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Effects of Pulsed and Continuous DC Electrofishing on Juvenile Rainbow Trout

Article  in  North American Journal of Fisheries Management · November 1998

DOI: 10.1577/1548-8675(1998)018<0905:EOPACD>2.0.CO;2

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North American Journal of Fisheries Management 18:905–918, 1998
q Copyright by the American Fisheries Society 1998

Effects of Pulsed and Continuous DC Electrofishing on

Juvenile Rainbow Trout
BARBARA J. AINSLIE, JOHN R. POST,* AND ANDREW J. PAUL
Division of Ecology, Department of Biological Sciences
University of Calgary, Calgary, Alberta, T2N 1N4, Canada

Abstract.—Three hundred fifty juvenile rainbow trout Oncorhynchus mykiss, reared under captive
conditions, were exposed to 300-V continuous DC or low-frequency (30-Hz) pulsed DC (PDC)
for one or three electroshocking passes to evaluate the effects of electroshock on mortality, injury,
and growth over 147 d. Mortality was negligible (;1%) in all treatments. Injury rates varied from
15% to 39%, with PDC causing a greater number (but typically less severe) injuries than analogous
DC sampling. Multiple-pass sampling designs caused more spinal injuries than single-pass designs.
Longer (heavier) fish sustained more spinal injuries. Electroshocking reduced mean growth rates,
but there were no statistically significant differences in growth between treatment groups. Growth
in length was significantly reduced with increasing severity of injuries. Thus, it appears that growth
was not directly impaired by electroshocking but rather by the occurrence of spinal injury, the
severity of which was directly proportional to the magnitude of the growth depression. Extrapo-
lation of the experimental data to field studies in which 20% or less of the population is sampled
suggested reductions of 3% or less in mean population growth with DC or low-frequency PDC
electroshocking.

