Fisheries Research 243 (2021) 106068
Available online 13 July 2021
0165-7836/© 2021 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).
Examining the effects of authentic C&R on the reproductive potential of
, Oscar Nordahl
, Marcus Hall
, Anton Rarysson
, Kristofer Bergstr¨
, Erik Petersson
, Juha Meril¨
, Petter Tibblin
Centre for Ecology and Evolution in Microbial Model Systems (EEMiS), Department of Biology and Environmental Science, Faculty of Health and Life Sciences, Linnaeus
University, SE-39231 Kalmar, Sweden
Swedish University of Agricultural Science, Department of Aquatic Resources, Institute of Freshwater Research, Drottningholm, Sweden
Ecological Genetics Research Unit, Organismal and Evolutionary Biology Research Programme, Faculty of Biological and Environmental Sciences, University of Helsinki,
Research Division of Ecology and Biodiversity, Faculty of Science, University of Hong Kong, Hong Kong Special Administrative Region
Handled by Dr. Steven X. Cadrin
The practice within recreational sheries to release captured sh back to the wild, known as catch-and-release
(C&R), is an increasingly important strategy to protect sh stocks from overexploitation. However, C&R is a
stressor and since animal reproduction is particularly sensitive to stress there is reason to suspect that such a
practice induces sublethal tness consequences. Here, we investigated whether and how C&R shing inuenced
the reproductive potential in an anadromous population of Northern pike (Esox lucius). First, female pike were
exposed to authentic C&R using rod-and-reel shing in a coastal foraging habitat prior to the spawning period.
Next, we observed the migration to the freshwater spawning habitat and compared both the timing of arrival and
maturity stage between C&R-treated and control individuals. Finally, to evaluate effects on the quality and
viability of eggs we stripped captured control and recaptured C&R-treated females, measured egg dry mass to
assess nutrient content, conducted articial fertilisations and incubated eggs in a controlled laboratory experi-
ment. We found no evidence of C&R causing alterations in either arrival time, maturity stage, or the quality and
viability of fertilised eggs. In combination, our results suggest that long-term effects of C&R-induced stress on key
reproductive traits of pike, if any, are minor.
The release of captured individuals within recreational sheries, a
practice known as catch-and-release (hereafter C&R), is becoming
increasingly popular to ensure sustainable use of sh populations and to
preserve the quality of sheries (Arlinghaus et al., 2007; Bartholomew
and Bohnsack, 2005). For C&R to be a sustainable alternative to the
traditional recreational shing (catch-and-kill), it requires sufcient
survival and successful reproduction of released sh. Many studies show
that mortality rates following C&R are generally low (Arlinghaus et al.,
2007; Cooke and Suski, 2005; Ferter et al., 2013). However, the con-
sequences of C&R on reproductive potential have rarely been
A typical C&R event will impose multiple stressors on sh including
hooking-related tissue damage, exercise during landing and exposure to
adverse air temperatures, oxygen deciency and gravity during
handling. These will trigger a full stress-response, similar to exhaustive
exercise, leading to energetic, ionic and hormonal changes (such as
elevated cortisol, lactate and glucose levels) (Arlinghaus et al., 2007).
Research in stress physiology recognise that animal reproduction is
particularly sensitive to stress, an overabundance of cortisol can due to
its anti-developmental, anti-growth and immunosuppressive traits have
negative long-term effects in sh and their progeny (Campbell et al.,
1992; Espmark et al., 2008; Giesing et al., 2011; Schreck et al., 2001).
For the period of gonadal development, stressed sh may have to alter
the allocation of energy between reproduction, maintenance or somatic
growth which can result in delayed ovulation, reduced gamete quality,
lowered progeny survival or, in worst case, complete spawning failure
* Corresponding author at: Centre for Ecology and Evolution in Microbial Model Systems (EEMiS), Department of Biology and Environmental Science, Faculty of
Health and Life Sciences, Linnaeus University, SE-39231 Kalmar, Sweden.
E-mail address: firstname.lastname@example.org (P. Tibblin).
