1
ARIZONA STATE UNIVERSITY
"
I
sol
a
tion of
Sa
lt
a
n
d Hea
vy
Me
t
a
l
T
ol
e
r
a
nt
E
n
d
ophyti
c
Bac
t
e
ri
a a
s
B
iopriming
A
g
e
nts for
R
i
ce Seed
"
JEFF M
.
2
BACKGROUND
S
oils with high l
e
v
e
ls of s
a
lt
a
n
d
h
ea
vy m
e
t
a
ls r
e
m
a
in
a
signifi
ca
nt
c
h
a
ll
e
ng
e
in
a
gri
c
ultur
e
.
T
his issu
e
is p
a
rtly
d
u
e
to th
e
c
onv
e
rsion of
a
gri
c
ultur
a
l l
a
n
d
into m
a
rgin
a
l l
a
n
d
,
su
c
h
a
s s
a
lin
e
soils
.
Sa
lin
e
soil is
c
h
a
r
ac
t
e
riz
ed b
y so
d
ium
c
ont
e
nt
e
x
ceed
ing
c
riti
ca
l or
pl
a
nt tol
e
r
a
n
ce
thr
e
shol
d
s
.
T
h
e acc
umul
a
tion of s
a
lt
a
n
d
h
ea
vy m
e
t
a
ls
in th
e
soil
ca
n r
ed
u
ce a
gri
c
ultur
a
l pro
d
u
c
tivity
,
su
c
h
a
s in ri
ce
c
ultiv
a
tion
.
T
h
e
limit
ed a
v
a
il
ab
ility of
a
r
ab
l
e
l
a
n
d
must
be ba
l
a
n
ced
with th
e
int
e
nsifi
ca
tion
a
n
d e
xt
e
nsifi
ca
tion of l
a
n
d
us
e
for ri
ce
c
ultiv
a
tion
,
whi
c
h s
e
rv
e
s
a
s
a
st
a
pl
e
foo
d
for th
e I
n
d
on
e
si
a
n popul
a
tion
.
R
i
ce
p
add
i
e
s
e
xp
e
ri
e
n
c
ing in
c
r
ea
s
ed
s
a
linity
a
r
e
oft
e
n thos
e
lo
ca
t
ed
n
ea
r
c
o
a
st
a
l
a
r
ea
s or with in
ade
qu
a
t
e
w
a
t
e
r
c
h
a
nn
e
ls
.
T
his p
a
ss
a
g
e
illustr
a
t
e
s th
e
gr
ea
tn
e
ss of
A
ll
a
h in r
e
viving
dead
or
ba
rr
e
n l
a
n
d
without lif
e b
y s
e
n
d
ing
d
own r
a
in
.
I
t is
a
lso
A
ll
a
h who
ca
us
e
s gr
a
ins lik
e
ri
ce a
n
d
wh
ea
t to grow
,
provi
d
ing st
a
pl
e
foo
d
for hum
a
ns on th
e
r
e
viv
ed
or f
e
rtil
e
l
a
n
d
.
P
oor irrig
a
tion m
a
n
a
g
e
m
e
nt
,
high
e
v
a
por
a
tion
r
a
t
e
s
,
a
n
d
low r
a
inf
a
ll
ca
n
a
ll
c
ontri
b
ut
e
to s
a
linity issu
e
s in
a
n
a
r
ea
.
Sa
linity r
e
f
e
rs to th
e
l
e
v
e
l of s
a
lt or solu
b
l
e
s
a
lts
(
NaC
l
,
Na
2
CO
3
,
Na
2
SO
4
)
in w
a
t
e
r
.
H
igh soil s
a
linity h
a
s
a
n
e
g
a
tiv
e
imp
ac
t on pl
a
nt
growth
a
n
d de
v
e
lopm
e
nt
,
ca
using pl
a
nts to struggl
e
in
ab
sor
b
ing
3
w
a
t
e
r
,
l
ead
ing to physiologi
ca
l
d
rought
d
u
e
to in
c
r
ea
s
ed
osmoti
c
pot
e
nti
a
l of th
e
soil solution
.
E
x
ce
ssiv
e
s
a
lt
c
ont
e
nt
a
lso r
e
sults in high
soil r
e
sist
a
n
ce
to p
e
n
e
tr
a
tion
,
e
l
e
v
a
t
ed c
on
ce
ntr
a
tions of
ca
tions su
c
h
a
s
ca
l
c
ium
(
Ca
),
m
a
gn
e
sium
(
M
g
),
so
d
ium
(
Na
),
a
n
d a
high
E
x
ce
ssiv
e
E
x
c
h
a
ng
eab
l
e S
o
d
ium
Pe
r
ce
nt
a
g
e
(
ESP
).
S
o
d
ium
c
hlori
de
(
NaC
l
)
is th
e
prim
a
ry s
a
lt foun
d
in s
a
lin
e
soils
.
Eac
h pl
a
nt sp
ec
i
e
s h
a
s
a d
iff
e
r
e
nt r
e
spons
e
to s
a
lt l
e
v
e
ls
.
F
or ri
ce
pl
a
nts
,
soils with s
a
lt
c
on
ce
ntr
a
tions
e
x
ceed
ing
a
n
e
l
ec
tri
ca
l
c
on
d
u
c
tivity
(
EC
)
of
4
dS
m
-
1
ca
n l
ead
to r
ed
u
ced c
rop yi
e
l
d
s
.
T
his is
ba
s
ed
on o
b
s
e
rv
a
tions th
a
t most
c
rops
e
xp
e
ri
e
n
ce dec
r
ea
s
ed
yi
e
l
d
s
a
t
s
a
linity l
e
v
e
ls
ab
ov
e
4
dS
m
-
1
,
whi
c
h
c
orr
e
spon
d
s to
40
m
M NaC
l
a
n
d
a
n osmoti
c
pr
e
ssur
e
of
0
.
2
MPa
.
H
igh s
a
lt
c
on
ce
ntr
a
tions
ca
n
ca
us
e
ion
toxi
c
ity
,
whi
c
h m
a
y
da
m
a
g
e
th
e c
h
e
mi
ca
l
b
on
d
s in pl
a
nt
c
hlorophyll
,
r
e
sulting in r
ed
u
ced c
hlorophyll l
e
v
e
ls
.
E
l
e
v
a
t
ed NaC
l l
e
v
e
ls in th
e
soil
a
lso l
ead
to nutri
e
nt im
ba
l
a
n
ce
s
,
oxi
da
tiv
e da
m
a
g
e
to pl
a
nt tissu
e
s
,
hin
d
r
a
n
ce
in th
e
synth
e
sis of photosynth
e
ti
c
pigm
e
nts
,
a
n
d d
isruptions
in th
e
photosynth
e
sis pro
ce
ss
,
ultim
a
t
e
ly imp
a
iring pl
a
nt m
e
t
ab
olism
.
I
n
add
ition to s
a
lt
-
l
ade
n soils
,
h
ea
vy m
e
t
a
ls lik
e
l
ead
(
Pb
)
a
n
d
cad
mium
(
Cd
)
ca
n
a
lso in
c
r
ea
s
e
soil toxi
c
ity
,
a
ff
ec
ting pl
a
nts
.
Lead
is
us
ed
in
ba
tt
e
ry pro
d
u
c
tion
,
c
osm
e
ti
c
s
,
m
e
t
a
l pro
d
u
c
ts su
c
h
a
s
a
mmunition
a
n
d
pip
e
s
,
whil
e cad
mium is us
ed
in
ba
tt
e
ri
e
s
,
pigm
e
nts
,
pl
a
sti
c
s
,
a
n
d
m
e
t
a
l
c
o
a
tings
,
a
n
d
is wi
de
ly
e
mploy
ed
in
e
l
ec
tropl
a
ting
.
H
um
a
n
ac
tiviti
e
s su
c
h
a
s mining
,
m
a
nuf
ac
turing
,
a
n
d
fossil fu
e
l
c
om
b
ustion h
a
v
e
r
e
sult
ed
in th
e acc
umul
a
tion of
Pb
,
Cd
,
a
n
d
th
e
ir
c
ompoun
d
s in th
e e
nvironm
e
nt
,
in
c
lu
d
ing
a
ir
,
w
a
t
e
r
,
a
n
d
soil
.
Pb a
n
d
4
Cd a
r
e
highly h
a
z
a
r
d
ous h
ea
vy m
e
t
a
ls
beca
us
e
th
e
y
a
r
e
non
-
de
gr
adab
l
e b
y living org
a
nisms
a
n
d ca
n
acc
umul
a
t
e
in th
e e
nvironm
e
nt
,
p
a
rti
c
ul
a
rly s
e
ttling in w
a
t
e
r
b
o
d
i
e
s
'
s
ed
im
e
nts
a
n
d
forming
c
ompl
e
x
e
s
with org
a
ni
c a
n
d
inorg
a
ni
c
m
a
t
e
ri
a
ls
.
T
h
e
m
a
ximum
a
llow
ab
l
e
limits for
h
ea
vy m
e
t
a
ls
,
acc
or
d
ing to th
e M
inistry of
E
nvironm
e
nt
,
a
r
e
100
ppm
for
Pb a
n
d
0
.
5
ppm for
Cd
in soil
.
O
n
e a
ppro
ac
h to
add
r
e
ssing th
e
c
h
a
ll
e
ng
e
s of ri
ce
growth in s
a
lin
e a
n
d
h
ea
vy m
e
t
a
l
-
c
ont
a
min
a
t
ed
soils
is through
b
iopriming
.
B
iopriming involv
e
s th
e
hy
d
r
a
tion of s
eed
s using
be
n
e
fi
c
i
a
l
b
iologi
ca
l
a
g
e
nts to
e
nh
a
n
ce
s
eed
g
e
rmin
a
tion qu
a
lity
.
T
h
e
s
e b
iologi
ca
l
a
g
e
nts not only poss
e
ss r
e
sist
a
n
ce
to s
a
linity
a
n
d
h
ea
vy m
e
t
a
ls
b
ut
a
lso h
a
v
e
th
e ab
ility to promot
e
pl
a
nt growth
.
E
n
d
ophyti
c bac
t
e
ri
a
,
whi
c
h
a
r
e c
ommonly foun
d
in n
a
tur
e
,
a
r
e
known for th
e
ir
ab
ility to improv
e
pl
a
nt qu
a
lity
.
T
h
e
s
e bac
t
e
ri
a
off
e
r
s
e
v
e
r
a
l
ad
v
a
nt
a
g
e
s
,
in
c
lu
d
ing prot
ec
tion from
ab
ioti
c
str
e
ss sin
ce
th
e
y
r
e
si
de
within th
e
pl
a
nt
,
o
cc
upy th
e
s
a
m
e
ni
c
h
e a
s p
a
thog
e
ns
,
ca
n
c
oloniz
e
pl
a
nt tissu
e
s
,
a
n
d
f
ac
ilit
a
t
e be
tt
e
r tr
a
nslo
ca
tion of m
e
t
ab
olit
e
s
into th
e
pl
a
nt
.
I
n lin
e
with
Ra
m
ad
h
a
n
'
s r
e
s
ea
r
c
h
,
it w
a
s shown th
a
t
e
n
d
ophyti
c bac
t
e
ri
a
positiv
e
ly influ
e
n
ce
th
e
h
e
ight of
I
np
a
ri
13
ri
ce
v
a
ri
e
ty
.
Acc
or
d
ing to
a
noth
e
r stu
d
y
b
y
F
itri
,
a
ll
e
n
d
ophyti
c bac
t
e
ri
a
l
isol
a
t
e
s pro
d
u
ce IAA
(
I
n
d
ol
e Ace
ti
c Ac
i
d
)
hormon
e
,
with on
e bac
t
e
ri
a
l
s
a
mpl
e
,
KSB
1
(
P
ot
a
ssium
S
olu
b
ilizing
Bac
t
e
ri
a
),
pro
d
u
c
ing
a
hormon
e
c
on
ce
ntr
a
tion of
57
ppm
.
E
n
d
ophyti
c bac
t
e
ri
a a
lso
e
nh
a
n
ce
ri
ce
pl
a
nt
growth
,
a
s
e
vi
de
n
ced b
y in
c
r
ea
s
ed
shoot h
e
ight
a
n
d
root l
e
ngth
.
KSB
1
is susp
ec
t
ed
to
be
long to th
e
g
e
nus
S
t
a
phylo
c
o
cc
us
.
B
iopriming of ri
ce
s
eed
s
ca
n mitig
a
t
e
th
e
n
e
g
a
tiv
e e
ff
ec
ts of s
a
linity str
e
ss
.
T
his w
a
s
5
de
monstr
a
t
ed
in
a
stu
d
y
b
y
Ra
w
a
t
,
wh
e
r
e b
iopriming with
T
ri
c
ho
de
rm
a
h
a
rzi
a
num isol
a
t
e
s un
de
r v
a
rious tr
ea
tm
e
nts signifi
ca
ntly r
ed
u
ced
str
e
ss
c
on
d
itions
a
n
d
improv
ed
shoot
a
n
d
root l
e
ngth
a
n
d
fr
e
sh w
e
ight
,
l
ea
f num
be
r
,
l
ea
f
a
r
ea
,
photosynth
e
ti
c
r
a
t
e
,
c
hlorophyll fluor
e
s
ce
n
ce
,
c
hlorophyll
c
ont
e
nt
,
a
n
d
soil
a
n
a
lysis
.
T
his r
e
s
ea
r
c
h
a
ims to isol
a
t
e
s
a
lt
-
a
n
d
h
ea
vy m
e
t
a
l
-
tol
e
r
a
nt
e
n
d
ophyti
c bac
t
e
ri
a a
s
b
iopriming
a
g
e
nts
.
E
n
d
ophyti
c bac
t
e
ri
a
w
e
r
e
sour
ced
from pl
a
nts with high s
a
lt
a
n
d
h
ea
vy m
e
t
a
l tol
e
r
a
n
ce
,
su
c
h
a
s
m
a
ngrov
e
s
.
Ma
ngrov
e
for
e
sts
a
r
e a
typ
e
of for
e
st foun
d a
long
c
o
a
stlin
e
s or riv
e
r
e
stu
a
ri
e
s
,
influ
e
n
ced b
y ti
da
l mov
e
m
e
nts
,
with s
ea
w
a
t
e
r inun
da
ting th
e a
r
ea d
uring high ti
de a
n
d
r
eced
ing
d
uring low
ti
de
.
