Академический Документы
Профессиональный Документы
Культура Документы
o
w
e
r
H
y
d
r
o
p
r
i
m
i
n
g
,
o
s
m
o
p
r
i
m
i
n
g
(
K
N
O
3
)
S
a
l
i
n
i
t
y
a
n
d
d
r
o
u
g
h
t
K
a
y
a
e
t
a
l
.
(
2
0
0
6
)
C
o
t
t
o
n
H
y
d
r
o
p
r
i
m
i
n
g
D
r
o
u
g
h
t
a
n
d
t
h
e
r
m
a
l
s
t
r
e
s
s
M
u
r
u
n
g
u
e
t
a
l
.
(
2
0
0
3
)
,
C
a
s
e
n
a
v
e
a
n
d
T
o
s
e
l
l
i
(
2
0
0
7
)
S
u
g
a
r
c
a
n
e
H
a
l
o
p
r
i
m
i
n
g
(
N
a
C
l
)
S
a
l
i
n
i
t
y
a
n
d
d
r
o
u
g
h
t
P
a
t
a
d
e
e
t
a
l
.
(
2
0
0
9
)
O
n
i
o
n
H
y
d
r
o
p
r
i
m
i
n
g
,
o
s
m
o
p
r
i
m
i
n
g
(
P
E
G
)
L
o
w
g
e
r
m
i
n
a
t
i
o
n
C
a
s
e
i
r
o
e
t
a
l
.
(
2
0
0
4
)
B
r
o
m
u
s
O
s
m
o
p
r
i
m
i
n
g
(
P
E
G
)
S
a
l
i
n
i
t
y
a
n
d
d
r
o
u
g
h
t
T
a
v
i
l
i
e
t
a
l
.
(
2
0
1
1
)
C
e
l
e
r
y
S
o
l
i
d
m
a
t
r
i
x
p
r
i
m
i
n
g
H
i
g
h
t
e
m
p
e
r
a
t
u
r
e
P
a
r
e
r
a
e
t
a
l
.
(
1
9
9
3
)
F
e
n
n
e
l
(
F
o
e
n
i
c
u
l
u
m
v
u
l
g
a
r
e
)
S
a
l
i
c
y
l
i
c
a
c
i
d
S
a
l
i
n
i
t
y
F
a
r
a
h
b
a
k
h
s
h
(
2
0
1
2
)
C
a
u
l
i
o
w
e
r
H
y
d
r
o
p
r
i
m
i
n
g
A
g
i
n
g
F
u
j
i
k
u
r
a
e
t
a
l
.
(
1
9
9
3
)
M
a
i
z
e
S
e
e
d
p
r
i
m
i
n
g
,
c
h
e
m
i
c
a
l
p
r
i
m
i
n
g
(
C
u
S
O
4
Z
n
S
O
4
)
,
o
n
-
f
a
r
m
s
e
e
d
p
r
i
m
i
n
g
S
a
l
i
n
i
t
y
a
n
d
d
r
o
u
g
h
t
M
u
r
u
n
g
u
e
t
a
l
.
(
2
0
0
3
)
,
F
i
n
c
h
-
S
a
v
a
g
e
e
t
a
l
.
(
2
0
0
4
)
,
F
o
t
i
e
t
a
l
.
(
2
0
0
8
)
,
J
a
n
m
o
h
a
m
m
a
d
i
e
t
a
l
.
(
2
0
0
8
)
T
o
m
a
t
o
O
s
m
o
p
r
i
m
i
n
g
(
N
a
C
l
)
,
K
N
O
3
S
a
l
i
n
i
t
y
a
n
d
l
o
w
t
e
m
p
e
r
a
t
u
r
e
s
.
H
e
y
d
e
c
k
e
r
e
t
a
l
.
(
1
9
7
3
)
,
O
z
b
i
n
g
o
l
e
t
a
l
.
(
1
9
9
8
)
,
N
a
w
a
z
e
t
a
l
.
(
2
0
1
1
)
B
a
s
i
l
H
y
d
r
o
p
r
i
m
i
n
g
S
a
l
i
n
i
t
y
F
a
r
a
h
a
n
i
a
n
d
M
a
r
o
u
(
2
0
1
1
)
B
a
r
l
e
y
O
n
-
f
a
r
m
p
r
i
m
i
n
g
S
a
l
t
s
t
r
e
s
s
R
a
s
h
i
d
e
t
a
l
.
(
2
0
0
6
)
W
a
t
e
r
m
e
l
o
n
H
y
d
r
o
p
r
i
m
i
n
g
,
N
a
C
l
p
r
i
m
i
n
g
L
o
w
t
e
m
p
e
r
a
t
u
r
e
S
u
n
g
a
n
d
C
h
i
u
(
1
9
9
5
)
,
D
e
m
i
r
a
n
d
M
a
v
i
(
2
0
0
4
)
A
l
f
a
a
l
f
a
O
s
m
o
p
r
i
m
i
n
g
(
m
a
n
n
i
t
o
l
)
,
h
y
d
r
o
p
r
i
m
i
n
g
S
a
l
t
A
m
o
o
a
g
h
a
i
e
(
2
0
1
1
)
R
i
c
e
H
y
d
r
o
p
r
i
m
i
n
g
,
P
E
G
,
K
C
l
,
C
a
C
l
2
,
a
s
c
o
r
b
a
t
e
,
p
r
i
m
i
n
g
D
r
o
u
g
h
t
B
a
s
r
a
e
t
a
l
.
(
2
0
0
5
)
,
F
a
r
o
o
q
e
t
a
l
.
(
2
0
1
0
)
,
Y
u
a
n
-
Y
u
a
n
e
t
a
l
.
(
2
0
1
0
)
,
Y
a
r
i
a
n
d
S
h
e
i
d
a
i
e
(
2
0
1
1
)
L
e
e
k
H
y
d
r
o
p
r
i
m
i
n
g
,
P
E
G
p
r
i
m
i
n
g
L
o
w
g
e
r
m
i
n
a
t
i
o
n
B
r
o
c
k
l
e
h
u
r
s
t
e
t
a
l
.
(
1
9
8
4
)
,
A
s
h
r
a
f
a
n
d
B
r
a
y
(
1
9
9
3
)
S
u
n
o
w
e
r
C
h
e
m
i
c
a
l
p
r
i
m
i
n
g
(
K
N
O
3
,
a
m
i
n
o
t
r
i
a
z
o
l
)
,
b
i
o
p
r
i
m
i
n
g
A
g
e
i
n
g
S
i
n
g
h
a
n
d
R
a
o
(
1
9
9
3
)
,
K
i
b
i
n
z
a
e
t
a
l
.
(
2
0
1
1
)
,
M
o
e
i
n
z
a
d
e
h
e
t
a
l
.
(
2
0
1
0
)
L
e
s
q
u
e
r
e
l
l
a
f
e
n
d
l
e
r
i
S
e
e
d
p
r
i
m
i
n
g
D
r
o
u
g
h
t
W
i
n
d
a
u
e
r
e
t
a
l
.