Contents lists available at ScienceDirect
journal homepage: www.elsevier.com/locate/ﬁshres
Received 1 March 2021; Received in revised form 2 July 2021; Accepted 6 July 2021
Fisheries Research 243 (2021) 106068
due to inhibited vitellogenesis or egg atresia (Campbell et al., 1992;
Roff, 1982; Schreck et al., 2001; Sopinka et al., 2016). However,
whether C&R-induced stress produce such severe effects in wild sh is
largely unexplored despite its clear implications for management and
sustainable use of sh stocks.
To date, most studies investigating C&R consequences on the
recruitment of wild sh have focused on parental care in black bass
(Micropterus spp.) rather than direct inuences of C&R on gonadal
development and reproduction. When nest-guarding black bass males is
temporarily removed from their nest by C&R it results in nest aban-
donment, brood predation and thereby reduced reproductive output (e.
g. Suski et al., 2003; Steinhart et al., 2004; Stein and Philipp, 2015).
Remaining studies have, with few exceptions, addressed the effects of
C&R on recruitment by studying movement behaviour in salmonids,
subjected to C&R during the stressful and energetically taxing spawning
migration rather than during the foraging season. These studies have
shown ambiguous results revealing either indirect evidence suggesting
impaired reproductive output due to alterations in movement behaviour
(Thorstad et al., 2007; Richard et al., 2014; Havn et al., 2015; Lennox
et al., 2015, 2016, 2017; Twardek et al., 2019; but see Jensen et al.,
2010; Smukall et al., 2019 for examples of no effects) or changes in
migratory timing (Thorstad et al., 2007; Havn et al., 2015; but see
Smukall et al., 2019 for examples of no effects). While there is a small
number of studies that have estimated C&R effects on reproductive
success more directly, results are disparate suggesting either none (Roth
et al., 2019; Smukall et al., 2019) or size dependent negative effects on
reproductive outcome with larger individuals being more susceptible
than smaller ones (Richard et al., 2013).
Here, we explore C&R effects on key reproductive traits in Northern
pike (Esox lucius, hereafter referred to as pike), an important target
species in recreational C&R sheries in the northern hemisphere as well
as an ecological and socioeconomical key species (Ekl¨
of et al., 2020;
Larsson et al., 2015). In contrast to most salmonids, which at time of
spawning migration have completed their reproductive investment
(King et al., 2003), female pike vitellogenesis and oocyte development
continuous up till spawning (Medford and Mackay, 1978). This has
consequences for the putative effects of C&R on reproduction since
stress-induced impairments are expected to compromise the develop-
ment of oocytes during vitellogenesis by energy deciencies and transfer
of stress hormones (Schreck et al., 2001). Further, we chose to assess
effects of authentic C&R in natural settings, rather than simulating C&R
events in controlled environments. While simulation of C&R is a com-
mon method and has some benets, it might obscure potential conse-
quences relevant for the actual implementation of C&R in management.
To the best of our knowledge, this is the rst non-salmonid study to
consider long-term sublethal effects of authentic C&R in natural settings
on migratory timing and reproductive potential. Our approach was to
subject female pikes to authentic C&R using rod-and-reel shing in the
coastal foraging habitat, prior to the spawning period. After this, we
observed the migratory timing to the dened spawning habitat
(wetland) using passive integrated transponder tags (PIT) and compared
both timing of arrival and maturity stage between C&R-treated and
naïve (control) individuals. Finally, to evaluate effects of C&R on the
quality and viability of eggs we stripped gametes from C&R-treated and
control females recaptured in the spawning habitat using fyke-nets. We
assessed the nutrient content in eggs by measuring dry mass and eval-
uated egg viability by conducting articial fertilisations and subsequent
incubation of eggs in a controlled laboratory experiment.