Ma
ngrov
e
s
a
r
e c
o
a
st
a
l pl
a
nts
c
h
a
r
ac
t
e
riz
ed b
y th
e
ir high s
a
lt
tol
e
r
a
n
ce
.
S
onn
e
r
a
ti
a a
l
ba
m
a
ngrov
e
s h
a
v
e a
strong stimul
a
tory
e
ff
ec
t
on growth
a
t high s
a
lt
c
on
ce
ntr
a
tions
c
omp
a
r
ed
to oth
e
r m
a
ngrov
e
sp
ec
i
e
s lik
e S
onn
e
r
a
ti
a ca
s
e
ol
a
ris
a
n
d S
onn
e
r
a
ti
a
ov
a
t
a
.
T
h
e
susp
e
nsion
ce
lls of
S
onn
e
r
a
ti
a a
l
ba a
r
e
highly h
a
lophili
c
,
not only to
NaC
l
b
ut
a
lso to oth
e
r s
ea
s
a
lt
c
ompon
e
nts su
c
h
a
s
KC
l
,
M
g
C
l
2
,
a
n
d
CaC
l
2
.
Add
ition
a
lly
,
th
e
m
a
ngrov
e
pl
a
nts
S
onn
e
r
a
ti
a a
l
ba a
n
d
R
yzophor
a a
pi
c
ul
a
t
a
h
a
v
e bee
n shown to
ab
sor
b
h
ea
vy m
e
t
a
ls lik
e Cd
a
n
d Pb
.
B
iopriming ri
ce
with
e
n
d
ophyti
c bac
t
e
ri
a
from
S
onn
e
r
a
ti
a a
l
ba
m
a
ngrov
e
s in s
a
lin
e a
n
d
h
ea
vy m
e
t
a
l
-
c
ont
a
min
a
t
ed
soil h
a
s not y
e
t
bee
n r
e
port
ed
,
m
a
king this
a
pot
e
nti
a
l st
a
rting point for improving ri
ce
pro
d
u
c
tion using s
a
lt
-
r
e
sist
a
nt m
a
ngrov
e bac
t
e
ri
a
.
Re
s
ea
r
c
h
P
ro
b
l
e
m
S
t
a
t
e
m
e
nt
W
h
a
t
a
r
e
th
e c
olor
,
c
olony morphology
,
a
n
d
physiologi
ca
l
-
b
io
c
h
e
mi
ca
l
c
h
a
r
ac
t
e
risti
c
s of
e
n
d
ophyti
c bac
t
e
ri
a
isol
a
t
ed
from th
e
m
a
ngrov
e S
onn
e
r
a
ti
a
6
a
l
ba
?
W
hi
c
h sp
ec
i
e
s of
e
n
d
ophyti
c bac
t
e
ri
a e
xhi
b
it th
e
high
e
st s
a
lt
a
n
d
h
ea
vy m
e
t
a
l
tol
e
r
a
n
ce
,
a
s w
e
ll
a
s th
e be
st pl
a
nt growth
-
promoting
e
ff
ec
ts on ri
ce
?
W
h
a
t is th
e e
ff
ec
t of
b
iopriming with
e
n
d
ophyti
c bac
t
e
ri
a
from
S
onn
e
r
a
ti
a a
l
ba
on ri
ce
pl
a
nt growth
?
H
ypoth
e
sis
E
n
d
ophyti
c bac
t
e
ri
a
isol
a
t
ed
from
S
onn
e
r
a
ti
a a
l
ba e
xhi
b
it signifi
ca
ntly
d
iff
e
r
e
nt
c
olor
,
c
olony morphology
,
a
n
d
physiologi
ca
l
-
b
io
c
h
e
mi
ca
l
c
h
a
r
ac
t
e
risti
c
s
.
E
n
d
ophyti
c bac
t
e
ri
a
l sp
ec
i
e
s from
S
onn
e
r
a
ti
a a
l
ba de
monstr
a
t
e
sup
e
rior s
a
lt
a
n
d
h
ea
vy m
e
t
a
l tol
e
r
a
n
ce a
n
d
th
e be
st growth
-
promoting
e
ff
ec
ts on ri
ce
c
omp
a
r
ed
to oth
e
r
bac
t
e
ri
a
.
B
iopriming with
e
n
d
ophyti
c bac
t
e
ri
a
from
S
onn
e
r
a
ti
a a
l
ba e
nh
a
n
ce
s ri
ce
pl
a
nt
growth
c
omp
a
r
ed
to untr
ea
t
ed
pl
a
nts
.
Re
s
ea
r
c
h
Ob
j
ec
tiv
e
s
o
T
o isol
a
t
e a
n
d c
h
a
r
ac
t
e
riz
e e
n
d
ophyti
c bac
t
e
ri
a
from th
e
m
a
ngrov
e
S
onn
e
r
a
ti
a a
l
ba
th
a
t
e
xhi
b
it tol
e
r
a
n
ce
to s
a
lt
a
n
d
h
ea
vy m
e
t
a
ls
.
o
T
o i
de
ntify th
e
sp
ec
i
e
s of
e
n
d
ophyti
c bac
t
e
ri
a
th
a
t
de
monstr
a
t
e
th
e
high
e
st tol
e
r
a
n
ce
to s
a
lt
a
n
d
h
ea
vy m
e
t
a
ls
,
a
s w
e
ll
a
s th
e be
st pl
a
nt
growth
-
promoting
e
ff
ec
ts on ri
ce
.
o
T
o
e
v
a
lu
a
t
e
th
e e
ff
ec
t of
b
iopriming with
e
n
d
ophyti
c bac
t
e
ri
a
from
S
onn
e
r
a
ti
a a
l
ba
on th
e
growth of ri
ce
pl
a
nts
.
7
LITERATURE REVIEW
2
.
1
Sa
linity
Sa
linity r
e
f
e
rs to th
e c
on
ce
ntr
a
tion of
d
issolv
ed
min
e
r
a
l s
a
lts
pr
e
s
e
nt in soil
(
soil solution
)
a
n
d
w
a
t
e
r
.
T
h
e
s
e d
issolv
ed
min
e
r
a
l s
a
lts
a
r
e a
mixtur
e
of
ca
tions
a
n
d a
nions
,
fun
c
tioning
a
s
e
l
ec
trolyt
e
s
.
T
h
e
prim
a
ry
ca
tions in s
a
lin
e
soil solutions in
c
lu
de Na
+,
Ca
2
+,
M
g
2
+,
a
n
d
K
+,
whil
e
th
e
m
a
in
a
nions
a
r
e C
l
-,
SO
42
-,
HCO
3
-,
CO
32
-,
NO
3
-,
a
long
with oth
e
r
c
onstitu
e
nts su
c
h
a
s
B
,
S
r
2
+,
S
i
O
2
,
M
o
,
Ba
2
+,
a
n
d A
l
3
+
th
a
t
c
ontri
b
ut
e
to th
e
s
a
linity in soil
a
n
d
hyp
e
rs
a
lin
e
w
a
t
e
rs
.
T
h
e
pr
ed
omin
a
nt s
a
lt in s
a
lin
e
soils is
NaC
l
.
A
n
e
x
ce
ssiv
e a
mount of s
a
lt in
th
e
soil
ca
n l
ead
to inhi
b
ition
a
n
d
r
ed
u
c
tion in pl
a
nt growth
a
n
d
de
v
e
lopm
e
nt
.
P
l
a
nts r
e
spon
d
to s
a
linity in two ph
a
s
e
s
.
D
uring th
e
initi
a
l ph
a
s
e
(
P
h
a
s
e I
),
th
e e
x
ce
ss s
a
lt in th
e e
xt
e
rn
a
l soil solution m
a
k
e
s
w
a
t
e
r l
e
ss
a
v
a
il
ab
l
e
for
ab
sorption
b
y th
e
pl
a
nt roots
d
u
e
to its
b
in
d
ing
with w
a
t
e
r mol
ec
ul
e
s
,
thus r
ed
u
c
ing pl
a
nt growth
.
T
h
e
s
a
linity
-
in
d
u
ced
r
ed
u
c
tion in growth
d
uring
P
h
a
s
e I
is follow
ed b
y th
e
gr
ad
u
a
l
acc
umul
a
tion of s
a
lt within th
e
pl
a
nt tissu
e
s
,
first in th
e
ol
de
r tissu
e
s
a
n
d
th
e
n in th
e
young
e
r on
e
s
,
a
long with th
e
upt
a
k
e
of s
a
lin
e
w
a
t
e
r
,
l
ead
ing to ion s
a
lt toxi
c
ity
.
P
h
a
s
e II
involv
e
s th
e de
v
e
lopm
e
nt of s
a
lt
str
e
ss
,
c
h
a
r
ac
t
e
riz
ed b
y th
e a
ging of tissu
e
s progr
e
ssing from ol
de
r to
young
e
r
(
l
ea
v
e
s
).
Wa
t
e
r
de
fi
c
i
e
n
c
y
ca
us
ed b
y s
a
linity str
e
ss
d
isrupts
th
e
photosynth
e
sis pro
ce
ss in pl
a
nts
.
T
h
e
r
a
t
e
of photosynth
e
sis
dec
r
ea
s
e
s
d
u
e
to th
e
imp
ac
t on
b
io
c
h
e
mi
ca
l pro
ce
ss
e
s within pl
a
nt
ce
lls
.
8
Sa
lt str
e
ss imp
ac
ts nutri
e
nt
ab
sorption in pl
a
nts
.
H
igh s
a
linity
l
e
v
e
ls l
ead
to
a
r
ed
u
c
tion in root
a
r
ea
,
r
e
sulting in
dec
r
ea
s
ed
nutri
e
nt
upt
a
k
e
from th
e
soil
,
whi
c
h
d
isrupts th
e
nutri
e
nt supply to th
e
shoots
a
n
d
l
ea
v
e
s
,
ultim
a
t
e
ly hin
de
ring th
e
photosynth
e
sis pro
ce
ss
.
I
n
c
r
ea
s
ed
Na
+
c
on
ce
ntr
a
tion in th
e
soil
ca
n
dec
r
ea
s
e K
+
a
n
d Ca
2
+
l
e
v
e
ls in pl
a
nt
tissu
e
s
.
L
ow
Ca
2
+
upt
a
k
e b
y pl
a
nts
ca
n imp
a
ir
ce
ll m
e
m
b
r
a
n
e
fun
c
tion
a
n
d
int
e
grity
,
l
ead
ing to
Na
+
acc
umul
a
tion in pl
a
nt tissu
e
s
.
P
ot
a
ssium
pl
a
ys
a c
ru
c
i
a
l rol
e a
s
a ca
t
a
lyst for v
a
rious
e
nzym
e
s
.
I
f pot
a
ssium
(
K
)
c
on
ce
ntr
a
tion in pl
a
nt tissu
e
s
dec
lin
e
s
,
e
nzym
e ac
tiviti
e
s su
c
h
a
s
nitr
a
t
e
r
ed
u
c
t
a
s
e
,
whi
c
h
c
onv
e
rts
NO
3
into
NH
3
(
a
pr
ec
ursor for
prot
e
in form
a
tion
),
will
dec
r
ea
s
e
.
Sa
linity l
ead
s to
a
r
ed
u
c
tion in
nitrog
e
n
(
N
)
a
n
d
phosphorus
(
P
)
c
ont
e
nt in
a
ll pl
a
nt tissu
e
s
.
T
h
e
dec
r
ea
s
e
in
N
upt
a
k
e
is
d
u
e
to th
e
int
e
r
ac
tion
be
tw
ee
n
Na
+
with
NH
4
+
a
n
d C
l
-
with
NO
3
-.
T
h
e
growth
a
n
d
yi
e
l
d
of
c
ultiv
a
t
ed
pl
a
nts g
e
n
e
r
a
lly
dec
lin
e
wh
e
n soil
EC
r
eac
h
e
s
4
dS
/
m
,
e
quiv
a
l
e
nt to
40
m
M NaC
l
.
I
n s
a
lin
e
e
nvironm
e
nts
,
pl
a
nts
acc
umul
a
t
e
su
b
st
a
n
ce
s su
c
h
a
s lysin
e
,
prolin
e
,
a
sp
a
r
a
gin
e
,
glut
a
min
e
,
a
sp
a
rti
c ac
i
d
,
glut
a
mi
c ac
i
d
,
a
l
a
nin
e
,
tyrosin
e
,
a
n
d
v
a
lin
e
.
T
h
e
in
c
r
ea
s
e
in
d
i
ca
r
b
oxyli
c a
mino
ac
i
d a
n
d a
mi
de
c
on
ce
ntr
a
tions in
d
i
ca
t
e
s
a d
isruption in prot
e
in synth
e
sis
d
u
e
to th
e
in
e
ffi
c
i
e
nt us
e
of nitrog
e
n
ab
sor
bed b
y th
e
pl
a
nt roots
.
2
.
2
Hea
vy
Me
t
a
l
Hea
vy m
e
t
a
ls
a
r
e
m
e
t
a
lli
c e
l
e
m
e
nts with high mol
ec
ul
a
r
w
e
ight
.
E
v
e
n in low
c
on
ce
ntr
a
tions
,
h
ea
vy m
e
t
a
ls
a
r
e
g
e
n
e
r
a
lly
h
a
z
a
r
d
ous
d
u
e
to th
e
ir toxi
c
n
a
tur
e
,
a
ff
ec
ting pl
a
nts
,
a
nim
a
ls
,
a
n
d
9
hum
a
ns
.
C
ommonly foun
d e
nvironm
e
nt
a
l pollut
a
nts in
c
lu
de H
g
,
C
r
,
Cd
,
A
s
,
a
n
d Pb
.
Hea
vy m
e
t
a
ls
ca
n
be d
istri
b
ut
ed
into th
e
hum
a
n
b
o
d
y through v
a
rious int
e
rm
ed
i
a
ri
e
s su
c
h
a
s
a
ir
,
foo
d
,
soil
,
a
n
d
w
a
t
e
r
,
with som
e acc
umul
a
ting within th
e b
o
d
y
.