(
2
0
0
7
)
C
a
c
t
i
H
y
d
r
o
p
r
i
m
i
n
g
L
o
w
g
e
r
m
i
n
a
t
i
o
n
D
u
b
r
o
v
s
k
y
(
1
9
9
6
)
C
a
p
s
i
c
u
m
H
2
O
2
a
n
d
A
B
A
,
N
a
C
l
p
r
i
m
i
n
g
C
o
l
d
a
n
d
s
a
l
t
s
t
r
e
s
s
Y
a
d
a
v
e
t
a
l
.
(
2
0
1
1
)
M
u
n
g
b
e
a
n
O
s
m
o
p
r
i
m
i
n
g
(
N
a
C
l
)
S
a
l
i
n
i
t
y
R
a
s
h
i
d
e
t
a
l
.
(
2
0
0
4
)
,
S
a
h
a
e
t
a
l
.
(
2
0
1
0
)
B
i
t
t
e
r
g
o
u
r
d
M
o
i
s
t
v
e
r
m
i
c
u
l
i
t
e
,
h
e
a
t
s
h
o
c
k
i
n
g
S
u
b
o
p
t
i
m
a
l
e
n
v
i
r
o
n
m
e
n
t
a
l
e
f
f
e
c
t
s
L
i
n
a
n
d
S
u
n
g
(
2
0
0
1
)
,
H
s
u
e
t
a
l
.
(
2
0
0
3
)
S
o
r
g
h
u
m
O
s
m
o
a
n
d
h
y
d
r
o
p
r
i
m
i
n
g
L
o
w
g
e
r
m
i
n
a
t
i
o
n
M
o
r
a
d
i
a
n
d
Y
o
u
n
e
s
i
(
2
0
0
9
)
M
e
l
o
n
(
C
u
c
u
m
i
s
m
e
l
o
)
N
a
C
l
p
r
i
m
i
n
g
S
a
l
i
n
i
t
y
S
i
v
r
i
t
e
p
e
e
t
a
l
.
(
2
0
0
3
,
2
0
0
5
)
S
p
i
n
a
c
h
O
s
m
o
p
r
i
m
i
n
g
(
P
E
G
)
C
h
i
l
l
i
n
g
a
n
d
d
r
o
u
g
h
t
s
t
r
e
s
s
C
h
e
n
a
n
d
A
r
o
r
a
(
2
0
1
1
)
S
a
l
i
c
o
r
n
i
a
u
t
a
h
e
n
s
i
s
G
r
o
w
t
h
r
e
g
u
l
a
t
o
r
s
S
a
l
i
n
i
t
y
G
u
l
a
n
d
K
h
a
n
(
2
0
0
3
)
Acta Physiol Plant (2013) 35:13811396 1389
1 3
plastic bag and the seeds are removed before radicle
emergence (Callan et al. 1990). Dual application of bene-
cial microorganisms to carrot and onion seed during drum
priming was demonstrated by Bennett and Whipps (2008).
Rhizobacteria are used as inoculants to enhance crop yield
and for biological control of fungal pathogens. Certain
strains of rhizosphere bacteria stimulate plant growth and
are, therefore, called plant growth-promoting rhizobacteria
(PGPR) (Tonelli et al. 2011). In the roots of rice and
tomato plants, mycorrhizal fungi were shown to induce the
accumulation of a number of transcripts and proteins,
respectively, many of which with a predicted function in
plant defense (Sharmila et al. 2000; Guimil et al. 2005;
Pozo and Azcon-Aguilar 2007). In Cicer arietinum L.,
application of PGPR improves the percentage of seed
germination under saline conditions and also increased the
shoot length, root length and dry matter (Mishra et al.
2010). In wheat, seed biopriming with different salinity-
tolerant isolates of Trichoderma were effective in
improving germination percentage and reducing reduction
percentage of germination during salinity stress (Rawat
et al. 2011). Biopriming of sunower seeds with Pseudo-
monas uorescens UTPf76 and UTPf86 enhanced the
ability of seeds to invigorate and seedlings to grow uni-
formly (Moeinzadeh et al. 2010).
Merits and demerits of priming
According to Bradford (1986), priming treatment of seeds
has been shown to improve the germination and emergence
of many species and the seed-priming techniques have
been found effective for better germination and seedling
establishment under controlled conditions (Basra et al.
2005). On-farm seed priming with water is used in the
semi-arid agricultural land of India to grow maize, rice and
chickpeas. Direct benets to all three crops included faster
emergence, better stands and lower incidence of re-sowing,
more vigorous plants, better drought tolerance, earlier
owering, earlier harvest and higher grain yield (Harris
et al. 1999). Interestingly, priming repairs damage of aged
seeds (Bailly et al. 1998; Butler et al. 2009) or seeds
exposed to abiotic stresses such as salinity (Ehsanfar et al.
2006) and also improves the germination performance.
Among various strategies, pre-sowing treatment and
priming of seeds are easy, low-cost, with low-risk and
effective approaches to overcome the environmental stress
problems (Ashraf and Foolad 2005).
Altogether, disadvantages of seed priming should be
lower than advantages in comparison. There are very few
reports about the demerits of seed priming. Induced resis-
tance by priming was found to be benecial against dif-
ferent abiotic stresses, but a precautious state developing in
T
a
b
l
e
1
c
o
n
t
i
n
u
e
d
P
l
a
n
t
P
r
i
m
i
n
g
m
e
t
h
o
d
T
o
l
e
r
a
n
c
e
d
e
v
e
l
o
p
e
d
f
o
r
R
e
f
e
r
e
n
c
e
s
W
h
e
a
t
H
y
d
r
o
p
r
i
m
i
n
g
,
h
a
l
o
p
r
i
m
i
n
g
(
C
a
S
O
4
)
,
c
h
e
m
i
c
a
l
p
r
i
m
i
n
g
(
K
H
2
P
O
4
,
H
2
O
2
,
S
N
P
)
,
G
A
3
p
r
i
m
i
n
g
,
a
u
x
i
n
p
r
i
m
i
n
g
S
a
l
t
a
n
d
d
r
o
u
g
h
t
D
a
s
a
n
d
C
h
o
u
d
h
u
r
y
(
1
9
9
6
)
,
G
h
a
n
a
a
n
d
S
c
h
i
l
l
i
n
g
e
r
(
2
0
0
3
)
,
A
k
b
a
r
i
e
t
a
l
.
(
2
0
0
7
)
,
W
a
h
i
d
e
t
a
l
.
(
2
0
0
7
)
D
u
n
a
l
p
l
a
n
t
(
W
i
t
h
a
n
i
a
s
o
m
n
i
f
e
r
a
)
C
h
e
m
i
c
a
l
p
r
i
m
i
n
g
(
t
r
i
a
d
i
m
e
f
o
n
)
S
a
l
i
n
i
t
y
J
a
l
e
e
l
e
t
a
l
.
(
2
0
0
8
)
P
e
r
i
w
i
n
k
l
e
C
h
e
m
i
c
a
l
p
r
i
m
i
n
g
(
p
a
c
l
o
b
u
t
r
a
z
o
l
)
S
a
l
i
n
i
t
y
J
a
l
e
e
l
e
t
a
l
.