Electroshocking is commonly used by fisheries
biologists to sample freshwater fish populations,
particularly in lotic systems, where many other
techniques are not viable (Bohlin et al. 1989).
However, studies have shown that high proportions
of sampled fish sustain skeletal injuries (Hauck
1949; Sharber and Carothers 1988; Fredenberg
1992; McMichael 1993; Hollender and Carline
1994; Sharber et al. 1994; Dalbey et al. 1996;
Thompson et al. 1997a) and external brands (Hor-
ak and Klein 1967; Fredenberg 1992) as a result
of electrofishing. Injuries from electrical shock in
fish can also occur with no visible external signs
(Reynolds 1996). These injuries include muscle,
nerve, and tissue damage (Hauck 1949; Mc-
Michael 1993; Hollender and Carline 1994;
Thompson et al. 1997a), physiological and behav-
ioral disturbances (Hauck 1949; Schreck et al.
1976; Mesa and Schreck 1989; Mitton and
McDonald 1994a), reduced swim stamina (Horak
and Klein 1967; Mitton and McDonald 1994b),
and reduced fertility (Marriott 1973; Muth and
Ruppert 1996).
The incidence and severity of electrofishing
damage is partly related to the type of equipment
chosen and the waveform produced (Sharber and
Carothers 1988; McMichael 1993; Sharber et al.
1994; Dalbey et al. 1996; Dwyer and White 1997).
Continuous DC or low-frequency (#30-Hz) pulsed
DC (PDC) have been recommended for electro-
* Corresponding author: jrpost@ucalgary.ca
905
fishing (Fredenberg 1992; Snyder 1992, 1995;
Sharber et al. 1994; Dalbey et al. 1996) because
lower spinal injury rates, particularly in salmonids,
occur with these waveforms (Fredenberg 1992;
Taube 1992; McMichael 1993; Sharber et al. 1994;
Dalbey et al. 1996). However, the potential cu-
mulative effects of commonly used multiple-pass
depletion or mark–recapture population estimation
techniques on the sampled individuals are not well
understood. Such electrofishing techniques alter
short-term behavior (Mesa and Schreck 1989),
cause spinal injuries (McMichael 1993), reduce
growth (Gatz et al. 1986; Dwyer and White 1997;
Thompson et al. 1997b), decrease body condition
(Thompson et al. 1997b), and increase short-term
mortality (Pratt 1955) among individual salmonids
compared with control fish.
Large (.300 mm) adult rainbow trout Onco-
rhynchus mykiss sustain high injury rates from di-
rect current electrofishing (Sharber and Carothers
1988; Sharber et al. 1994; Thompson et al. 1997a);
however, the effects on smaller juvenile fish are
less well known. Smaller fish intercept a smaller
head-to-tail potential than larger fish (Sharber and
Carothers 1988) and may therefore be subject to
lower injury rates (e.g., Hollender and Carline
1994; Dalbey et al. 1996; Thompson et al. 1997a;
but see Jesien and Hocutt 1990). The long-term
effects of electroshocking damage on juvenile fish
growth also need to be quantified. Taube (1992)
and Dalbey et al. (1996) found that large injured
rainbow trout adults experienced impaired long-
906 AINSLIE ET AL.
term (182 d and 335 d, respectively) growth rates
after electroshocking, though long-term survival
was not affected in either study.
We compared the effects of single-pass and tri-
ple-pass sampling designs and low-frequency (30-
Hz) PDC and DC waveforms on different groups
of juvenile rainbow trout electroshocked in a
hatchery raceway. Our study addressed the effects
of electroshock on (1) acute mortality, (2) spinal
injury and, (3) long-term growth. We also esti-
mated population-level growth effects of electro-
shock by extrapolating from observed individual
effects.
Methods
Three hundred and fifty rainbow trout (Mount
Lassen Livingston strain; mean fork length, FL 6
SD 5 13.34 6 1.56 cm, mean mass 5 27.06 6
9.84 g) were randomly assigned to one of five
treatment groups: (1) single-pass continuous direct
current (DC*1), (2) single-pass pulsed direct cur-
rent (PDC*1), (3) triple-pass DC (DC*3), (4) triple-
pass PDC (PDC*3) and, (5) control (handled only).
Each experimental treatment was applied with a
Smith-Root model 15-C Programmable Output
Waveform backpack electrofishing unit with a gas-
oline-powered Honda EX350 generator. The gen-
erator outputs alternating current (AC) at a fre-
quency of 60 Hz to the electrofishing unit; the
electrofishing unit was set at 300 V to output either
PDC (frequency of 30 Hz and pulse width of 4
ms) or DC. The anode was a 0.28-m-diameter alu-
minum loop (10-mm tubing, outside diameter),
and the cathode was a 3.02-m cable (1.78 m was
bared and 1.24 m insulated; the bared cable had a
diameter of 5 mm). Electrofishing was conducted
on September 15, 1995, in a barricaded concrete
raceway. The barricaded area was 6.22 m long,
2.45 m wide, and had a water depth of 0.18 m.
Specific conductivity was 285 mS/cm, and water
temperature was 10.58C.
Field intensity and output waveform of the elec-
trofishing unit were measured on March 12, 1997,
in the same raceway under similar conditions of
water depth (0.18 m), water temperature (128C),
and specific conductivity (284 mS/cm). Voltage
gradient was measured with a voltage gradient
probe consisting of two insulated wires separated
by a distance of 10 mm and with approximately 2
mm of bared wire exposed at the tips (Kolz 1993).
The voltage gradient probe was attached, at sep-
arate times, to both a digital voltage meter and a
cathode-ray oscilloscope (CRO). The CRO was
used to confirm voltage gradient readings for the
pulsed waveform (ordinary multimeters are not de-
signed to measure peak values for PDC; Jesien and
Hocutt 1990). We measured voltage gradient start-
ing from the anode ring and moving at 0.10-m
intervals directly in front of the anode. The cath-
ode was behind the anode and was directly op-
posite the voltage gradient meter. Maximum volt-
age gradient, which occurs at very close proximity
to the anode, could not be interpreted with our
methods (Kolz 1993). The CRO was used to de-
scribe the output waveform for both PDC and DC.
This was done with the voltage gradient probe po-
sitioned 1 m directly in front of the anode ring,
with the cathode situated behind the anode, as in
the electrofishing design. The two waveforms pro-
duced by the electrofishing unit were traced from
the CRO onto graph paper by hand.
At the start of each electrofishing pass, all fish
were released into the raceway and were allowed
to acclimate for 15 min. During each pass, one
person operated the electrofishing unit, and anoth-
er dipnetted shocked fish. Fish were removed after
they showed narcosis in response to electroshock-
ing. Limits on the rate at which fish could be dip-
netted and removed may have caused some fish to
remain in the electric field long enough to expe-
rience tetany. An electroshocking pass was con-
sidered complete only after all 70 fish in a treat-
ment group were transferred from the raceway to
a recovery vessel. All electroshocking was com-
pleted within 1 d. The triple-pass treatment groups
were allowed approximately 3 h to recover in sep-
arate holding pens between sequential shocking
passes.
After the application of the experimental treat-
ments, individual fish were netted, anaesthetized
with tricaine methanesulfonate (MS-222, 0.03 g/
L), weighed to the nearest 0.1 g, measured for fork
length to the nearest millimeter, and individually
marked with a passive integrated transponder
(PIT) tag (Avid Manufacturing, Norco, Califor-
nia). All visually apparent brand marks were also
documented. After processing, each treatment
group was transferred into a separate mesh holding
container (1.81 3 1.02 3 1.00 m) within a cement
raceway and held for 72 h to assess short-term
mortality.
Following the 72-h holding period, equal num-
bers of rainbow trout from each of the five treat-
ment groups were randomly assigned to each of
six tanks where they were held for 147 d. Tanks
were assigned a number of fish approximately pro-
portional to their volume. There were four rect-
angular tanks (one tank, 3.18 3 1.52 3 0.25 m
 ELECTROFISHING JUVENILE RAINBOW TROUT
with 90 fish; one tank, 1.37 3 1.52 3 0.43 m with
64 fish; and two tanks, 2.08 3 0.55 3 0.38 m with
39 fish each) and two circular tanks (1.22-m di-
ameter, 0.50 m deep with 59 fish each).
Fish were fed once daily with a type and quantity
of food designed to optimize growth as a function
of the previous week’s water temperature and the
average size of the fish in each tank. Feeding
schedules were updated weekly. A 10-h light : 14-h
dark cycle was maintained throughout the exper-
iment. Tanks were checked daily for dead fish.
The fork length (nearest mm) and mass (nearest
0.1 g) of all individuals, as identified by their PIT
tag, were recorded 14 times throughout the ex-
periment. Individuals were lightly anaesthetized
(MS-222, 0.03 g/L) before handling. This sam-
pling was done at weekly intervals for the first 7
weeks of the experiment, at biweekly intervals for
the next 8 weeks, and at triweekly intervals for
the last 6 weeks. At the conclusion of the exper-
iment, all fish were killed with MS-222 and frozen
