FINAL REPORT OF FISH GENETICS PRACTICUM
HYBRIDIZATION IN CARP
(
C
yprinus
ca
rpio
)
AND NILEM FISH
(
O
st
e
o
c
hilus h
a
ss
e
lti
CV
)
P
r
e
p
a
r
ed a
s on
e
of th
e
r
e
quir
e
m
e
nts to fulfill th
e
fin
a
l pr
ac
ti
c
um r
e
port
a
ssignm
e
nt
F
ish
Ge
n
e
ti
c
s
C
ours
e
,
e
v
e
n s
e
m
e
st
e
r
BRADFORD H
.
DEPARTEMENT OF BIOLOGY
ARIZONA STATE UNIVERSITY
INTRODUCTION
1
.
1
Bac
kgroun
d
Se
l
ec
tion
ca
n
a
lt
e
r th
e a
v
e
r
a
g
e
v
a
lu
e
of qu
a
ntit
a
tiv
e
tr
a
its in
a
popul
a
tion
b
y utilizing
add
itiv
e
g
e
n
e
ti
c
v
a
ri
a
n
ce
(
VA
),
whi
c
h is
de
t
e
rmin
ed b
y
a
ll
e
l
e
s
.
T
his
add
itiv
e
g
e
n
e
ti
c
v
a
ri
a
n
ce
r
e
fl
ec
ts th
e
sup
e
riority of
a
ll
e
l
e
s in th
e
p
a
r
e
nt fish
a
n
d ca
n
be
fully inh
e
rit
ed b
y th
e
offspring
,
a
s
VA
r
e
m
a
ins un
a
ff
ec
t
ed b
y
a
ll
e
l
e
s
e
gr
e
g
a
tion or
c
rossov
e
r
d
uring m
e
iosis
.
T
h
e
proportion of
a
utosom
a
l g
e
n
e
s with
add
itiv
e e
ff
ec
ts
on ph
e
notypi
c
v
a
ri
a
n
ce
(
VP
)
d
i
c
t
a
t
e
s th
e
h
e
rit
ab
ility of
a
tr
a
it
.
I
f th
e
h
e
rit
ab
ility
(
h
²)
is l
e
ss th
a
n
15
%
,
improving
VA
through s
e
l
ec
tion
bec
om
e
s
mor
e c
h
a
ll
e
nging
.
W
h
e
n
VA
is limit
ed
,
hy
b
ri
d
iz
a
tion
ca
n
be e
mploy
ed
to
c
r
ea
t
e
n
e
w
a
ll
e
l
e c
om
b
in
a
tions
,
pot
e
nti
a
lly l
ead
ing to th
e
inh
e
rit
a
n
ce
of
d
omin
a
nt sup
e
rior
a
ll
e
l
e
s
.
T
his
e
xploit
a
tion of
d
omin
a
nt g
e
n
e
ti
c
v
a
ri
a
n
ce
(
VD
)
in hy
b
ri
d
iz
a
tion
ca
n r
e
sult in n
e
w str
a
ins
e
xhi
b
iting h
e
t
e
rosis
,
th
e
r
eb
y
e
nh
a
n
c
ing sp
ec
ifi
c
tr
a
its in fish
.
1
.
2
Ide
ntifi
ca
tion of pro
b
l
e
ms
H
y
b
ri
d
iz
a
tion r
e
f
e
rs to th
e
pro
ce
ss of
c
om
b
ining
d
iff
e
r
e
nt g
e
n
e
ti
c
m
a
t
e
ri
a
l from two
d
istin
c
t sp
ec
i
e
s or str
a
ins to pro
d
u
ce
offspring with
de
sir
ab
l
e
tr
a
its
.
T
h
e
m
e
tho
d
of hy
b
ri
d
iz
a
tion involv
e
s s
e
l
ec
ting p
a
r
e
nt
org
a
nisms with sp
ec
ifi
c c
h
a
r
ac
t
e
risti
c
s
a
n
d
int
e
ntion
a
lly
b
r
eed
ing th
e
m
to
c
r
ea
t
e
hy
b
ri
d
s
.
T
h
e
purpos
e
of
a
hy
b
ri
d
iz
a
tion progr
a
m is to
e
nh
a
n
ce ce
rt
a
in qu
a
liti
e
s in th
e
offspring
,
su
c
h
a
s improv
ed
r
e
sili
e
n
ce
,
pro
d
u
c
tivity
,
or
ada
pt
a
tion to sp
ec
ifi
c e
nvironm
e
nt
a
l
c
on
d
itions
.
I
n fish
e
ggs tr
ea
t
ed
with hy
b
ri
d
iz
a
tion
,
FR
(
f
e
rtiliz
a
tion r
a
t
e
),
SR
(
surviv
a
l r
a
t
e
),
a
n
d HR
(
h
a
t
c
hing r
a
t
e
)
a
r
e
k
e
y p
a
r
a
m
e
t
e
rs th
a
t
a
r
e
m
ea
sur
ed
to
e
v
a
lu
a
t
e
th
e
su
cce
ss
a
n
d e
ff
ec
tiv
e
n
e
ss of th
e
hy
b
ri
d
iz
a
tion pro
ce
ss
.
1
.
3
Ob
j
ec
tiv
e
T
h
e a
im of this pr
ac
ti
c
um is th
a
t stu
de
nts
ca
n
a
pply
a
hy
b
ri
d
iz
a
tion
progr
a
m th
a
t
c
oll
ec
ts
d
omin
a
nt g
e
n
e
ti
c
v
a
ri
a
nts if th
e
p
a
r
e
nt s
e
l
ec
tion
progr
a
m
d
o
e
s not
ac
hi
e
v
e
th
e e
xp
ec
t
ed
r
e
sults
beca
us
e
th
e SD a
n
d CV
v
a
lu
e
s of
a
tr
a
it
a
r
e
r
e
l
a
tiv
e
ly sm
a
ll
.
1
.
4
U
s
e
s
T
h
e
us
e
fuln
e
ss of this pr
ac
ti
c
um is th
a
t pr
ac
tition
e
rs
ca
n know
,
un
de
rst
a
n
d
,
a
n
d a
pply hy
b
ri
d
iz
a
tion progr
a
ms to gol
d
fish
a
n
d
nil
e
m fish
b
y
c
oll
ec
ting
d
omin
a
nt g
e
n
e
ti
c
v
a
ri
a
nts
.
CHAPTER II
LITERATURE REVIEW
2
.
1
G
ol
d
fish
G
ol
d
fish
a
r
e
highly
ada
pt
ab
l
e
fish th
a
t thriv
e
in v
a
rious
e
nvironm
e
nts
,
l
ead
ing to th
e
ir wi
de
spr
ead d
istri
b
ution
ac
ross
a
lmost
e
v
e
ry p
a
rt of th
e
worl
d
.