O
v
e
r tim
e
,
a
n
d
in
signifi
ca
nt
a
mounts
,
this
acc
umul
a
tion
ca
n pos
e
h
ea
lth risks
.
T
h
e
ab
sorption of h
ea
vy m
e
t
a
ls
b
y soil is influ
e
n
ced b
y th
e
soil
'
s
ad
sorption
ca
p
ac
ity to form
c
ompl
e
x
e
s with org
a
ni
c
m
a
t
e
ri
a
ls lik
e
humus
,
a
s w
e
ll
a
s th
e
pr
e
s
e
n
ce
of insolu
b
l
e c
ompoun
d
s un
de
r
r
ed
u
c
ing
c
on
d
itions
.
I
n soil solutions
,
h
ea
vy m
e
t
a
ls
ca
n
e
xist
a
s ions
or
c
ompl
e
x
e
s
.
T
h
e
y t
e
n
d
to form
c
ompl
e
x
b
on
d
s with org
a
ni
c
m
a
tt
e
r in th
e
soil
,
l
ead
ing to high
e
r
c
on
ce
ntr
a
tions in th
e
topsoil
l
a
y
e
rs
.
T
h
e
pr
e
s
e
n
ce
of h
ea
vy m
e
t
a
ls
c
ont
a
min
a
ting th
e
soil
ca
n
d
isrupt pl
a
nt growth
b
y r
ed
u
c
ing soil f
e
rtility
a
n
d
low
e
ring
c
rop
qu
a
lity
d
u
e
to th
e acc
umul
a
tion of pollut
a
nts
.
Lead
(
Pb
)
is
a
h
ea
vy m
e
t
a
l with
a
high risk pot
e
nti
a
l
d
u
e
to
its
ab
ility to
ea
sily
acc
umul
a
t
e
in w
a
t
e
r
,
s
ed
im
e
nts
,
a
n
d
pl
a
nts
.
W
h
e
n l
ead acc
umul
a
t
e
s in tissu
e
,
its
c
on
ce
ntr
a
tion in
c
r
ea
s
e
s with
th
e
rising l
e
v
e
ls of
Pb
in th
e
surroun
d
ing
e
nvironm
e
nt
.
Lead ca
n
e
nt
e
r pl
a
nts through two m
a
in m
ec
h
a
nisms
:
root
ab
sorption
a
n
d
foli
a
r
ab
sorption
.
O
n
ce
insi
de
th
e
pl
a
nt syst
e
m
,
Pb b
in
d
s to
ce
ll
m
e
m
b
r
a
n
e
s
,
mito
c
hon
d
ri
a
,
a
n
d c
hloropl
a
sts
.
I
t
ca
n
be
qui
c
kly
ab
sor
bed
wh
e
n tr
a
nsf
e
rr
ed
or wh
e
n th
e
pl
a
nt
'
s roots h
a
v
e d
i
ed
.
T
h
e
pot
e
nti
a
l
e
nvironm
e
nt
a
l
a
n
d
hum
a
n h
a
z
a
r
d
s pos
ed b
y h
ea
vy
m
e
t
a
ls in
c
lu
de
:
1
.
Hea
vy m
e
t
a
ls
a
r
e d
iffi
c
ult to
de
gr
ade
,
l
ead
ing to th
e
ir
c
ontinu
a
l
10
acc
umul
a
tion in th
e e
nvironm
e
nt
.
2
.
T
h
e
y h
a
v
e
th
e ab
ility to
acc
umul
a
t
e
in org
a
nisms
,
r
e
sulting in
in
c
r
ea
s
ed c
on
ce
ntr
a
tion ov
e
r tim
e
,
a
pro
ce
ss known
a
s
b
io
acc
umul
a
tion
a
n
d b
iom
a
gnifi
ca
tion
.
3
.
Hea
vy m
e
t
a
ls
ea
sily
acc
umul
a
t
e
in s
ed
im
e
nts
,
l
ead
ing to high
e
r
c
on
ce
ntr
a
tions in s
ed
im
e
nts
c
omp
a
r
ed
to w
a
t
e
r
.
Lead
(
Pb
)
Lead
,
c
ommonly known
a
s
b
l
ac
k l
ead
,
be
longs to th
e G
roup
IV
-
A
m
e
t
a
ls in
th
e
p
e
rio
d
i
c
t
ab
l
e
.
I
t h
a
s
a
n
a
tomi
c
num
be
r of
82
a
n
d a
mol
ec
ul
a
r w
e
ight of
207
.
2
.
Lead
is
a
soft m
e
t
a
l with
a
r
edd
ish
-
b
rown
c
olor
.
I
t h
a
s
a
low m
e
lting
point
,
is
ea
sily sh
a
p
ed
,
a
n
d e
xhi
b
its
ac
tiv
e c
h
e
mi
ca
l prop
e
rti
e
s
.
D
u
e
to th
e
s
e
c
h
a
r
ac
t
e
risti
c
s
,
it is oft
e
n us
ed
to
c
o
a
t m
e
t
a
ls to pr
e
v
e
nt
c
orrosion
.
T
h
e
l
ead
c
ont
e
nt in pl
a
nts is non
-
e
ss
e
nti
a
l
,
typi
ca
lly r
a
nging from
0
.
1
to
10
ppm
.
Ge
n
e
r
a
lly
,
th
e
l
ead c
ont
e
nt in v
a
rious typ
e
s of pl
a
nts r
a
ng
e
s
be
tw
ee
n
0
.
5
a
n
d
3
.
0
ppm
.
H
ow
e
v
e
r
,
som
e
pl
a
nts
e
xhi
b
it v
e
ry high l
e
v
e
ls of toxi
c
ity tow
a
r
d
s
l
ead
.
T
his situ
a
tion
ca
n
be da
ng
e
rous
beca
us
e
,
a
lthough pl
a
nts m
a
y
a
pp
ea
r
h
ea
lthy
a
n
d
show no signs of l
ead
poisoning
,
th
e
y
ca
n still pos
e a
pot
e
nti
a
l
h
a
z
a
r
d
if
c
onsum
ed b
y hum
a
ns
.
Cad
mium
(
Cd
)
Cad
mium is
a b
ypro
d
u
c
t of th
e
zin
c
pro
d
u
c
tion pro
ce
ss
.
I
t is
a
soft
,
whit
e
m
e
t
a
l with
a b
luish tint
,
a
n
d be
longs to
G
roup
VI
.
Cad
mium h
a
s
a
n
a
tomi
c
num
be
r of
48
a
n
d a
n
a
tomi
c
w
e
ight of
112
.
41
.
I
ts m
e
lting point is
321
º
C
,
a
n
d
it
h
a
s
a de
nsity of
8
.
65
g
/
m
L
.
11
Cad
mium
a
n
d
its
c
ompoun
d
s h
a
v
e
high
e
r solu
b
ility in w
a
t
e
r
c
omp
a
r
ed
to
oth
e
r m
e
t
a
ls
.
T
h
e
ir high
b
iologi
ca
l
a
v
a
il
ab
ility m
ea
ns th
e
y t
e
n
d
to
acc
umul
a
t
e
in org
a
nisms
.
L
ong
-
t
e
rm
e
xposur
e
to
cad
mium
ca
n
ca
us
e
morphop
a
thologi
ca
l
c
h
a
ng
e
s in th
e
ki
d
n
e
ys
,
a
s
cad
mium int
e
r
ac
ts through nutrition
a
n
d
in
d
u
ce
s
toxi
c
ity
.
E
n
d
ophyti
c Bac
t
e
ri
a
E
n
d
ophyti
c bac
t
e
ri
a a
r
e a
typ
e
of
bac
t
e
ri
a
th
a
t liv
e
within th
e
ir host
tissu
e
s without
ca
using
a
ny symptoms of
d
is
ea
s
e
.
I
t h
a
s
bee
n r
e
port
ed
th
a
t
th
e
pr
e
s
e
n
ce
of
e
n
d
ophyti
c bac
t
e
ri
a
in pl
a
nt tissu
e
s pl
a
ys
a
signifi
ca
nt rol
e
in
promoting pl
a
nt growth
d
u
e
to th
e
ir
ab
ility to pro
d
u
ce
growth
-
promoting
su
b
st
a
n
ce
s
,
fix nitrog
e
n
,
mo
b
iliz
e
phosph
a
t
e
,
a
n
d c
ontri
b
ut
e
to pl
a
nt h
ea
lth
.
E
n
d
ophyti
c bac
t
e
ri
a
h
a
v
e
th
e
pot
e
nti
a
l to pro
d
u
ce
s
ec
on
da
ry m
e
t
ab
olit
e
s
foun
d
in th
e
ir host pl
a
nts through g
e
n
e
ti
c
r
ec
om
b
in
a
tion
.
T
h
e
s
e bac
t
e
ri
a ca
n
be
foun
d
in th
e
roots
,
l
ea
v
e
s
,
a
n
d
st
e
ms of pl
a
nts
.
R
ol
e
of
E
n
d
ophyti
c Bac
t
e
ri
a
in
P
l
a
nt
G
rowth
E
n
d
ophyt
e
s pl
a
y
a
n import
a
nt rol
e a
s pl
a
nt growth stimul
a
nts
.
Bac
t
e
ri
a
su
c
h
a
s
E
nt
e
ro
bac
t
e
r
a
n
d K
luyv
e
r
a a
s
c
or
ba
t
a SUD
165
a
r
e
known to
stimul
a
t
e
pl
a
nt growth
a
n
d
in
c
r
ea
s
e
r
e
sist
a
n
ce
to h
ea
vy m
e
t
a
ls
.
S
om
e
e
n
d
ophyti
c bac
t
e
ri
a
h
a
v
e
th
e ab
ility to pro
d
u
ce I
n
d
ol
e Ace
ti
c Ac
i
d
(
IAA
),
c
ommonly known
a
s
a
uxin
.
A
uxin pl
a
ys
a
rol
e
in stimul
a
ting pl
a
nt growth
a
n
d
is g
e
n
e
r
a
lly foun
d
in m
e
rist
e
m
a
ti
c
tissu
e
s
.
E
n
d
ophyti
c bac
t
e
ri
a
h
a
v
e
pot
e
nti
a
l
a
s
b
io
c
ontrol
a
g
e
nts
beca
us
e
th
e
ir pr
e
s
e
n
ce
within pl
a
nt tissu
e
s
e
n
ab
l
e
s th
e
m
to withst
a
n
d b
ioti
c a
n
d ab
ioti
c
str
e
ss
e
s
.
I
n
add
ition to
ac
ting
a
s
b
io
c
ontrol
a
g
e
nts
,
ce
rt
a
in typ
e
s of
e
n
d
ophyti
c bac
t
e
ri
a a
lso fun
c
tion
a
s pl
a
nt growth
12
stimul
a
nts
a
n
d e
nh
a
n
ce
pl
a
nt immunity
a
g
a
inst p
a
thog
e
ns lik
e B
urkhol
de
ri
a
sp
.
2
.
3
S
onn
e
r
a
ti
a a
l
ba Ma
ngrov
e
Ma
ngrov
e
s
a
r
e a
group of pl
a
nts th
a
t inh
ab
it tropi
ca
l
c
o
a
st
a
l
r
e
gions
,
grow on mu
dd
y su
b
str
a
t
e
s
,
a
n
d
thriv
e
in
a
r
ea
s with high s
a
lt
c
ont
e
nt
.
T
h
e
m
a
ngrov
e ec
osyst
e
m is oft
e
n r
e
f
e
rr
ed
to
a
s
a b
r
ac
kish
for
e
st
beca
us
e
it is foun
d
in
b
r
ac
kish r
e
gions
(
e
stu
a
ri
e
s
),
whi
c
h h
a
v
e
s
a
linity l
e
v
e
ls r
a
nging from
0
.
5
%
to
30
%
.
T
his
ec
osyst
e
m is
a
lso known
a
s
a
ti
da
l
ec
osyst
e
m
beca
us
e
it is lo
ca
t
ed
in
a
r
ea
s influ
e
n
ced b
y ti
da
l
s
ea
mov
e
m
e
nts
.
Ma
ngrov
e
s
a
r
e c
onsi
de
r
ed a
qu
a
ti
c
pl
a
nts
ca
p
ab
l
e
of
ab
sor
b
ing min
e
r
a
l s
a
lts
,
in
c
lu
d
ing h
ea
vy m
e
t
a
ls
.
T
h
e
s
e
h
ea
vy m
e
t
a
ls
a
r
e
t
a
k
e
n up from soil s
ed
im
e
nt or w
a
t
e
r through th
e
root syst
e
m
a
n
d
d
istri
b
ut
ed
to oth
e
r p
a
rts of th
e
pl
a
nt
,
in
c
lu
d
ing th
e
fruit
a
n
d
l
ea
v
e
s
.
T
r
ee
s in m
a
ngrov
e
for
e
sts
ca
n
ada
pt to low oxyg
e
n l
e
v
e
ls
a
n
d
tol
e
r
a
t
e
high s
a
lt
c
on
ce
ntr
a
tions
.
T
his
ada
ptiv
e ab
ility is
d
u
e
to th
e
de
v
e
lopm
e
nt of stru
c
tur
e
s in th
e
l
ea
v
e
s th
a
t
ca
n
e
xp
e
l s
a
lt through th
e
stom
a
t
a
.
Add
ition
a
lly
,
m
a
ngrov
e
l
ea
v
e
s
a
r
e
thi
c
k
e
nough to stor
e a
signifi
ca
nt
a
mount of w
a
t
e
r
.
Acc
or
d
ing to
a
stu
d
y
c
on
d
u
c
t
ed b
y th
e Fac
ulty of
F
or
e
stry
a
t
IPB
in
c
oll
ab
or
a
tion with th
e D
ir
ec
tor
a
t
e Ge
n
e
r
a
l of
La
n
d Re
h
ab
ilit
a
tion
a
n
d
S
o
c
i
a
l
F
or
e
stry in
1999
,
th
e a
r
ea
of m
a
ngrov
e
for
e
sts in
I
n
d
on
e
si
a
is
e
stim
a
t
ed
to
c
ov
e
r
9
.
2
million h
ec
t
a
r
e
s
,
with
3
.