for X-ray analysis.
X-ray assessment.—All fish were X-rayed to de-
termine the number and location of spinal defor-
mities. Fish were partially thawed, then X-rayed
(right lateral view) at a focal distance of 91.4 cm
with a Philips Medical System X-ray machine
(1.0–4.0 mA, 2.5–10.0 ms, 40-kV potential). The
X rays were taken with 35.6 3 43.2-cm Fuji
SHRS-30 X-ray plates. There were approximately
six fish radiographed per X-ray plate.
The X-ray photographs were analyzed by a sin-
gle reviewer, and the locations of all vertebral ab-
normalities were identified by number. The first
well-defined vertebra behind the skull was des-
ignated as vertebra 1. The treatment groups to
which the fish belonged were unknown to the re-
viewer during this analysis. X rays were of good
quality and all were readable because they were
taken from fish that had grown through the 147-d
experiment (mean FL 6 SE 5 22.47 6 1.15 cm).
It was not possible to rate injury severity by tra-
ditional methods (e.g., Hollender and Carline
1994: Table 2; Dalbey et al. 1996: Table 1); rather,
injury severity was quantified by the number of
malformed vertebrae per fish.
We compared frequency distributions of the
number of malformed vertebrae per injured fish
per treatment between pairs of treatment groups
with the Kolmogorov–Smirnov two-sample test, as
modified for large sample sizes (Sokal and Rohlf
1995). Treatment groups were each compared with
the control group, waveforms were compared be-
tween single-pass and multiple-pass treatments,
907
and single and multiple passes were compared be-
tween DC and PDC treatments. Functional rela-
tionships between the number of malformed ver-
tebrae and the initial length and mass of treatment
fish were assessed with model-1 linear regressions
(Sokal and Rohlf 1995). The hypothesis that the
pattern in position of injured vertebrae was not
significantly different from uniform was assessed
with the G-test for goodness of fit of single-clas-
sification data (Sokal and Rohlf 1995). The hy-
pothesis that the pattern in position of injured ver-
tebrae did not differ among treatments was eval-
uated with the Kolmogorov–Smirnov two-sample
test, as modified for large sample sizes (Sokal and
Rohlf 1995). Numbers of vertebrae damaged per
fish were compared between branded and non-
branded injured fish with a two-sample t-test (So-
kal and Rohlf 1995).
Growth analysis.—Standardized growth in
length and mass was calculated for each treatment–
tank combination for the 13 sampling intervals.
Growth for each treatment–tank over a time in-
terval was calculated as the mean from individual
fish within that treatment–tank. Standardized
growth for each treatment–tank was then calcu-
lated by subtracting the mean growth of control
fish from the mean growth of treatment fish. The
resulting data set is a time series of standardized
growth for six replicates of each of four treatment
groups. A negative standardized growth indicated
the electroshocked fish grew at a slower rate than
the control fish. This method of assessing growth
avoids the potential influence of tank effects by
scaling observed growth rates of treated fish to
untreated fish raised in the same tank under iden-
tical conditions. These data were analyzed with
repeated-measures analysis of variance (ANOVA)
to test the hypotheses that waveform, frequency of
shocking, and their interactions affected standard-
ized growth rate.
The hypothesis that growth rate was modified
by the severity of injury, rather than simply by
electrofishing, was tested by regressing the number
of vertebrae damaged for an individual fish in the
experiment against its standardized growth over
the 147 d. Growth was assessed by using length
and mass. These measures were standardized by
subtracting the mean growth of control fish from
the mean growth of treatment fish in the same tank.
We used t-tests to test the regression hypotheses
that (1) severity of spinal injury affects growth, as
evidenced by a nonzero slope and (2) electro-
shocking itself directly affects growth, as evi-
denced by a nonzero intercept.
908 AINSLIE ET AL.
Population-level growth effects.—The data from
the laboratory experiment were used to model the
effects of six commonly used field electroshocking
protocols on population growth rates. To accom-
plish this, a weighted-average model, similar to
that used by Schill and Beland (1995) was pro-
posed. The factors considered in the model were
(1) waveform (DC versus 30-Hz PDC), and (2)
potential number of exposures to electrofishing in
each of three different abundance estimates (sin-
gle-pass removal, triple-pass depletion removal,
and capture–mark–recapture with three passes).
For each waveform, we estimated the standard-
ized growth of fish as the mean daily growth rate
of fish electroshocked a particular number of times
divided by the mean daily growth rate of control
fish. Mean daily growth rates, as a function of
electroshocking protocol, were determined from
the total growth in mass over the 147-d period,
averaged over the six tank replicates. Growth rates
for two electrofishing passes were estimated by
taking the arithmetic average of the single-pass
and triple-pass growth rates for a particular wave-
form. The change in growth within the sampled
stream section was modeled as the percentage
change of observed standardized growth from the
ideal case of standardized growth, in which elec-
troshocked and control fish grew at the same rate.
For multiple-pass removal treatments, we assumed
that a proportion of the population equal to the
capture efficiency of the pass was removed after
each pass.
In our population extrapolation, we assumed
that (1) when using a capture–mark–recapture pro-
tocol, the captured fish are returned to the sampling
area after each electrofishing pass, whereas fish
captured under a depletion–removal protocol will
only be returned to the sampling area once all elec-
trofishing is completed, (2) capture efficiency is
identical for both waveform types, (3) all fish in
an area are exposed to the electric current during
an electroshocking pass, (4) all fish exposed to a
particular number of electroshocking passes will
experience an alteration in their growth pattern
consistent with that number of passes, (5) growth
rates experienced by domestic rainbow trout in a
hatchery are indicative of the growth of wild trout
in nature, (6) the probability of capture decreases
during sequential electroshocking passes (Schnute
1983; Riley and Fausch 1992) from an estimated
50% on the first pass to 30% on the second pass
and 10% on the third and, (7) there is no immi-
gration into or emigration out of the electrofished
area of stream during the sampling procedure.
When a stream population assessment is done,
typically only a small proportion of the total
stream will be electroshocked (e.g., Schill and Be-
land 1995); thus, our model results for the sampled
section were extrapolated to the population of the
entire stream. When extrapolating, we assumed
that if x% of the whole stream population was
electrofished, then it would experience a decrease
in growth equal to x% of the growth alteration for
the sampled section. We modeled x 5 5% and x
5 20%.
Results
Electroshocker Output
Voltage gradients for both DC and PDC were
similar and ranged from 1 V/cm near the anode to
less than 0.1 V/cm at distances greater than 0.5 m
in front of the anode (Figure 1). The CRO trace
for the PDC waveform showed that the frequency
of pulses was 30 Hz (i.e., wavelength 5 33 ms)
with a pulse width of approximately 4 ms (Figure
1). However, the pulsed waveform was not square
but showed a leading spike of about 1 ms duration
(Figure 1). The DC waveform showed a pattern of
rippled DC with an embedded pulse having a fre-
quency of approximately 120 Hz and pulse width
of 3–4 ms (Figure 1).
Mortality
Mortality was negligible in all treatments over
the 147 d of the experiment. No fish died in the
PDC*1, DC*1, and control groups, and only one
fish died in each of the DC*3 and PDC*3 treatment
groups. Therefore, the mortality rate was only
about 1% in each of those groups.
Spinal Injury
The injury rate was greater for triple-pass than
for single-pass electroshocking and for PDC than
for DC designs (Figure 2). Of the electrofishing
treatments, PDC*3 had the highest injury rate
(39%), and DC*1 had the lowest (15%).
Electrofishing had a significant affect on the fre-
quency of injury intensity in three out of four treat-
ment groups. Frequency distributions of vertebral
malformities differed significantly between un-
treated and all electrofished groups, except the
DC*1 group (Figure 3; Table 1). Sampling design
significantly affected the frequency of injury in-
tensity for the DC treatments only. Waveform did
not significantly affect the frequency of injury in-
tensity for either sampling design. The injuries
caused by DC tended to be more severe than the
analogous PDC treatments as the mean numbers
 ELECTROFISHING JUVENILE RAINBOW TROUT 909