T
h
e
y
a
r
e
known
b
y
d
iff
e
r
e
nt n
a
m
e
s in v
a
rious r
e
gions
.
I
n
E
nglish
,
th
e
y
a
r
e
r
e
f
e
rr
ed
to
a
s
c
ommon
ca
rp
,
whil
e
in
Ja
v
a
,
th
e
y
a
r
e ca
ll
ed
m
a
sm
a
s
a
n or us
ed a
s
a
gol
d
fish
d
ish
.
I
n
S
um
a
tr
a
,
gol
d
fish
a
r
e
known
a
s
r
a
yo fish or m
a
m
e
h fish
.
C
l
a
ssifi
ed
un
de
r th
e A
nim
a
li
a
king
d
om
,
gol
d
fish
be
long to th
e C
hor
da
t
a
phylum
,
th
e Ac
tinopt
e
rygii
c
l
a
ss
,
th
e
C
ypriniform
e
s or
de
r
,
th
e C
yprini
dae
f
a
mily
,
a
n
d
th
e C
yprinus g
e
nus
,
2
.
1
.
1
G
ol
d
fish
M
orphology
G
ol
d
fish
e
xhi
b
it
d
istin
c
t morphologi
ca
l f
ea
tur
e
s
,
in
c
lu
d
ing
a
slightly
e
long
a
t
ed a
n
d
fl
a
tt
e
n
ed b
o
d
y with
a
t
e
rmin
a
l mouth th
a
t
ca
n protru
de
.
T
wo p
a
irs of
a
nt
e
nn
ae a
r
e
lo
ca
t
ed
in front of th
e
mouth
,
a
n
d
th
e
inn
e
r
e
n
d
of th
e
mouth h
a
s thr
ee
rows of ph
a
ryng
ea
l t
ee
th
.
T
h
e
ir
b
o
d
i
e
s
a
r
e
mostly
c
ov
e
r
ed
with r
e
l
a
tiv
e
ly l
a
rg
e c
y
c
loi
d
s
ca
l
e
s
.
T
h
e
gol
d
fish poss
e
ss
fiv
e
typ
e
s of fins
:
d
ors
a
l
,
p
ec
tor
a
l
,
p
e
lvi
c
,
a
n
a
l
,
a
n
d ca
u
da
l
,
with
a
ri
b
lin
e
running
a
long th
e
mi
dd
l
e
of th
e b
o
d
y from th
e
gills to th
e
t
a
il
.
G
ol
d
fish
a
r
e
prim
a
rily foun
d
in fr
e
shw
a
t
e
r
e
nvironm
e
nts su
c
h
a
s pon
d
s
a
n
d
l
a
k
e
s
,
pr
e
f
e
rring
a
r
ea
s with mo
de
r
a
t
e a
ltitu
de
s
be
tw
ee
n
150
-
600
m
e
t
e
rs
a
n
d
t
e
mp
e
r
a
tur
e
s r
a
nging from
25
-
30
°
C
.
De
spit
e be
ing fr
e
shw
a
t
e
r fish
,
th
e
y
a
r
e
o
cca
sion
a
lly s
ee
n in
b
r
ac
kish w
a
t
e
rs
.
I
n
d
on
e
si
a
is hom
e
to
e
ight v
a
ri
e
ti
e
s of gol
d
fish
,
a
mong whi
c
h
Ma
j
a
l
a
y
a
,
S
inyony
a
,
Ta
iw
a
n
,
a
n
d
hy
b
ri
d
typ
e
s
a
r
e
p
a
rti
c
ul
a
rly popul
a
r
.
T
h
e
P
unt
e
n gol
d
fish st
a
n
d
s out with its
da
rk gr
ee
n s
ca
l
e
s
,
c
omp
ac
t
b
o
d
y
,
high
bac
k
,
a
n
d a
gil
e
mov
e
m
e
nt
,
d
ispl
a
ying
a
l
e
ngth
-
to
-
h
e
ight r
a
tio of
2
.
3
:
1
.
Ma
j
a
l
a
y
a
gol
d
fish
,
r
ec
ogniz
ab
l
e b
y th
e
ir gr
a
yish
-
gr
ee
n s
ca
l
e
s with
da
rk
e
r
ed
g
e
s
,
poss
e
ss
a
short
e
r
b
o
d
y
a
n
d a
slow
e
r p
ace
,
oft
e
n s
ee
n swimming
n
ea
r th
e
w
a
t
e
r
'
s surf
ace
,
with
a
3
.
2
:
1
l
e
ngth
-
to
-
h
e
ight r
a
tio
.
T
h
e
Ca
ngkring
a
n v
a
ri
e
ty
,
with its r
edd
ish
-
y
e
llow s
ca
l
e
s
a
n
d
r
ed
fins
,
f
ea
tur
e
s
a
n
e
long
a
t
ed b
o
d
y
a
n
d a
l
e
ngth
-
to
-
h
e
ight r
a
tio of
2
.
87
:
1
.
S
inyony
a
gol
d
fish
a
r
e
i
de
ntifi
ed b
y th
e
ir light y
e
llow s
ca
l
e
s
a
n
d e
long
a
t
ed b
o
d
i
e
s
,
moving slowly
a
n
d
pr
e
f
e
rring th
e
w
a
t
e
r
'
s surf
ace
,
with
a
r
a
tio of
3
.
6
:
1
.
La
stly
,
th
e Ta
iw
a
n
ca
rp
,
with y
e
llowish
-
gr
ee
n s
ca
l
e
s
,
d
ispl
a
ys
a
long
b
o
d
y
,
roun
ded bac
k
,
a
n
d
mor
e ac
tiv
e
mov
e
m
e
nts
,
with
a
3
.
5
:
1
l
e
ngth
-
to
-
h
e
ight
r
a
tio
.
2
.
2
N
il
e
m
F
ish
I
n
I
n
d
on
e
si
a
,
nil
e
m fish
a
r
e
r
e
f
e
rr
ed
to
b
y v
a
rious n
a
m
e
s su
c
h
a
s
nil
e
m
,
l
e
h
a
t
,
m
a
gut
,
r
e
gis
,
mil
e
m
,
muntu
,
p
a
lung
,
p
a
l
a
u
,
p
a
w
a
s
,
puy
a
u
,
a
s
a
ng
,
a
n
d
p
e
nop
a
,
a
n
d a
r
e
oft
e
n r
ec
ogniz
ed a
s
ca
rp
.
