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Animal
Reproduction
Science
132 (2012) 213–
222
Contents
lists
available
at
SciVerse
ScienceDirect
Animal
Reproduction
Science
jou
rn
al
h
om
epa
ge:
www
.elsevier.com/locate/anireprosci
Effects
of
different
photoperiods
and
handling
stress
on
spawning
and
reproductive
performance
of
pikeperch
Sander
lucioperca
Sara
Pourhosein
Sarameha,
Bahram
Falahatkarb,∗,
Ghobad
Azari
Takamic,
Iraj
Efatpanahd
aDepartment
of
Fisheries,
Faculty
of
Natural
Resources,
Islamic
Azad
University,
Lahijan
Branch,
Lahijan,
Guilan,
Iran
bFisheries
Department,
Faculty
of
Natural
Resources,
University
of
Guilan,
Sowmeh
Sara,
Guilan,
Iran
cAquatics
Health
and
Disease
Department,
Faculty
of
Veterinary
Medicine,
University
of
Tehran,
Tehran,
Iran
dDr.
Yousefpour
Fish
Hatchery
Center,
Siahkal,
Guilan,
Iran
a
r
t
i
c
l
e
i
n
f
o
Article
history:
Received
19
November
2011
Received
in
revised
form
14
May
2012
Accepted
16
May
2012
Available online 24 May 2012
Keywords:
Light
regimen
Reproduction
Sex
steroids
Stress
Pikeperch
a
b
s
t
r
a
c
t
The
objective
of
this
study
was
to
control
the
reproductive
cycle
of
pikeperch
(Sander
luciop-
erca)
through
determining
the
effects
of
different
photoperiods
and
handling
stress
on
the
reproduction
quality,
timing
and
quality
of
spawning,
fertilization,
sex
steroids,
and
cor-
tisol
concentrations.
In
this
study,
72
pikeperch
broodstocks
with
an
average
weight
of
1367
±
55.3
g
were
exposed
to
different
photoperiods
including
constant
light
(24L:0D),
constant
darkness
(0L:24D),
and
12
h
of
light,
12
h
of
darkness
(12L:12D)
for
40
days.
Half
of
the
broodstocks
of
each
photoperiod
treatment
were
exposed
to
handling
stress
at
a
specific
time
of
the
day.
Applying
different
photoperiods
caused
changes
in
the
timing
of
broodstocks’
spawning,
so
that
fish
under
24L:0D
spawned
earlier
than
those
of
other
photoperiods,
and
stressed
fish
of
the
0L:24D
photoperiod
had
a
delayed
spawning
com-
pared
to
others.
Also,
the
spawning
of
the
broodstocks
at
different
photoperiods
which
were
exposed
to
handling
stress
was
either
delayed
or
did
not
occur
at
all.
The
highest
and
lowest
spawnings
were
observed
in
the
morning
and
at
night,
respectively.
Fertilization
percentage,
number
of
eggs
per
gram,
sex
steroids
including
estradiol,
progesterone,
and
testosterone,
as
well
as
cortisol
and
calcium
concentrations
did
not
show
any
significant
difference
in
different
photoperiods
and
handling
stress.
In
stressed
males
of
the
24L:0D
photoperiod,
there
only
was
a
significant
decrease
of
testosterone
concentration
compared
to
the
beginning
of
the
experiment.
Results
indicated
that
the
spawning
performance
of
pikeperch
broodstocks
could
be
considerably
stimulated
using
an
effective
photoperiod.
Similarly,
pikeperch
broodstocks
in
culture
systems
are
usually
affected
by
handling
stress,
and
this
stress
could
lead
to
a
poor
reproductive
performance
and
inhibition
of
spawning.
© 2012 Elsevier B.V. All rights reserved.
1.
Introduction
In
some
northern
and
central
European
countries,
pikeperch
Sander
lucioperca
is
mainly
known
as
an
edi-
ble
and
commercial
fish
and,
therefore,
is
considered
∗Corresponding
author
at:
Fisheries
Department,
Faculty
of
Natural
Resources,
University
of
Guilan,
Sowmeh
Sara,
1144,
Guilan,
Iran.
Tel.:
+98
182
322
3599;
fax:
+98
182
322
2102.
E-mail
address:
falahatkar@guilan.ac.ir
(B.
Falahatkar).
important
for
aquaculture
purposes
(Muller-Belecke
and
Zienert,
2008).
Because
the
spawning
season
of
these
fish
is
only
once
a
year
from
April
to
early
May
(Muller-Belecke
and
Zienert,
2008),
out-of-season
spawning
induction
methods
for
this
species
seems
important.
In
fact,
develop-
ing
pikeperch
culture
requires
controlling
their
reproduc-
tive
cycle
so
that
an
out-of-season
spawning
is
achieved.
Therefore,
induction
of
the
reproductive
cycle
should
be
done
irrespective
of
season
(Migaud
et
al.,
2004b).
In
fish
of
temperate
regions,
inducing
this
cycle
is
usually
specified
by
annual
changes
of
photoperiod
and
temperature
0378-4320/$
–
see
front
matter ©
2012 Elsevier B.V. All rights reserved.
http://dx.doi.org/10.1016/j.anireprosci.2012.05.011
Author's personal copy
214 S.
Pourhosein
Sarameh
et
al.
/
Animal
Reproduction
Science
132 (2012) 213–
222
(Bromage
et
al.,
2001).
The
reproductive
cycle
of
pikeperch
also
depends
on
annual
photoperiod
and
temperature
(Migaud
et
al.,
2003,
2004b;
Wang
et
al.,
2010).
Related
researches
have
proven
that
photoperiod
and
daily
changes
are
involved
in
the
environmental
induction
and
quality
of
perch
reproduction
which
includes
the
timing
and
rate
of
spawning,
fertilization
and
the
mortality
of
broodstock
(Fontaine
et
al.,
2006;
Migaud
et
al.,
2006).
As
a
result,
with
photoperiod
as
one
of
the
main
environ-
mental
cues
of
the
timing
of
reproduction
(Bromage
et
al.,
1993;
Craig,
2000)
taken
into
account
which
works
through
the
hypothalamus–pituitary
axis
(Borg,
1994;
Nagahama,
1994),
and
because
illumination
can
affect
the
reproductive
performance
of
perch
(Fontaine
et
al.,
2006),
it
is
probable
that
photoperiod
changes
during
the
final
stages
of
repro-
duction
and
near
the
time
of
spawning
would
have
an
effect
on
broodstocks’
breeding
and
spawning
as
well.
Besides,
handling
stress
is
another
factor
which
has
critical
effects
on
the
reproduction
success.
In
fact,
it
is
assumed
that
in
fish
cultured
under
stressful
condi-
tions,
a
kind
of
exchange
between
the
allocation
of
energy
for
reproduction
and
maintenance
could
happen
(Schreck
et
al.,
2001;
Wang
et
al.,
2006).