(
2
0
0
7
)
C
a
n
o
l
a
H
y
d
r
o
p
r
i
m
i
n
g
S
a
l
t
a
n
d
d
r
o
u
g
h
t
s
t
r
e
s
s
O
m
i
d
i
e
t
a
l
.
(
2
0
0
9
)
B
e
a
n
B
i
o
p
r
i
m
i
n
g
(
P
G
E
R
)
,
h
y
d
r
o
p
r
i
m
i
n
g
S
a
l
t
s
t
r
e
s
s
B
a
r
a
s
s
i
e
t
a
l
.
(
2
0
0
6
)
,
A
b
e
b
e
a
n
d
M
o
d
i
(
2
0
0
9
)
L
e
t
t
u
c
e
C
h
e
m
i
c
a
l
p
r
i
m
i
n
g
(
K
H
2
P
O
4
)
,
b
i
o
p
r
i
m
i
n
g
(
P
G
E
R
)
S
a
l
t
s
t
r
e
s
s
K
o
r
k
m
a
z
a
n
d
P
i
l
l
(
2
0
0
3
)
,
B
a
r
a
s
s
i
e
t
a
l
.
(
2
0
0
6
)
1390 Acta Physiol Plant (2013) 35:13811396
1 3
the seed right before the germination may adversely affect
the seedling emergence vigor.
Summary
Over the past few years, seed priming has emerged as a
promising strategy in modern stress management because it
protects plants against various abiotic stresses without
heavily affecting tness. Moreover, seed priming offers a
smart, effective and realistic option for effective plant
protection. Drought, salinity, extreme temperatures and
oxidative stress are often interconnected, and may induce
similar damage. As a consequence, these diverse environ-
mental stresses often activate similar cell signaling path-
ways and cellular responses. It is known that seed priming
can activate these signaling pathways in the early stages of
growth and result in faster plant defense responses. The
exact molecular mechanism behind priming is not com-
pletely known, it is speculated that sensitization was
associated with accumulation of inactive signaling proteins
in primed cells. Upon subsequent exposure to abiotic
stresses, a second signaling event could hyperactive the
signaling proteins thereby amplifying signal transduction,
and thus leading to more rapid and/or more intense acti-
vation of defense responses (Conrath et al. 2006).
Besides the above, there are various other views put
forward by other authors on the mechanism behind prim-
ing. According to Nascimento and West (1998), the
increase in germination percentage/seed vigor of primed
seeds is due to reserve mobilization of food materials,
activation and resynthesis of some enzymes and also due to
the increased DNA and RNA synthesis. Priming is also
capable of repairing some of the damages due to seed
erosion which in turn results in increased vigor of primed
seeds (Arif et al. 2008). Seed priming affects the lag phase
of seed germination and thus causes early DNA replication
(Bray et al. 1989). It is desirable to design suitable seed-
priming methods for different crop plants to meet the
challenges of the environment. The current knowledge of
different seed-priming methods practiced in various crops
is summarized in the Table 1.
Author contribution The idea on the writing of this
review and its common structure and organization came
from Jos T. Puthur, who wrote the draft. K.C. Jisha wrote
the main text of the manuscript and collected the necessary
references. K. Vijayakumari supported with supplementary
data and in preparing the table. All authors have read and
approved the nal manuscript.
Acknowledgments J.T.P. would like to acknowledge the funding
received fromUniversity Grants Commission (India) (39-367/2010(SR)
and KSCSTE, Govt. of Kerala (India) (011/SRSLS/2010/CSTE).
References
Abebe AT, Modi AT (2009) Hydropriming in dry bean (Phaseolus
vulgaris L.). Res J Seed Sci 2:2331
Afzal I, Basra SMA, Ahmad N, Farooq M (2005) Optimization of
hormonal priming techniques for alleviation of salinity stress in
wheat (Triticum aestivum L.). Caderno de Pesquisa Serie
Biologia 17:95109
Agarwal VK, Sinclair JB (1997) Principles of seed pathology. CRC,
Boca Raton
Akbari G, Sanavy SA, Yousefzadeh S (2007) Effect of auxin and salt
stress (NaCl) on seed germination of wheat cultivars (Triticum
aestivum L.). Pak J Biol Sci 10:25572561
Ali NI, Siddiqui IA, Shaukat SS, Zaki MJ (2001) Survival of
Pseudomonas aeruginosa in various carriers for the inhibition of
root rotroot knot disease complex of mungbean. Phytopathol
Mediterr 40:108112
Almansouri M, Kinet JM, Lutts S (2001) Effect of salt and osmotic
stresses on germination in durum wheat (Triticum durum Desf.).
Plant Soil 231:243254
Al-Mudaris MA, Jutzi SC (1999) The inuence of fertilizer based
seed priming treatment on emergence and seedling growth of
Sorghum bicolor and Pennisetum glaucum in pot trials under
greenhouse conditions. J Agron Crop Sci 182:135141
Amooaghaie R (2011) The effect of hydro and osmopriming on
alfalfa seed germination and antioxidant defenses under salt
stress. Afr J Biotechnol 10:62696275
Amzallag GN, Lerner HR, Poljakoff-Mayber A (1990) Exogenous
ABA as a modulator of the response of sorghum to high salinity.
J Exp Bot 54:15291534
Angrish R, Kumar B, Datta KS (2001) Effect of gibberellic acid and
kinetin on nitrogen content and nitrate reductase activity in
wheat under saline condition. Indian J Plant Physiol 6:172177
Arif M, Ali S, Shah A, Javed N, Rashid A (2005) Seed priming maize
for improving emergence and seedling growth. Sarhad J Agric
21:239243
Arif M, Waqas M, Nawab K, Shahid M (2007) Effect of seed priming
in Zn solutions on chickpea and wheat. Afr Crop Sci Conf Proc
8:237240
Arif M, Jan MT, Marwat KB, Khan MA (2008) Seed priming
improves emergence and yield of soybean. Pak J Bot
40(3):11691177
Ashraf M, Bray CM (1993) DNA synthesis in osmoprimed leek
(Allium porrum L.) seeds and evidence for repair and replication.