7
million h
ec
t
a
r
e
s
lo
ca
t
ed
within for
e
st
a
r
ea
s
a
n
d
5
.
5
million h
ec
t
a
r
e
s outsi
de
for
e
st
a
r
ea
s
.
T
h
e
stru
c
tur
e a
n
d c
omposition of m
a
ngrov
e
s in
I
n
d
on
e
si
a
show
mor
e d
iv
e
rsity
c
omp
a
r
ed
to oth
e
r r
e
gions
.
13
I
n
I
n
d
on
e
si
a
,
v
a
rious typ
e
s of m
a
ngrov
e
st
a
n
d
s
ca
n
be
foun
d
,
r
a
nging from
A
vi
ce
nni
a
m
a
rin
a
,
whi
c
h grows to
ab
out
1
-
2
m
e
t
e
rs
a
n
d
is
foun
d
on
beac
h
e
s th
a
t
a
r
e c
ontinuously su
b
m
e
rg
ed b
y s
ea
w
a
t
e
r
,
to
mix
ed
st
a
n
d
s of
B
rugui
e
r
a
-
R
hizophor
a
-
Ce
riops th
a
t r
eac
h h
e
ights of
ov
e
r
30
m
e
t
e
rs
.
O
n op
e
n
beac
h
e
s
,
sp
ec
i
e
s su
c
h
a
s
A
vi
ce
nni
a a
l
ba a
n
d
S
onn
e
r
a
ti
a a
l
ba a
r
e
oft
e
n foun
d
,
whil
e a
long riv
e
rs with low
e
r s
a
linity
,
p
a
lm sp
ec
i
e
s lik
e N
yp
a
fruti
ca
ns
a
n
d S
onn
e
r
a
ti
a ca
s
e
ol
a
ris
a
r
e
c
ommon
.
I
n s
ec
on
da
ry m
a
ngrov
e a
r
ea
s
,
m
a
ngrov
e
s
eed
lings
d
omin
a
t
e
,
a
long with v
a
rious typ
e
s of shru
b
s or h
e
r
b
s lik
e Aca
nthus ili
c
ifolius
a
n
d
Ac
rosti
c
hum
a
ur
e
um
.
2
.
4
B
iopriming
P
riming is
a
gr
ad
u
a
l hy
d
r
a
tion pro
ce
ss
ca
rri
ed
out
be
for
e
s
eed
s
a
r
e
pl
a
nt
ed
to
e
nsur
e
th
a
t th
e
w
a
t
e
r
c
ont
e
nt within th
e
s
eed
s
r
eac
h
e
s
a ba
l
a
n
ce
th
a
t
a
llows m
e
t
ab
oli
c ac
tiv
a
tion
.
P
riming tr
ea
tm
e
nts
ca
n
be c
om
b
in
ed
with th
e a
ppli
ca
tion of
b
iologi
ca
l
a
g
e
nts th
a
t h
a
v
e
th
e ab
ility to improv
e
s
eed
g
e
rmin
a
tion qu
a
lity
.
F
or
e
x
a
mpl
e
,
using
nitrog
e
n
-
fixing mi
c
ro
be
s
,
mi
c
ro
be
s th
a
t pro
d
u
ce
growth hormon
e
s
,
or
mi
c
ro
be
s th
a
t
e
nh
a
n
ce
str
e
ss r
e
sist
a
n
ce
.
T
his
c
om
b
in
a
tion tr
ea
tm
e
nt is
known
a
s
b
io
-
priming
.
Seed b
iopriming is
a
n
ad
v
a
n
ced
s
eed
tr
ea
tm
e
nt
t
ec
hniqu
e
involving th
e a
ppli
ca
tion of
be
n
e
fi
c
i
a
l mi
c
roorg
a
nisms on th
e
s
eed
surf
ace
,
follow
ed b
y s
eed
hy
d
r
a
tion
.
Seed b
iopriming
e
nh
a
n
ce
s
th
e
initi
a
l st
a
g
e
s of pl
a
nt
de
v
e
lopm
e
nt
b
y improving s
eed
g
e
rmin
a
tion
a
n
d
provi
d
ing prot
ec
tion
be
for
e
s
eed
ling
e
m
e
rg
e
n
ce
.
B
iopriming
14
tr
ea
tm
e
nt
ca
n promot
e c
h
a
ng
e
s in pl
a
nt
c
h
a
r
ac
t
e
risti
c
s
a
n
d
f
ac
ilit
a
t
e
uniform s
eed
g
e
rmin
a
tion
a
n
d
growth
a
sso
c
i
a
t
ed
with mi
c
ro
b
i
a
l
ino
c
ul
a
tion
.
2
.
5
S
p
e
ktrofotom
e
t
e
r
UV
-
V
is
T
h
e UV
-
V
is sp
ec
trophotom
e
t
e
r is
a
sp
ec
tros
c
opy
a
n
a
lysis
m
e
tho
d
th
a
t utiliz
e
s
e
l
ec
trom
a
gn
e
ti
c
r
ad
i
a
tion sour
ce
s in th
e
n
ea
r
-
ultr
a
viol
e
t r
a
ng
e
(
190
-
380
nm
)
a
n
d
visi
b
l
e
light
(
380
-
780
nm
)
with
a
sp
ec
trophotom
e
t
e
r
de
vi
ce
.
T
his
de
vi
ce
is us
ed
to m
ea
sur
e
th
e
r
e
l
a
tiv
e
e
n
e
rgy
a
sso
c
i
a
t
ed
with tr
a
nsmission
,
r
e
fl
ec
tion
,
a
n
d e
mission of
e
n
e
rgy
a
s
a
fun
c
tion of w
a
v
e
l
e
ngth
.
T
h
e
r
e a
r
e
s
e
v
e
r
a
l import
a
nt
c
onsi
de
r
a
tions in ultr
a
viol
e
t sp
ec
trophotom
e
tri
c a
n
a
lysis
:
-
Se
l
ec
tion of
Ma
ximum
Wa
v
e
l
e
ngth
:
T
h
e
w
a
v
e
l
e
ngth us
ed
for
qu
a
ntit
a
tiv
e a
n
a
lysis is wh
e
r
e
m
a
ximum
ab
sorption o
cc
urs
.
T
o
de
t
e
rmin
e
this w
a
v
e
l
e
ngth
,
a c
urv
e
is
c
r
ea
t
ed
th
a
t
c
onn
ec
ts
ab
sor
ba
n
ce
with w
a
v
e
l
e
ngth from
a
st
a
n
da
r
d
solution
a
t
a
sp
ec
ifi
c
c
on
ce
ntr
a
tion
.
-
Ca
li
b
r
a
tion
C
urv
e C
r
ea
tion
:
A
s
e
ri
e
s of st
a
n
da
r
d
solutions is pr
e
p
a
r
ed
from th
e
su
b
st
a
n
ce
to
be a
n
a
lyz
ed
with v
a
rying
c
on
ce
ntr
a
tions
.
T
h
e
ab
sor
ba
n
ce
of
eac
h solution with
d
iff
e
r
e
nt
c
on
ce
ntr
a
tions is m
ea
sur
ed
,
a
n
d
th
e
n
a c
urv
e
is
c
r
ea
t
ed
to illustr
a
t
e
th
e
r
e
l
a
tionship
be
tw
ee
n
ab
sor
ba
n
ce a
n
d c
on
ce
ntr
a
tion
.
I
f th
e Bee
r
-
La
m
be
rt l
a
w is follow
ed
,
th
e
ca
li
b
r
a
tion
c
urv
e
will
be a
str
a
ight lin
e
.
21
METHOD
Ma
t
e
ri
a
ls
T
h
e
m
a
t
e
ri
a
ls us
ed
in
c
lu
de S
onn
e
r
a
ti
a a
l
ba
m
a
ngrov
e
l
ea
v
e
s
from
Te
luk
Ra
t
a
i
,
Pe
s
a
w
a
r
a
n
Re
g
e
n
c
y
,
S
outh
La
mpung
;
Pad
j
a
j
a
r
a
n
A
grit
a
n ri
ce
s
eed
s
(
O
ryz
a
s
a
tiv
a L
.);
soil o
b
t
a
in
ed
from th
e BATAN
B
iology
Lab
or
a
tory
;
ri
ce
husk
a
sh
;
2
%
so
d
ium hypo
c
hlorit
e
;
70
%
e
th
a
nol
;
80
%
m
e
th
a
nol
;
N
utri
e
nt
B
roth
(
NB
);
so
d
ium
c
hlori
de
;
l
ead
m
e
t
a
l solution
(
Pb
);
cad
mium m
e
t
a
l solution
(
Cd
);
d
istill
ed
w
a
t
e
r
;
a
g
a
r
;
su
c
ros
e
;
d
ipot
a
ssium phosph
a
t
e
;
monopot
a
ssium phosph
a
t
e
;
h
e
pt
a
hy
d
r
a
t
e
m
a
gn
e
sium sulf
a
t
e
;
f
e
rrous sulf
a
t
e
;
so
d
ium moly
bda
t
e
;
c
opp
e
r
ca
r
b
on
a
t
e
;
m
a
gn
e
sium
c
hlori
de
;
ca
l
c
ium phosph
a
t
e
;
pot
a
ssium
c
hlori
de
;
a
mmonium sulf
a
t
e
;
glu
c
os
e
;
ca
l
c
ium sulf
a
t
e
;
f
e
rri
c c
hlori
de
;
7
.
5
%
so
d
ium
ca
r
b
on
a
t
e
solution
;
nitri
c ac
i
d
;
hy
d
ro
c
hlori
c ac
i
d
;
30
%
hy
d
rog
e
n p
e
roxi
de
(
J
t
Ba
k
e
r
);
sulfuri
c ac
i
d
;
ace
ton
e
;
g
a
lli
c ac
i
d
;
IAA
;
1
%
tryptoph
a
n
;
Sa
lkowski r
ea
g
e
nt
;
Ne
ssl
e
r r
ea
g
e
nt
;
10
%
F
olin
-
C
io
ca
lt
e
u
;
a
mmonium moly
bda
t
e
;
a
n
d
oth
e
r
c
h
e
mi
ca
ls
.
E
quipm
e
nts
T
h
e e
quipm
e
nt us
ed
in
c
lu
de
s
a bea
k
e
r
,
stirring ro
d
,
E
rl
e
nm
e
y
e
r
fl
a
sk
,
gr
ad
u
a
t
ed c
ylin
de
r
,
Pe
tri
d
ish
,
ino
c
ul
a
ting n
eed
l
e
,
B
uns
e
n
b
urn
e
r
,
gr
ad
u
a
t
ed
pip
e
tt
e
,
mi
c
ropip
e
tt
e
,
d
ropp
e
r
,
t
e
st tu
be
s
,
sh
a
k
e
r
,
gl
a
ss sli
de
s
,
a
n
a
lyti
ca
l
ba
l
a
n
ce
,
mi
c
ros
c
op
e
,
a
uto
c
l
a
v
e
,
hotpl
a
t
e
,
th
e
rmom
e
t
e
r
,
ce
ntrifug
e
,
w
a
t
e
r
ba
th
,
fum
e
hoo
d
,
l
a
min
a
r flow
cab
in
e
t
,
furn
ace
,
ov
e
ns
a
t
60
°
C a
n
d
100
°
C
,
in
c
u
ba
tor
,
de
si
cca
tor
,
UV
-
V
is sp
ec
trophotom
e
t
e
r
,
a
n
d a
tomi
c ab
sorption
sp
ec
trophotom
e
t
e
r
.
22
Re
s
ea
r
c
h
P
ro
ced
ur
e
I
sol
a
tion of
E
n
d
ophyti
c Bac
t
e
ri
a
from
Ma
ngrov
e S
onn
e
r
a
ti
a a
l
ba
23
Bac
t
e
ri
a
l
P
urifi
ca
tion
Bac
t
e
ri
a
isol
a
t
ed
from m
ed
i
a B
with
d
ilutions of
10
^-
9
a
n
d
10
^-
10
w
e
r
e
purifi
ed
on
Pe
tri
d
ish
e
s using th
e
str
ea
k pl
a
t
e
m
e
tho
d
in
NA
m
ed
i
a
.
T
h
e d
ish
e
s
c
ont
a
ining th
e
bac
t
e
ri
a
w
e
r
e
th
e
n in
c
u
ba
t
ed
in
a
n in
c
u
ba
tor for
3
da
ys to o
b
t
a
in pur
e c
ultur
e
s or
singl
e
isol
a
t
e
s
.
E
n
d
ophyti
c Bac
t
e
ri
a C
h
a
r
ac
t
e
riz
a
tion
M
orphologi
ca
l
C
h
a
r
ac
t
e
riz
a
tion
T
h
e bac
t
e
ri
a
l isol
a
t
e
s w
e
r
e c
h
a
r
ac
t
e
riz
ed b
y o
b
s
e
rving th
e c
olor
a
n
d
sh
a
p
e
of
c
oloni
e
s
.
Bac
t
e
ri
a
l
c
oloni
e
s w
e
r
e e
x
a
min
ed
using
a
mi
c
ros
c
op
e
with
G
r
a
m st
a
ining
m
e
tho
d
.
L
iqui
d bac
t
e
ri
a
l isol
a
t
e
s w
e
r
e
pl
aced
on
a
gl
a
ss sli
de a
n
d a
ir
-
d
ri
ed
,
follow
ed
b
y th
e add
ition of
c
ryst
a
l viol
e
t
a
n
d
in
c
u
ba
tion for
2
minut
e
s
.
T
h
e
sli
de
w
a
s th
e
n rins
ed
with
d
istill
ed
w
a
t
e
r
,
follow
ed b
y th
e add
ition of
L
ugol
'
s io
d
in
e a
n
d
in
c
u
ba
tion for
2
minut
e
s
.
T
h
e
sli
de
w
a
s rins
ed a
g
a
in with
d
istill
ed
w
a
t
e
r
a
n
d
th
e
n
a
l
c
ohol w
a
s
a
ppli
ed
until th
e c
olor
d
is
a
pp
ea
r
ed
.
Ne
xt
,
th
e
sli
de
w
a
s rins
ed
with
d
istill
ed
w
a
t
e
r
a
n
d
st
a
in
ed
with s
a
fr
a
nin for
2
minut
e
s
.