T
h
e
ir
d
istri
b
ution
sp
a
ns s
e
v
e
r
a
l
c
ountri
e
s in
c
lu
d
ing
Ma
l
a
ysi
a
,
T
h
a
il
a
n
d
,
V
i
e
tn
a
m
,
Ca
m
b
o
d
i
a
,
a
n
d
p
a
rts of
I
n
d
on
e
si
a
lik
e Ja
v
a
,
S
um
a
tr
a
,
Ka
lim
a
nt
a
n
,
a
n
d S
ul
a
w
e
si
.
T
h
e
nil
e
m fish
be
longs to th
e P
is
ce
s
c
l
a
ss
a
n
d
is
c
l
a
ssifi
ed
un
de
r th
e
or
de
r
O
st
a
riophysi
,
su
b
-
or
de
r
C
yprinoi
dea
,
within th
e C
yprini
dae
f
a
mily
,
a
n
d
th
e
su
b
-
f
a
mily
C
yprinin
ae
.
I
ts g
e
nus is
O
sth
e
o
c
hilus
,
a
n
d
th
e
sp
ec
i
e
s is
i
de
ntifi
ed a
s
O
sth
e
o
c
hilus h
a
ss
e
lti
.
2
.
2
.
1
N
il
e
m
F
ish
M
orphology
T
h
e N
il
e
m fish poss
e
ss
e
s
a
fl
a
t
b
o
d
y with
a
t
e
rmin
a
l mouth th
a
t
ca
n
protru
de
outw
a
r
d
,
situ
a
t
ed a
t th
e
tip of its nos
e
.
I
ts p
e
lvi
c
fins
a
r
e
lo
ca
t
ed
be
hin
d
th
e
p
ec
tor
a
l fins
,
a
n
d
it f
ea
tur
e
s
c
y
c
loi
d
s
ca
l
e
s
.
T
h
e
upp
e
r j
a
w is
a
s
long
a
s
,
or long
e
r th
a
n
,
th
e e
y
e
'
s
d
i
a
m
e
t
e
r
,
whil
e
its snout whisk
e
rs
a
r
e
short
e
r th
a
n th
e
h
ead
.
T
h
e d
ors
a
l fin st
a
rts in lin
e
with th
e
8
th to
10
th
l
a
t
e
r
a
l lin
e
s
ca
l
e
s
,
a
n
d
th
e a
n
a
l fin is slightly upright
,
position
ed
opposit
e
th
e
22
n
d
or
23
r
d
l
a
t
e
r
a
l lin
e
s
ca
l
e
s
.
B
oth th
e
p
e
lvi
c a
n
d
p
ec
tor
a
l fins
a
r
e
n
ea
rly
e
qu
a
l in l
e
ngth
,
with th
e
p
e
lvi
c
fins st
a
rting s
e
v
e
r
a
l s
ca
l
e
s
a
w
a
y
from th
e
l
a
t
e
r
a
l lin
e a
n
d
not r
eac
hing th
e a
nus
.
T
h
e N
il
e
m h
a
s
a
fork
ed
ca
u
da
l fin
,
a
n
d
its
ca
u
da
l p
ed
un
c
l
e
is
a
lmost
a
s long
a
s it is high
,
c
ov
e
r
ed
b
y
16
s
ca
l
e
s
.
De
p
e
n
d
ing on s
ca
l
e c
olor
,
N
il
e
m fish
a
r
e c
l
a
ssifi
ed
into two
typ
e
s
:
b
l
ac
kish
-
b
rown
a
n
d
r
ed
.
T
h
e
fish thriv
e
in fr
e
shw
a
t
e
r
e
nvironm
e
nts
with
d
issolv
ed
oxyg
e
n l
e
v
e
ls
be
tw
ee
n
5
-
8
mg
/
L
.
T
h
e
y
a
r
e
w
e
ll
-
suit
ed
for
tropi
ca
l
a
r
ea
s
a
t
a
ltitu
de
s
be
tw
ee
n
150
-
1000
m
e
t
e
rs
,
with th
e
optim
a
l
a
ltitu
de a
roun
d
800
m
e
t
e
rs
ab
ov
e
s
ea
l
e
v
e
l
.
S
p
a
wning o
cc
urs wh
e
n
oxyg
e
n l
e
v
e
ls r
a
ng
e
from
5
-
6
mg
/
L
,
with
a
suit
ab
l
e
t
e
mp
e
r
a
tur
e
r
a
ng
e
of
18
-
28
°
C a
n
d
p
H be
tw
ee
n
6
-
8
.
6
,
whil
e a
mmoni
a c
ont
e
nt shoul
d
r
e
m
a
in
be
low
0
.
5
mg
/
L
.
2
.
3
A
rtifi
c
i
a
l
S
p
a
wning
A
rtifi
c
i
a
l fish sp
a
wning involv
e
s in
d
u
c
ing fish to sp
a
wn
b
y
ad
minist
e
ring hormon
a
l stimul
a
tion to h
a
st
e
n th
e
m
a
turity of th
e
gon
ad
s
a
n
d
trigg
e
r ovul
a
tion using
a
rtifi
c
i
a
l m
e
tho
d
s
,
su
c
h
a
s stripping or sorting
t
ec
hniqu
e
s
.
F
ish sp
ec
i
e
s lik
e Pa
tin
,
gol
d
fish
,
a
n
d ca
tfish
ca
n un
de
rgo this
pro
ce
ss
.
H
ormon
a
l stimul
a
tion is
ac
hi
e
v
ed
using su
b
st
a
n
ce
s su
c
h
a
s
O
v
a
prim
a
n
d HCG
.
F
or f
e
m
a
l
e
fish
,
a
st
a
n
da
r
d d
os
a
g
e
of
0
.
5
ml
/
kg is
giv
e
n
,
whil
e
for m
a
l
e
s
,
a d
os
e
of
0
.
2
ml
/
kg is
ad
minist
e
r
ed
wh
e
n n
eeded
.
T
h
e d
os
a
g
e
is typi
ca
lly split
,
with on
e
-
thir
d
of th
e
tot
a
l
a
mount giv
e
n
d
uring th
e
first inj
ec
tion
a
n
d
th
e
r
e
m
a
ining two
-
thir
d
s in
a
su
b
s
e
qu
e
nt
inj
ec
tion
.
2
.
4
H
y
b
ri
d
iz
a
tion
Me
tho
d
H
y
b
ri
d
iz
a
tion is
a b
r
eed
ing t
ec
hniqu
e a
ppli
cab
l
e
to fish
,
shrimp
,
sh
e
llfish
,
a
n
d
s
ea
w
eed
,
a
iming to pro
d
u
ce
sup
e
rior in
d
ivi
d
u
a
ls
,
though
som
e
m
a
y
be
st
e
ril
e a
n
d c
r
ea
t
e
n
e
w str
a
ins
.