Seed Sci Res 3:1523
Ashraf M, Foolad RM (2005) Pre-sowing seed treatmenta shotgun
approach to improve germination, plant growth and crop yield
under saline and non-saline conditions. Adv Agron 88:223271
Bailly C, Benamar A, Corbineau C, Come D (1998) Free radical
scavenging as affected by accelerated ageing and subsequent
priming in sunower seeds. Physiol Plant 104:646652
Bajehbaj AA (2010) The effects of NaCl priming on salt tolerance in
sunower germination and seedling grown under salinity
conditions. Afr J Biotechnol 9:17641770
Barassi CA, Ayrault G, Creus CM, Sueldo RJ, Sobrero MT (2006)
Seed inoculation with Azospirillum mitigates NaCl effects on
lettuce. Sci Hortic 109:814
Barba-Esp n G, Hernandez JA, Diaz-Vivancos P (2012) Role of H
2
O
2
in pea seed germination. Plant Signal Behav 7:193195
Bartlett DW, Clough JM, Godwin JR, Hall AA, Hamer M, Parr-
Dobrzanski B (2002) The strobilurin fungicides. Pest Manag Sci
58:649662
Basra SMA, Pannu IA, Afzal I (2003) Evaluation of seedling vigor of
hydro and matriprimed wheat (Triticum aestivum L.) seeds. Int J
Agric Biol 5:121123
Acta Physiol Plant (2013) 35:13811396 1391
1 3
Basra SMA, Farooq M, Tabassum R (2005) Physiological and
biochemical aspects of seed vigour enhancement treatments in
ne rice (Oryza sativa L.). Seed Sci Technol 33:623628
Beckers GJM, Conrath U (2007) Priming for stress resistance: from
the lab to the eld. Curr Opin Plant Biol 10:425431
Bennett MA, Waters L (1987) Seed hydration treatments for
improved sweet maize germination and stand establishment.
J Am Soc Hortic Sci 112:4549
Bennett AJ, Whipps JM (2008) Dual application of benecial micro-
organisms to seed during drum priming. Appl Soil Ecol 38:8389
Bisht C, Badoni A, Vashishtha RK, Nautiyal MC (2009) Photope-
riodic effect on seed germination in pyrethrum (Chrysanthemum
cinerariaefolium Vis.) under the inuence of some growth
regulators. J Am Sci 5:147150
Bourgne S, Job C, Job D (2000) Sugarbeet seed priming: solubili-
zation of the basic subunit of 11S globulin in individual seeds.
Seed Sci Res 10:153161
Bradford KJ (1986) Manipulation of seed water relations via osmotic
priming to improve germination under stress conditions. Hortic
Sci 21:11051112
Bray EA (1997) Plant responses to water decit. Trends Plant Sci
2:4854
Bray CM, Davison PA, Ashraf M, Taylor MR (1989) Biochemical
events during osmopriming of leek seed. Ann Appl Biol
102:185193
Brocklehurst PA, Dearman J, Drew RLK (1984) Effects of osmotic
priming on seed germination and seedling growth in leek. Sci
Hortic 24:201210
Bruggink GT, Ooms JJJ, Vander Toorn P (1999) Induction of
longevity in primed seeds. Seed Sci Res 9:4953
Butler LH, Hay FR, Ellis RH, Smith RD, Murray TB (2009) Priming
and re-drying improve the survival of mature seeds of Digitalis
purpurea during storage. Ann Bot 103:12611270
Cakmak I (2005) The role of potassium in alleviating detrimental
effects of abiotic stresses in plants. J Plant Nutr Soil Sci
168:521530
Callan NW, Mathre DE, Miller JB (1990) Bio-priming seed treatment
for biological control of Pythium ultimum pre emergence
damping off in sh2 sweet corn. Plant Dis 74:368372
Capanoglu E (2010) The potential of priming in food production.
Trends Food Sci Technol 21:399407
Caseiro R, Bennett MA, Marcos-Filho J (2004) Comparison of three
priming techniques for onion seed lots differing in initial seed
quality. Seed Sci Technol 32:365375
Casenave EC, Toselli ME (2007) Hydropriming as a pre-treatment for
cotton germination under thermal and water stress conditions.
Seed Sci Technol 35:8898
Cayuela E, Perez-Alfocea F, Caro M, Bolarin MC (1996) Priming of
seeds with NaCl induces physiological changes on tomato plants
grown under salt stress. Physiol Plant 96:231236
Chandler PM, Robertson M (1994) Gene expression regulation by
abscisic acid and its relation to stress tolerance. Ann Rev Plant
Physiol Plant Mol Biol 45:113141
Cha-um S, Supaibulwatana K, Kirdmanee C (2006) Water relations,
photosynthetic ability and growth of Thai rice (Oryza sativa L. ssp.
Indica cv. KDML 105) to salt stress by application of exogenous
glycinebetaine and choline. J Agron Crop Sci 192:2536
Chen K, Arora R(2011) Dynamics of the antioxidant systemduring seed
osmopriming, post-priming germination, and seedling establish-
ment in spinach (Spinacia oleracea). Plant Sci 180:212220
Chen R, Murata N (2002) Enhancement of tolerance of abiotic stress
by metabolic engineering of betaines and other compounds. Curr
Opin Plant Biol 5:250257
Chen CC, Sung JM (2001) Priming bitter gourd seeds with selenium
solution enhances germinability and antioxidative responses
under sub-optimal temperature. Physiol Plant 111:916
Chen TY, Lei MG, Suzuki T, Morrison DC (1992) Lipopolysaccha-
ride receptors and signal transduction pathways in mononuclear
phagocytes. Curr Top Micro Immunol 181:169188
Chiu KY, Chuang SJ, Sung JM (2006) Both anti-oxidation and lipid-
carbohydrate conversion enhancements are involved in priming-
improved emergence of Echinacea purpurea seeds that differ in
size. Sci Hortic 108:220226
Cohen Y, Baider A, Gotilieb D, Rubin E (2007) Control of Bremia
lectucae in eld-grown lettuce by DL-3-amino-n-butanoic acid
(BABA). In: 3rd QLIF congress, Hohenheim, Germany,
pp 2023
Conrath U, Beckers GJM, Flors V, Garcia-Agustin P, Jakab G, Mauch
F, Newman MA, Pieterse CMJ, Poinssot B, Pozo MJ, Pugin A,
Schaffrath U, Ton J, Wendehenne D, Zimmerli L, Mauch-Mani
B (2006) Priming: getting ready for battle. Mol Plant Microbe
Interact 19:10621071
Copeland LO, McDonald MB (1995) Principles of seed science and
technology, 3rd edn. Chapmann and Hall, USA
Cuartero J, Fernandez-Munoz R (1999) Tomato and salinity. Sci
Hortic 78:83125
Cushman JC, Bohnert HJ (2000) Genomic approaches to plant stress
tolerance. Curr Opin Plant Biol 3:117124
Das JC, Choudhury AK (1996) Effect of seed hardening, potassium
fertilizer, and paraquat as anti-transpirant on rainfed wheat
(Triticum aestivum L.). Indian J Agron 41:397400
Demir I, Mavi K (2004) The effect of priming on seedling emergence
of differentially matured watermelon (Citrullus lanatus (Thunb.)