Handling
is
among
the
avoidably
stressful
factors
at
breeding
and
culture
centers
that
have
destructive
effects
on
the
physiological
systems
of
fish
including
a
decrease
in
broodstock
fecundity,
inci-
dence
of
abnormal
behaviors
and
a
decrease
in
the
growth.
Studying
responses
of
the
fish
to
handling
stress
with
regard
to
the
prevalence
of
intensive
fish
culture
and
an
increased
probability
of
exposure
to
stress
would
result
in
improving
management
methods
for
fish
breeding
and
ultimately
for
their
health
and
enhanced
growth
(Belanger
et
al.,
2001).
In
fact
changing
the
reproductive
techniques
under
stressful
situations
can
have
important
effect
on
the
improvement
reproductive
fitness
for
fish
in
the
wild.
Clearly,
understanding
of
such
processes
is
helpful
for
man-
agement
of
hatchery
and
broodstocks
(Schreck
et
al.,
2001).
Moreover,
little
is
known
about
factors
which
facilitate
induced
spawning
in
pikeperch
and,
practically,
there
is
no
information
about
the
interaction
between
different
pho-
toperiods
and
handling
stress
and
its
effect
on
spawning.
Consequently,
since
determining
the
induction
of
repro-
duction
and
spawning
might
be
a
multi-factor
process,
conducting
studies
with
the
purpose
of
identifying
differ-
ent
photoperiods
and
handling
stress
as
effective
factors
in
the
reproduction
success,
and
finding
the
best
option
to
induce
spawning
along
with
its
effect
on
the
quality
of
reproduction
are
necessary.
To
study
the
effect
of
light
severity
or
the
direction
of
the
change
of
photoperiod
in
the
timing
of
spawn-
ing,
in
the
final
stages
of
pikeperch
reproduction,
fish
with
the
same
photoperiod
history
were
exposed
to
three
photoperiods:
half
days
of
light
followed
by
an
abrupt
change
to
half
days
of
darkness
(12L:12D)
photoperiod
that
represents
normal
photoperiod,
and
continuous
light
(24L:0D)
and
continuous
darkness
(0L:24D)
to
study
if
increasing
or
decreasing
light
in
the
final
stages
of
repro-
duction
can
alter
spawning
times
by
advancing
or
delaying
spawning
period.
Additionally,
stress
can
have
suppres-
sive
effect
on
reproduction.
Therefore,
the
objective
of
this
study
was
to
enhance
the
understanding
for
environmental
controlling
of
the
pikeperch
reproductive
cycle
along
with
determining
the
effects
of
different
photoperiods
and
handling
stress
on
the
quality
of
reproduction,
the
timing
and
quality
of
spawning,
fertilization,
and
sex
steroids
con-
centrations
during
spawning
season.
Hatcheries
can
use
this
knowledge
to
manipulate
maturation
and
spawning
time
to
produce
all-year-round
supplies
of
eggs
and
fry.
2.
Materials
and
methods
2.1.
Fish
and
facilities
In
autumn,
broodstocks
were
captured
from
a
reservoir
lake
in
Aras
dam,
located
in
northwestern
Iran
between
Iran
and
Nakhjavan
Republic
as
an
important
pikeperch
habitat
at
the
southern
basin
of
the
Caspian
Sea.
Then,
they
were
transferred
to
the
Dr.
Yousefpour
Fish
Hatchery
Center
at
Siahkal,
Guilan,
Iran.
For
3
months,
fish
were
kept
in
win-
tering
earth
ponds
while
being
fed
with
bait
fish
(carp
fries,
etc.).
In
March
2009,
they
were
transferred
to
18
tanks
[4
fish
per
tank,
age
4–5
years
(two
females
and
two
males
as
a
sex
ratio
of
1:1)]
with
1490
l
in
capacity,
with
50
cm
depth
and
an
average
water
flow
of
20
±
0.88
l
min−1at
a
non-recirculation
system.
The
age
of
the
broodstocks
was
determined
by
the
scales
taken
from
the
line
of
demarcation
between
the
lateral
line
and
dorsal
fin.
The
scales
were
examined
under
microscope
with
magnification
of
20×
and
the
age
was
determined
by
counting
the
dark
and
light
circles
(Biswas,
1993).
Broodstocks
(including
36
males
and
36
females)
had
an
average
weight
of
1367
±
55.3
g
and
an
average
length
of
53.7
±
0.6
cm
(mean
±
SE).
At
the
beginning
of
the
exper-
iment,
one
fish
(one
male
or
female;
totally
9
males
and
9
females)
was
randomly
selected
for
blood
sampling
from
each
tank.
Temperature
and
dissoluble
oxygen
were
checked
using
a
thermometer
and
an
oxymeter,
respectively,
three
times
a
day
at
8:30
AM,
14:30
PM
and
20:30
PM.
In
a
40-day
period,
temperature
was
kept
at
13.1
±
0.5 ◦C
and
dissolved
oxygen
was
at
9.7
±
0.4
mg
l−1(mean
±
SE).
2.2.
Experimental
design
In
this
study,
three
different
photoperiods
were
applied
for
40
days.
To
provide
light
during
24L:0D
and
12L:12D
photoperiods,
a
100
W
lamp
(Pars
Shahab,
Guilan,
Iran)
with
630
lx
was
used
40
cm
above
the
water
surface
for
each
tank.
Also,
black
plastic
was
used
for
covering
the
tanks
during
darkness
times
for
0L:24D
and
12L:12D
photoperiods.
In
addition,
due
to
low
temperature
and
pre-
vention
of
eggs
eaten
by
baitfish,
broodstocks
were
not
fed
during
the
experimental
period.
For
each
photoperiod,
there
were
two
groups
of
fish.
In
one
group,
fish
were
not
exposed
to
any
stress
and
were
called
the
fish
with-
out
stress
photoperiods.
In
order
to
evaluate
the
effect
of
sampling
or
capture
on
the
reproductive
performance
of
the
other
group,
the
water
level
in
the
tank
was
reduced
to
10
cm
from
the
bottom
at
9:15
AM
every
day.
Then,
the
fish
were
captured
by
net
and
kept
out
of
water
for
20
s
and
were
transferred
back
to
the
tank
with
its
water
level
increased
back
to
the
first
level.
These
fish
were
called
the
Author's personal copy
S.
Pourhosein
Sarameh
et
al.
/
Animal
Reproduction
Science
132 (2012) 213–
222 215
fish
of
stressful
photoperiods.
Each
photoperiod
consisted
of
three
stressful
and
three
stress
free
replications.
The
broodstocks
of
the
stressful
photoperiod’s
conditions
were
under
the
influence
of
handling
stress
for
40
days.
2.3.
Induction
of
spawning
and
sampling
To
induce
spawning,
10
days
after
applying
different
photoperiods,
inputting
nest
in
tanks
was
done.
In
this
pro-
cess,
nests
(53
cm
×
53
cm
×
5
cm)
were
placed
inside
tanks
such
that
there
were
two
nests
per
tank.