T
his pro
ce
ss is f
ac
ilit
a
t
ed b
y
a
rtifi
c
i
a
l r
e
pro
d
u
c
tion m
e
tho
d
s
,
su
c
h
a
s thos
e
us
ed
for gol
d
fish
a
n
d
til
a
pi
a
,
wh
e
r
e e
gg
a
n
d
sp
e
rm stripping
a
r
e
possi
b
l
e
.
H
y
b
ri
d
iz
a
tion involv
e
s
m
a
ting
be
tw
ee
n
d
iff
e
r
e
nt sp
ec
i
e
s to
ac
hi
e
v
e
offspring with
e
nh
a
n
ced
tr
a
its
.
F
ish
e
ri
e
s g
e
n
e
ti
c
ists
c
l
a
ssify hy
b
ri
d
iz
a
tion into thr
ee
typ
e
s
:
intr
a
sp
ec
ifi
c
hy
b
ri
d
iz
a
tion
,
whi
c
h o
cc
urs
be
tw
ee
n th
e
s
a
m
e
sp
ec
i
e
s from
d
iff
e
r
e
nt popul
a
tions
;
int
e
rsp
ec
ifi
c
hy
b
ri
d
iz
a
tion
,
whi
c
h o
cc
urs within th
e
s
a
m
e
g
e
nus
b
ut
d
iff
e
r
e
nt sp
ec
i
e
s
;
a
n
d
int
e
rg
e
n
e
ri
c
hy
b
ri
d
iz
a
tion
,
whi
c
h
o
cc
urs
be
tw
ee
n
d
iff
e
r
e
nt g
e
n
e
r
a
.
T
h
e
fun
da
m
e
nt
a
l prin
c
ipl
e
of
hy
b
ri
d
iz
a
tion is to g
e
n
e
r
a
t
e
n
e
w
a
ll
e
l
e c
om
b
in
a
tions
,
wh
e
r
e d
omin
a
nt
sup
e
rior
a
ll
e
l
e
s
e
nh
a
n
ce
pro
d
u
c
tivity
.
S
u
cce
ssful hy
b
ri
d
iz
a
tion r
e
quir
e
s
r
e
pro
d
u
c
ing th
e
p
a
r
e
nt fish
c
ross
e
s to pro
d
u
ce
high
-
qu
a
lity hy
b
ri
d
offspring
.
T
his hy
b
ri
d
iz
a
tion initi
a
tiv
e
involv
e
s
c
ross
-
b
r
eed
ing
Ma
j
a
l
a
y
a
str
a
in gol
d
fish
(
da
rk gr
ee
n
)
or koi fish with
S
i
N
yony
a
str
a
in gol
d
fish
(
y
e
llow
)
or
c
om
e
t fish using
a
rtifi
c
i
a
l f
e
rtiliz
a
tion t
ec
hniqu
e
s
.
T
h
e a
rtifi
c
i
a
l
f
e
rtiliz
a
tion pro
ce
ss is impl
e
m
e
nt
ed acc
or
d
ing to th
e
outlin
ed
hy
b
ri
d
iz
a
tion progr
a
m
.
T
h
e
progr
a
m
de
t
a
ils in
c
lu
de c
om
b
in
a
tions su
c
h
a
s
Ma
j
a
l
a
y
a
with
T
h
e Lad
y
,
T
h
e Lad
y with
Ma
j
a
l
a
y
a
,
Ma
j
a
l
a
y
a
with
Ma
j
a
l
a
y
a
,
a
n
d T
h
e Lad
y with
C
om
e
t
,
a
s w
e
ll
a
s
T
h
e Lad
y with
K
oi
.
2
.
5
P
urpos
e
of
H
y
b
ri
d
iz
a
tion
T
h
e
purpos
e
of
c
on
d
u
c
ting hy
b
ri
d
iz
a
tion on gol
d
fish or koi is to
e
n
ab
l
e
stu
de
nts to impl
e
m
e
nt
a
hy
b
ri
d
iz
a
tion progr
a
m th
a
t
ca
ptur
e
s
d
omin
a
nt g
e
n
e
ti
c
v
a
ri
a
nts if th
e
p
a
r
e
nt s
e
l
ec
tion progr
a
m
d
o
e
s not yi
e
l
d
th
e de
sir
ed
out
c
om
e
s
,
p
a
rti
c
ul
a
rly wh
e
n th
e
st
a
n
da
r
d de
vi
a
tion
(
SD
)
a
n
d
c
o
e
ffi
c
i
e
nt of v
a
ri
a
tion
(
CV
)
v
a
lu
e
s for
a
tr
a
it
a
r
e
r
e
l
a
tiv
e
ly low
.
H
y
b
ri
d
iz
a
tion
a
ims to
e
nh
a
n
ce
s
eed
qu
a
lity
b
y improving growth r
a
t
e
s
,
de
l
a
ying s
e
xu
a
l m
a
turity for optim
a
l growth
,
a
n
d b
oosting r
e
sist
a
n
ce
to
d
is
ea
s
e
s
a
n
d ad
v
e
rs
e c
on
d
itions
.
T
his
e
nh
a
n
ce
m
e
nt
a
ris
e
s from th
e
r
e
sulting tr
a
its
.
I
ntr
a
sp
ec
ifi
c
hy
b
ri
d
iz
a
tion
,
a
s
ca
t
e
goriz
ed b
y
d
iff
e
r
e
nt
typ
e
s
,
ca
n
be
within th
e
s
a
m
e
sp
ec
i
e
s
,
be
tw
ee
n sp
ec
i
e
s of th
e
s
a
m
e
g
e
nus
,
or
be
tw
ee
n
d
iff
e
r
e
nt g
e
n
e
r
a
.
CHAPTER III
MATERIALS AND METHODS
3
.
2
T
ools
a
n
d Ma
t
e
ri
a
ls
I
n th
e
pr
ac
ti
ca
l
ac
tiviti
e
s
,
th
e
following tools
a
n
d
m
a
t
e
ri
a
ls
a
r
e
utiliz
ed
.
T
h
e
pr
ac
ti
ca
l tools in
c
lu
de a
1
m
³
fi
be
rgl
a
ss t
a
nk for
b
r
eed
ing
b
roo
d
fish
,
a
n
ae
r
a
tor for oxyg
e
n supply
,
a
n
e
t for
ca
pturing p
a
r
e
nt fish
,
rul
e
rs
a
n
d
s
ca
l
e
s for m
ea
suring fish l
e
ngth
a
n
d
w
e
ight
,
sm
a
ll
a
n
d
l
a
rg
e
a
qu
a
riums for r
ea
ring l
a
rv
ae a
n
d
s
eed
fish
,
h
ea
t
e
rs for m
a
int
a
ining w
a
t
e
r
t
e
mp
e
r
a
tur
e
,
a
n
d a b
u
c
k
e
t for t
e
mpor
a
rily hol
d
ing fish
d
uring inj
ec
tion
a
n
d
f
e
rtiliz
a
tion
.