Matsum and Nakai) seeds. Sci Hortic 102:467473
Demir AO, Goksoy AT, Buyukcangaz H, Turan ZM, Koksal ES
(2006) Decit irrigation of sunower (Helianthus annuus L.) in a
sub-humid climate. Irrig Sci 24:279289
Demir I, Ozuaydn I, Yasar F, Staden JV (2012) Effect of smoke-
derived butenolide priming treatment on pepper and salvia seeds
in relation to transplant quality and catalase activity. S Afr J Bot
78:8387
Draganic I, Lekic S (2012) Seed priming with antioxidants improves
sunower seed germination and seedling growth under unfavor-
able germination conditions. Turk J Agric For 36:421428
Dubrovsky JG (1996) Seed hydration memory in Sonoran desert cacti
and its ecological implications. Am J Bot 68:227233
Ehsanfar S, Modarres-Sanavy SA, Tavakkol-Afshari R(2006) Effects of
osmopriming on seed germination of canola (Brassica napus L.)
under salinity stress. Commun Agric Appl Biol Sci 71:155159
Eisvand HR, Tavakkol-Afshari R, Sharifzadeh F, Maddah Are H,
Hesamzadeh Hejazi SM (2010) Effects of hormonal priming and
drought stress on activity and isozyme proles of antioxidant
enzymes in deteriorated seed of tall wheatgrass (Agropyron
elongatum Host). Seed Sci Technol 38:280297
Elkoca E, Haliloglu K, Esitken A, Ercisli S (2007) Hydro- and
osmopriming improve chickpea germination. Acta Agric Scand
Sect B Soil Plant Sci 57:193200
Farahani HA, Marou K (2011) Effect of hydropriming on seedling
vigour in basil (Ocimum basilicum L.) under salinity conditions.
Adv Environ Biol 5:828833
Farahbakhsh H (2012) Germination and seedling growth in unprimed
and primed seeds of fennel as affected by reduced water
potential induced by NaCl. Int Res J Appl Basic Sci 3:737744
Farhoudi R, Sharifzadeh F (2006) The effects of NaCl priming on salt
tolerance in canola (Brassica napus L.) seedlings grown under
saline conditions. Indian J Crop Sci 1:7478
Farooq M, Basra SMA, Hafeez K (2005) Seed invigoration by
osmohardening in Indica and Japonica rice. Seed Sci Technol
34(1):181187
Farooq M, Basra SMA, Hafeez K (2006a) Seed invigoration by
osmohardening in ne and course rice. Seed Sci Technol
34:181186
1392 Acta Physiol Plant (2013) 35:13811396
1 3
Farooq M, Basra SMA, Hefeez-ue-Rehman, Mehmood T (2006b)
Germination and early seedling growth as affected by pre sowing
ethanol seed treatments in ne rice. Int J Agric Biol 8:1922
Farooq M, Basra SMA, Wahid A (2006c) Priming of eld sown rice
seed enhances germination, seedling establishment, allometry
and yield. Plant Growth Regul 44:28294
Farooq M, Basra SMA, Wahid A, Ahamad N (2010) Changes in
nutrient homeostasis and reserve metabolism during rice seed
priming: consequences for seedling emergence and growth.
Agric Sci China 9:191198
Finch-Savage WE, Dent KC, Clark LJ (2004) Soak conditions and
temperature following sowing inuence the response of maize
(Zea mays L.) seeds to on-farm priming (pre-sowing seed soak).
Field Crops Res 90:361374
Flowers TJ, Garcia A, Koyama M, Yeo AR (1997) Breeding for salt
tolerance in crop plants-the role of molecular biology. Acta
Physiol Plant 19:427433
Flowers TJ, Koyama ML, Flowers SA, Sudhakar C, Singh KP, Yeo
AR (2000) QTL: their place in engineering tolerance of rice to
salinity. J Exp Bot 51:99106
Foti R, Aburenia K, Tigerea A, Gotosab J, Gerec J (2008) The
efcacy of different seed priming osmotica on the establishment
of maize (Zea mays L.) caryopses. J Arid Environ 72:11271130
Foyer CH, Noctor G (2005) Redox homeostasis and antioxidant
signaling: a metabolic interface between stress perception and
physiological responses. Plant Cell 17:18661875
Fujikura Y, Kraak HL, Basra AS, Karssen CM (1993) Hydropriming,
a simple and in expensive priming method. Seed Sci Technol
21:639642
Fujita M, Fujita Y, Noutoshi Y, Takahashi F, Narusaka Y, Yamag-
uchi-Shinozaki K, Shinozak K (2006) Crosstalk between abiotic
and biotic stress responses: a current view from the points of
convergence in the stress signaling networks. Curr Opin Plant
Biol 9:436442
Gao YP, Bonham-Smith PC, Gusta LV (2002) The role of peroxire-
doxin antioxidant and calmodulin in ABA-primed seeds of
Brassica napus exposed to abiotic stresses during germination.
J Plant Physiol 159:951958
Ghana SG, Schillinger WF (2003) Seed priming winter wheat for
germination, emergence, and yield. Crop Sci 43:21352141
Ghassemi-Golezani K, Esmaeilpour B (2008) The effect of salt
priming on the performance of differentially matured cucumber
(Cucumis sativus) seeds. Not Bot Hort Agrobot Cluj 36:6770
Guan YJ, Hu J, Wang XJ, Shao CX (2009) Seed priming with
chitosan improves maize germination and seedling growth in
relation to physiological changes under low temperature stress.
J Zhejiang Univ Sci B 10:427433
Guimil S, Chang HS, Zhu T, Sesma A, Osbourn A, Roux C, Ionnidis
V, Oakeley EJ, Docquier M, Descombes P, Briggs SP,
Paszkowski U (2005) Comparative transcriptomics of rice
reveals an ancient pattern of response to microbial colonization.
Proc Natl Acad Sci USA 102:80668070
Gul B, Khan MA (2003) Effect of growth regulators and osmotic in
alleviating salinity effects on the germination of Salicornia
utahensis. Pak J Bot 35:877886
Gust AA, Brunner F, Nurnberger T (2010) Biotechnological concepts
for improving plant innate immunity. Curr Opin Biotechnol
21:204210
Haghpanah A, Younesi O, Moradi A (2009) The effect of priming on
seedling emergence of differentially matured sorghum (Sorghum
bicolor L.) seeds. J Appl Sci Res 5:729732
Hancock JT, Neill SJ, Wilson ID (2011) Nitric oxide and ABA in the
control of plant function. Plant Sci 181:555559
Harb EZ (1992) Effect of soaking seeds in some growth regulators
and micronutrients on growth, some chemical constituents and
yield of faba bean and cotton plants. Bull Faculty Agric Univ
Cairo 43:429452
Harris D, Joshi A, Kham PA, Khan PA, Gothkar P, Sodhi PS (1999)
On-farm seed priming in semi arid agriculture: development and
evaluation in maize, rice and chick pea in India using partic-
ipatory methods. Exp Agric 35:1529
Hasanuzzaman M, Anwar Hossain M, Fujita M (2010) Selenium in
higher plants: physiological role, antioxidant metabolism and
abiotic stress tolerance. J Plant Sci 5:354375
Heydecker W, Higgins J, Gulliver RL (1973) Accelerated germina-
tion by osmotic seed treatment. Nature 246:4244
Hsu CC, Chen CL, Chen JJ, Sung JM (2003) Accelerated aging-
enhanced lipid peroxidation in bitter gourd seeds and effects of
priming and hot water soaking treatments. Sci Hortic 98:201212
Ingram J, Bartels D (1996) The molecular basis of dehydration
tolerance in plants. Annu Rev Plant Physiol Plant Mol Biol
47:377403
Iqbal M, Ashraf M (2005) Changes in growth, photosynthetic
capacity and ionic relations in spring wheat (Triticum aestivum
L.) due to presowing seed treatment with polyamines. Plant
Growth Regul 46:1930
Jain N, Kulkarni MG, Van Staden J (2006) A butenolide isolated from
smoke can overcome the detrimental effect of extreme temper-
atures during tomato seed germination. Plant Growth Regul
49:263267
Jakab G, Ton J, Flors V, Zimmerli L, Metraux JP, Mauch-Mani B
(2005) Enhancing Arabidopsis salt and drought stress tolerance
by chemical priming for its abscisic acid responses. Plant Physiol
139:267274
Jaleel CA, Manivannan P, Gopi R, Panneerselvam R (2007)
Responses of antioxidant defense system of Catharanthus roseus
(L.) G. Don. to paclobutrazol treatment under salinity. Acta
Physiol Plant 29:205209
Jaleel CA, Lakshmanan GMA, Gomathinayagam M, Panneerselvam
R (2008) Triadimefon induced salt stress tolerance in Withania
somnifera and its relationship to antioxidant defense system.