T
h
e
pr
e
p
a
r
a
tion w
a
s th
e
n rins
ed
with
d
istill
ed
w
a
t
e
r
,
d
ri
ed
,
a
n
d
o
b
s
e
rv
ed
un
de
r
a
1000
x m
a
gnifi
ca
tion mi
c
ros
c
op
e
.
P
hysiologi
ca
l
a
n
d B
io
c
h
e
mi
ca
l
C
h
a
r
ac
t
e
riz
a
tion
T
h
e e
n
d
ophyti
c bac
t
e
ri
a
l isol
a
t
e
s w
e
r
e
th
e
n
c
h
a
r
ac
t
e
riz
ed
for physiologi
ca
l
a
n
d
b
io
c
h
e
mi
ca
l prop
e
rti
e
s
,
in
c
lu
d
ing nitrog
e
n fix
a
tion
,
phosph
a
t
e
solu
b
iliz
a
tion
,
a
n
d IAA
pro
d
u
c
tion using
a UV
-
V
is sp
ec
trophotom
e
t
e
r
.
T
h
e bac
t
e
ri
a
us
ed
for th
e
s
e
t
e
sts w
e
r
e
c
ultur
ed
in liqui
d
form
.
Bac
t
e
ri
a
from pl
a
t
e
s w
e
r
e
ino
c
ul
a
t
ed
into st
e
riliz
ed NB
m
ed
i
a
a
n
d
in
c
u
ba
t
ed
for
3
da
ys
a
t room t
e
mp
e
r
a
tur
e
with
c
ontinuous stirring
a
t
100
rpm
.
T
o t
e
st th
e bac
t
e
ri
a
’
s
ab
ility to fix
a
tmosph
e
ri
c
nitrog
e
n
,
th
e bac
t
e
ri
a
l liqui
d
24
c
ultur
e
w
a
s
added
to
B
urk
’
s m
ed
ium
(
N
fr
ee
m
ed
ium
).
B
urk
’
s m
ed
ium w
a
s pr
e
p
a
r
ed
with
M
g
SO
4 1
g
,
K
2
HPO
4 4
g
,
KH
2
PO
4 1
g
,
CaSO
4 0
.
65
g
,
FeC
l
3 0
.
00725
g
,
a
n
d Na
moly
bda
t
e
0
.
001265
g in
1
lit
e
r of w
a
t
e
r
.
T
h
e bac
t
e
ri
a
w
e
r
e
in
c
u
ba
t
ed
for
7
da
ys
a
t
27
°
C
.
N
itrog
e
n
c
ont
e
nt w
a
s
de
t
e
rmin
ed
using th
e Ne
ssl
e
r m
e
tho
d
.
A
1
ml
a
liquot of th
e
s
a
mpl
e
sup
e
rn
a
t
a
nt w
a
s tr
a
nsf
e
rr
ed
to
a
mi
c
rotu
be a
n
d ce
ntrifug
ed a
t
12
,
000
rpm for
10
minut
e
s
.
T
h
e c
l
ea
r sup
e
rn
a
t
a
nt w
a
s th
e
n tr
a
nsf
e
rr
ed
to
a
r
eac
tion tu
be
,
a
n
d
50
µ
L
of
Ne
ssl
e
r
’
s r
ea
g
e
nt w
a
s
added a
n
d
in
c
u
ba
t
ed
for
30
minut
e
s
.
T
h
e
sup
e
rn
a
t
a
nt
a
n
d
r
ea
g
e
nt solution w
e
r
e
th
e
n
d
ilut
ed
with
4
.
5
ml
d
istill
ed
w
a
t
e
r
,
a
n
d ab
sor
ba
n
ce
w
a
s
m
ea
sur
ed a
t
425
nm
.
T
o t
e
st th
e bac
t
e
ri
a
’
s
ab
ility to solu
b
iliz
e
phosph
a
t
e
,
th
e bac
t
e
ri
a
w
e
r
e
grown in
NBRIP
m
ed
ium
.
NBRIP
m
ed
ium
c
ont
a
in
ed
glu
c
os
e
10
g
,
Ca
3
(
PO
4
)
2 5
g
,
M
g
C
l
2
.
6
H
2
O
5
g
,
M
g
SO
4
.
7
H
2
O
0
.
25
g
,
KC
l
0
.
2
g
, (
NH
4
)
2
SO
4 0
.
1
g
,
a
n
d d
istill
ed
w
a
t
e
r
to
1
L
with
a
p
H
of
7
.
T
h
e bac
t
e
ri
a
w
e
r
e
in
c
u
ba
t
ed
for
7
da
ys
a
t
27
°
C
.
A
1
ml
a
liquot of
sup
e
rn
a
t
a
nt w
a
s
ce
ntrifug
ed a
t
12
,
000
rpm for
10
minut
e
s
.
T
h
e c
l
ea
r sup
e
rn
a
t
a
nt w
a
s
th
e
n tr
a
nsf
e
rr
ed
to
a
r
eac
tion tu
be
,
a
n
d
1
ml
HNO
3
a
n
d
500
µ
L
of
AB
solution
(
a
mmonium moly
bda
t
e a
n
d a
mmonium m
e
t
a
v
a
n
ada
t
e
in
a
1
:
1
r
a
tio
)
w
e
r
e added
.
T
h
e
solution w
a
s in
c
u
ba
t
ed
for
30
minut
e
s
,
a
n
d ab
sor
ba
n
ce
w
a
s m
ea
sur
ed a
t
430
nm
.
T
o
t
e
st th
e
pro
d
u
c
tion of
IAA
,
bac
t
e
ri
a
w
e
r
e
grown in
NB
m
ed
i
a
suppl
e
m
e
nt
ed
with
tryptoph
a
n
a
t
0
.
2
mg
/
m
L
.
T
h
e bac
t
e
ri
a
w
e
r
e
in
c
u
ba
t
ed
for
7
da
ys
a
t
27
°
C
.
T
h
e
liqui
d
c
ultur
e
w
a
s
ce
ntrifug
ed a
t
12
,
000
rpm for
10
minut
e
s
.
A
1
m
L a
liquot of th
e
sup
e
rn
a
t
a
nt w
a
s th
e
n
added
to
2
m
L
of
Sa
lkowski r
ea
g
e
nt
a
n
d
in
c
u
ba
t
ed
for
30
minut
e
s
.
T
h
e
form
a
tion of
a
r
ed c
olor in th
e
sup
e
rn
a
t
a
nt in
d
i
ca
t
ed a
positiv
e
r
e
sult for
IAA
pro
d
u
c
tion
.
T
h
e
int
e
nsity of th
e
r
ed c
olor w
a
s m
ea
sur
ed
using
a
sp
ec
trophotom
e
t
e
r
a
t
530
nm
,
a
n
d ab
sor
ba
n
ce
v
a
lu
e
s w
e
r
e c
omp
a
r
ed
with
a
st
a
n
da
r
d
IAA c
urv
e
.
T
h
e
st
a
n
da
r
d IAA
solutions h
ad c
on
ce
ntr
a
tions of
0
,
5
,
10
,
15
,
20
,
25
,
30
,
a
n
d
35
ppm
.
25
R
i
ce Seed S
tr
e
ss
Te
sting
R
i
ce
s
eed
s w
e
r
e
w
a
sh
ed
with
2
%
NaOC
l for
30
minut
e
s
a
n
d
th
e
n rins
ed
with w
a
t
e
r
thr
ee
tim
e
s
.
S
tr
e
ss solutions w
e
r
e
pr
e
p
a
r
ed b
y
add
ing
NaC
l
500
ppm
,
Pb
500
ppm
,
a
n
d Cd
500
ppm to
d
istill
ed
w
a
t
e
r in
a
100
m
L
volum
e
tri
c
fl
a
sk
.
T
h
e
solutions w
e
r
e
d
ilut
ed
to
c
on
ce
ntr
a
tions of
0
,
25
,
50
,
100
,
200
,
300
,
400
,
a
n
d
500
ppm
.
C
l
ea
n ri
ce
s
eed
s w
e
r
e
so
a
k
ed
in vi
a
ls
acc
or
d
ing to th
e
str
e
ss v
a
ri
a
tions for
24
hours
.
T
h
e
s
eed
s
w
e
r
e
th
e
n pl
a
nt
ed
using th
e b
lott
e
r t
e
st m
e
tho
d
on
Pe
tri
d
ish
e
s
,
with
20
s
eed
s p
e
r
d
ish
a
n
d
thr
ee
r
e
pli
ca
t
e
s
,
for
6
-
7
da
ys in
a da
rk
c
h
a
m
be
r
.
T
h
e e
n
d
ophyti
c bac
t
e
ri
a
us
ed
w
e
r
e
r
e
sist
a
nt to s
a
linity
a
n
d
h
ea
vy m
e
t
a
l str
e
ss
a
n
d
h
ad
th
e ab
ility to promot
e
ri
ce
pl
a
nt growth
,
in
c
lu
d
ing nitrog
e
n fix
a
tion
,
phosph
a
t
e
solu
b
iliz
a
tion
,
a
n
d IAA
pro
d
u
c
tion
.
T
h
e
ri
ce
v
a
ri
e
ty us
ed
w
a
s
Pad
j
a
j
a
r
a
n
A
grit
a
n from
PAIR
-
BATAN
.
E
n
d
ophyti
c bac
t
e
ri
a
w
e
r
e
in
d
u
ced
into ri
ce
s
eed
s using
a C
ompl
e
t
e
ly
Ra
n
d
omiz
ed De
sign
(
CRD
)
with
3
tr
ea
tm
e
nts
(
NaC
l str
e
ss
,
Pb
-
Cd
str
e
ss
,
a
n
d
no str
e
ss
)
a
n
d
3
r
e
pli
ca
t
e
s
.
B
iopriming of
m
a
ngrov
e e
n
d
ophyti
c bac
t
e
ri
a S
onn
e
r
a
ti
a a
l
ba
into ri
ce
s
eed
s w
a
s
d
on
e b
y
d
issolving
th
e bac
t
e
ri
a
in
250
m
L NB
m
ed
i
a
.
F
ourt
ee
n s
eed
s p
e
r pot w
e
r
e
so
a
k
ed
for
24
hours
,
with s
eed
s th
a
t sink
be
ing
c
onsi
de
r
ed
goo
d
.
P
l
a
nting m
ed
i
a c
onsist
ed
of w
a
sh
ed a
n
d
d
ri
ed
soil
,
fill
ed
with
5
kg soil p
e
r tu
b
.
Sa
lt str
e
ss
(
NaC
l
)
a
t
6
dS
.
m
^-
1
/
60
m
M
,
h
ea
vy
m
e
t
a
ls
Pb a
t
100
ppm
,
a
n
d Cd a
t
25
ppm w
e
r
e added
21
da
ys
a
ft
e
r pl
a
nting
(
DAP
).
Seed
s
b
ioprim
ed
with
e
n
d
ophyti
c bac
t
e
ri
a
w
e
r
e
th
e
n pl
a
nt
ed
in th
e
soil
.
P
l
a
nt
m
a
int
e
n
a
n
ce
in
c
lu
ded
w
a
t
e
ring with w
a
t
e
r
e
v
e
ry
2
da
ys to k
ee
p th
e
soil moist
a
n
d
w
a
t
e
ring with
1
%
bac
t
e
ri
a
l solution
e
v
e
ry
7
da
ys
.
Weed
r
e
mov
a
l w
a
s
a
lso p
e
rform
ed
.
P
l
a
nting
c
ontinu
ed
for
42
da
ys until th
e
ri
ce
s
eed
s
be
g
a
n to grow
.
S
oil
a
n
d c
ompost
s
a
mpl
e
s
be
for
e
pl
a
nting
(
0
DAP
)
w
e
r
e
t
e
st
ed
for moistur
e c
ont
e
nt
acc
or
d
ing to
ASTM
26
D
-
2216
-
71
.
Sa
mpl
e
s w
e
r
e
pl
aced
in pr
e
-
w
e
igh
ed
por
ce
l
a
in
d
ish
e
s
,
d
ri
ed
in
a
n ov
e
n
a
t
105
°
C
for
24
hours until
a
ll w
a
t
e
r
e
v
a
por
a
t
ed
,
a
n
d
th
e
n w
e
igh
ed a
g
a
in to
de
t
e
rmin
e
moistur
e c
ont
e
nt
.
R
i
ce G
rowth
E
v
a
lu
a
tion
P
hysiologi
ca
l qu
a
lity of s
eed
s w
a
s
a
ss
e
ss
ed b
y pl
a
nt h
e
ight
,
b
iom
a
ss w
e
ight
,
vigor
in
de
x
,
ca
nopy
a
n
d
root moistur
e c
ont
e
nt
,
tot
a
l
c
hlorophyll
,
a
n
d
tot
a
l ph
e
noli
c
s
.
Da
t
a
w
e
r
e a
n
a
lyz
ed
using
ANOVA
follow
ed b
y th
e BNT
t
e
st
a
t
a
5
%
signifi
ca
n
ce
l
e
v
e
l
.
R
i
ce
pl
a
nts from
4
s
eed
s p
e
r tu
b
w
e
r
e ca
r
e
fully h
a
rv
e
st
ed
,
c
l
ea
n
ed
with
d
istill
ed
w
a
t
e
r
,
a
n
d
m
ea
sur
e
m
e
nts for h
e
ight
a
n
d b
iom
a
ss w
e
ight p
e
r pl
a
nt w
e
r
e
r
ec
or
ded
.
C
l
ea
n
a
n
d
d
ri
ed
ri
ce
pl
a
nt s
a
mpl
e
s
a
r
e ca
r
e
fully s
e
p
a
r
a
t
ed
into th
e
shoot
a
n
d
root s
ec
tions
.
T
h
e
moistur
e c
ont
e
nt of th
e
shoot
a
n
d
root of th
e
ri
ce
pl
a
nts is t
e
st
ed
using th
e
s
a
m
e
pro
ced
ur
e a
s for soil
a
n
d c
ompost moistur
e c
ont
e
nt
be
for
e
ri
ce
pl
a
nting
.
T
ot
a
l
c
hlorophyll is
a
ss
e
ss
ed b
y
e
xtr
ac
ting
c
hlorophyll from fr
e
sh l
ea
v
e
s
.