T
h
e
pr
ac
ti
ca
l m
a
t
e
ri
a
ls
c
onsist of
O
v
a
prim hormon
e
to
acce
l
e
r
a
t
e
ovul
a
tion
a
n
d
sp
a
wning
,
1
ml
a
n
d
2
ml syring
e
s for hormon
e
inj
ec
tions
,
syring
e
s
a
n
d
por
ce
l
a
in
c
ups for
c
oll
ec
ting fish sp
e
rm
a
n
d e
ggs
,
Na
upli
A
rt
e
mi
a a
n
d T
u
b
if
e
x
a
s foo
d
for l
a
rv
ae a
n
d
s
eed
fish
,
p
e
ll
e
ts with
30
%
prot
e
in for f
eed
ing p
a
r
e
nt fish
,
Ma
l
ac
hyt
e
gr
ee
n to pr
e
v
e
nt mol
d
on
e
ggs
,
physiologi
ca
l
NaC
l solution for thinning fish sp
e
rm
,
f
e
m
a
l
e C
om
e
t
fish
a
n
d
m
a
l
e K
oi fish
a
s t
e
st s
a
mpl
e
s
,
G
ol
d
fish r
e
pr
e
s
e
nting thr
ee
str
a
ins
,
a
n
d A
qu
ade
s for
d
iluting
O
v
a
prim
.
3
.
3
P
r
ac
ti
c
um
S
t
a
g
e
s
P
r
ac
ti
ca
l pro
ced
ur
e
s for hy
b
ri
d
iz
a
tion of gol
d
fish
a
n
d
nil
e
m fish go
12
through th
e
pr
e
p
a
r
a
tion
a
n
d
impl
e
m
e
nt
a
tion st
a
g
e
s
.
3
.
3
.
1
P
r
e
p
a
r
a
tion
S
t
a
g
e
T
h
e
pr
e
p
a
r
a
tion pro
ce
ss for th
e
hy
b
ri
d
iz
a
tion pr
ac
ti
c
um involv
e
s
s
e
v
e
r
a
l st
e
ps
:
first
,
c
l
ea
n th
e a
qu
a
rium thoroughly
.
Ne
xt
,
s
e
t up th
e
ae
r
a
tion syst
e
m to
e
nsur
e
it op
e
r
a
t
e
s
c
orr
ec
tly
a
n
d
fill th
e a
qu
a
rium with
w
a
t
e
r up to thr
ee
-
qu
a
rt
e
rs of its
ca
p
ac
ity
.
C
hoos
e
th
e b
roo
d
sto
c
k for th
e
pr
ac
ti
c
um
,
a
n
d
th
e
n s
e
p
a
r
a
t
e
th
e
m
a
l
e a
n
d
f
e
m
a
l
e
p
a
r
e
nts
.
I
nj
ec
t th
e
f
e
m
a
l
e a
n
d
m
a
l
e b
roo
d
sto
c
k with
O
v
a
prim
a
t
d
os
a
g
e
s of
0
.
4
ml
/
kg
a
n
d
0
.
1
ml
/
kg
,
r
e
sp
ec
tiv
e
ly
.
P
ro
ceed
with sp
e
rm stripping for th
e
m
a
l
e
p
a
r
e
nt
a
n
d e
gg stripping for th
e
f
e
m
a
l
e
p
a
r
e
nt on
ce
ovul
a
tion o
cc
urs
.
3
.
3
.
2
I
mpl
e
m
e
nt
a
tion
S
t
a
g
e
T
h
e
hy
b
ri
d
iz
a
tion pr
ac
ti
c
um involv
e
s s
e
v
e
r
a
l st
e
ps
:
first
,
th
e
sp
e
rm
pro
d
u
ced b
y th
e
m
a
l
e
p
a
r
e
nt is
d
ilut
ed
with
a
physiologi
ca
l
NaC
l solution
b
y
a
n
a
ssist
a
nt
.
T
his sp
e
rm is th
e
n pl
aced
into
a
p
e
tri
d
ish or
bea
k
e
r
b
y
th
e a
ssist
a
nt
.
Ne
xt
,
stripping is p
e
rform
ed
on f
e
m
a
l
e c
om
e
t fish
,
a
n
d
th
e
ir
e
ggs
a
r
e c
oll
ec
t
ed a
n
d
stor
ed
in p
e
tri
d
ish
e
s
.
T
h
e e
ggs
a
r
e
th
e
n f
e
rtiliz
ed
with koi fish sp
e
rm
.
F
ollowing f
e
rtiliz
a
tion
,
th
e e
ggs
a
r
e
monitor
ed
,
a
n
d
th
e
num
be
r of f
e
rtiliz
ed e
ggs
a
n
d
f
e
rtiliz
a
tion r
a
t
e
s
a
r
e
r
ec
or
ded
.
T
h
e
f
e
rtiliz
ed e
ggs
a
r
e
th
e
n
d
istri
b
ut
ed
into th
e
provi
ded a
qu
a
rium
.
A
ft
e
r two
da
ys
,
th
e e
ggs
a
r
e e
x
a
min
ed
,
a
n
d
h
a
t
c
hing r
a
t
e
s
a
n
d
surviv
a
l r
a
t
e
s
a
r
e
ca
l
c
ul
a
t
ed
.
13
3
.
4
Me
tho
d
T
h
e
m
e
tho
d
us
ed
in this pr
ac
ti
c
um is
e
xp
e
rim
e
nt
a
l using s
e
v
e
r
a
l
tr
ea
tm
e
nts
.
T
h
e
tr
ea
tm
e
nts giv
e
n in
c
lu
de
inj
ec
tion of th
e
hormon
e
ov
a
prim
,
stripping
a
n
d d
ilution
.
3
.
5
Ob
s
e
rv
ed Pa
r
a
m
e
t
e
rs
3
.
5
.
1
FR ca
l
c
ul
a
tion
T
h
e
f
e
rtiliz
a
tion r
a
t
e
(
FR
)
r
e
pr
e
s
e
nts th
e e
xt
e
nt of
e
gg f
e
rtiliz
a
tion
.
Ob
s
e
rv
a
tions of this f
e
rtiliz
a
tion r
a
t
e
w
e
r
e c
on
d
u
c
t
ed a
ft
e
r
c
ompl
e
ting th
e
hy
b
ri
d
iz
a
tion pro
ce
ss
.
Fe
rtiliz
ed e
ggs
e
xhi
b
it
a b
right
c
olor
,
wh
e
r
ea
s
unf
e
rtiliz
ed e
ggs
a
pp
ea
r
d
ull
.
T
h
e FR
is
c
omput
ed ba
s
ed
on th
e e
ggs
pr
e
s
e
nt in th
e a
qu
a
rium
.