S Afr J Bot 74:126132
Janmohammadi M, Moradi Dezfuli P, Sharifzadeh F (2008) Seed
invigoration techniques to improve germination and early growth
of inbred line of maize under salinity and drought stress. Gen
Appl Plant Physiol 34:215226
Jett LW, Welbaum GE, Morse RD (1996) Effect of matric and
osmotic priming treatments on broccoli seed germination. J Am
Soc Hortic Sci 121:423429
Jeun YC, Park KS, Kim CH, Flower WD, Kloepper JW (2004)
Cytological observations of cucumber plants during induced
resistance elicited by rhizobacteria. Biol Control 29:3442
Job D, Capron I, Job C, Dacher F, Corbineau F, Come D (2000)
Identication of germination- specic protein markers and their
use in seed priming technology. In: Black M, Bradford KJ,
Vazguez-Ramos J (eds) Seed biology: advances and applica-
tions. CAB International, Wallingford, pp 449459
Kaur S, Gupta AK, Kaur N (2002) Effect of osmo- and hydropriming
of chickpea on seedling growth and carbohydrate metabolism
under water decit stress. Plant Growth Regul 37:1722
Kaur S, Gupta AK, Kaur N (2005) Seed priming increases crop yield
possibly by modulating enzymes of sucrose metabolism in
chickpea. J Agron Crop Sci 19:8187
Kaya MD, Okcu G, Atak M, Cikili Y, Kolsarici O (2006) Seed
treatments to overcome salt and drought stress during germina-
tion in sunower (Helianthus annuus L.). Eur J Agron
24:291295
Khajeh-Hosseini M, Powell AA, Bingham IJ (2003) The interaction
between salinity stress and seed vigour during germination of
soyabean seeds. Seed Sci Technol 31:715725
Acta Physiol Plant (2013) 35:13811396 1393
1 3
Khalil SK, Mexal JG, Murray LW (2001) Germination of soybean
seed primed in aerated solution of polyethylene glycol (8000).
J Biol Sci 1:105107
Khan MA, Gul B, Weber DJ (2002) Improving seed germination of
Salicornia rubra (Chenopodiaceae) under saline conditions using
germination-regulating chemicals. West N Am Nat 62:101105
Khan MB, Gurchani MA, Hussain M, Freed S, Mahmood K (2011)
Wheat seed enhancement by vitamin and hormonal priming. Pak
J Bot 43:14951499
Kibinza S, Bazin J, Bailly C, Farrant JM, Corbineau F, El-Maarouf-
Bouteau H (2011) Catalase is a key enzyme in seed recovery
from ageing during priming. Plant Sci 181:309315
Korkmaz A, Pill WG (2003) The effect of different priming
treatments and storage conditions on germination performance
of lettuce seeds. Eur J Hortic Sci 68:260265
Kubik KK, Eastin JA, Eskridge KM (1988) Solid matrix priming of
tomato and pepper. In: Proceedings of the international confer-
ence on stand establishment for horticultural crops. Lancaster,
Pennsylvania, USA, pp 8696
Kulkarni MG, Sparg SG, Light ME, Van Staden J (2006) Stimulation
of rice (Oryza sativa L.) seedling vigour by smoke-water and
butenolide. J Agron Crop Sci 192:395398
Lee SS, Kim JH, Hong SB, Yuu SH, Park EH (1998) Priming effect
of rice seeds on seedling establishment under adverse soil
conditions. K J Crop Sci 43:194198
Lee-suskoon KM, Hyeum J, Beom HS, Minkyeong K, Euiho P (1998)
Optimum water potential, temperature and duration for priming
of rice seeds. J Crop Sci 43:15
Li J, Yin LY, Jongsma MA, Wang CY (2011) Effects of light,
hydropriming and abiotic stress on seed germination, and shoot
and root growth of pyrethrum (Tanacetum cinerariifolium). Ind
Crop Prod 34:15431549
Lin JM, Sung JM (2001) Pre-sowing treatments for improving
emergence of bitter gourd seedlings under optimal and sub-
optimal temperatures. Seed Sci Technol 29:3950
Lorenzo MCB (1991) Seed invigoration of soybean and corn through
solid matrix priming. BS, Philippines
Manish K, Geetika S, Renu B, Sandeep K, Gaganodeep J (2010)
Effect of exogenous H
2
O
2
on antioxidant enzymes of Brassica
juncea L. seedlings in relation to 24-epibrassinolide under
chilling stress. Indian J Biochem Biophys 47:378382
Manjunatha G, Raj SN, Shetty NP, Shetty HS (2008) Nitric oxide
donor seed priming enhances defense responses and induces
resistance against pearl millet downy mildew disease. Pestic
Biochem Physiol 91:111
Margot P, Huggenberger F, Amrein J, Weiss B (1998) A new
broadspectrum strobilurin fungicide. Brighton Crop Prot Conf
Pest Disease 2:375382
Marschner H (1995) Mineral nutrition of higher plants, 2nd edn.
Academic press, San Diego
Maughan S, Foyer CH (2006) Engineering and genetic approaches to
modulating the glutathione network in plants. Physiol Plant
126:382397
McDonald MB (2000) Seed priming. In: Black M, Bewley JD (eds)
Seed technology and its biological basis. Shefeld Academic
Press, Shefeld, pp 287325
Mercado MFO, Fernandez PG (2002) Solid matrix priming of
soybean seeds. Philipp J Crop Sci 27:2735
Miller G, Suzuki N, Ciftci-Yilmaz S, Mittler R (2009) Reactive
oxygen species homeostasis and signaling during drought and
salinity stresses. Plant Cell Environ 33:453467
Mishra M, Kumar U, Mishra PK, Prakash V (2010) Efciency of
plant growth promoting rhizobacteria for the enhancement of
Cicer arietinum L. growth and germination under salinity. Adv
Biol Res 4:9296
Mittler R (2002) Oxidative stress, antioxidants and stress tolerance.