T
h
e
l
ea
v
e
s
a
r
e c
ut
into sm
a
ll pi
ece
s
a
n
d
w
e
igh
ed
using
a
n
a
n
a
lyti
ca
l
ba
l
a
n
ce
to
1
g p
e
r pl
a
nt
.
T
h
e
l
ea
v
e
s
a
r
e
th
e
n pl
aced
in
a
n
E
rl
e
nm
e
y
e
r fl
a
sk with
20
m
L
of
ace
ton
e
.
T
h
e ace
ton
e e
xtr
ac
t is
ce
ntrifug
ed
,
a
n
d
1
m
L
is tr
a
nsf
e
rr
ed
to
a
t
e
st tu
be
.
T
h
e
solution is mix
ed
with
2
m
L
of
d
istill
ed
w
a
t
e
r
,
a
n
d ab
sor
ba
n
ce
is m
ea
sur
ed a
t w
a
v
e
l
e
ngths of
646
nm
a
n
d
663
nm
.
A
ll
pro
ced
ur
e
s
a
r
e ca
rri
ed
out in th
e ab
s
e
n
ce
of sunlight
.
T
ot
a
l ph
e
noli
c c
ont
e
nt is
m
ea
sur
ed
using th
e F
olin
-
C
io
ca
lt
e
u r
ea
g
e
nt
.
F
r
e
sh l
ea
v
e
s
a
r
e d
ri
ed
in
a
n ov
e
n
a
t
60
°
C
for
24
hours
,
th
e
n
c
hopp
ed
into sm
a
ll pi
ece
s
.
1
g of l
ea
v
e
s is mix
ed
with
20
m
L
of
80
%
m
e
th
a
nol in
a
n
E
rl
e
nm
e
y
e
r fl
a
sk
.
T
h
e e
xtr
ac
t solution is sh
a
k
e
n on
a
sh
a
k
e
r for
24
hours
.
T
h
e
m
e
th
a
nol
e
xtr
ac
t is
ce
ntrifug
ed
,
a
n
d
500
µ
L
is tr
a
nsf
e
rr
ed
to
a
t
e
st tu
be
.
T
h
e
solution is th
e
n mix
ed
with
2
.
5
m
L
of
10
%
F
olin
a
n
d
2
m
L
of
7
.
5
%
Na
2
CO
3
.
T
h
e
e
xtr
ac
t solution is h
ea
t
ed a
t
45
°
C
for
15
minut
e
s
.
Ab
sor
ba
n
ce
is m
ea
sur
ed a
t
a
27
w
a
v
e
l
e
ngth of
750
nm using
a
sp
ec
trophotom
e
t
e
r
.
T
ot
a
l ph
e
noli
c c
ont
e
nt is o
b
t
a
in
ed
b
y
c
omp
a
ring it to
a
ph
e
noli
c
st
a
n
da
r
d c
urv
e
.
T
h
e
st
a
n
da
r
d
solution us
ed
is g
a
lli
c ac
i
d
solution
.
Te
sting for
Pb a
n
d Cd
in soil us
ed
for ri
ce c
ultiv
a
tion is
c
on
d
u
c
t
ed
on
da
ys
0
a
n
d
42
.
T
h
e
soil is
d
ri
ed
in
a
n ov
e
n
a
t
60
°
C
for
24
hours
.
3
g of
d
ry soil is w
e
igh
ed a
n
d
pl
aced
in
a bea
k
e
r
.
T
h
e
soil is mix
ed
with
10
m
L
of
1
:
1
HC
l
,
10
m
L
of
1
:
1
HNO
3
,
15
m
L
of
d
istill
ed
w
a
t
e
r
,
a
n
d
4
-
5
d
rops of
30
%
H
2
O
2
,
th
e
n stirr
ed
.
T
h
e
mixtur
e
is
d
ri
ed
in
a
w
a
t
e
r
ba
th
,
th
e
n
10
m
L
of
1
:
1
HC
l
a
n
d
40
m
L
of
d
istill
ed
w
a
t
e
r
a
r
e added
.
T
h
e
mixtur
e
is h
ea
t
ed
in th
e
w
a
t
e
r
ba
th for
10
minut
e
s
,
th
e
n filt
e
r
ed a
n
d
rins
ed
with
30
m
L
of
0
.
3
N
HC
l
.
T
h
e
filtr
a
t
e
is
d
ri
ed
in
a
w
a
t
e
r
ba
th
,
th
e
n
10
m
L
of
1
:
1
HC
l is
added a
n
d
stirr
ed
.
T
h
e
solution is tr
a
nsf
e
rr
ed
to
a
100
m
L
volum
e
tri
c
fl
a
sk
,
ad
just
ed
with
0
.
3
N HC
l
,
a
n
d
homog
e
niz
ed
.
T
h
e
pr
e
p
a
r
ed
s
a
mpl
e
is th
e
n t
e
st
ed
for
Pb a
n
d Cd c
ont
e
nt using
a
n
a
tomi
c ab
sorption sp
ec
trophotom
e
t
e
r
(
AAS
)
a
t
a
w
a
v
e
l
e
ngth of
283
.
3
nm
.
28
RESULT
E
n
d
ophyti
c Bac
t
e
ri
a
E
n
d
ophyti
c bac
t
e
ri
a a
r
e
mi
c
roorg
a
nisms th
a
t liv
e
sym
b
ioti
ca
lly
within th
e
ir host
'
s tissu
e
s without
ca
using
d
is
ea
s
e
.
T
h
e
s
e bac
t
e
ri
a a
r
e
known to
be
n
e
fit th
e
ir host pl
a
nts
b
y in
c
r
ea
sing tol
e
r
a
n
ce
to
e
nvironm
e
nt
a
l str
e
ss
,
e
nh
a
n
c
ing nutrition
,
or promoting pl
a
nt growth
.
I
sol
a
tion of
E
n
d
ophyti
c Bac
t
e
ri
a
I
n this stu
d
y
,
e
n
d
ophyti
c bac
t
e
ri
a
w
e
r
e
isol
a
t
ed
from
S
onn
e
r
a
ti
a
a
l
ba
m
a
ngrov
e
l
ea
v
e
s
c
oll
ec
t
ed
from
Te
luk
Ra
t
a
i
.
Te
luk
Ra
t
a
i
,
lo
ca
t
ed
in
S
outh
La
mpung
Ba
y
,
is r
ec
ogniz
ed a
s
a
n optim
a
l
a
r
ea
for m
a
ngrov
e
growth
.
T
h
e
m
a
ngrov
e
l
ea
f s
a
mpl
e
s w
e
r
e
tr
a
nsport
ed
to th
e PAIR
-
BATAN
l
ab
or
a
tory for
e
n
d
ophyti
c bac
t
e
ri
a
isol
a
tion
.
I
sol
a
tion w
a
s p
e
rform
ed
using two m
ed
i
a
:
Med
i
a A
(
N
utri
e
nt
B
roth
a
n
d d
istill
ed
w
a
t
e
r
)
a
n
d Med
i
a B
(
N
utri
e
nt
B
roth
,
NaC
l
,
Pb
,
a
n
d
Cd
).
T
h
e
purpos
e
of using two m
ed
i
a
w
a
s to
a
ss
e
ss th
e bac
t
e
ri
a
’
s
ab
ility to surviv
e
un
de
r
b
oth str
e
ss
-
fr
ee a
n
d
str
e
ss
ed c
on
d
itions
.
Bac
t
e
ri
a
l isol
a
tion involv
ed d
ilutions up to
10
tim
e
s to f
ac
ilit
a
t
e
su
b
s
e
qu
e
nt purifi
ca
tion
.
Re
sults show
ed d
iff
e
r
e
n
ce
s
be
tw
ee
n th
e
two m
ed
i
a
.
Med
i
a A
,
whi
c
h
d
i
d
not in
c
lu
de
s
a
lt or h
ea
vy m
e
t
a
l str
e
ss
,
support
ed
th
e
growth
of num
e
rous
bac
t
e
ri
a
l
c
oloni
e
s
.
C
onv
e
rs
e
ly
,
Med
i
a B
,
whi
c
h in
c
lu
ded
NaC
l
,
Pb
,
a
n
d Cd
str
e
ssors
,
support
ed
only
a
f
e
w
c
oloni
e
s
,
in
d
i
ca
ting
th
a
t s
a
lt
a
n
d
h
ea
vy m
e
t
a
l str
e
ss inhi
b
it
bac
t
e
ri
a
l surviv
a
l
.
C
oloni
e
s in
29
b
oth m
ed
i
a
w
e
r
e
o
b
s
e
rv
ed
visu
a
lly
a
n
d c
ount
ed
using th
e T
ot
a
l
P
l
a
t
e
C
ount
(
TPC
)
m
e
tho
d
.
TPC
r
e
sults for
d
ilutions
1
through
8
e
x
ceeded
363
bac
t
e
ri
a
l
c
oloni
e
s
,
surp
a
ssing th
e
m
a
ximum
c
ounting thr
e
shol
d a
n
d
w
e
r
e
not
c
ount
ed
furth
e
r
.
T
h
e
num
be
r of
c
oloni
e
s p
e
r s
a
mpl
e a
n
d d
ilution f
ac
tor
a
lign
ed
with th
e d
ilution prin
c
ipl
e
—
high
e
r
d
ilution f
ac
tors r
e
sult
ed
in
f
e
w
e
r
bac
t
e
ri
a
l
c
oloni
e
s
.
I
n
Med
i
a A
,
d
ilution
9
yi
e
l
ded
363
c
oloni
e
s
,
a
n
d d
ilution
10
h
ad
200
c
oloni
e
s
.
I
n
Med
i
a B
,
d
ilution
9
pro
d
u
ced
4
c
oloni
e
s
,
a
n
d d
ilution
10
h
ad
5
c
oloni
e
s
.
T
h
e
purifi
ed e
n
d
ophyti
c
bac
t
e
ri
a
l isol
a
t
e
w
a
s from
Med
i
a B
,
a
s it surviv
ed
un
de
r s
a
lt
a
n
d
h
ea
vy
m
e
t
a
l str
e
ss
c
on
d
itions
.
P
urifi
ca
tion w
a
s only p
e
rform
ed
on isol
a
t
e
s from
d
ilutions
9
a
n
d
10
,
tot
a
ling
9
isol
a
t
e
s
.
C
h
a
r
ac
t
e
riz
a
tion of
E
n
d
ophyti
c Bac
t
e
ri
a
Bac
t
e
ri
a
l
M
orphology
N
in
e e
n
d
ophyti
c bac
t
e
ri
a
l isol
a
t
e
s with
c
o
de
s
B
91
,
B
92
,
B
93
,
B
94
,
B
101
,
B
102
,
B
103
,
B
104
,
a
n
d B
105
w
e
r
e
o
b
s
e
rv
ed
morphologi
ca
lly
.
Bac
t
e
ri
a
l morphology w
a
s
e
x
a
min
ed
using
a
mi
c
ros
c
op
e
with
1000
x
m
a
gnifi
ca
tion
,
e
mploying th
e G
r
a
m st
a
ining m
e
tho
d
.
T
h
e G
r
a
m
st
a
ining pro
ce
ss involv
e
s
a
pplying
c
ryst
a
l viol
e
t
,
s
a
fr
a
nin
,
io
d
in
e
,
a
l
c
ohol
,
a
n
d d
istill
ed
w
a
t
e
r
.
C
ryst
a
l viol
e
t s
e
rv
e
s
a
s th
e
prim
a
ry st
a
in
,
imp
a
rting
a
purpl
e c
olor to th
e bac
t
e
ri
a
,
whil
e
s
a
fr
a
nin
ac
ts
a
s
a
s
ec
on
da
ry st
a
in
,
giving
a
r
ed c
olor
.
I
o
d
in
e e
nh
a
n
ce
s
c
olor
b
in
d
ing
,
a
l
c
ohol rins
e
s
a
w
a
y
e
x
ce
ss prim
a
ry st
a
in
,
a
n
d d
istill
ed
w
a
t
e
r
c
l
ea
ns up
30
r
e
si
d
u
a
l
c
ryst
a
l viol
e
t
,
io
d
in
e
,
a
n
d
s
a
fr
a
nin
a
ft
e
r st
a
ining
.
Bac
t
e
ri
a
with simil
a
r sh
a
p
e
s
a
n
d c
olors w
e
r
e
group
ed
tog
e
th
e
r
.
T
his stu
d
y i
de
ntifi
ed
four groups of
e
n
d
ophyti
c bac
t
e
ri
a
,
with on
e
r
e
pr
e
s
e
nt
a
tiv
e
from
eac
h group s
e
l
ec
t
ed ba
s
ed
on
c
olor
a
n
d
sh
a
p
e
.
Bac
t
e
ri
a
with
c
o
de B
102
r
e
s
e
m
b
l
ed
thos
e
of
B
91
,
B
103
a
n
d B
104
r
e
s
e
m
b
l
ed B
92
,
a
n
d B
93
a
n
d B
94
r
e
s
e
m
b
l
ed B
105
.
T
h
e bac
t
e
ri
a
s
e
l
ec
t
ed
for furth
e
r
c
h
a
r
ac
t
e
riz
a
tion w
e
r
e B
91
,
B
92
,
B
101
,
a
n
d B
105
.
B
io
c
h
e
mistry
T
h
e
t
e
sts in
c
lu
de
nitrog
e
n fix
a
tion
,
phosph
a
t
e
solu
b
iliz
a
tion
,
a
n
d IAA
pro
d
u
c
tion
.
T
h
e
s
e
t
e
sts
a
im to i
de
ntify th
e be
st
e
n
d
ophyti
c bac
t
e
ri
a
th
a
t
ca
n
be
us
ed a
s s
eed b
iopriming
a
g
e
nts for ri
ce
pl
a
nts
.
T
h
e
first
t
e
st is th
e
nitrog
e
n
-
fixing
ab
ility of
e
n
d
ophyti
c bac
t
e
ri
a
.
T
his t
e
st
de
t
e
rmin
e
s th
e ab
ility of
e
n
d
ophyti
c bac
t
e
ri
a
to fix nitrog
e
n from th
e
a
ir
(
N
2
).