T
o
de
t
e
rmin
e
th
e
f
e
rtiliz
a
tion
de
gr
ee
of fish
e
ggs
,
a
sp
ec
ifi
c
formul
a
is us
ed
,
wh
e
r
e FR de
not
e
s th
e
p
e
r
ce
nt
a
g
e
of f
e
rtiliz
ed
e
ggs
,
P
is th
e
tot
a
l num
be
r of s
a
mpl
e e
ggs
,
a
n
d P
o r
e
pr
e
s
e
nts th
e
num
be
r
of f
e
rtiliz
ed e
ggs
.
3
.
5
.
2
HR ca
l
c
ul
a
tion
T
h
e
h
a
t
c
hing r
a
t
e
(
HR
)
r
e
f
e
rs to th
e e
xt
e
nt to whi
c
h
e
ggs
su
cce
ssfully h
a
t
c
h
.
T
his is
a
ss
e
ss
ed b
y monitoring wh
e
n
e
m
b
ryos
de
v
e
lop
into l
a
rv
ae
.
T
h
e HR
is
ca
l
c
ul
a
t
ed ba
s
ed
on th
e
num
be
r of
e
ggs th
a
t h
a
t
c
h
14
within th
e a
qu
a
rium
.
T
o
de
t
e
rmin
e
th
e
h
a
t
c
hing
de
gr
ee
of fish
e
ggs
,
a
sp
ec
ifi
c
formul
a ca
n
be
us
ed
wh
e
r
e HR
r
e
pr
e
s
e
nts th
e
h
a
t
c
hing r
a
t
e
,
P
t
de
not
e
s th
e
num
be
r of
e
ggs th
a
t h
a
v
e
h
a
t
c
h
ed
,
a
n
d P
o in
d
i
ca
t
e
s th
e
num
be
r of f
e
rtiliz
ed e
ggs
.
3
.
5
.
3
SR ca
l
c
ul
a
tion
S
urviv
a
l r
a
t
e
(
SR
)
m
ea
sur
e
s th
e e
xt
e
nt of fish surviv
a
l
.
E
v
a
lu
a
tions of fish surviv
a
l w
e
r
e c
on
d
u
c
t
ed
sol
e
ly for th
e
pro
ce
ss
e
s of
gynog
e
n
e
sis
,
hy
b
ri
d
iz
a
tion
,
a
n
d
triploi
d
iz
a
tion on
ce
th
e
fish l
a
rv
ae
r
eac
h
ed
s
e
v
e
n
da
ys of
a
g
e
.
3
.
6
Da
t
a a
n
a
lysis
FR ca
l
c
ul
a
tion
FR
(
%
) =
P
o
P
x
100
%
FR
(
%
) =
155
305
x
100
%
FR
(
%
) =
50
.
80
%
HR ca
l
c
ul
a
tion
15
HR
(
%
) =
P
t
P
ox
100
%
HR
(
%
) =
0
155
x
100
%
HR
(
%
) =
0
%
SR ca
l
c
ul
a
tion
SR
(
%
) =
N
t
N
ox
100
%
SR
(
%
) =
0
0
x
100
%
SR
(
%
) =
0
%
CHAPTER IV
RESULTS AND DISCUSSION
4
.
1
C
l
a
ss
Ob
s
e
rv
a
tion
Re
sults
Ba
s
ed
on th
e
r
e
sults of o
b
s
e
rv
a
tions r
e
g
a
r
d
ing hy
b
ri
d
iz
a
tion
ca
rri
ed
out in
c
l
a
ss
C
,
th
e
following r
e
sults w
e
r
e
o
b
t
a
in
ed
:
4
.
1
.
1
FR
Ba
s
ed
on th
e
o
b
s
e
rv
a
tion r
e
sults of f
e
rtiliz
a
tion r
a
t
e
s
(
FR
),
th
e
hy
b
ri
d
iz
a
tion
c
on
d
u
c
t
ed
in
C
l
a
ss
C
yi
e
l
ded
th
e
following out
c
om
e
s
:
I
n th
e
first t
e
st
,
hy
b
ri
d
iz
a
tion r
e
sults w
e
r
e
57
%
,
34
.
55
%
,
a
n
d
52
%
.
F
or th
e
s
ec
on
d
t
e
st
,
th
e
r
a
t
e
s w
e
r
e
50
.
80
%
,
62
%
,
a
n
d
68
%
.
I
n th
e
thir
d
t
e
st
,
th
e
r
a
t
e
s w
e
r
e
79
.
16
%
,
73
%
,
a
n
d
72
%
.
T
h
e a
v
e
r
a
g
e FR
for th
e
s
e
t
e
sts w
a
s
64
.
74
%
,
58
.
13
%
,
a
n
d
60
.
5
%
,
r
e
sp
ec
tiv
e
ly
.
4
.
1
.
2
HR
T
h
e
fin
d
ings from th
e HR
(
h
ac
king r
a
t
e
)
hy
b
ri
d
iz
a
tion o
b
s
e
rv
a
tions
c
on
d
u
c
t
ed
in
C
l
a
ss
C
r
e
v
ea
l
ed
th
e
following r
e
sults
:
Tab
l
e
5
shows th
e HR
da
t
a
from th
e
hy
b
ri
d
iz
a
tion in
C
l
a
ss
C
.
F
or
eac
h tr
ea
tm
e
nt t
e
st
,
th
e
hy
b
ri
d
iz
a
tion r
a
t
e
s w
e
r
e c
onsist
e
ntly
0
%
ac
ross
a
ll thr
ee
t
e
sts
,
r
e
sulting in
a
n
a
v
e
r
a
g
e
of
0
%
.
17
4
.
1
.
3
SR
Ob
s
e
rv
a
tions of th
e
surviv
a
l r
a
t
e
(
SR
)
from hy
b
ri
d
iz
a
tion
e
xp
e
rim
e
nts in
c
l
a
ss
C
yi
e
l
ded
th
e
following r
e
sults
:
th
e SR
p
e
r
ce
nt
a
g
e
s
for hy
b
ri
d
iz
a
tion t
e
sts
ac
ross
a
ll tr
ea
tm
e
nts w
e
r
e c
onsist
e
ntly
0
%
.
F
or
w
a
t
e
r qu
a
lity m
ea
sur
e
m
e
nts
d
uring th
e
hy
b
ri
d
iz
a
tion pr
ac
ti
c
um in th
e
s
a
m
e c
l
a
ss
,
th
e da
t
a
show
ed
th
a
t th
e
t
e
mp
e
r
a
tur
e
w
a
s g
e
n
e
r
a
lly
a
roun
d
26
°
C
with slight v
a
ri
a
tions
,
p
H
l
e
v
e
ls w
e
r
e c
onsist
e
ntly
7
,
a
n
d d
issolv
ed
oxyg
e
n
(
DO
)
r
a
ng
ed
from
4
.