2. Material and methods
2.1. Study species
Pike is a large-bodied, iteroparous and long-lived species, important
as a top predator in freshwater lakes and brackish waters (Donadi et al.,
of et al., 2020). Anadromous pike in the Baltic Sea migrate to
spawn in freshwater streams and wetlands between February to May,
forming subpopulations geographically separated during early life
stages (Nilsson et al., 2014; Tibblin et al., 2015). Since the 1990s many
Baltic Sea coastal pike populations have declined, essentially due to
recruitment problems, and this has resulted in regulations of recrea-
tional sheries such as shery closure during spawning season, bag-limit
and size window with mandatory C&R (Ljunggren et al., 2010; Nilsson
et al., 2019, 2004). The species serve as an established model organism
in ecology and evolution (Forsman et al., 2015) and its homing behav-
iour support studies of reproductive potential in wild sh by enabling
recapture of ripe individuals prior to spawning (Berggren et al., 2016;
Tibblin et al., 2016a, 2015). Pike is one of the most popular target
species in recreational sheries in Fennoscandia and pike shing during
late winter is particularly popular (Swedish Agency for Marine and
Water Management, 2019). One reason for this is that large numbers of
pike aggregate in shallow waters during winter and thereby become
easily accessible for anglers. Also, this is prior to spawning and the fe-
males are then close to their maximum weight. Pike is considered to be
relatively resilient to C&R shing and previous studies have found low
direct mortality (Arlinghaus et al., 2008, 2009; Baktoft et al., 2013;
Klefoth et al., 2008; Muoneke and Childress, 1994; Stålhammar et al.,
2.2. Study area
We conducted C&R shing in three adjacent bays at the Baltic Sea
shore of ¨
Oland, Sweden (Fig. 1). This area is known to harbour a pop-
ulation of anadromous pike that use the coastal habitat for foraging and,
at time of spawning, migrate through a small stream (~350 m long, <3
m wide and average depth of <50 cm) to reach their spawning habitat:
the wetland Harfj¨
arden (N 56◦49′15.6′′; E 16◦48′32.7′′). We chose this
study area based on low recreational shing pressure such that pike will
be naïve to prior C&R experience, in combination with extensive
background knowledge on population dynamics and migratory behav-
iour from previous studies (Nordahl et al., 2019; Sunde et al., 2018a,
2018b, 2019). In addition, the small stream allows robust detection of
migrating PIT-tagged sh and can be entirely closed off with a fyke-net.
To conrm our assumption of low recreational shing pressure in the
area we performed continuous monitoring (35 visits) of the study area
between October 2019 to February 2020. No pike shing was observed
in the study area during these visits. Throughout the sampling period,
two temperature loggers (HOBO Pendant) were placed in the wetland at
potential spawning grounds to hourly track the water temperature
2.3. Catch-and-release treatment
We C&R a total of 87 female pike (total length 66.7 ±9.0 cm, mean ±
s.d.) between 30 October 2019 to 18 February 2020 in 15 separate
shing efforts. We shed either from boat or the shore, by actively
casting with spinning rod and reel, using articial hard- or soft lures. All
sh were captured in depths less than 1.5 m, thus excluding potential
effects of barotrauma. C&R followed the common procedure regarding
landing (Cooke and Suski, 2005); captured pike were landed swiftly
(average landing time was 42 s) in a rubber net and, if possible, handled
under water whilst unhooked with pliers.
Following unhooking, focal sh was handled above water to mimic
the procedure of documentation (photographs and measurements of
length and weight). The handling time was on average 2.5 min, which is
similar (but in the higher end) to previous C&R studies on pike
(Arlinghaus et al., 2009; Dubois et al., 1994). During this time, we also
recorded length (total length, nearest cm) and inserted a PIT-tag (23.1
mm long and 3.85 mm in diameter, model HDX23, Biomark, Boise,
Idaho, USA). To minimise the risk of damaging reproductive organs,
PIT-tags were inserted under the pelvic girdle rather than in the body
cavity, using a sterile needle mounted on an injector. After the marking
H. Flink et al.
Fisheries Research 243 (2021) 106068
procedure all pike were released back to the water. There were cases of
multiple C&R events for individual sh. Eight pikes were subjected to
C&R twice and one pike was C&R three times. Out of the individuals
C&R-treated twice, three were included in the analyses of migration
timing and maturity stage, the rest were not detected at the wetland
(except for one individual partially detected, see below for explanation).
All other analyses were only based on single C&R individuals.
2.4. Swim-through PIT-tag reader to monitor spawning migratory timing
To evaluate inuences of C&R on migratory timing, we placed a PIT-
tag reader station directly downstream from the outlet pool of the
wetland (Fig. 1). The PIT-tag reader was installed February 14, 2020,
but a proportion (n =20) of tagged and C&R-treated females had
already migrated to the wetland or failed to be detected at arrival as
evidenced by their subsequent downstream migration. The PIT-tag
reader was removed April 24, thirteen days after the last upstream
movement had been detected. To estimate timing of arrival of C&R-
treated individuals, we included only females for which we could
conrm an exact date of upstream migration (n =39). The PIT-tag
reader station consisted of two antennas that enabled determination of
migration direction, however due to partial detections of sh (detected
by only one of the two antennas), we know that an additionally 6 C&R-
treated females migrated to the wetland during the spawning season but
the direction of movement could not be determined, and they were
subsequently not included in any comparisons.