FIGURE 1.—Voltage gradient measured with a cathode-ray oscilloscope or digital multimeter at given distances in
front of the anode for either DC or pulsed DC (PDC). Insets are cathode-ray oscilloscope tracings of the pulsed and
continuous waveforms measured with a voltage gradient probe at 1 m in front of the anode (power was initiated at time
5 5 ms).

of vertebral malformities per malformed fish
(6SE) were greater: PDC*1 5 9.0 6 0.8, DC*1
5 10.6 6 2.8, PDC*3 5 11.2 6 1.0, DC*3 5 11.9
6 1.1.
The number of malformed vertebrae was posi-
tively related to the initial fork length of the fish
in the PDC treatment groups but not related to
length in the DC treatments (Figure 4). The num-
ber of vertebrae damaged was also related to the
initial mass of the treatment fish in the PDC*1 (r 2
5 0.35, N 5 23, P 5 0.003), PDC*3 (r 2 5 0.16,
N 5 26, P 5 0.045) and DC*3 (r 2 5 0.19, N 5
26, P 5 0.028) treatments but unrelated to initial
mass in the DC*1 (r 2 5 0.10, N 5 10, P 5 0.373)
treatment.
Vertebral abnormalities were detected over most
of the spine and were indistinguishable from nat-
ural abnormalities. The majority (;60%) of all
abnormalities were anterior to the dorsal fin (Fig-
ure 5). Vertebra 21 was injured most often, with
52% of all injured electroshocked fish sustaining
an injury in this position. Few injuries were de-
tected in either the far anterior or posterior regions
of the spine.
Electroshocking had a selective effect on the
position of spinal damage in three out of four treat-
ment groups, with certain vertebrae sustaining
higher injury rates than others. Frequency distri-
butions of injury position for the control and DC*1
groups did not significantly differ from an even
910 AINSLIE ET AL.