5
to
4
.
7
mg
/
L ac
ross
d
iff
e
r
e
nt groups
.
4
.
2
G
roup
Ob
s
e
rv
a
tion
Re
sults
Acc
or
d
ing to th
e
fin
d
ings from th
e
o
b
s
e
rv
a
tions of hy
b
ri
d
iz
a
tion
c
on
d
u
c
t
ed b
y
G
roup
2
,
th
e
r
e
sults
a
r
e
summ
a
riz
ed a
s follows
:
Tab
l
e
8
pr
e
s
e
nts th
e da
t
a
on
G
roup
2
'
s hy
b
ri
d
iz
a
tion out
c
om
e
s
,
showing
a FR
p
e
r
ce
nt
a
g
e
of
82
.
08
%
,
with
b
oth
HR a
n
d SR
p
e
r
ce
nt
a
g
e
s
a
t
0
%
.
4
.
2
.
1
FR
FR
(
%
) =
P
o
P
x
100
%
FR
(
%
) =
155
305
x
100
%
FR
(
%
) =
50
.
80
%
4
.
2
.
2
HR
18
HR
(
%
) =
P
t
P
ox
100
%
HR
(
%
) =
0
305
x
100
%
HR
(
%
) =
0
%
4
.
2
.
3
SR
SR
(
%
) =
N
t
N
ox
100
%
SR
(
%
) =
0
0
x
100
%
SR
(
%
) =
0
%
4
.
2
.
4
Wa
t
e
r
Q
u
a
lity
Mea
sur
e
m
e
nt
Acc
or
d
ing to th
e
o
b
s
e
rv
a
tions
c
on
d
u
c
t
ed b
y
G
roup
2
on
hy
b
ri
d
iz
a
tion
,
th
e
w
a
t
e
r qu
a
lity w
a
s m
ea
sur
ed a
s follows
:
G
roup
2
r
ec
or
ded
th
e
t
e
mp
e
r
a
tur
e
,
p
H
,
a
n
d d
issolv
ed
oxyg
e
n
(
DO
)
l
e
v
e
ls
.
4
.
3
G
roup
D
is
c
ussion
Ba
s
ed
on th
e
r
e
sults of o
b
s
e
rv
a
tions r
e
g
a
r
d
ing hy
b
ri
d
iz
a
tion
ca
rri
ed
out
b
y group
2
,
th
e FR
,
HR a
n
d SR
v
a
lu
e
s o
b
t
a
in
ed ca
n
be e
xpl
a
in
ed a
s
follows
:
19
4
.
3
.
1
FR
Fe
rtiliz
a
tion r
a
t
e
(
FR
)
m
ea
sur
e
s th
e e
xt
e
nt of f
e
rtiliz
a
tion in
e
ggs
.
Ob
s
e
rv
a
tions of
FR
w
e
r
e c
on
d
u
c
t
ed a
ft
e
r th
e
hy
b
ri
d
iz
a
tion pro
ce
ss
,
d
istinguishing f
e
rtiliz
ed e
ggs
b
y th
e
ir
b
right
c
olor
a
n
d dead e
ggs
b
y th
e
ir
d
ull
c
olor
.
FR
is
ca
l
c
ul
a
t
ed b
y
e
x
a
mining th
e
num
be
r of
e
ggs in th
e
a
qu
a
rium
a
n
d
i
de
ntifying thos
e
th
a
t h
a
v
e
h
a
t
c
h
ed
.
F
or inst
a
n
ce
,
G
roup
2
o
b
s
e
rv
ed
305
e
gg s
a
mpl
e
s with
155
be
ing f
e
rtiliz
ed
,
r
e
sulting in
a
n
FR
of
50
.
80
%
.
T
his v
a
lu
e
is
c
onsi
de
r
ed
low
,
a
s
a
f
e
rtiliz
a
tion r
a
t
e ab
ov
e
70
%
is
typi
ca
lly r
e
g
a
r
ded a
s high
.
Se
v
e
r
a
l f
ac
tors influ
e
n
ce
th
e
low
FR
in koi fish
,
in
c
lu
d
ing th
e
su
cce
ss of sp
e
rm p
e
n
e
tr
a
tion into th
e e
gg
ce
ll
'
s mi
c
ropyl
e
,
th
e
siz
e d
is
c
r
e
p
a
n
c
y
be
tw
ee
n
e
gg
a
n
d
sp
e
rm from
d
iff
e
r
e
nt sp
ec
i
e
s
,
a
n
d
th
e
physiologi
ca
l
c
on
d
ition of
b
oth
e
ggs
a
n
d
sp
e
rm
.
Ke
y influ
e
n
c
ing
f
ac
tors in
c
lu
de
g
e
n
e
ti
c
,
physiologi
ca
l
(
e
.
g
.,
sp
e
rm qu
a
lity
),
a
n
d
morphologi
ca
l
a
sp
ec
ts
(
e
.
g
.,
a
lignm
e
nt of th
e e
gg
'
s mi
c
ropyl
e
with th
e
sp
e
rm h
ead
).
4
.
3
.
1
HR
Ha
t
c
hing r
a
t
e
(
HR
)
r
e
f
e
rs to th
e
proportion of
e
ggs th
a
t su
cce
ssfully h
a
t
c
h
.
T
o
de
t
e
rmin
e HR
,
th
e
num
be
r of
e
ggs th
a
t h
a
t
c
h into l
a
rv
ae
is o
b
s
e
rv
ed
.
I
n this
ca
s
e
,
HR
is
ca
l
c
ul
a
t
ed ba
s
ed
on th
e e
ggs th
a
t h
a
t
c
h
ed
in th
e
a
qu
a
rium
.
O
ut of
305
e
ggs
,
155
w
e
r
e
f
e
rtiliz
ed
,
r
e
sulting in
a Fe
rtiliz
a
tion
Ra
t
e
(
FR
)
of
50
.
80
%
.
HR
is
c
omput
ed b
y
d
ivi
d
ing th
e
num
be
r of h
a
t
c
h
ed
20
e
ggs
b
y th
e
num
be
r of f
e
rtiliz
ed e
ggs
.
G
roup
2
'
s h
a
t
c
hing r
a
t
e
w
a
s
0
%
,
in
d
i
ca
ting th
a
t no
e
ggs h
a
t
c
h
ed
,
whi
c
h is
c
onsi
de
r
ed
v
e
ry low
a
n
d
sugg
e
sts th
a
t th
e e
ggs
d
i
d
not surviv
e
.
T
ypi
ca
lly
,
fish h
a
t
c
hing r
a
t
e
s r
a
ng
e
from
50
to
80
%
.