Then,
nests
were
checked
every
2
h
for
checking
the
spawning
with
very
short
and
rapid
removal
of
the
sheet
especially
during
the
darkness.
Broodstocks
which
had
spawned
were
biometri-
cally
recorded
for
weight
(g)
and
length
(mm)
of
fish
with
their
blood
samples
taken.
At
the
end
of
40
days
of
experi-
ment,
the
blood
samples
of
the
remaining
broodstocks
that
did
not
spawn
were
also
taken
as
non
respondent
fish.
2.4.
Spawning
time
and
quality
The
time
of
spawning
was
set
as
follows:
morning
(spawning
which
occurred
during
4–12
AM),
afternoon
(spawning
that
occurred
during
12–20
PM)
and
night
(spawning
which
occurred
between
20
PM
and
4
AM).
In
order
to
compare
the
quality
of
eggs
produced
by
the
broodstocks
at
different
treatments,
the
quality
of
spawn-
ing
was
measured
based
on
the
spawning
rate
which
included
good
spawning
(eggs
covering
100%
surface
of
the
nest)
the
eggs
that
were
round,
transparent,
pale
yellow
to
colorless,
average
spawning
(eggs
covering
50%
surface
of
the
nest),
and
the
eggs
that
were
relatively
transparent,
and
poor
spawning
(eggs
covering
less
than
25%
surface
of
the
nest),
the
eggs
were
relatively
turbid.
2.5.
Reproduction
indices
To
determine
the
fertilization
percentage
and
the
num-
ber
of
eggs
per
gram,
two
batches
of
400–500
eggs
were
randomly
sampled.
Egg
samples
were
weighed
using
a
digi-
tal
scale
with
a
0.01
g
precision.
The
fertilization
percentage
of
the
obtained
eggs
was
specified
in
two
stages:
blastula
(about
5
h
after
fertilization)
and
gastrula
(nearly
48
h
after
fertilization).
Fertilized
eggs
with
a
translucent
appearance
were
counted
and
differentiated
from
unfertilized
eggs
(whitish
appearance).
Eggs
were
observed
daily
to
check
for
embryogenesis
and
signs
of
hatching.
When
hatching
completed,
hatching
rate
was
determined
by
counting
the
larvae
and
remaining
dead
eggs.
2.6.
Blood
sampling,
plasma
assessment
and
analysis
All
parts
of
sampling
and
fish
care
during
the
experi-
ments
were
conducted
in
accordance
with
the
Guide
for
the
Care
and
Use
of
Agricultural
Animals
in
Agricultural
Research
and
blood
sampling
was
done
after
anesthetiz-
ing
the
brooders
with
a
300
mg
l−1dosage
of
clove
powder.
The
first
blood
sampling
of
18
broodstocks
(one
broodstock
per
tank)
was
performed
a
day
prior
to
applying
different
photoperiods
and
handling
stress.
The
second
blood
sam-
pling
was
conducted
immediately
after
spawning
in
which
2
ml
blood
was
taken
from
their
caudal
vein
using
5
ml
hep-
arinized
syringes.
Blood
was
centrifuged
at
1500
×
g
for
10
min,
and
the
plasma
was
kept
in
tubes
at
−25 ◦C
until
the
time
of
analysis.
Testosterone
(T),
17-estradiol
(E2),
and
progesterone
(P)
concentrations
were
determined
after
two
extractions
with
cyclohexane/ethylacetate
(v/v)
using
the
radioim-
munoassay
(RIA)
method
according
to
Fostier
and
Jalabert
(1986).
Samples
were
tested
as
duplicates
while
the
stan-
dards
were
tested
as
triples.
All
samples
were
analyzed
in
a
separate
RIA
for
each
steroid
(Migaud
et
al.,
2004b).
Because
of
the
positive
relation
between
calcium
and
vitellogenin
concentrations,
plasma
calcium
concentra-
tions
were
measured
through
spectrophotometry
(Norberg
et
al.,
2004)
using
the
Arsenazo
III
calcium
commercial
kit
(Sigma,
St.
Louis,
MO,
USA)
in
this
experiment.
Plasma
cortisol
concentrations
were
measured
by
RIA
using
a
commercial
antibody
kit
(cortisol-3-OCMO-
antiserum,
Immunotech,
Prague,
Czech
Republic)
as
described
by
Ruane
et
al.
(2001).
Plasma
glucose
concen-
tration
was
measured
using
analytical
kits
(Wako
Pure,
Chemical
Ind.
Ltd.,
Osaka,
Japan)
(Ruane
et
al.,
2001).
Plasma
lactate
was
enzymatically
determined
using
Sigma
Diagnostic
Kits
(St
Louis,
MO,
USA).
2.7.
Statistical
analyses
All
data
collected
at
the
place
of
experiment
along
with
those
of
the
laboratory
were
recorded
in
Excel
software
and
analyzed
by
the
SPSS
software
(Version
13,
Inc.,
Chicago,
IL,
USA).
Then,
these
were
displayed
as
mean
±
SE.
The
normality
of
the
data
was
measured
through
Kolmogorov–Smirnov
test.
All
percent
data
were
Arc-sin
transformed
before
analysis.
Two-way
analysis
of
variance
(ANOVA)
test
was
used
to
study
the
effect
of
photoperiod
(three
levels)
and
stress
(two
levels)
as
inde-
pendent
variables
on
all
biochemical
parameters,
while
fertilization
percentage
and
the
number
of
eggs
per
gram
were
separately
used
as
dependent
variables.
When
there
was
an
interaction,
Tukey’s
test
as
post
hoc
test
was
used
for
determining
differences
among
means.
Compar-
ison
of
the
two
stresses
condition
(stress
and
no
stress
fish)
in
each
photoperiod
treatment
was
performed
using
the
Independent
Samples
t-test.
For
nonparametric
data,
Mann–Whitney
U
test
was
used
to
find
significant
effect
of
different
photoperiods
or
stress
conditions.
The
signifi-
cance
level
in
this
study
was
considered
as
P
<
0.05.
3.
Results
3.1.
Number,
timing
and
the
quality
of
spawning
Spawning
occurred
in
a
4-week
period
after
nesting
(March
19
to
April
14;
Fig.
1).
Out
of
a
total
number
of
36
females,
29
spawned
and
only
seven
brooders
of
dif-
ferent
photoperiods
did
not
spawn
due
to
handling
stress.
The
ratio
of
spawning
was
not
significant.
The
first
spawn-
ing
was
observed
2
days
after
nesting
in
the
broodstocks
of
24L:0D
photoperiod.
The
first
spawning
of
broodstocks
which
were
exposed
to
handling
stress
during
differ-
ent
photoperiods
was
observed
14
days
after
nesting
and
Author's personal copy
216 S.
Pourhosein
Sarameh
et
al.