FIGURE 2.—Proportion of rainbow trout with at least

one damaged vertebra in each of five electrofishing treat-
ment groups. Sample sizes were as follows: control 5 69;
DC, one pass (DC*1) 5 69; pulsed DC, one pass (PDC*1)
5 71; DC, three passes (DC*3) 5 71; and pulsed DC,
three passes (PDC*3) 5 67.

distribution (Gcontrol 5 63.94, GDC*1 5 77.93; P .

0.05), whereas the other electrofished groups had
uneven distributions of injuries (GPDC*1 5 283.99,
GPDC*3 5 209.92, GDC*35 443.59, P # 0.05; Figure
5). Frequency distributions of injury position for
DC*3 and PDC*1 treatments did not differ sig-
FIGURE 3.—Frequencies of spinal injuries to rainbow
nificantly from one another(D 5 0.108, P . 0.05), trout for each of five electrofishing treatment groups: con-
whereas PDC*3 and DC*3 treatments differed sig- trol; DC, one pass (DC*1); pulsed DC, one pass (PDC*1);
nificantly (D 5 0.154, P # 0.05), as did PDC*1 DC, three passes (DC*3); and pulsed DC, three passes
and PDC*3 (D 5 0.155, P # 0.05). (PDC*3). Injury frequency is expressed as the number of
malformed vertebrae per fish.
External Signs of Injury
External signs of injury (brands) were indicators
of internal spinal damage. Of the 85 injured fish,
22 (26%) had visible brands, whereas no uninjured TABLE 1.—Comparisons between frequency distribu-
tions of electroshocking injuries for selected pairs of treat-
fish were branded. Branded fish also had signifi-
ment groups, determined by the Kolmogorov–Smirnov
cantly more vertebrae damaged per fish (13.0 6 two-sample test, as modified for large sample sizes. Sig-
1.2) than nonbranded injured fish (9.9 6 0.6; t 5 nificantly different distributions (D) are indicated by an
2.3487, N 5 22,63, P # 0.05). Brands were found asterisk. Treatment groups are as follows: DC, one pass 5
to be temporary indicators of electrofishing-related DC*1; DC, three passes 5 DC*3; pulsed DC, one pass 5
spinal injury because only one brand (of 22) re- PDC*1; pulsed DC, three passes 5 PDC*3. Parenthetical
mained visible 1 week after the initial electro- values are sample sizes.
shocking. A comparison of approximate brand po- Comparison D P
sitions with the frequency distribution of vertebral
Electroshocking effect
malformities for branded fish indicated that brands Control (69) versus PDC*1 (71) 0.280* 0.009
were usually posterior to the position of spinal Control (69) versus DC*1 (69) 0.101 .0.9
injury (Figure 6). Control (69) versus PDC*3 (67) 0.344* 0.0007
Control (69) versus DC*3 (71) 0.337* 0.0007
Electroshocking Effects on Growth Waveform effect
Standardized rates of growth in length were not PDC*1 (71) versus DC*1 (69) 0.180 0.21
PDC*3 (67) versus DC*3 (71) 0.047 .0.9
significantly affected by electroshocking wave-
form, number of shocking passes, or their inter- Sampling-design effect
action (repeated-measures ANOVA: F 5 0.33, df PDC*1 (71) versus PDC*3 (67) 0.103 .0.9
DC*1 (69) versus DC*3 (71) 0.264* 0.015
5 1, P 5 0.858; F 5 3.749, df 5 1, P 5 0.067;
 ELECTROFISHING JUVENILE RAINBOW TROUT 911