Fac
tors influ
e
n
c
ing h
a
t
c
hing in
c
lu
de
t
e
mp
e
r
a
tur
e a
n
d
light int
e
nsity
,
whi
c
h
a
ff
ec
t th
e
mov
e
m
e
nt n
ece
ss
a
ry for h
a
t
c
hing
.
A
low
h
a
t
c
hing r
a
t
e ca
n
be a
ttri
b
ut
ed
to v
a
rious f
ac
tors
,
su
c
h
a
s zygot
e
mort
a
lity
d
u
e
to insuffi
c
i
e
nt
d
issolv
ed
oxyg
e
n
,
improp
e
r t
e
mp
e
r
a
tur
e
s
,
m
ec
h
a
ni
ca
l
d
istur
ba
n
ce
s lik
e
sho
c
k or fri
c
tion
,
or p
a
r
a
sit
e
inf
e
st
a
tions
from
bac
t
e
ri
a
,
fungi
,
or ins
ec
t l
a
rv
ae
.
4
.
3
.
1
SR
S
urviv
a
l r
a
t
e
(
SR
)
r
e
f
e
rs to th
e
proportion of fish th
a
t r
e
m
a
in
a
liv
e
.
I
n this
c
ont
e
xt
,
SR
w
a
s
a
ss
e
ss
ed
sp
ec
ifi
ca
lly for gynog
e
n
e
sis
,
hy
b
ri
d
iz
a
tion
,
a
n
d
triploi
d
iz
a
tion pro
ce
ss
e
s st
a
rting from wh
e
n fish l
a
rv
ae
w
e
r
e
s
e
v
e
n
da
ys
ol
d
.
I
t is
ca
l
c
ul
a
t
ed a
s th
e
p
e
r
ce
nt
a
g
e
of fish surviving
a
t th
e e
n
d
of
r
ea
ring
c
omp
a
r
ed
to th
e
tot
a
l num
be
r of fish initi
a
lly pl
aced
in th
e
r
ea
ring
e
nvironm
e
nt
.
Acc
or
d
ing to
da
t
a
from group
2
,
a
n
SR
of
0
%
w
a
s r
ec
or
ded
,
whi
c
h o
cc
urr
ed beca
us
e
th
e
s
a
mpl
e
s us
ed
w
e
r
e
h
a
t
c
h
ed e
ggs
a
n
d d
i
d
not
r
e
fl
ec
t th
e
num
be
r of fish
.
Add
ition
a
lly
,
th
e
fin
a
l num
be
r of fish
a
t th
e e
n
d
of th
e
pr
ac
ti
ca
l s
e
ssion w
a
s unknown
,
l
ead
ing to
a
n
SR
of
0
%
.
F
or nil
e
m
fish
,
this
SR
in
d
i
ca
t
e
s
a
n
e
xtr
e
m
e
ly low surviv
a
l r
a
t
e
,
e
ss
e
nti
a
lly non
e
,
d
u
e
to th
e ab
norm
a
lity of l
a
rv
ae
r
e
sulting from th
e
hy
b
ri
d
iz
a
tion pro
ce
ss
,
21
whi
c
h imp
a
irs th
e
ir
ab
ility to
c
onsum
e
foo
d e
ff
ec
tiv
e
ly
.
T
his
c
on
c
lusion is
support
ed b
y th
e
notion th
a
t l
a
rv
a
l mov
e
m
e
nt
a
n
d acce
ss to goo
d
foo
d
a
r
e c
riti
ca
l for surviv
a
l
.
CHAPTER V
CONCLUSIONS AND RECOMMENDATIONS
5
.
1
C
on
c
lusion
F
ollowing th
e
o
b
s
e
rv
a
tion
b
y group
2
a
ft
e
r
e
gg f
e
rtiliz
a
tion in th
e
hy
b
ri
d
iz
a
tion pro
ce
ss
,
th
e
s
a
mpl
e
siz
e
of
e
ggs
(
P
)
w
a
s
305
,
with
155
f
e
rtiliz
ed e
ggs
(
P
o
),
r
e
sulting in
a
f
e
rtiliz
a
tion r
a
t
e
(
FR
)
of
50
.
80
%
.
H
ow
e
v
e
r
,
no
e
ggs h
a
t
c
h
ed
(
P
t
),
l
ead
ing to
a
h
a
t
c
hing r
a
t
e
(
HR
)
of
0
%
a
n
d
a
surviv
a
l r
a
t
e
(
SR
)
of
0
%
.
T
h
e
low
FR
v
a
lu
e
is
a
ttri
b
ut
ed
to g
e
n
e
ti
c a
n
d
physiologi
ca
l f
ac
tors
,
su
c
h
a
s th
e
qu
a
lity of th
e
m
a
l
e
p
a
r
e
nt
'
s sp
e
rm
,
a
s
w
e
ll
a
s stru
c
tur
a
l
a
n
d
morphologi
ca
l f
ac
tors
.
T
h
e
f
a
ilur
e
in
e
gg h
a
t
c
hing
a
n
d
surviv
a
l r
a
t
e
s w
a
s
d
u
e
to l
a
rv
a
l mort
a
lity
,
influ
e
n
ced b
y int
e
rn
a
l
f
ac
tors lik
e
g
e
n
e
s
a
n
d
hormon
e
s
,
a
n
d e
xt
e
rn
a
l f
ac
tors su
c
h
a
s
e
nvironm
e
nt
a
l
c
on
d
itions
,
d
issolv
ed
oxyg
e
n
,
w
a
t
e
r qu
a
lity
,
p
H
l
e
v
e
ls
,
a
n
d
in
ade
qu
a
t
e ae
r
a
tion
.
l
.
5
.
2
S
ugg
e
stions
P
r
ac
tition
e
rs g
a
in
a dee
p
e
r un
de
rst
a
n
d
ing of hy
b
ri
d
iz
a
tion
m
a
t
e
ri
a
ls
be
for
e e
ng
a
ging in pr
ac
ti
ca
l hy
b
ri
d
iz
a
tion
e
x
e
r
c
is
e
s to
e
nsur
e
th
e ac
tiviti
e
s pro
ceed
without issu
e
s or
e
rrors
.
Add
ition
a
lly
,
th
e
y fo
c
us
c
los
e
ly on th
e a
ssist
a
nt
'
s
e
xpl
a
n
a
tions of th
e
pr
ac
ti
c
um pro
ced
ur
e
s to
e
nsur
e
th
e
pro
ce
ss is
c
on
d
u
c
t
ed
prop
e
rly
a
n
d e
ff
ec
tiv
e
ly
,
thus minimizing
pot
e
nti
a
l f
ac
tors th
a
t
c
oul
d
l
ead
to f
a
ilur
e
.
23