/
Animal
Reproduction
Science
132 (2012) 213–
222
0
20
40
60
80
100
stressed unstressed
stressed unstresse
d
stressed unstressed
24L 12L-12D 24D
Days after inputting nest
Spawned fish (%)
1-14Day
15-28Day
Fig.
1.
Percentage
of
pikeperch
Sander
lucioperca
broodstocks
which
spawned
during
the
different
photoperiods
including
constant
light
(24L:0D),
12
h
of
light:12
h
of
darkness
(12L:12D),
constant
darkness
(0L:24D),
constant
light
with
stress
(24L:0D-s),
12
h
of
light:12
h
of
darkness
with
stress
(12L:12D-s),
and
constant
darkness
with
stress
(0L:24D-s).
n
=
6
females
in
each
photoperiod
by
stress
combination.
No
difference
(P
>
0.05)
was
observed
in
terms
of
handling
stress
or
photope-
riod.
during
24L:0D-s
photoperiod.
Totally,
broodstocks
of
the
24L:0D
photoperiod,
stressed
or
unstressed,
spawned
ear-
lier
than
those
of
other
photoperiods.
Also,
the
spawning
of
stressed
broodstocks
compared
to
those
in
the
unstressed
photoperiods
was
either
delayed
or
did
not
occur.
Among
different
stressed
and
unstressed
photoperiods,
24L:0D
and
0L:24D-s
had
the
earliest
and
latest
spawning,
respec-
tively.
All
broodstocks
exposed
to
unstressed
photoperiods
spawned;
however,
in
stressful
photoperiods,
1/3
of
brood-
stocks
exposed
to
24L:0D-s
and
12L:12D-s
photoperiods
and
half
of
those
kept
in
0D:24D-s
did
not
spawn.
Regarding
the
time
of
spawning,
no
significant
dif-
ference
was
observed
among
different
photoperiod
treatments,
while
the
greatest
and
least
rates
of
spawning
were
that
of
the
morning
and
night,
respectively.
More-
over,
broodstocks
of
0L:24D
and
12L:12D
photoperiods
did
not
spawn
at
night
and
broodstocks
exposed
to
0L:24D-s
spawned
only
in
the
morning
(Fig.
2).
No
interaction
was
also
found
between
light
and
stress.
The
quality
of
broodstocks’
spawning
did
not
reveal
any
significant
difference
during
different
photoperiods,
while
in
0L:24D
a
good
quality
spawning
(100%
covered
the
0
20
40
60
80
100
24L
12L-12D
24D
24L
12L-12D 24
D
24L
12L-12D
24D
Morning
Afternoon Night
Spawning time
Brood spawned (%)
unstressed
stressed
Fig.
2.
Comparison
of
the
pikeperch
Sander
lucioperca
broodstocks
spawn-
ing
during
different
times:
morning
(4–12
AM),
afternoon
(12–20
PM)
and
night
(20
PM
to
4
AM)
in
different
photoperiods
including
constant
light
(24L:0D),
12
h
of
light:12
h
of
darkness
(12L:12D),
constant
darkness
(0L:24D),
with
and
without
stress.
No
significant
difference
(P
>
0.05)
was
observed
in
terms
of
handling
stress
or
photoperiod.
0
20
40
60
80
100
24L 12L-12D 24D 24L 12L-12D 24D 24L 12L-12D 24D
good average poor
Photoperiod
Spawning quality (%)
unstressed
stressed
Fig.
3.
Comparison
the
quality
of
spawning
pikeperch
Sander
lucioperca
broodstocks
in
three
condition;
good,
average
and
poor
in
different
pho-
toperiods
including
constant
light
(24L:0D),
12
h
of
light:12
h
of
darkness
(12L:12D),
constant
darkness
(0L:24D),
with
and
without
stress.
No
differ-
ence
(P
>
0.05)
was
observed
in
terms
of
handling
stress
or
photoperiod.
nest)
was
not
observed
either
under
stressful
or
unstressed
condition.
In
addition,
during
24L:0D-s,
no
poor
quality
spawning
was
observed
(Fig.
3),
but
no
interaction
was
found
between
light
and
stress.
Number
of
eggs
per
gram
did
not
show
any
significant
difference
in
different
pho-
toperiods
and
stress
conditions
(Fig.
4).
Hatching
efficiency
and
fertilization
percentage
during
blastula
and
gastrula
stages
showed
no
significant
difference
in
different
pho-
toperiod
treatments
(Figs.
5
and
6)
as
well.
No
interaction
was
also
found
between
light
and
stress
while
in
all
stress-
ful
photoperiods,
lower
fertilization
percentages
during
the
gastrula
stage
were
observed
compared
to
photoperiods
without
any
stress
(P
>
0.05).
The
body
weights
of
the
broodstocks
were
1195
±
200.7
g
and
1662
±
220.4
g
for
males
and
females,
respectively,
in
the
beginning
of
the
experiment
before
applying
different
photoperiod
and
handling
stress,
and
attained
to
1004
±
125.5
g
and
1535
±
200.1
g
for
males
and
females
at
the
end
of
40
days
of
fasting
period.
No
significant
different
was
observed
for
both
sexes
during
this
condition.
3.2.
Sex
steroids
Plasma
testosterone
concentration,
in
both
males
and
females,
showed
no
significant
difference
during
different
0
200
400
600
800
1000
1200
24L 12L-12D 24D
Photoperiod
Number of egg per gram
unstressed
stressed
Fig.
4.
Number
of
egg
pikeperch
Sander
lucioperca
broodstocks/gram
in
different
photoperiods
including
constant
light
(24L:0D),
12
h
of
light:12
h
of
darkness
(12L:12D),
constant
darkness
(0L:24D),
with
and
without
stress.
No
differece
(P
>
0.05)
was
observed
in
terms
of
handling
stress
or
photoperiod.
Author's personal copy
S.
Pourhosein
Sarameh
et
al.
/
Animal
Reproduction
Science
132 (2012) 213–
222 217
0
10
20
30
40
50
60
70
80
90
100
24L
12L-12D
24D
Photoperiod
Fertilization in gastrula (%) Fertilization in blastula (%)
unstressed
stressed
B
0
10
20
30
40
50
60
70
80
90
100
24L
12L-12D
24D
Photoperiod
unstressed
stressed
A
Fig.
5.
Fertilization
percentage
of
pikeperch
Sander
lucioperca
brood-
stocks
in
two
stages:
(A)
blastula
and
(B)
gastrula
in
different
photoperiods
including
constant
light
(24L:0D),
12
h
of
light:12
h
of
darkness
(12L:12D),
constant
darkness
(0L:24D),
with
and
without
stress.
No
difference
(P
>
0.05)
was
observed
in
terms
of
handling
stress
or
photoperiod.
photoperiods,
while
24L:0D-s
resulted
in
lesser
concentra-
tions
of
this
hormone
and
indicated
a
significant
difference
compared
to
that
of
the
beginning
of
the
experiment
in
males.