Tagging of individuals has been conducted in the study area since
2017. Here, we take advantage of previously tagged female pike, with no
recent (focal year) experience of C&R, to estimate timing of arrival in
control sh (n =32) during 2020.
2.5. Assessment of maturity stage and stripping of gametes
To capture focal sh for assessment of maturity stage and to strip
gametes for egg quality assessment and the articial fertilisation
experiment (see below), we used a stream-wide fyke net that completely
shut-off the outlet pool of the wetland between February 23 to March 27,
2020 (Fig. 1). Since a high proportion of females caught during up-
stream migration were not yet ovulating, the stream-wide fyke net were
only used periodically. Additional fyke nets (n =3) were placed inside
the wetland, where the proportion of ovulating females were higher.
Fyke nets were emptied daily. Maturity stage, categorised as either pre-
spawn condition (not ovulating) or spawn (ovulating)/spent condition,
was assessed for all females by applying a gentle pressure on the
Fig. 1. Study site. Map of the study area at the Baltic Sea shore of ¨
Oland, Sweden. Map includes the three bays in which pike were caught-and-released, the
freshwater wetland to which pike migrate and spawn as well as the outlet pool of the wetland were a swim-through PIT-tag reader was installed.
H. Flink et al.
Fisheries Research 243 (2021) 106068
In total, 10 ovulating C&R-treated females were recaptured and
stripped for eggs. Simultaneously to each recapture, ovulating C&R-
naïve females (to be used as control) as well as C&R-naïve males were
stripped for gametes to produce crossings by articial fertilisation (de-
tails, see below). There was an incident of shortage of ovulating C&R-
naïve females, resulting in one less stripped control female and male.
Before release back to the wetland, all stripped sh were measured for
body length (total length, nearest cm) and, if unmarked, PIT-tagged to
ensure that no sh were stripped twice. Body length was similar in C&R-
treated (n =10, total length 69.3 ±10.0 cm) and control females (n =9,
total length 66.6 ±9.0 cm) stripped for articial fertilisation (two
sample t-test, t
=0.62, p =0.54). The rst batch of eggs from each
female was discarded in order to avoid contact with water and a
resulting premature opening of the micropyle. Eggs were collected in 10
mL Falcon tubes and milt in 1.25 mL Micro tubes that were put on ice
and transported to the laboratory.
2.6. Articial fertilisation experiment and measurements of egg quality
The fertilisation experiment to evaluate effects of C&R on egg
viability was carried out at the Linnaeus University, Kalmar Sweden
between March 2 to April 2 in a temperature-controlled room, set such
that the water (tap water aerated for 48 h) temperature was ~6 ◦C to
mimic natural conditions during spawning according to previous studies
(Sunde et al., 2019). Fertilisation was conducted no more than three
hours after gamete stripping. To account for potential effects of male
sperm quality on fertilisation success and egg viability, milt from each
individual male was used to fertilise eggs from both a C&R-treated and
the respective control female, resulting in 19 crosses. Each cross was
done in two replicates, which were produced by independent articial
fertilisations to reduce effects of random errors on fertilisation and in-
cubation, resulting in a total of 38 experimental units. The fertilisation
process for all experimental units followed the method established in
Sunde et al. (2018, 2019) by placing ~30 eggs (31 ±2 eggs) from the
focal female in a small porcelain bowl, pipetting an excess of milt (~50
l) from the focal male on top and adding 1 mL of water from the
experimental system to the bowl. The gametes were mixed by a gentle
whirl for 2 min, rinsed with water three times to remove excess milt and
then immediately transferred to an 800 mL plastic cup lled with water.