Bac
t
e
ri
a ca
p
ab
l
e
of fixing
N
2
from th
e a
ir
c
onv
e
rt it into
NO
3
-
within pl
a
nt tissu
e
s through th
e e
nzym
e
nitrog
e
n
a
s
e
.
T
h
e ab
ility of
e
n
d
ophyti
c bac
t
e
ri
a
l isol
a
t
e
s to fix nitrog
e
n v
a
ri
e
s
,
with
c
on
ce
ntr
a
tions
r
a
nging from
35
.
38
to
37
.
12
ppm
.
T
h
e
isol
a
t
e B
105
show
ed
th
e
high
e
st
nitrog
e
n
c
on
ce
ntr
a
tion
a
t
37
.
12
ppm
.
T
his
c
on
ce
ntr
a
tion is signifi
ca
ntly
high
e
r th
a
n th
a
t foun
d
in
e
n
d
ophyti
c bac
t
e
ri
a
from oth
e
r pl
a
nts
,
su
c
h
a
s th
e
high
e
st nitrog
e
n
c
on
ce
ntr
a
tion of
15
.
18
ppm foun
d
in
B
1
isol
a
t
e
from sw
ee
t pot
a
to roots
.
T
h
e
s
ec
on
d
t
e
st
e
v
a
lu
a
t
e
s th
e
phosph
a
t
e
solu
b
ilizing
ab
ility of
e
n
d
ophyti
c bac
t
e
ri
a
l isol
a
t
e
s
.
P
hosph
a
t
e
-
solu
b
ilizing
bac
t
e
ri
a
h
e
lp provi
de
phosphorus
b
y
c
onv
e
rting
b
oun
d
phosph
a
t
e
s with
e
l
e
m
e
nts lik
e Ca
,
A
l
,
a
n
d Fe
into solu
b
l
e
forms
31
through org
a
ni
c ac
i
d
s
,
m
a
king phosphorus
a
v
a
il
ab
l
e
to pl
a
nts
.
P
hosphorus
(
P
)
is
a
m
ac
ro nutri
e
nt
e
ss
e
nti
a
l for pl
a
nt growth
,
though
its
a
v
a
il
ab
ility in soil is g
e
n
e
r
a
lly low
e
r
c
omp
a
r
ed
to nitrog
e
n
(
N
),
pot
a
ssium
(
K
),
a
n
d ca
l
c
ium
(
Ca
).
T
h
e
r
e
sults show th
a
t
a
ll t
e
st
ed
e
n
d
ophyti
c bac
t
e
ri
a
l isol
a
t
e
s
ca
n solu
b
iliz
e
phosph
a
t
e
.
T
h
e B
91
isol
a
t
e
h
ad
th
e
high
e
st solu
b
l
e
phosphorus
c
on
ce
ntr
a
tion
(
101
.
21
ppm
)
c
omp
a
r
ed
to oth
e
r isol
a
t
e
s su
c
h
a
s
B
92
,
B
101
,
a
n
d B
105
.
T
h
e
low
e
st
solu
b
l
e
phosphorus
c
on
ce
ntr
a
tion w
a
s foun
d
in isol
a
t
e B
101
a
t
66
.
67
ppm
.
T
h
e
thir
d
t
e
st
a
ss
e
ss
e
s
IAA
pro
d
u
c
tion in
e
n
d
ophyti
c bac
t
e
ri
a
l
isol
a
t
e
s
.
I
n
d
ol
e Ace
ti
c Ac
i
d
(
IAA
)
is
a c
ru
c
i
a
l pl
a
nt growth hormon
e
in
th
e a
uxin group
,
pl
a
ying rol
e
s in
ce
ll
d
ivision
,
ce
ll
e
long
a
tion
,
root
growth
,
a
pi
ca
l
d
omin
a
n
ce
,
flow
e
ring
,
l
ea
f
ab
s
c
ission
,
a
n
d
tropi
c
mov
e
m
e
nts
.
T
h
e
r
e
sults in
d
i
ca
t
e
th
a
t
a
ll t
e
st
ed e
n
d
ophyti
c bac
t
e
ri
a
l
isol
a
t
e
s
ca
n pro
d
u
ce IAA
.
T
h
e B
91
isol
a
t
e
h
ad
th
e
high
e
st
IAA
c
on
ce
ntr
a
tion
c
omp
a
r
ed
to
B
92
,
B
101
,
a
n
d B
105
.
T
h
e
high
e
st
IAA
c
on
ce
ntr
a
tion w
a
s
1
.
15
ppm in
B
91
,
whil
e
th
e
low
e
st w
a
s
0
.
58
ppm in
B
101
a
n
d B
105
.
T
h
e
s
e IAA
v
a
lu
e
s
a
r
e
low
e
r
c
omp
a
r
ed
to
a
stu
d
y wh
e
r
e
e
n
d
ophyti
c bac
t
e
ri
a
of th
e
g
e
nus
S
t
a
phylo
c
o
cc
us h
ad IAA
l
e
v
e
ls of
57
ppm
.
T
h
e IAA
t
e
st us
ed a
nutri
e
nt
b
roth m
ed
ium suppl
e
m
e
nt
ed
with
L
-
tryptoph
a
n
,
a
n
a
mino
ac
i
d
pr
ec
ursor in
IAA b
iosynth
e
sis
.
T
his
add
ition
a
ims to stimul
a
t
e bac
t
e
ri
a
l
IAA
pro
d
u
c
tion
.
L
-
tryptoph
a
n is
a
pr
ec
ursor
in
IAA b
iosynth
e
sis in
b
oth
bac
t
e
ri
a a
n
d
pl
a
nts
.
T
h
e
r
ed c
olor form
ed
a
ft
e
r
add
ing
Sa
lkowski r
ea
g
e
nt in
d
i
ca
t
e
s th
e
int
e
r
ac
tion of
IAA
with
Fe
,
forming th
e c
ompl
e
x
[
Fe
2
(
OH
)
2
(
IA
)
4
],
wh
e
r
e IA
is in
d
ol
e
-
3
-
ace
t
a
t
e
.
A
32
mor
e
int
e
ns
e
r
ed c
olor in
d
i
ca
t
e
s high
e
r
IAA c
on
ce
ntr
a
tion
.
Ba
s
ed
on th
e b
io
c
h
e
mi
ca
l physiologi
ca
l
c
h
a
r
ac
t
e
riz
a
tion
(
nitrog
e
n
,
solu
b
l
e P
,
a
n
d IAA
t
e
sts
)
c
on
d
u
c
t
ed
on th
e
four isol
a
t
e
s
(
B
91
,
B
92
,
B
101
,
a
n
d B
105
),
th
e be
st
b
iopriming
a
g
e
nt is isol
a
t
e B
91
.
B
91
h
a
s th
e
high
e
st
solu
b
l
e P a
n
d IAA c
on
ce
ntr
a
tions
a
t
101
.
21
ppm
a
n
d
1
.
15
ppm
,
r
e
sp
ec
tiv
e
ly
.
S
olu
b
l
e P a
ff
ec
ts s
eed
growth
a
n
d
root stru
c
tur
e
,
whil
e
IAA
influ
e
n
ce
s
ce
ll
a
n
d
root
e
long
a
tion
,
whi
c
h
ca
n
be
o
b
s
e
rv
ed
in th
e
shoot
a
n
d
root l
e
ngths
.
N
itrog
e
n
a
ff
ec
ts
c
hlorophyll form
a
tion
,
m
ea
sur
ab
l
e b
y th
e
tot
a
l
c
hlorophyll in l
ea
v
e
s
.
T
h
e E
ff
ec
ts of
Sa
linity
a
n
d Hea
vy
Me
t
a
l
S
tr
e
ss on
R
i
ce Seed
s
T
h
e
thr
e
shol
d
s for s
a
linity
a
n
d
h
ea
vy m
e
t
a
l str
e
ss w
e
r
e
de
t
e
rmin
ed
through str
e
ss v
a
ri
a
tion t
e
sting
.
T
h
e
m
e
tho
d
us
ed
for this
t
e
st is th
e b
lott
e
r t
e
st
,
a
t
ec
hniqu
e
for isol
a
ting fungi
b
y
a
llowing th
e
m
to grow on filt
e
r p
a
p
e
r
(
b
lott
e
r
)
moist
e
n
ed
with st
e
ril
e d
istill
ed
w
a
t
e
r
.
T
h
e b
lott
e
r t
e
st is
ad
v
a
nt
a
g
e
ous
beca
us
e
it not only i
de
ntifi
e
s surf
ace
fungi
b
ut
a
lso off
e
rs
a
pr
ac
ti
ca
l
a
ppro
ac
h for th
e
pro
ced
ur
e
.
S
tr
e
ss
v
a
ri
a
tions t
e
st
ed
in
c
lu
ded NaC
l s
a
lt
a
n
d
h
ea
vy m
e
t
a
ls
Pb a
n
d Cd a
t
c
on
ce
ntr
a
tions of
0
,
25
,
50
,
100
,
200
,
300
,
400
,
a
n
d
500
ppm
.
S
t
e
riliz
ed
ri
ce
s
eed
s
,
tr
ea
t
ed
with
2
%
NaOC
l
,
w
e
r
e
th
e
n so
a
k
ed
in
str
e
ss solutions
a
t
eac
h
c
on
ce
ntr
a
tion for
24
hours to
a
ss
e
ss th
e
e
ff
ec
ts of s
a
lts
a
n
d
h
ea
vy m
e
t
a
ls on ri
ce
s
eed
g
e
rmin
a
tion
a
t sp
ec
ifi
c
c
on
ce
ntr
a
tions
.
R
i
ce
s
eed
s so
a
k
ed
in str
e
ss solutions w
e
r
e
pl
a
nt
ed
on
filt
e
r p
a
p
e
r in
Pe
tri
d
ish
e
s
,
with
20
s
eed
s p
e
r
d
ish to
a
voi
d
int
e
rf
e
r
e
n
ce
d
uring g
e
rmin
a
tion
.
T
h
e Pe
tri
d
ish
e
s w
e
r
e
l
abe
l
ed a
n
d
k
e
pt in
a c
los
ed
,
33
da
rk
e
nvironm
e
nt to promot
e
growth
,
a
s pl
a
nts grow f
a
st
e
r
d
u
e
to
in
c
r
ea
s
ed a
uxin hormon
e
l
e
v
e
ls
,
whi
c
h influ
e
n
ce
st
e
m
e
long
a
tion
.
O
n
th
e
5
th
da
y
,
th
e
l
e
ngth of
eac
h g
e
rmin
a
t
ed
s
eed
ling from root to
sprout tip w
a
s m
ea
sur
ed
using
a
20
c
m rul
e
r
.
T
h
e
num
be
r of
g
e
rmin
a
t
ed
s
eed
s w
a
s
a
lso
c
ount
ed
to
de
t
e
rmin
e
th
e
s
eed
vigor in
de
x
.
Mea
sur
e
m
e
nts of s
eed
ling l
e
ngth
a
n
d
g
e
rmin
a
tion
c
ounts
a
r
e
shown in
A
pp
e
n
d
ix
3
.
Da
t
a a
n
a
lysis in
d
i
ca
t
e
s th
a
t ri
ce
s
eed
s
be
g
a
n to struggl
e
with st
e
m
e
long
a
tion un
de
r s
a
lt
a
n
d
h
ea
vy m
e
t
a
l str
e
ss
c
on
ce
ntr
a
tions
ab
ov
e
200
ppm
,
sp
ec
ifi
ca
lly
a
t
300
,
400
,
a
n
d
500
ppm
.
T
h
e da
t
a a
lso
shows th
a
t ri
ce
s
eed
s h
ad
signifi
ca
nt
d
iffi
c
ulty g
e
rmin
a
ting
a
t
400
a
n
d
500
ppm
.
A
t
400
ppm str
e
ss
,
only
4
out of
20
s
eed
s g
e
rmin
a
t
ed
ac
ross thr
ee
r
e
p
e
titions
.
A
t
500
ppm
,
non
e
of th
e
20
s
eed
s g
e
rmin
a
t
ed
.
T
h
e
vigor in
de
x r
e
f
e
rs to th
e
uniformity
a
n
d
sp
eed
of s
eed
g
e
rmin
a
tion ov
e
r
a
sp
ec
ifi
c
p
e
rio
d
.
Ob
s
e
rv
a
tions for
ca
l
c
ul
a
ting th
e
vigor in
de
x
a
r
e
m
ade
on th
e
first o
b
s
e
rv
a
tion
(
da
y
5
)
b
y
c
ounting th
e
num
be
r of norm
a
l s
eed
lings
.
P
l
a
nts su
b
j
ec
t
ed
to high
c
on
ce
ntr
a
tions of
NaC
l s
a
lt str
e
ss fin
d
it mor
e c
h
a
ll
e
nging to grow
.
T
his
d
iffi
c
ulty
a
ris
e
s
beca
us
e e
x
ce
ss s
a
lt r
ed
u
ce
s w
a
t
e
r
a
v
a
il
ab
ility for pl
a
nt growth
d
u
e
to
osmoti
c
pr
e
ssur
e
on th
e
roots
,
d
isrupting th
e
photosynth
e
sis pro
ce
ss
.
Add
ition
a
lly
,
e
x
ce
ss s
a
lt
ca
n l
ead
to ion toxi
c
ity
.
I
n
c
r
ea
s
ed Na
+
l
e
v
e
ls
ca
n r
ed
u
ce
th
e ab
sorption of nitrog
e
n
(
N
)
a
n
d
phosphorus
(
P
),
whi
c
h
a
r
e e
ss
e
nti
a
l for s
eed
ling growth
.
Hea
vy m
e
t
a
ls lik
e Pb a
n
d Cd a
t high
c
on
ce
ntr
a
tions
ca
n int
e
r
ac
t
with
e
nzym
e
s
,
d
isrupting pl
a
nt m
e
t
ab
olism
.
Cd ca
n
b
in
d
with
e
ss
e
nti
a
l
nutri
e
nts su
c
h
a
s
P
,
pot
a
ssium
(
K
),
a
n
d
iron
(
Fe
),
limiting th
e
ir
34
ab
sorption
.