Furthermore,
no
interaction
was
found
between
light
and
stress
in
females,
an
interaction
was
only
found
in
testosterone
concentration
in
males.
Progesterone
con-
centrations
in
males
and
females
were
not
significantly
different
during
different
photoperiods,
and
no
interac-
tion
was
found
between
light
and
stress.
Estradiol
in
both
sexes
was
not
significantly
different
during
different
pho-
toperiods
with
no
interaction
between
light
and
stress
(Tables
1
and
2).
0
5
10
15
20
25
30
35
40
24L 12L-12D 24D
Photoperiod
Hatching efficiency (%)
unstr
ess e
d
stress ed
Fig.
6.
Hatching
efficiency
(%)
of
pikeperch
Sander
lucioperca
brood-
stocks
in
different
photoperiods
including
constant
light
(24L:0D),
12
h
of
light:12
h
of
darkness
(12L:12D),
constant
darkness
(0L:24D),
with
and
without
stress.
No
difference
(P
>
0.05)
was
observed
in
terms
of
handling
stress
or
photoperiod.
Table
1
Sex
steroids,
calcium
and
stress
indicators
(mean
±
SE)
of
pikeperch
male
broodstocks
Sander
lucioperca
in
different
photoperiods
including
constant
light
(24L:0D),
12
h
of
light
and
12
h
of
darkness
(12L:12D),
constant
darkness
(0L:24D)
and
start
(a
day
before
the
experiment
was
started;
n
=
9),
with
and
without
stress
(n
=
6
for
each).
Samples
were
taken
after
40
days
of
photoperiod/stress
applying.
Start 0L:24D 12L:12D 24L:0D 2-Way
ANOVA
Unstressed
Stressed
Unstressed
Stressed
Unstressed
Stressed
Stress
×
photoperiod
Stress
Photoperiod
Progesterone
(ng
ml−1)
0.13
±
0.01
0.14
±
0.01
0.14
±
0.01
0.15
±
0.02
0.11
±
0.01
0.14
±
0.00
0.13
±
0.02
NS
NS
NS
Estradiol
(ng
ml−1)
35.44 ±
8.32
57.51 ±
13.89
37.01 ±
15.72
44.71 ±
14.13
41.33
±
8.30
43.01
±
13.45
30.01
±
7.41
NS
NS
NS
Testosterone
(ng
ml−1)
2.74 ±
0.63a6.91 ±
3.32
0.38 ±
0.19b0.70 ±
0.45
1.20 ±
0.82ab 2.68 ±
1.63
1.23 ±
0.10ab
*S
>
LNS
Calcium
(mg
dl−1)
10.76
±
0.17
10.83
±
0.53
10.88
±
0.71
11.22
±
1.24
9.53
±
0.43
10.55
±
0.27
10.16
±
0.73
NS
NS
NS
Cortisol
(ng
ml−1)
64.61 ±
29.31
155 ±
39.6
96.6 ±
30.47
149 ±
34.9
97.48 ±
38.66
124.5 ±
28.07
165.87 ±
56.1
NS NS NS
Glucose
(mg
dl−1)
97.89
±
8.04
150.83
±
35.88
115
±
22.52
158.5
±
41.18
87.17
±
17.76
136.83
±
22.97
124.66
±
31.5
NS
NS
NS
Lactate
(mg
dl−1)
28.89 ±
5.07
51.83 ±
5.63
41.8
±
4.56
50
±
14.66
41.66
±
6.05
38.83
±
9.08
48.33
±
6.1
NS
NS
NS
S:
start
(the
beginning
of
experiment);
L:
constant
light;
NS:
no
difference
(P
>
0.05)
in
terms
of
handling
stress
or
photoperiod;
S
>
L:
start
(the
beginning
of
experiment)
had
a
greater
testosterone
concentration
and
there
was
no
difference
with
constant
light
(24L:0D)
in
terms
of
handling
stress. a,
b,
ab Difference
(P
<
0.05)
in
terms
of
handling
stress.
*Interaction
between
light
and
stress.
Author's personal copy
218 S.
Pourhosein
Sarameh
et
al.
/
Animal
Reproduction
Science
132 (2012) 213–
222
Table
2
Sex
steroids
and
calcium
concentrations
(mean
±
SE)
of
pikeperch
female
broodstocks
Sander
lucioperca
in
different
photoperiods
including
constant
light
(24L:0D),
12
h
of
light
and
12
h
of
darkness
(12L:12D),
constant
darkness
(0L:24D)
and
start
(a
day
before
the
experiment
was
started;
n
=
9),
with
and
without
stress
(n
=
6
for
each).
Samples
were
taken
after
40
days
of
photoperiod/stress
applying.
Start
0L:24D
12L:12D
24L:0D
2-Way
ANOVA
Unstressed
Stressed
Unstressed
Stressed
Unstressed
Stressed
Stress
×
photoperiod
Stress
Photoperiod
Progesterone
(ng
ml−1)
0.12
±
0.01
0.13
±
0.01
0.12
±
0.02
0.14
±
0.01
0.13
±
0.01
0.16
±
0.01
0.13
±
0.02
NS
NS
NS
Estradiol
(ng
ml−1)
68.56
±
9.72
50.01
±
50.22
63.57
±
9.36
78.86
±
5.29
66.33
±
9.47
49.01
±
12.74
62.67
±
11.90
NS
NS
NS
Testosterone
(ng
ml−1)
4.67
±
0.67
0.09
±
0.03
3.45
±
1.86
0.57
±
0.42
5.34
±
4.07
2.91
±
2.81
1.65
±
1.02
NS
NS
NS
Calcium
(mg
dl−1)
11.28
±
0.33
11.33
±
0.43
10.27
±
0.46
11.75
±
0.51
11.31
±
0.73
12.26
±
0.77
11.46
±
0.51
NS
NS
NS
Cortisol
(ng
ml−1)
45.24
±
20.99
73.5
±
20.24
110.5
±
32.08
109.03
±
22.42
103.83
±
30.25
200.67
±
55.76
116.66
±
30.09
NS
NS
NS
Glucose
(mg
dl−1)
83.78
±
6.09b190.17
±
28.76a* 84.29
±
17.98*161.86
±
27.64ab 95.33
±
13.74
161.33
±
21.82ab 139.33
±
21.87
NS
D
>
Ds
D
>
S
Lactate
(mg
dl−1)
30
±
4.33
43.83
±
5.86
41.86
±
5.05
32.86
±
2.43
36.83
±
3.66
32
±
2.21
38.33
±
6.1
NS
NS
NS
An
asterisk
shows
significant
difference
between
with
and
without
stress
conditions
in
a
photoperiod.