All cups/experimental units were randomly distributed within the room
and two temperature loggers (Hobo Pendant) were placed in unused
cups lled with water to get hourly measurements of water temperature
during the experiment (Fig. 2b). Photographs of each experimental unit
was taken immediately after fertilisation so that the exact number of
eggs at start could be quantied. A partial water exchange (75 % of
volume) was performed once a day across the experimental system by an
automated water drip system and identication and removal of dead
eggs were done daily in each replicate to avoid potential confounding
effects through effects on the water quality. Dead pike eggs become
opaque and are easy to discern, however, unfertilised eggs are discern-
ible rst after ~3−5 days when incubated in freshwater (Sunde et al.,
2018a). Consequently, it is difcult to visually discriminate between
unfertilised eggs and eggs that die during early development. Viability of
incubated eggs (estimated as still being alive) was thus analysed by
counting viable eggs from a second photograph taken 10–21 days post
fertilisation, including also potential unfertilised eggs. Due to the paired
design (a single male fertilising eggs from both control and C&R-treated
females at same occasion/day) as well as replication within family
crossings, this measurement captures the ability of the egg to be fertil-
ised and subsequently develop for a dened period. Although the in-
cubation time until the count of viable eggs varied between families
there was no variation in incubation time between treatments due to the
Dry mass per egg, a proxy of nutrient content and egg quality
(Berggren et al., 2016; Murry et al., 2008), was estimated by drying 1 mL
of unfertilised eggs from each female in the fertilisation experiment
overnight at 60 ◦C in a heating cabinet. However, two paired samples,
one from each treatment, failed (resulting in 9 samples from
C&R-treated females and 8 samples from control females). To increase
the sample size and robustness of the comparison, eggs from additional
C&R-naïve females (n =9), sampled concurrently, was added resulting
in a total of 17 control samples. Body length was similar in C&R-treated
(n =9, total length 67.7 ±8.9 cm) and control females (n =17, total
length 68.9 ±9.7 cm) used for estimating egg quality (two sample t-test,
=0.34, p =0.74). Before drying, eggs from each female were spread
Fig. 2. a) Migration timing. Arrival of caught-
and-released and control pike females to the
spawning wetland according to PIT-tag de-
tections. The grey background show 25-75 %
quartile of arrivals. b) Water temperature.
Hourly water temperature in the wetland (grey
line) and in the incubation experiment (black
line) during the study period. The hourly water
temperature for each location is calculated as
an average from two temperature loggers.
Dashed vertical line show the median arrival
date of all females to the wetland.
H. Flink et al.
Fisheries Research 243 (2021) 106068
out in a single layer on an aluminium baking tin and photographed for
quantication of number of eggs (1 mL of eggs amounted to 131 ±22
eggs). Following drying, each sample of eggs was weighed to the nearest
0.1 mg (BL 210 S Analytical balance, Sartorius, Goettingen) and divided
by the number of eggs to calculate dry mass/egg.
2.7. Data analysis and statistics
The software R 4.0.3 (R Core Team, 2020) was used for all statistical
analyses. Figures were prepared in R using the packages ggplot2
(Wickham, 2016) and ggmap (Kahle and Wickham, 2013). Photographs
of eggs for measurements of dry egg mass and egg viability were ana-
lysed using ImageJ software (1.52q) (Schneider et al., 2012). An alpha
level of 0.05 was used for all statistical tests.
Timing of arrival to the wetland was examined using median, 25th
and 75th percentiles of calendar date for C&R-treated and control fe-
males. In addition, a Kolmogorov-Smirnov (KS) test was used to evaluate
if timing of arrival, measured as number of days from start of monitoring
until arrival of focal sh, differed between C&R-treated and control
Contingency tables with Fisher´s exact test for count data were used
to determine whether stage of maturity, dened as either pre-spawn or
spawn/spent, differed between C&R-treated and control females. Ana-
lyses were separated for females caught arriving to the wetland and
females caught inside the wetland.
ANCOVA was used to test the possible effect of C&R treatment on
resource allocation in eggs (dry mass per egg) with female body length
treated as a covariate. The nonsignicant interaction term between
treatment (C&R vs control) and body length was removed from the
model. However, the covariate body length was signicant and thus kept
in the model (ANCOVA, effect of total length: F
=5.09, p =0.03).