Trends Plant Sci 9:405410
Moeinzadeh A, Sharif-Zadeh F, Ahmadzadeh M, Heidari Tajabadi F
(2010) Biopriming of sunower (Helianthus annuus L.) seeds
with Pseudomonas uorescens for improvement of seed invig-
oration and seedling growth. Aust J Crop Sci 4:564570
Mohanan C, Sharma JK (1991) Seed pathology of forest tree species
in India- present status, practical problems and future prospects.
Commonw For Rev 70:133151
Moradi A, Younesi O (2009) Effects of osmo- and hydropriming on
seed parameters of grain sorghum (Sorghum bicolor L.). Aust J
Basic Appl Sci 3:16961700
Murungu FS, Nyamugafata P, Chiduza C, Clark LJ, Whalley WR
(2003) Effects of seed priming, aggregate size and soil matric
potential on emergence of cotton (Gossypium hirsutum L.) and
maize (Zea mays L.). Soil Tillage Res 74:161168
Nascimento WM, West SH (1998) Priming and seed orientation affect
emergence and seed coat adherence and seedling development of
muskmelon transplants. Hortic Sci 33:847848
Nawaz A, Amjad M, Pervez MA, Afzal I (2011) Effect of
halopriming on germination and seedling vigor of tomato. Afr
J Agric Res 6:35513559
Niinemets U (2009) Mild versus severe stress and BVOCs: thresh-
olds, priming and consequences. Trends Plant Sci 15:145153
Omidi H, Khazaei F, Hamzi Alvanagh S, Heidari-Sharifabad H
(2009) Improvement of seed germination traits in canola
(Brassica napus L.) as affected by saline and drought stresses.
Plant Ecol Physiol 3:151158
Ozbingol N, Corbineau F, Come D (1998) Response of tomato seeds
to osmoconditioning as related to temperature and oxygen. Seed
Sci Res 8:377384
Pandita VK, Anand A, Nagarajan S, Seth R, Sinha SN (2010) Solid
matrix priming improves seed emergence and crop performance
in okra. Seed Sci Technol 38:665674
Parera CA, Cantliffe DJ (1991) Improved germination and modied
imbibitions of shrunken-2 sweet corn by seed disinfection and
solid matrix priming. J Am Soc Hortic Sci 116:942945
Parera CA, Cantliffe DJ (1994) Presowing seed priming. Hortic Rev
16:109114
Parera CA, Qiao P, Cantliffe DJ (1993) Enhanced celery germination
at stress temperature via solid matrix priming. Hortic Sci
28:2022
Passam HC, Kakouriotis D (1994) The effects of osmoconditioning
on the germination, emergence and early plant growth of
cucumber under saline conditions. Sci Hortic 57:233240
Patade VY, Sujata B, Suprasanna P (2009) Halopriming imparts
tolerance to salt and PEG induced drought stress in sugarcane.
Agric Ecosyst Environ 134:2428
Posmyk MM, Janas KM (2007) Effects of seed hydropriming in
presence of exogenous proline on chilling injury limitation in
Vigna radiata L. seedlings. Acta Physiol Plant 29:509517
Pozo MJ, Azcon-Aguilar C (2007) Unraveling mycorrhiza-induced
resistance. Curr Opin Plant Biol 10:393398
Radi AF, Shaddad MAK, El-Enany AE, Omran FM (2001) Interactive
effects of plant hormones (GA
3
or ABA) and salinity on growth
and some metabolites of wheat seedlings. In: Horst WJ, Schenk
MK, Burkert A, Claassen N, Flessa H, Frommer WB, Goldbach
H, Olfs HW, Romheld V (eds) Plant nutrition, food security and
sustainability of agro-ecosystems through basic and applied
research. 14th international plant nutrition colloquium, Han-
nover, Germany, pp 436437
Raetz CR, Ulevitch RJ, Wright SD, Sibley CH, Ding A, Nathan CF
(1991) Gram-negative endotoxin: an extraordinary lipid with
profound effects on eukaryotic signal transduction. Fed Am Soc
Exp Biol J 5:26522660
1394 Acta Physiol Plant (2013) 35:13811396
1 3
Randhir R, Shetty K (2005) Developmental stimulation of total
phenolics and related antioxidant activity in light- and dark-
germinated corn by natural elicitors. Proc Biochem
40:17211732
Rashid A, Harris D, Hollington P, Ali S (2004) On-farm seed priming
reduces yield losses of mungbean (Vigna radiata) associated
with mungbean yellow mosaic virus in the North West Frontier
Province of Pakistan. Crop Prot 23:11191124
Rashid A, Hollington PA, Harris D, Khan P (2006) On-farm seed
priming for barley on normal, saline and salinesodic soils in
North West Frontier Province, Pakistan. Eur J Agron
24:276281
Rathinasabapathi B (2000) Metabolic engineering for stress tolerance:
installing osmoprotectant synthesis pathways. Ann Bot
86:709716
Rawat L, Singh Y, Shukla N, Kumar J (2011) Alleviation of the
adverse effects of salinity stress in wheat (Triticum aestivum L.)
by seed biopriming with salinity tolerant isolates of Trichoderma
harzianum. Plant Soil 347:387400
Reddy CS, Smith JD (1978) Effects of delayed post treatment of
gamma-irradiated seed with cysteine on the growth of Sorghum
bicolor seedlings. Environ Exp Bot 18:241243
Reddy MVB, Arul J, Angers P, Couture L (1999) Chitosan treatment
of wheat seeds induces resistance to Fusarium graminearum and
improves seeds quality. J Agric Food Chem 47:6772
Reyes LF, Cisneros-Zevallos L (2007) Electron-beam ionizing
radiation stress affects on mango fruit (Mangifera indica L.)
antioxidant constituents before and during post harvest storage.
J Agric Food Chem 55:61326139
Rithichai P, Sampantharat P, Jirakiattikul Y (2009) Coriander
(Coriandrum sativum L.) seed quality as affected by accelerated
aging and subsequent hydropriming. As J Food Ag-Ind Special
Issue:S217S221
Rock CD (2000) Pathways to abscisic acid regulated gene expression.
New Phytol 148:357384
Rouhi HR, Afshari RT, Moosavi SA, Gharineh MH (2010) Effects of
osmopriming on germination and vigour traits of bersim clover
(Trifolium alexandrinum L.). Not Sci Biol 2:5963
Rouhi HR, Aboutalebian MA, Sharif-Zadeh F (2011) Effects of hydro
and osmopriming on drought stress tolerance during germination
in four grass species. Int J Agric Sci 1:701774
Ruan SL, Xue QZ (2002) Effects of chitosan coating on seed
germination and salt-tolerance of seedlings in hybrid rice (Oryza
sativa L.). Acta Agron Sin 28:803808
Rubluo A (1982) The effect of L-cysteine on presoaked barley seeds
treated with methyl methanesulfonate. Experientia 38:326327
Saha P, Chatterjee P, Biswas AK (2010) NaCl pretreatment alleviates
salt stress by enhancement of antioxidant defense system and
osmolyte accumulation in mungbean (Vigna radiata L. Wil-
czek). Indian J Exp Biol 48:593600
Sahu MP, Singh D (1995) Role of thiourea in improving productivity
of wheat (Triticum aestivum L.). J Plant Growth Regul
14:169173
Sahu MP, Kumawat SM, DSouza SF, Ramaswamy NK, Singh G
(2005) Sulphydryl bioregulator technology for increasing mus-
tard production. Res Bull RAUBARC, pp 152
Sakamoto M, Murata N (2002) The role of glycinebetaine in the
protection of plants from stress: clues from transgenic plants.