S:
start
(the
beginning
of
experiment);
D:
constant
darkness;
Ds:
constant
darkness
with
stress;
NS:
no
differece
(P
<
0.05)
in
terms
of
handling
stress
or
photoperiod;
D
>
S:
constant
darkness
(0L:24D)
had
a
greater
glucose
concentration
and
difference
from
the
beginning
of
the
experiment
in
terms
of
photoperiod;
D
>
Ds:
constant
darkness
(0L:24D)
had
a
greater
glucose
concentrations
and
difference
with
constant
darkness
with
stress
(0L:24D-s)
in
terms
of
handling
stress. a,
b,
ab:
Difference
(P
<
0.05)
in
terms
of
handling
stress.
3.3.
Calcium
concentration
Average
calcium
concentration
of
males
was
not
sig-
nificantly
different
while
applying
different
photoperiods
and
handling
stress.
No
significant
difference
was
observed
in
females
regarding
average
calcium
concentrations
when
exposed
to
different
photoperiods
and
handling
stress.
In
addition,
there
was
no
interaction
between
light
and
stress
treatments
(Tables
1
and
2).
3.4.
Cortisol,
glucose
and
lactate
concentrations
Average
plasma
cortisol
concentrations
were
not
sig-
nificantly
different
in
male
broodstocks
during
different
photoperiods
and
handling
stress
(Table
1).
Females
did
not
show
any
such
difference
in
terms
of
average
cor-
tisol
concentrations
during
the
photoperiods
and
stress
handling
as
well
(Table
2).
Maximum
average
cortisol
concentration
of
male
broodstocks
was
observed
dur-
ing
the
24L:0D-s
photoperiod,
and
in
females,
it
was
seen
in
the
24L:0D.
These
maximum
concentrations
were
165
±
57
and
309
±
181
ng
ml−1in
males
and
females,
respectively.
Average
glucose
concentrations
in
male
broodstocks
showed
no
significant
difference
in
the
different
photope-
riods
and
handling
stress
(Table
1).
Although
no
significant
difference
of
glucose
concentrations
was
observed
in
dif-
ferent
photoperiods
in
female
broodstocks,
0L:24D
had
a
greater
glucose
concentration
and
there
was
a
significant
difference
from
the
beginning
of
the
experiment
(P
<
0.05).
Furthermore,
this
concentration
in
stressed
fish
during
the
0L:24D-s
(constant
darkness
with
stress)
photoperiod
had
a
significant
difference
from
that
of
the
unstressed
fish
(Table
2).
Average
lactate
concentrations
were
not
significantly
different
in
male
broodstocks
which
were
exposed
to
dif-
ferent
photoperiods
and
handling
stress
(Table
1).
Females
did
not
show
any
significant
difference
in
this
regard
either
(Table
2).
Overall,
the
interaction
of
light
and
stress
in
terms
of
stress
responses
was
not
observed
in
male
and
female
broodstocks.
4.
Discussion
Results
obtained
from
this
study
indicated
that
different
photoperiods
affect
the
induction
of
spawning
of
pikeperch
broodstocks
during
their
final
stages
of
reproduction
and
lead
to
changes
in
the
timing
of
their
spawning.
Moreover,
the
greatest
and
least
number
of
spawnings
was
observed
in
the
morning
and
at
night,
respectively.
Some
researches
have
stated
that
the
timing
of
spawning
depends
on
the
harmony
of
a
rhythm
or
an
annual
internal
clock
by
chang-
ing
photoperiods
(Bromage
et
al.,
1992;
Bon
et
al.,
1999;
Bonnet
et
al.,
2007).
Results
from
the
present
study
were
consistent
with
those
of
Migaud
et
al.
(2004a,
2006)
showing
that
daily
changes
of
illumination
are
important
to
control
the
spawning
of
Eurasian
perch
Perca
fluviatilis.
In
addition,
in
the
present
study,
the
greatest
and
least
amount
of
spawn-
ing
that
occurred
in
the
morning
and
at
night,
respectively
indicated
that
illumination
changes
function
as
an
Author's personal copy
S.
Pourhosein
Sarameh
et
al.
/
Animal
Reproduction
Science
132 (2012) 213–
222 219
important
stimulant
for
determining
the
time
of
spawn-
ing
in
pikeperch.
So,
it
confirmed
the
results
obtained
by
Migaud
et
al.
(2006),
who
had
observed
the
greatest
amount
of
spawning
in
the
morning
in
Eurasian
perch
exposed
to
different
photoperiods.
The
present
study
also
indicated
that
continuous
illumination
(24L:0D)
dur-
ing
the
final
stage
of
reproduction
accelerates
the
time
of
spawning,
while
0L:24D
delays
spawning.
Lengthen-
ing
photoperiods
during
the
final
stage
of
reproduction,
therefore,
induces
spawning
in
these
fish
and
shortening
photoperiods
delays
their
spawning.
The
findings
of
the
present
study
confirm
those
obtained
by
Muller-Belecke
and
Zienert
(2008),
that
they
stimulated
the
fish
by
photo-
thermal
and
photoperiod
regimens
and
made
it
possible
for
pikeperch
to
spawn
2
months
prior
to
the
spawning
season.
In
the
present
study,
number
of
eggs
per
gram,
percent-
age
of
fertilization
during
blastula
and
gastrula
stages,
sex
steroids,
and
calcium
concentrations
were
the
same
among
different
photoperiods
and
handling
stress.
Also,
cortisol
concentrations
in
treatments
with
different
photoperiods
and
handling
stress
did
not
show
a
significant
difference
which
could
be
explained
by
the
fish
being
adapted
to
daily
stress
because,
according
to
reports
by
researchers,
long-
term
exposure
to
a
stressor
could
lead
to
allostasis,
which
is
practically
considered
as
the
ability
to
return
to
the
physio-
logical
concentrations
observed
prior
to
exposure
to
stress
(McEwen,
1998;
Schreck,
2000).
No
increase
in
cortisol
concentrations
of
brooders
which
were
stressed
for
a
long
time
during
the
present
research
could
be
the
result
of
a
negative
feedback
of
a
daily
increase
of
the
cortisol
concentration
in
the
hypothalamus–pituitary–interrenal
axis
(Pickering
and
Pottinger,
1987;
Fast
et
al.,
2008),
and
fish
were
adapted
with
the
stress
after
a
long
period
of
exposure.
As
mentioned,
adaptation
to
stress
during
the
experiment
seems
possible.
However,
the
effect
of
stress
on
cortisol
concentrations
may
be
detectable
while
comparing
con-
centrations
with
those
obtained
before
treatments
began
at
“start”.
Biswas
et
al.
(2006a,b)
found
that
different
pho-
toperiods
do
not
cause
a
considerably
significant
stress
response
(glucose–cortisol)
in
red
sea
bream
Pagrus
major
which
confirms
results
here.
Moreover,
results
obtained
from
a
study
conducted
by
Fast
et
al.
(2008)
on
salmons
(Salmo
salar)
and
those
of
Wang
et
al.
(2004)
on
striped
bass
(Morone
saxatilis)
brooders
showed
that
continuous
acute
handling
stressor
did
not
result
in
a
long-term
increase
in
glucose
and
cortisol
concentrations.
