifically avoid sta-
tion-holding behaviour in salmonids, centrarchids, and scor-
paenids. Thus, station-holding behaviour might not be
possible for fish swimming during endurance protocols, cre-
ating larger swimming performance variability between indi-
viduals possessing different swimming abilities. In contrast,
high TBF variability in Ucrit methodology seems to reflect
more consistent swimming performance values.
Because individual fish were followed through time for the
Ucrit swimming test, it was possible to test whether individu-
als that exhibited lower TBF had higher Ucrit values. Fish
with the lowest TBF values had the highest Ucrit values
(Fig. 3). These lower TBF values can be interpreted as a re-
sult of increased station-holding behaviour. Thus, the Ucrit for
station-holding individuals differs from active swimmers.
This station-holding strategy might provide to be a selective
advantage for fish in the wild, because A. brevirostrum live
in large tidal rivers, and might be able to reduce their overall
metabolic costs in environments of inconsistent water veloc-
ities (Kieffer et al. 2009).
Acknowledgements
We thank C. Ceapa for providing the fish and making ex-
perimental space available, J. Chase for technical assistance,
and J. Houlahan and K. Munkittrick for constructive com-
ments on earlier versions of the manuscript. Funding for this
research was provided by a Natural Sciences and Engineering
Research Council of Canada (NSERC) discovery grant to
J.D.K. Support was also provided by the MADSAM fish
group.
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