The inuence of C&R treatment on egg viability was analysed using a
generalised linear mixed model (GLMM) in the lme4 package with a
binomial t and a logit-link function. The response variable was the
number of viable eggs at the end of incubation out of the total number of
eggs at start of incubation. The treatment of females (C&R vs control)
was treated as a xed factor. To account for replication within family
crossings, individual families were included in the model as a random
effect. Parameter estimates and associated statistical signicance levels
of the random effect are not reported since we were not interested in
quantifying these effects as such.
3.1. Migration timing
In total, 60 out of 87 (69 %) C&R-treated females were conrmed by
detection of their PIT-tags to arrive at their spawning habitat in the
wetland. C&R-treated females had their peak arrival, i.e. the time period
when 50 % of the individuals arrived, between February 25 to March 4,
2020 (n =39, median =March 2) (Fig. 2a). In comparison, PIT-tagged
females that had not been subjected to C&R in this study (control fe-
males) had an overlapping peak of distribution at March 1–5 (n =32,
median =March 4). There was no signicant difference between C&R-
treated and control females in time of arrival (two sample KS-test, D =
0.25, p =0.20).
3.2. Maturity stage
The maturity stage of C&R-treated and control females did neither
differ signicantly at arrival to the wetland (C&R-treated: n =10, 80 %
pre-spawn, control: n =66, 72 % pre-spawn, Fisher´s exact test: odds
ratio =0.67, p =1) nor inside the wetland (C&R-treated: n =14, 7% pre-
spawn, control: n =74, 24 % pre-spawn, Fisher´s exact test: odds ratio:
4.13, p =0.29) (Fig. 3).
3.3. Egg quality
Egg quality, measured as dry mass per egg, of C&R-treated (~2.30
mg, condence interval: 2.14–2.47 mg) and control female pikes (~2.28
mg, condence interval: 2.16–2.39 mg) did not differ signicantly
(ANCOVA, effect of C&R: F
=0.09, p =0.76) (Fig. 4a).
3.4. Egg viability
Egg viability was high in both C&R-treated (~93 % viable eggs,
Fig. 3. Maturity stage. Proportion females in pre-spawn and spawn/spent condition, caught either on arrival to the wetland (C&R: n =10, control: n =64) or inside
the wetland (C&R: n =14, control: n =74).
H. Flink et al.
Fisheries Research 243 (2021) 106068
GLMM condence interval: 89–96 %) and control females (~94 % viable
eggs, GLMM condence interval: 90–97 %). Egg viability did not differ
signicantly between the groups (GLMM, effect of C&R treatment: z =
0.38, p =0.71 (Fig. 4b).
Fish reproduction is sensitive to stress, accordingly researchers have
advocated C&R sheries to avoid capturing sh immediately before or
during the spawning period, until there is rm evidence that there is no
C&R-induced negative impacts on recruitment success (Arlinghaus
et al., 2007; Cooke and Suski, 2005). Yet, to evaluate reproductive
consequences of C&R-induced stress in natural settings is challenging,
hence the knowledge and understanding on such effects are inadequate
and, adding to the complexity, the few existing studies show disparate
results. Our study provides unique and important knowledge by testing
such long-term effects of authentic C&R in natural settings in pike
during the period of vitellogenesis, whilst oocyte development is sus-
ceptible to stress. Although our results suggest that C&R did not impair
any of the focal components of reproductive potential in pike, continued
cautiousness is required in the implementation of these ndings in
Arrival timing at spawning sites have clear bearings on the tness of
sh by inuencing the survival of both juvenile and adult life-stages
(Einum and Fleming, 2000; Tamario et al., 2019; Tibblin et al.,
2016b). We anticipated C&R to alter arrival time and maturity stage in
anadromous pike due to the anti-developmental and inhibitory repro-
ductive effects of stress hormones. For example, rainbow trout (Onco-
rhynchus mykiss) exposed to acute stress by air exposure have been
shown to delay ovulation several weeks (Campbell et al., 1992) and
exposure to C&R in rainbow trout leads to a reduction in plasma levels of
oestrogens (Pankhurst and Dedualj, 1994). Further, C&R have shown to
induce downstream movement and delay upstream spawning migration
in Atlantic salmon (Salmo salar). However, this is suggested to result
from short-term effects on physical recovery, loss of orientation or
escape behaviour rather than due to suspended gamete development
(Havn et al., 2015; Thorstad et al., 2007). Our results show an over-
lapping peak arrival of C&R-treated and control females, without any
tendency of C&R delaying spawning migration. Similarly, we found no
evidence of delayed ovulation in C&R-treated females. Upon arrival to
the wetland, 80 % of C&R-treated females were in pre-spawn condition,
similar proportion as in control sh. Among sh caught in the spawning
grounds only 7% of C&R-treated sh were found in pre-spawn condi-
tion, the rest were either ovulating or spent, as expected during a normal
spawning event. Our results are in accordance with pond-based studies
on Australian bass (Macquaria novemaculeata) and golden perch (Mac-
quaria ambigua), demonstrating normal gonadal development following
simulated C&R, with exception for harshly angled sh (Hall et al., 2017,
In oviparous (and viviparous) sh, all nutrients essential for
embryonal development must be incorporated in the oocyte before
ovulation, hence nutrient content of an egg together with the genetic
material determines egg quality (Brooks et al., 1997). Rainbow trout
exposed to acute stress experience reduced egg size and viability
(Campbell et al., 1992). However, consequences of C&R-induced stress
on egg nutrient content has prior to our study never been considered.