It
is
also
probable
that
a
significant
increase
in
glucose
concentrations
of
female
brooders
of
the
0L:24D
photope-
riod
compared
to
that
of
the
beginning
of
the
experiment
and
the
0L:24D-s
photoperiod
is
due
to
the
fact
that
long
exposures
to
darkness
(considering
the
delayed
spawning
during
this
photoperiod
compared
to
other
photoperiods)
would
cause
a
stress
response
through
increasing
the
glu-
cose
concentration.
Also,
it
seems
that
pikeperch
brooders
are
sensitive
to
darkness
and,
in
the
long
run,
react
to
it
by
an
increased
glucose
concentration
as
a
secondary
stress
response.
In
as
much
as
stressor
responses
might
change
in
different
photoperiods,
these
may
cause
greater
or
lesser
sensitivity
(Biswas
et
al.,
2006a).
In
addition,
the
exposure
time
during
which
pikeperch
are
exposed
to
stressors
could
affect
the
magnitude
of
the
physiolog-
ical
response,
and
the
sensitivity
of
their
response
depends
on
stressor
type
(Acerete
et
al.,
2004),
which
justifies
the
reason
for
a
significant
decrease
in
the
glucose
concentra-
tion
in
brooders
during
0L:24D-s
compared
to
that
of
the
0L:24D.
In
the
present
study,
handling
stress
resulted
in
a
delay
or
lack
of
spawning
in
pikeperch
which
could
be
due
to
the
fact
that
in
fish
kept
under
stressful
conditions
some
kind
of
exchange
between
allocations
of
energy
for
repro-
duction
and
maintenance
occurs
(Schreck
et
al.,
2001;
Wang
et
al.,
2006).
Wang
et
al.
(2006)
demonstrated
that
handling
stress
is
an
important
modulating
factor
in
the
reproductive
cycle
of
fish
and
can
be
of
paramount
impor-
tance
during
the
reproductive
cycle.
Results
of
Campbell
et
al.
(1992)
showed
that
applying
acute
stress
during
the
completion
of
the
reproduction
process
delays
the
time
of
spawning
in
rainbow
trout
(Oncorhynchus
mykiss).
They
suggested
that
reproduction
is
a
physiological
pro-
cess
which
is
very
sensitive
to
the
harmful
effects
of
stress.
In
the
present
study,
the
reason
why
over
half
of
the
fish
exposed
to
handling
stress
spawned
could
be
related
to
their
relative
ripeness
for
spawning
or,
in
course
of
time,
these
fish
became
adapted
to
stressful
conditions
because,
according
to
other
researchers,
long
periods
of
exposure
to
a
stressor
could
lead
to
allostasis
which
is
prac-
tically
the
body’s
ability
to
return
to
its
physiological
state
observed
before
applying
stress
(McEwen,
1998;
Schreck,
2000).
Morehead
et
al.
(2000)
also
showed
that
striped
trumpeter
Latris
lineata,
in
spite
of
frequent
handling
with
stressful
conditions,
became
adapted
to
9
or
12
months
of
consecutive
photo-thermal
cycles
and
handling
stress
and
completed
their
spontaneous
maturity
and
spawning
cycles
which
is
a
confirmation
of
the
findings
of
the
present
study.
Additionally,
absence
of
spawning
in
about
1/3
of
brood-
stocks
to
which
handling
stress
was
applied
in
comparison
with
the
spawning
of
over
half
of
the
handled
fish
could
be
due
to
individual
differences
among
fish.
Schreck
(1981)
stated
that
the
fish
performance,
when
exposed
to
stress,
is
regulated
with
their
ability
to
have
individual
perform-
ances.
Depending
on
the
fact
that
in
which
stage
of
life
and
with
what
intensity
a
stress
is
experienced
and
for
how
long
a
stressor
performance
lasts,
these
factors
may
affect
reproduction
in
different
ways
(Schreck,
2009).
Also,
nutri-
tional
factors
(Bromage
et
al.,
1992;
Carrillo
et
al.,
1995;
Zohar
et
al.,
1995;
Pereira
et
al.,
1998;
Siddiqui
et
al.,
1998),
fish
age
and
spawning
rank
(Kjorsvik,
1994;
Navas
et
al.,
1995;
Brooks
et
al.,
1997),
the
amount
of
stress
(Campbell
et
al.,
1992;
Contreras-Sanchez
et
al.,
1998;
Schreck
et
al.,
2001),
strain
(Bromage
et
al.,
1990),
and,
probably,
their
final
sexual
maturity
can
directly
influence
fish
reaction
and
reproduction
quality.
Because
teleosts
use
relatively
different
reproduction
strategies
to
overcome
stress,
it
is
possible
that
different
fish
species,
in
terms
of
the
nature
of
their
physiological
reaction,
would
have
different
repro-
ductive
outcomes
in
relation
to
a
stressful
factor,
and
their
reaction
to
stress
depending
on
their
species,
stage
of
matu-
rity,
and
type
and
intensity
of
a
stressor
could
be
different
(Schreck
et
al.,
2001).
Author's personal copy
220 S.
Pourhosein
Sarameh
et
al.
/
Animal
Reproduction
Science
132 (2012) 213–
222
During
early
oogenesis
or
maturation,
the
reproductive
clock
could
buffer
some
eggs
from
energetic
or
nutritional
deficits
by
causing
atresia
of
others.
Near
complete
matu-
rity,
reproduction
success
is
optimized
by
regulating
the
timing
of
spawning.
Thus,
depending
on
the
fact
that
in
which
stage
of
the
reproduction
stressful
factors
are
expe-
rienced,
they
may
have
different
effects,
and
when
the
fish
are
exposed
to
these
factors,
the
period
of
matura-
tion
seems
important
(Schreck
et
al.,
2001).
As
a
result,
since
in
this
study,
pikeperch
were
exposed
to
different
photoperiods
in
their
final
stages
of
maturity,
the
effect
of
handling
stress
was
observed
on
the
timing
of
spawning,
but
it
did
not
have
any
significant
effect
on
the
number
of
eggs
per
gram,
fertilization
percentage,
sex
steroids,
and
calcium
concentrations.
A
study
conducted
by
Contreras-Sanchez
et
al.
(1998)
showed
that
rainbow
trout
exposed
to
mild
stress
dur-
ing
early
vitellogenesis
produced
smaller
eggs
which
were
different
in
size,
while
in
those
stressed
during
late
vitel-
logenesis,
no
changes
in
average
egg
size
was
observed.
Besides,
rainbow
trout
which
were
exposed
to
mild
stress,
showed
no
changes
in
average
egg
size,
but
egg
sizes
were
more
heterogeneous
and
consistent
with
the
results
of
this
research.
In
a
study
conducted
by
Fontaine
et
al.
(2006)
early
plasma
testosterone
and
estradiol
concentrations
in
male
and
female
Eurasian
perch
during
different
photoperiods
8L:16D,
12L:12D,
and
16L:8D
showed
no
significant
differ-
ence.