FIGURE 4.—Number of malformed vertebrae versus initial fork length of rainbow trout for each of four electrofishing
treatment groups: DC, one pass (DC*1); pulsed DC, one pass (PDC*1); DC, three passes (DC*3); and pulsed DC, three
passes (PDC*3). Also shown are regression lines with slopes significantly different from zero.

F 5 0.339, df 5 1, P 5 0.567, respectively; Figure
7). Standardized rates of growth in mass were also
not significantly affected by electroshocking
waveform, number of shocking passes, or their in-
teraction (repeated-measures ANOVA: F 5 0.123,
df 5 1, P 5 0.730; F 5 3.668, df 5 1, P 5 0.070;
F 5 0.338, df 5 1, P 5 0.567, respectively; Figure
7). However, mean standardized growth rates in
length and mass of the triple-pass treatments tend-
ed to be lower than those of the single-pass treat-
ments (Figure 7), and sampling design was only
marginally nonsignificant for both length and
mass.
Standardized growth in length was inversely re-
lated to the number of injured vertebrae in exper-
imental fish (Figure 8). Growth in mass was un-
related to the number of vertebrae damaged but
was significantly less for electroshocked fish than
for the control fish.
Population-Level Growth Effects
Electrofishing a section of stream can be ex-
pected to change the natural growth rates of the
juvenile salmonids in that section of stream by
11.54% to 214.29%, depending on the sampling
protocol chosen (Table 2). Obviously, the mag-
nitude of the electrofishing-associated growth al-
teration was lower at the whole-stream population
level. In our model, the largest population level
growth reduction was caused when 20% of the
juvenile rainbow trout population was electro-
shocked (22.86%), and the smallest change was
caused when 5% of the population was sampled
(10.08%). For both DC and 30-Hz PDC wave-
912 AINSLIE ET AL.

FIGURE 5.—Frequencies of injuries to vertebrae of rainbow trout subjected to five electrofishing treatments: control;
DC, one pass (DC*1); pulsed DC, one pass (PDC*1); DC, three passes (DC*3); and pulsed DC, three passes (PDC*3).
The locations of abnormal vertebrae were recorded from X rays of all fish. Vertebrae were numbered sequentially,
beginning with the first clearly defined vertebra behind the skull. Arrows indicate the middle of the dorsal and anal
fins, respectively.