Our results demonstrate no difference in egg dry mass, a proxy of
nutrient content in pike eggs (Murry et al., 2008), between C&R-treated
and control females. Egg dry mass from C&R-treated individuals was
around 2.14–2.47 mg per egg (95 % condence interval) compared to
2.16–2.39 mg in control females. The observed quality of eggs coheres to
the results from our laboratory experiment of fertilised egg viability that
showed over 90 % overall egg viability in both C&R-treated and control
females. Previous studies on the effects of C&R on fertilised egg viability
have been conducted in salmonids using simulated C&R which have
generally demonstrated no impacts of C&R (Booth et al., 1995; Pettit,
1977; Smukall et al., 2019). By using authentic C&R in a natural setting
rather than simulated C&R we provide additional and unique support of
that neither quality nor viability of eggs seem to be signicantly reduced
Admittedly, the lack of signicant statistical effects of C&R on the
recruitment potential of pike, in combination with the focal sample
sizes, require cautious inferences and implementation of the results.
Still, that there were no signicant negative effects in any of the
investigated responses to C&R (arrival timing, maturity stage, egg
quality and egg viability) points to that the negative effects of C&R on
Fig. 4. a) Egg quality. Least square means
(adjusted for female body length effect) of dry
mass per egg from C&R and control females.
Black point represents mean and error bar
represents 95 % condence interval. Note that
the scale on the y-axis does not start at zero. b)
Egg viability. Percentage egg viability for C&R
and control females as predicted from the
GLMM model t, black point represents mean
and error bar represents 95 % condence in-
terval. Note that the scale on the y-axis does not
start at zero.
H. Flink et al.
Fisheries Research 243 (2021) 106068
pike reproduction are minor although negative effects mediated through
other pathways such as atresia cannot be excluded. Moreover, in
experimental studies demonstrating reproductive consequences of
stress, focal sh have been exposed to repeated acute stress (Campbell
et al., 1992; Giesing et al., 2011). In our study only a few sh were
captured multiple times and thus we cannot exclude potential additive
effects of multiple C&R which may occur in intense C&R sheries. In the
same vein, it is important to note that sh handling in this study was
conducted by experts and extreme caution was exercised when handling
the sh. It is possible that different handling methods and/or prolonged
retention time would have visible effects on reproductive traits.
A possible explanation to that C&R did not result in discernible
reproductive impairments could be that female pike is capable of pro-
tecting embryos from negative effects of stress by regulating the transfer
of stress hormones to oocytes during vitellogenesis, as demonstrated in
zebrash (Danio rerio) (Faught et al., 2016). Alternatively, stress
induced by C&R might inuence the trade-off between reproductive
output and somatic growth such that C&R-treated females allocated
energy to uphold their reproductive potential but deprioritised
self-maintenance and growth (Schreck et al., 2001). To some extent this
is supported by a whole-lake C&R experiment that showed reduced
growth rates across seven months in C&R-treated pikes (Klefoth et al.,
2011). However, such negative effects of C&R on the growth rates of
pike could potentially also result in lowered fecundity (Berggren et al.,
2016) which emphasis the need of further studies to disentangle the
effects of C&R on the growth-reproduction trade-off.