Sulistyo
et
al.
(1998)
showed
that
fluctuations
in
sex
steroids
in
the
reproductive
cycle
of
Eurasian
perch
com-
pared
to
other
species
(e.g.
cyprinids
and
salmonids)
are
less,
which
can
justify
the
lack
of
significant
differences
between
sex
steroid
concentrations
in
different
photope-
riod
treatments
of
the
present
research.
In
the
present
study,
males
of
the
24L:0D-s
photoperiod
had
lesser
concentrations
of
testosterone
and
showed
a
significant
difference
with
those
of
the
beginning
of
the
experiment.
This
finding
suggested
the
negative
effect
of
stress
on
the
reduction
of
testosterone.
It
is
probable
that
more
reduc-
tions
in
testosterone
in
this
photoperiod
is
due
to
the
fact
that
fish
kept
in
24L:0D-s
use
more
energy
because
of
releasing
some
corticostroids
during
constant
light
condi-
tion.
Moreover,
because
glucocorticostroid
rhythms
reflect
changes
in
the
balance
of
energy
in
animals,
when
needs
for
energy
are
more
than
the
existing
energy
resources,
the
glucocorticostroid
concentrations
increase
(Goymann
and
Wingfeld,
2004).
In
addition,
chronic
stress
reduces
the
circulation
of
sex
steroids
through
the
cortisol
function
(Pankhurst
and
Van
Der
Kraak,
1997).
Greater
concentra-
tions
of
cortisol,
therefore,
result
in
more
reductions
in
testosterone
concentrations
(our
unpublished
data).
Sim-
ilarly,
Acerete
et
al.
(2004)
found
that
the
intensity
of
response
in
Eurasian
perch
depended
on
the
type
of
stres-
sor
and
that
differences
in
the
nature,
duration
of
a
stress
along
with
species
differences
could
lead
to
differences
in
the
timing
and
magnitude
of
the
cortisol
response
(Ramsay
et
al.,
2009).
The
previous
findings
indicated
that
stress
response
was
hereditary
and
individual
responses
through
the
passage
of
time
were
a
stable
characteristic
(Fevolden
et
al.,
1991;
Pottinger
and
Carrick,
1999;
Wang
et
al.,
2004).
Therefore,
changes
which
occur
due
to
stress
may
be
rapid
delayed,
long-term,
or
short-term,
depending
on
fish
species,
their
life
stage,
the
nature
of
stressor,
and
other
environmental
factors
(Schreck,
2000;
Barton,
2000,
2002).
In
view
of
these
results
and
given
that
fasting
can
induce
stress
and
other
endocrine
changes
(e.g.
hypoglycemia)
it,
in
the
present
experiment,
created
unique
conditions
under
which
different
photoperiods
and
another
stress-
inducing
factor,
handling,
were
tested.
Some
studies
have
shown
that
food
deprivation
modifies
cortisol
response
to
handling
disturbance
in
rainbow
trout
(Vijayan
and
Moon,
1992;
Reddy
et
al.,
1995).
Similarly,
food
deprivation
also
alters
stressor-induced
hyperglycemia
in
fish
(Vijayan
and
Moon,
1992).
In
a
study
conducted
by
Jorgensen
et
al.
(2002),
the
pre-handling
plasma
cortisol
concentrations
were
much
greater
in
food-deprived
than
in
fed
fish.
Their
results
are
in
contrast
to
other
studies
showing
that
food
deprivation
either
suppressed
(Barton
et
al.,
1988;
Sumpter
et
al.,
1991;
Vijayan
and
Moon,
1994;
Reddy
et
al.,
1995)
or
did
not
affect
plasma
cortisol
concentration
in
rainbow
trout
(Farbridge
and
Leatherland,
1992;
Vijayan
and
Moon,
1992).
Jorgensen
et
al.
(2002)
expressed
that
the
reason
for
these
differences
may
be
related
to
the
longer
dura-
tion
of
the
food
deprivation
(∼140
days)
compared
to
other
studies,
up
to
6
weeks.
But
in
the
present
study,
during
the
experimental
period
of
40
days,
the
fish
were
fasting
and
had
greater
plasma
cortisol
concentrations
than
the
beginning
of
the
experiment.
Also,
in
Jorgensen
et
al.
(2002)
study,
there
was
no
significant
difference
in
plasma
corti-
sol
concentrations
between
the
fed
and
food-deprived
fish
at
6
and
23
h
post-handling
disturbance.
Although
fasting
was
applied
to
all
groups
in
the
present
study,
it
is
important
to
consider
and
monitor
the
effects
of
fasting
on
endocrine
changes
and
other
aspects
of
the
physiology
and
reproductive
performance.
More
detailed
studies,
like
controlled
challenge
of
fish
fed
at
different
photoperiod
are
needed.
The
present
study
increases
the
knowledge
of
repro-
ductive
processes
in
pikeperch
and
found
photoperiod
manipulation
at
the
final
stage
of
reproduction
in
fish
leads
to
changes
in
the
timing
of
their
spawning
and
does
not
have
any
effect
on
other
fertilization
parameters.
In
addi-
tion,
despite
the
acceleration
of
spawning
during
24L:0D
compared
to
other
photoperiods,
no
changes
were
found
in
the
number
of
eggs
per
gram
and
other
fertilization
parameters.
Thus,
an
enhanced
spawning
performance
in
the
final
stage
of
reproduction
can
be
attributed
to
this
photoperiod.
However,
in
order
to
ensure
that
whether
applying
constant
light
that
during
the
final
stage
of
repro-
duction
caused
the
spawning
of
pikeperch
to
accelerate
compared
to
other
photoperiods
is
a
species-related
char-
acteristic,
or
it
changes
during
longer
periods
or
at
early
stages
of
reproduction
and
maturity
requires
more
stud-
ies
to
be
done.
Additionally,
with
consideration
of
the
delayed
or
absence
of
spawning
in
half
of
the
fish
exposed
to
0L:24D-s
and
1/3
of
those
kept
in
24L:0D-s
and
12L:12D-
s,
it
is
concluded
that
the
handling
process
induces
stress
in
pikeperch
and
leads
to
a
poor
reproductive
perfor-
mance.
Thus,
the
importance
of
preventing
harmful
effects
of
handling
stress
and
identifying
methods
which
can
minimize
the
adverse
effects
of
handling
needs
to
be
con-
sidered.
Author's personal copy
S.
Pourhosein
Sarameh
et
al.
/
Animal
Reproduction
Science
132 (2012) 213–
222 221
Acknowledgements
Thanks
are
extended
to
the
staff
at
the
Dr.
Yousefpour
Fish
Hatchery
Center
for
their
help
during
the
experiments
and
M.
Maleki
for
analyzing
the
steroids
samples.
Also,
we
are
especially
grateful
to
Dr.
Fariborz
Jamalzad
for
his
help.
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