forms, the smallest alterations in natural growth
rate occurred with a single-pass sampling design;
a capture–mark–recapture design with three passes
caused the largest magnitude alterations.
Discussion
Mortality
Factors thought to be important in causing elec-
trofishing deaths, such as respiratory failure, hem-
orrhaging, internal injuries, and trauma (Hauck
1949; Schreck et al. 1976; Lamarque 1990), were
not important in our experiment as we observed
negligible short- and long-term (147-d) mortality.
High short-term survivorship (#20 d; $90%) is
commonly reported in electroshocking studies on
hatchery trout (Pratt 1955; Horak and Klein 1967;
Maxfield et al. 1971; Hudy 1985; McMichael
1993; Mitton and McDonald 1994a) and wild trout
(Dalbey et al. 1996; Habera et al. 1996). Longer-
term studies (7–33 months) have reported some-
what lower survival of hatchery trout in the lab-
oratory (65–93%; Maxfield et al. 1971; Taube
 ELECTROFISHING JUVENILE RAINBOW TROUT
FIGURE 6.—Comparison of the frequency distributions of brand positions (line) and injury positions (bars) for all
branded, electroshocked fish (combined over treatment groups, N 5 22). Arrows indicate the mid-points of the dorsal
and anal fins, respectively.
1992) and of wild trout in a natural pond (54–
60%; Dalbey et al. 1996). Despite these studies,
it is not clear whether natural mortality rates are
compensatory or cumulative with electrofishing-
related mortality rates.
Spinal Injury
The rate of injury we observed (15–39%) was
similar to that reported for experiments that used
similar waveforms and pulse frequencies (Taube
1992; McMichael 1993; Sharber et al. 1994; Dal-
bey et al. 1996). In our study, DC treatments
caused lower injury rates than 30-Hz PDC treat-
ments, which, in turn, were lower than injury rates
caused by higher pulse frequencies (Sharber and
Carothers 1988; Fredenberg 1992; Taube 1992;
McMichael 1993; Sharber et al. 1994; Dalbey et
al. 1996; Thompson et al. 1997a). When other fac-
tors are held constant, injury rates tend to increase
with pulse frequency (Taube 1992; McMichael
1993; Sharber et al. 1994).
Injury rates were greater for triple-pass than for
single-pass electrofishing. In the DC treatments,
22% more fish were injured with triple-pass sam-
pling than with single-pass sampling, which was
9–27% higher than has been reported in single-
pass electrofishing with this waveform (Freden-
berg 1992; Taube 1992; McMichael 1993; Dalbey
et al. 1996). In the PDC treatments, triple-pass
sampling only caused a 6% increase in injury rate
over single-pass sampling.
Injury rates were directly related to initial fork
length for both single-pass and triple-pass PDC
sampling and were directly related to initial mass
for single-pass PDC and triple-pass DC sampling.
Given that we had a small range of initial fish sizes,
our experiment was not designed to adequately test
913
this hypothesis. Other studies have also reported
increases in both injury occurrence and severity
with size (Hollender and Carline 1994; Dalbey et
al. 1996; Thompson et al. 1997a), but no studies
have been conducted to specifically address this
hypothesis.
In our study, most injuries were anterior to the
dorsal fin, which contrasts other studies that found
that most injuries occurred posterior to the dorsal
fin (Sharber and Carothers 1988; Fredenberg 1992;
Hollender and Carline 1994). The reason for this
difference may be due to the smaller size of our
experimental fish or to some unknown effect of
environmental conditions during shocking. How-
ever, our results and those of others (Sharber and
Carothers 1988; Fredenberg 1992) have found few
damaged vertebrae in either the far posterior or far
anterior regions of the spine. This is expected be-
cause spinal injury tends to occur where the white
muscle tissue on both sides of the spinal column
simultaneously excites; the spinal column com-
pensates for this stress by compressing, fracturing
or misaligning (Lamarque 1990). The central re-
gion of the body contains more muscle mass than
the far anterior or posterior spine regions (Lam-
arque 1990) and is therefore more susceptible to
damage.
We may have underestimated the frequency and
severity of spinal injuries because we only used
one X-ray view and may have missed some injuries
(e.g., Thompson et al. 1997a). On the other hand,
spinal injury rates may have been overestimated
as we found spinal malformities in unshocked con-
trol fish that were impossible to distinguish from
electroshock-related injuries. The 4% malformity
rate in the unshocked control group, an estimate
of the natural malformity rate, was similar to mal-
914 AINSLIE ET AL.

FIGURE 7.—Standardized growth of rainbow trout by length (mm/d) and mass (g/d), for four electroshocked treatment
groups over 147 d. The treatment groups were DC, one pass (DC*1); pulsed DC, one pass (PDC*1); DC, three passes
(DC*3); and pulsed DC, three passes (PDC*3).

formity rates reported for other unshocked salmo-
nids (McCrimmon and Bidgood 1965; Gill and
Fisk 1966).
The presence of brands was related to the pres-
ence and severity of spinal damage, but lack of
branding did not indicate a lack of spinal injury.
Even though brands accounted for only 25% of all
injured electroshocked fish, all branded fish sus-
tained injuries and were injured the most severely
overall. Fredenberg (1992) also found that brand
presence was related to more serious spinal inju-
ries. However, brands remained visible for a very
short period (#1 week postelectroshock) and are
thus not reliable indicators of past electroshocking
injury. Brands are thought to be caused by erratic
stimulation of chromatophores when nerve fibers
are damaged by vertebral dislocation (Lamarque
1990; Fredenberg 1992) or by hemorrhaging of
damaged capillaries near the skin (Reynolds
1996).
Growth
Triple-pass sampling did not significantly re-
duce growth rates of electroshocked fish when
compared with single-pass sampling. However,
our inability to detect a difference in growth rates
between triple-pass and single-pass sampling may
have been because treatment groups contained dif-
 ELECTROFISHING JUVENILE RAINBOW TROUT 915

FIGURE 8.—Relationships between the number of vertebral malformities and standardized growth in length and mass.
Growth rates were standardized to the mean growth of control fish in the same tank over the 147-d experiment. The
sample size in all regressions is 277 fish. Fitted regressions and zero growth lines are shown. Intercepts (a) and slopes
(b) are reported for all plots; those significantly different from zero are marked with an asterisk, and those not signifi-
cantly different are marked by ‘‘NS.’’