Pb e
nt
e
ring pl
a
nt tissu
e
s
ca
n
b
in
d
with
ce
ll m
e
m
b
r
a
n
e
s
,
mito
c
hon
d
ri
a
,
a
n
d c
hloropl
a
sts
,
int
e
rf
e
ring with s
eed
ling growth
.
B
iopriming
A
ppli
ca
tion of
E
n
d
ophyti
c Bac
t
e
ri
a B
iopriming on
R
i
ce Seed
s
T
h
e
soil us
ed
for ri
ce
pl
a
nting
c
om
e
s from th
e PAIR
-
BATAN
l
ab
or
a
tory
.
T
h
e
soil is fin
e
ly groun
d
so th
a
t s
a
lt
a
n
d
h
ea
vy m
e
t
a
l str
e
ss
ca
n
be e
v
e
nly
d
istri
b
ut
ed a
mong
eac
h ri
ce
s
eed
in th
e c
ont
a
in
e
r
.
C
ompost is
added
to th
e
soil
a
s
add
ition
a
l nutrition for th
e
ri
ce
s
eed
s
.
B
iopriming of ri
ce
s
eed
s is
ca
rri
ed
out
b
y in
d
u
c
ing th
e B
91
bac
t
e
ri
a
l isol
a
t
e
for
24
hours
.
Bac
t
e
ri
a B
91
is
c
h
a
r
ac
t
e
riz
ed b
y its
sup
e
rior
ab
ility to
e
nh
a
n
ce
growth
ba
s
ed
on
IAA
,
phosph
a
t
e
,
a
n
d
nitrog
e
n t
e
sts
.
Sa
lt str
e
ss with
NaC
l
a
n
d
h
ea
vy m
e
t
a
ls
Pb a
n
d Cd a
r
e
a
ppli
ed a
t
21
Da
ys
A
ft
e
r
P
l
a
nting
(
DAP
).
T
his is to
e
nsur
e
th
a
t th
e
pl
a
nts
ca
n go through th
e ea
rly v
e
g
e
t
a
tiv
e
ph
a
s
e
so th
a
t ri
ce
e
v
a
lu
a
tion un
de
r str
e
ss
c
on
d
itions
ca
n
be
p
e
rform
ed
.
T
h
e
s
a
lt str
e
ss
us
ed
is
a
t th
e
m
a
ximum limit of
6
dS
.
m
-
1
/
60
m
M
.
T
h
e
h
ea
vy m
e
t
a
l
str
e
ss l
e
v
e
ls
a
r
e
100
ppm for
Pb a
n
d
25
ppm for
Cd
.
R
i
ce
s
eed
s
a
r
e
pl
a
nt
ed
in
18
c
ont
a
in
e
rs
,
eac
h with
14
s
eed
s
.
T
h
e
pl
a
nting is
d
on
e
in
a
gr
ee
nhous
e
to
be
tt
e
r
c
ontrol f
ac
tors su
c
h
a
s t
e
mp
e
r
a
tur
e
,
humi
d
ity
,
a
n
d
light
.
T
his
a
ims to improv
e
pl
a
nt yi
e
l
d a
n
d
qu
a
lity
,
r
ed
u
ce
p
e
st
a
n
d
d
is
ea
s
e
risks
,
a
n
d
minimiz
e
th
e
imp
ac
t of
ad
v
e
rs
e
w
ea
th
e
r
c
on
d
itions
.
35
E
v
a
lu
a
tion of
R
i
ce G
rowth
R
i
ce
s
eed
s pl
a
nt
ed
in soil using
a C
ompl
e
t
e
ly
Ra
n
d
omiz
ed De
sign
(
CRD
)
w
e
r
e e
v
a
lu
a
t
ed
to
de
t
e
rmin
e
th
e
physiologi
ca
l qu
a
lity of th
e
pl
a
nts
.
I
n this stu
d
y
,
ri
ce
pl
a
nt growth w
a
s
a
ss
e
ss
ed ba
s
ed
on s
e
v
e
r
a
l
p
a
r
a
m
e
t
e
rs in
c
lu
d
ing ri
ce
h
e
ight
a
t
7
–
42
Da
ys
A
ft
e
r
P
l
a
nting
(
DAP
),
vigor in
de
x
a
t
7
DAP
,
l
e
ngth
a
n
d
w
e
ight of whol
e
pl
a
nts
,
p
a
ni
c
l
e
s
,
a
n
d
roots
a
t
21
a
n
d
42
DAP
,
a
n
d
moistur
e c
ont
e
nt of p
a
ni
c
l
e
s
a
n
d
roots
a
t
21
a
n
d
42
DAP
,
a
s w
e
ll
a
s tot
a
l
c
hlorophyll
a
n
d
ph
e
noli
c c
ont
e
nt of ri
ce
l
ea
v
e
s
a
t
42
DAP
.
R
i
ce He
ight
a
t
7
,
14
,
21
,
28
,
35
,
a
n
d
42
DAP
P
l
a
nt h
e
ight is on
e
of th
e
growth in
d
i
ca
tors us
ed
to m
ea
sur
e
growth influ
e
n
ced b
y
e
nvironm
e
nt
a
l f
ac
tors
.
T
his is
beca
us
e
pl
a
nt
h
e
ight is th
e
most visi
b
l
e
p
a
r
a
m
e
t
e
r th
a
t
ca
n
be
m
ea
sur
ed
without
da
m
a
ging th
e
s
a
mpl
e
pl
a
nts
.
T
h
e
h
e
ight of ri
ce
pl
a
nts with priming
tr
ea
tm
e
nt
a
n
d
str
e
ss
add
ition th
a
t gr
e
w up to
42
DAP ca
n
be
s
ee
n in
F
igur
e
7
.
T
h
e
t
a
ll
e
st ri
ce
h
e
ight
a
t
21
DAP
w
a
s
B
1
C
2
a
t
38
.
15
c
m
,
whil
e
th
e
short
e
st h
e
ight w
a
s
B
0
C
0
a
t
32
.
48
c
m
(
F
igur
e
7
).
T
his is
c
onsist
e
nt
with stu
d
i
e
s in
d
i
ca
ting th
a
t
e
n
d
ophyti
c bac
t
e
ri
a ca
n in
c
r
ea
s
e
pl
a
nt
h
e
ight through
b
iopriming m
e
tho
d
s
.
T
h
e
in
c
r
ea
s
e
in pl
a
nt h
e
ight is
susp
ec
t
ed
to
be d
u
e
to th
e IAA
phytohormon
e
pro
d
u
ced b
y
e
n
d
ophyti
c
bac
t
e
ri
a
in th
e
pl
a
nt tissu
e
s
.
Sa
lt
a
n
d
h
ea
vy m
e
t
a
l str
e
ss
e
s w
e
r
e added
a
t
21
DAP
,
so th
e
h
e
ight of th
e
ri
ce
w
a
s only influ
e
n
ced b
y
b
iopriming
with
bac
t
e
ri
a B
91
in th
e
s
eed
s
a
n
d
w
a
t
e
ring with
a
1
%
bac
t
e
ri
a
l
solution
a
t
7
a
n
d
14
DAP
.
36
T
h
e
t
a
ll
e
st ri
ce
h
e
ight
a
t
42
DAP
w
a
s
B
0
C
0
a
t
63
.
67
c
m
,
whil
e
th
e
short
e
st w
a
s
B
0
C
2
a
t
59
.
33
c
m
(
F
igur
e
7
).
T
h
e
ri
ce
h
e
ight from s
eed
s
with
b
iopriming tr
ea
tm
e
nts
B
91
,
in
c
lu
d
ing
B
1
C
0
,
B
1
C
1
,
a
n
d B
1
C
2
,
d
i
d
not
d
iff
e
r signifi
ca
ntly from
B
0
C
0
,
whi
c
h us
ed
only w
a
t
e
r priming
,
a
t
a
5
%
signifi
ca
n
ce
l
e
v
e
l
.
T
his l
ac
k of
d
iff
e
r
e
n
ce
is susp
ec
t
ed
to
be d
u
e
to
th
e
r
ed
u
ced ac
tivity of
bac
t
e
ri
a B
91
on
da
y
42
.
V
igor
I
n
de
x of
R
i
ce a
t
7
DAP
T
h
e
vigor in
de
x is
a
qu
a
lity p
a
r
a
m
e
t
e
r for s
eed
s th
a
t
in
d
i
ca
t
e
s th
e
sp
eed a
n
d
uniformity of s
eed
growth
.
A
high
e
r vigor
in
de
x r
e
fl
ec
ts f
a
st
e
r s
eed
growth
.
T
h
e
r
e
sults of s
eed
g
e
rmin
a
tion
,
st
e
m l
e
ngth
,
g
e
rmin
a
tion
,
a
n
d
vigor in
de
x of ri
ce
s
eed
s
a
t
7
DAP ca
n
be
s
ee
n in
Tab
l
e
5
.
T
h
e
high
e
st vigor in
de
x s
a
mpl
e
w
a
s
B
1
C
1
a
t
894
.
29
c
omp
a
r
ed
to oth
e
r s
a
mpl
e
s
(
Tab
l
e
5
).
T
h
e
low
e
st vigor in
de
x w
a
s
B
0
C
0
a
t
763
.
63
.
T
his in
d
i
ca
t
e
s th
a
t
bac
t
e
ri
a B
91
in
d
u
ced
into ri
ce
s
eed
s positiv
e
ly
a
ff
ec
t
ed
g
e
rmin
a
tion or th
e
num
be
r of s
eed
s
g
e
rmin
a
t
ed a
n
d
th
e
vigor in
de
x
a
t
7
DAP
.
T
his improv
e
m
e
nt
,
though
signifi
ca
nt
,
w
a
s not v
e
ry pronoun
ced
in oth
e
r ri
ce
s
eed
s
a
mpl
e
s
.
E
n
d
ophyti
c bac
t
e
ri
a B
91
ca
n pro
d
u
ce
nitrog
e
n
,
solu
b
l
e
phosphorus
,
a
n
d IAA
in ri
ce
pl
a
nt tissu
e
s
ba
s
ed
on its
b
io
c
h
e
mi
ca
l physiologi
ca
l
c
h
a
r
ac
t
e
risti
c
s
.
T
his r
e
sults in in
c
r
ea
s
ed a
v
a
il
ab
ility of nutri
e
nts n
eeded
for s
eed
g
e
rmin
a
tion
a
n
d
growth
,
thus
e
nh
a
n
c
ing s
eed
uniformity
a
n
d
growth sp
eed
.
I
n
c
ontr
a
st
,
ri
ce
s
eed
s
a
mpl
e
s without
e
n
d
ophyti
c
bac
t
e
ri
a
in
d
u
c
tion
d
i
d
not r
ece
iv
e add
ition
a
l nitrog
e
n
,
solu
b
l
e
37
phosphorus
,
a
n
d IAA
,
so th
e
sp
eed a
n
d
uniformity of s
eed
growth
d
i
d
not improv
e
.
Seed
g
e
rmin
a
tion is
a c
ompl
e
x pro
ce
ss involving
morphologi
ca
l
,
physiologi
ca
l
,
a
n
d b
io
c
h
e
mi
ca
l
c
h
a
ng
e
s
.
Ge
rmin
a
tion
be
gins with w
a
t
e
r
ab
sorption
b
y th
e
s
eed
s
a
n
d e
n
d
s with th
e
e
long
a
tion of th
e e
m
b
ryo
a
xis
.
Seed
g
e
rmin
a
tion is influ
e
n
ced b
y
int
e
rn
a
l f
ac
tors
(
g
e
n
e
ti
c
s
,
d
orm
a
n
c
y
,
e
m
b
ryo
)
a
n
d e
xt
e
rn
a
l f
ac
tors
(
t
e
mp
e
r
a
tur
e
,
light
,
a
n
d
soil moistur
e
) (
Ha
n
da
y
a
ni
,
2021
).
38
C
on
c
lusion
II
sol
a
tion of
e
n
d
ophyti
c bac
t
e
ri
a
from
S
onn
e
r
a
ti
a a
l
ba
m
a
ngrov
e
l
ea
v
e
s pro
d
u
ced
four isol
a
t
e
groups
:
B
91
,
B
92
,
B
101
,
a
n
d B
105
,
whi
c
h
a
r
e
r
ed
-
c
olor
ed
(
G
r
a
m
-
n
e
g
a
tiv
e
),
ro
d
-
sh
a
p
ed
,
a
n
d e
xhi
b
it
r
e
sist
a
n
ce
to
NaC
l s
a
lt
a
n
d
h
ea
vy m
e
t
a
ls
Pb a
n
d Cd
.
T
h
e
e
n
d
ophyti
c bac
t
e
ri
a
with th
e be
st r
e
sist
a
n
ce
to s
a
lt
a
n
d
h
ea
vy
m
e
t
a
ls
,
a
s w
e
ll
a
s th
e
high
e
st growth
-
promoting
ca
p
ab
ility for ri
ce
pl
a
nts
,
is
A
l
ca
lig
e
n
e
s f
aeca
lis
(
B
91
),
ba
s
ed
on
a P
solu
b
ility t
e
st of
1
.
15
ppm
a
n
d IAA
of
101
.
21
ppm
.
Seed b
iopriming h
a
s
a
n
e
ff
ec
t on
in
c
r
ea
sing s
eed
vigor in
de
x
a
t
7
Da
ys
A
ft
e
r
P
l
a
nting
(
DAP
),
ri
ce
h
e
ight
a
t
21
DAP
,
shoot
a
n
d
root w
a
t
e
r
c
ont
e
nt
a
t
21
DAP
,
a
n
d
tot
a
l
l
ea
f
c
hlorophyll
a
t
42
DAP
.
Rec
omm
e
n
da
tion
I
t is
ad
vis
ed
to
add a
1
%
solution of
e
n
d
ophyti
c bac
t
e
ri
a
,
25
m
L
eac
h w
ee
k
,
d
uring th
e ea
rly v
e
g
e
t
a
tiv
e
ph
a
s
e
of th
e
pl
a
nt
(
ri
ce a
t
35
da
ys
)
to optimiz
e
th
e ac
tivity of
e
n
d
ophyti
c bac
t
e
ri
a
in
e
nh
a
n
c
ing ri
ce
pl
a
nt growth
.