From a sheries management point of view, our results represent an
important nding suggesting that C&R do not represent a stressor with
substantial long-term negative consequences on the reproductive po-
tential in pike and subsequent impacts on recruitment and population
dynamics. However, such inference of our results rest upon that the
direct mortality of C&R is low, that negative effects of somatic growth do
not translate into effects on reproduction or that survival in later juve-
nile life-stages is impaired, and, nally, that there are no additive effects
of multiple C&R events or unfavourable handling and must thus be
implemented with caution. While pike has shown to be relatively
resilient to C&R, there is considerable variation among species in the
response to C&R shing (Muoneke and Childress, 1994) that stress for
species-specic management (Cooke and Suski, 2005). As an alternative
to traditional catch-and-kill shing, C&R can play an important role as
management tool to aid sustainable sheries, as shown in wild pop-
ulations of Atlantic salmon where C&R-treated sh contributes signi-
cantly to the total reproductive output (Richard et al., 2013; Thorstad
et al., 2003; Whoriskey et al., 2000). As participation in recreational
shing is popular and the practice of C&R increases in many countries
(Arlinghaus et al., 2007; Bartholomew and Bohnsack, 2005), future
research on this topic is warranted. For successful management, there is
a need to better understand sublethal effects of C&R in general, for
example on reproductive traits not covered here such as total repro-
ductive investment, absolute fecundity, spawning competition, hatching
rate and fry performance. Future research should also address the effects
of C&R on the trade-off between reproduction and somatic growth and
try to disentangle how it changes depending on gonadal maturation in
females during C&R exposure.
To conclude, we found no evidence of that C&R of pike during pre-
spawn season cause alterations upon spawning in either arrival time,
maturity stage, egg quality or egg viability. In combination, our results
suggest that C&R-induced stress do not cause signicant long-term ef-
fects on key reproductive traits. The results are of general interest
because we, unlike related studies, exposed females to C&R during
vitellogenesis while there is an additional demand of energy and oocytes
are susceptible to hormonal impairment. In addition, the study reects
authentic C&R on wild sh in their foraging habitat and evaluates the
consequences by eld observations in the spawning ground as well as in
a controlled laboratory experiment. Although our results are promising
in the context of the sustainability of C&R, there is still large gaps of
knowledge in our understanding of the consequences of C&R on
recruitment, and species- and context-dependence thereof, that needs to
be answered for C&R to be unconditionally implemented in sheries
All applicable international, national, and/or institutional guidelines
for the care and use of animals were followed. The laboratory was
approved as research facility (Dnr 5.2.18–17988/18) and the study was
granted ethical approvals (Dnr 168677-2018; Dnr 19359-2019) by the
Ethical Committee on Animal Experiments, Swedish Board of Agricul-
ture, in Link¨
oping and Stockholm respectively.
CRediT authorship contribution statement
Henrik Flink: Conceptualization, Project administration, Method-
ology, Writing - original draft, Writing - review & editing, Formal
analysis, Visualization, Investigation. Oscar Nordahl: Investigation,
Writing - review & editing. Marcus Hall: Investigation, Writing - review
& editing. Anton Rarysson: Investigation, Writing - review & editing.
om: Investigation, Writing - review & editing. Per
Larsson: Investigation, Resources, Writing - review & editing. Erik
Petersson: Conceptualization, Writing - review & editing. Juha Meril¨
Conceptualisation, Writing - review & editing. Petter Tibblin:
Conceptualization, Project administration, Resources, Methodology,
Writing - original draft, Writing - review & editing, Funding acquisition,
Declaration of Competing Interest
The authors declare that they have no known competing nancial
interests or personal relationships that could have appeared to inuence
the work reported in this paper.
We thank Jasper Münnich, Johanna Sunde, Per Koch-Schmidt,
en and Jonas Jakobsson for excellent help in the eld and
the laboratory. We are grateful to the landowners granting us access to
the spawning pike sampling site. We thank two anonymous reviewers
or valuable comments on earlier drafts of the manuscript. This work
was supported by funds kindly provided by the Swedish Research
Council FORMAS(grant 2018-00605 to P.T.) the Crafoord Foundation
(grant 20190636 to P.T.) and G¨
oms Foundation(grant to P.
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