ferent proportions of injured fish, which grew dif-
ferently from uninjured and control fish. We ex-
pected a priori that electrofishing-related injuries
would decrease growth rates because (1) injured
fish may need to use (potentially substantial) por-
tions of their energy budget to repair damaged tis-
sues and bones (Gatz et al. 1986; Gatz and Adams
1987), (2) recovery from electroshock causes a
short-term decrease in feeding behaviors (Bouck
and Ball 1966; Mesa and Schreck 1989), and (3)
the heavy calcification and fusion of injured ver-
tebrae during healing (Sharber and Carothers
1988; Fredenberg 1992) may cause abnormal
swimming behavior, which could interfere with
foraging effectiveness and predator avoidance.
In our study, fish with a large number of dam-
aged vertebrae gained less length than did less
severely injured fish. Dalbey et al. (1996) also re-
ported that moderate to severely injured fish (i.e.,
fish which sustained spinal misalignment or frac-
ture) had significantly lower growth and condition
after 335 d than did uninjured and slightly injured
fish. Previously injured fish recaptured with elec-
troshock have likely grown more slowly than other
individuals within the general population.
Population-Level Growth Effects
The population-level effect of electrofishing de-
pends on the type of abundance estimate, the num-
ber of sampling passes, and the proportion of the
population sampled. If one-fifth or less of the pop-
ulation is sampled according to the techniques
used in this study, the growth effects would prob-
ably be undetectable in the field. However, these
effects may confound long-term monitoring stud-
ies that use electroshock as a sampling tool be-
cause the sampling method is creating growth al-
terations in individuals that, over time, could ac-
cumulate and affect the population as a whole. The
only long-term (.1 year) studies examining the
effects of electroshocking on salmonid populations
found that salmonid abundance was stable or had
916 AINSLIE ET AL.

TABLE 2.—Mean growth effects expected within an Management Implications

electroshocked stream section and when various propor-
tions of the total juvenile salmonid population are elec- Electrofishing waveform and sampling design
troshocked. Growth is expressed as the percentage change (number of passes) should be taken into account
in mass (g/d) expected with a particular waveform and when designing studies of juvenile stream salmo-
type of abundance estimate. In the sampled section, all fish nids. Our results support those of others (Freden-
are electroshocked; whereas in the two extrapolations to berg 1992; Snyder 1992, 1995; Sharber et al. 1994;
whole populations, only 20% or 5% of the population is
Dalbey et al. 1996) who have recommended the
exposed to electroshock.
use of low-frequency pulsed DC or continuous DC
Sample Sample to reduce injury rates. However, injury rates and
Sampled 20% of 5% of growth alterations increased with the number of
Abundance estimate section population population
passes. Investigators must balance the need for in-
Continuous waveform
creasing the number of passes to improve the pre-
Single pass 1.54% 0.31% 0.08%
Depletion–removal with
cision of population estimates against the impacts
3 passes 25.19% 21.04% 20.26% created by improving those estimates. Growth ef-
Capture–mark–recapture fects may not be detectable in the field when sam-
with 3 passes 214.29% 22.86% 20.71% pling juvenile rainbow trout with the methods used
Low-frequency pulsed waveform in this study and when one-fifth or less of a pop-
Single pass 23.34% 20.67% 20.17% ulation is sampled but could be a factor when re-
Depletion–removal with
3 passes 27.77% 21.55% 20.39%
peated electroshockings are used to monitor long-
Capture–mark–recapture term growth trends. Some fraction of detected
with 3 passes 213.77% 22.75% 20.69% growth alterations will probably have been caused
by the sampling procedure. With these consider-
ations in mind, electrofishing with continuous DC
or low-frequency pulsed DC will remain a valuable
increased after 4 years (Habera et al. 1996) or 8 tool for sampling juvenile salmonids in streams.
years (Kocovsky et al. 1997) of annual electro-
shock sampling. Both studies reported relatively Acknowledgments
low (3–26%) injury rates. These results suggest
Funding for this study was provided through the
that these stream salmonid populations were not
Alberta Fisheries Management Enhancement Pro-
adversely affected by multiple years of electro-
gram Trust Fund. We thank the Sam Livingston
shock sampling, although the effect of electrofish-
Fish Hatchery (Alberta Department of Environ-
ing on growth was not assessed in either study
mental Protection, Natural Resources Service) for
(Habera et al. 1996; Kocovsky et al. 1997).
providing additional funds, fish, experimental fa-
Direct extrapolation of our results to wild salmo-
cilities, and technical expertise. The University of
nids in natural streams must be made cautiously.
Calgary Health Sciences Center provided the X-
Low mortality rates of hatchery rainbow trout after
ray facilities and staff for the X-ray analysis. J.
electroshocking may be due to the favorable living
Doll, D. DePape, H. Finch, T. Hamor, M. Lutz, S.
environment of hatchery fish (Taube 1992;
Shima, and W. Schrenk all contributed valuable
McMichael 1993). Cultured fish do not have to
assistance.
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Received July 7, 1997
Accepted March 13, 1998