Plant Cell Environ 25:163171
Sakhabutdinova AR, Fatkhutdinova DR, Beazrukova MV, Shakirova
FM (2003) Salicylic acid prevents the damaging action of stress
factor of wheat plants. Bulg J Plant Physiol Special Issue:314319
Salama KHA, Mansour MMF, Hassan NS (2011) Choline priming
improves salt tolerance in wheat (Triticum aestivum L.). Aust J
Basic Appl Sci 5:126132
Sarwar N, Yousaf S, Jamil FF (2006) Induction of salt tolerance in
chickpea by using simple and safe chemicals. Pak J Bot
38:325329
Sedghi M, Nemati A, Amanpour-Balaneji B, Gholipouri A (2010)
Inuence of different priming materials on germination and
seedling establishment of milk thistle (Silybum marianum) under
salinity stress. World Appl Sci J 11:604609
Sha M, Bakht J, Hassan MJ, Raziuddin M, Zhang G (2009) Effect of
cadmium and salinity stresses on growth and antioxidant enzyme
activities of wheat (Triticum aestivum L.). Bull Environ Contam
Toxicol 82:772776
Shahrokhi M, Tehranifar A, Hadizadeh H, Selahvarzi Y (2011) Effect
of drought stress and paclobutrazol-treated seeds on physiolog-
ical response of Festuca arundinacea L. Master and Lolium
perenne L. Barrage. J Biol Environ Sci 5:7785
Shakirova FM, Sakhabutdinova AR, Bezrokuva MV, Fatkhutdinova
RA, Fatkhutdinova DR (2003) Changes in the hormonal status of
wheat seedlings induced by salicylic acid and salinity. Plant Sci
164:317322
Shao CX, Hu J, Song WJ, Hu WM (2005) Effects of seed priming
with chitosan solutions of different acidity on seed germination
and physiological characteristics of maize seedling. J Zhejiang
Univ (Agric & Life Sci) 31:705708
Sharmila P, Puthur JT, Pardha Saradhi P (2000) Vesicular arbuscular
mycorrhizal fungi improves establishment of micropropagated
plants. In: Mukerji KG, Chamola BP, Singh J (eds) Mycorrhizal
Biology. Kluwer Academic/Plenum, New York, pp 236250
Singh BG, Rao G (1993) Effects of chemical soaking of sunower
(Helianthus annuus L.) seed on vigour index. Indian J Agric Sci
63:232233
Sivritepe N, Sivritepe HO, Eris A (2003) The effects of NaCl priming
on salt tolerance in melon seedlings grown under saline
conditions. Sci Hortic 97:229237
Sivritepe HO, Sivritepe N, Eris A, Turhan E (2005) The effects of
NaCl pre-treatments on salt tolerance of melons grown under
long-term salinity. Sci Hortic 106:568581
Srivastava AK, Ramaswamy NK, Mukopadhyaya R, Chiramal Jincy
MG, DSouza SF (2009) Thiourea modulates the expression and
activity prole of mtATPase under salinity stress in seeds of
Brassica juncea L. Ann Bot (Lond) 103:403410
Srivastava AK, Lokhande VH, Patade VY, Suprasanna P, Sjahril R,
DSouza SF (2010a) Comparative evaluation of hydro-, chemo-,
and hormonal priming methods for imparting salt and PEG stress
tolerance in Indian mustard (Brassica juncea L.). Acta Physiol
Plant 32:11351144
Srivastava AK, Suprasanna P, Srivastava S, DSouza SF (2010b)
Thiourea mediated regulation in the expression prole of
aquaporins and its impact on water homeostasis under salinity
stress in Brassica juncea roots. Plant Sci 178:517522
Su J, Hirji R, Zhang L, He C, Selvaraj G, Wu R (2006) Evaluation of
the stress-inducible production of choline oxidase in transgenic
rice as a strategy for producing the stress-protectant glycine
betaine. J Exp Bot 57:11291135
Sudisha J, Niranjana SR, Sukanya SL, Girijamba R, Lakshmi Devi N,
Shekar Shetty H (2010) Relative efcacy of strobilurin formu-
lations in the control of downy mildew of sunower. J Pest Sci
83:461470
Sung JM, Chiu KY (1995) Hydration effects on seedling emergence
strength of watermelon seed differing in ploidy. Plant Sci
110:2126
Tavili A, Zare S, Enayati A (2009) Hydropriming, ascorbic and
salicylic acid inuence on germination of Agropyron elongatum
Host. seeds under salt stress. Res J Seed Sci 2:1622
Tavili A, Zare S, Moosavi SA, Enayati A (2011) Effects of seed
priming on germination characteristics of Bromus species under
Acta Physiol Plant (2013) 35:13811396 1395
1 3
salt and drought conditions. American-Eurasian J Agric Environ
Sci 10:163168
Taylor AG, Harman GE (1990) Concepts and technologies of selected
seed treatments. Ann Rev Phytopathol 28:321329
Thomas UC, Varughese K, Thomas A, Sadanandan S (2000) Seed
primingfor increased vigour, viability and productivity of
upland rice. Leisa India 4:14
Tiryaki I, Korkmaz A, Nas MN, Ozbay N (2005) Priming combined
with plant growth regulators promotes germination and emer-
gence of dormant Amaranthus cruentus L. seeds. Seed Sci
Technol 33:571579
Tonelli ML, Furlan A, Taurian T, Castro S, Fabra A (2011) Peanut
priming induced by biocontrol agents. Physiol Mol Plant
75:100105
Uchida A, Jagendorf AT, Hibino T, Takabe T (2002) Effects of
hydrogen peroxide and nitric oxide on both salt and heat stress
tolerance in rice. Plant Sci 163:515523
Van Hulten M, Pelser M, van Loon LC, Pieterse CMJ, Ton J (2006)
Costs and benets of priming for defense in Arabidopsis. Proc
Natl Acad Sci USA 103:56025607
Vickers CE, Gershenzon J, Lerdau MT, Loreto F (2009) A unied
mechanism of action for volatile isoprenoids in plant abiotic
stress. Nat Chem Biol 5:283291
Wahid A, Perveen M, Gelani S, Shahzad MA, Basra SMA (2007)
Pretreatment of seed with H
2
O
2
improves salt tolerance of wheat
seedlings by alleviation of oxidative damage and expression of
stress proteins. J Plant Physiol 164:283294
Windauer L, Altuna A, Benech-Arnold R (2007) Hydrotime analysis
of Lesquerella fendleri seed germination responses to priming
treatments. Ind Crop Prod 25:7074
Xiong L, Zhu JK (2002) Molecular and genetic aspects of plant
responses to osmotic stress. Plant Cell Environ 25:131139
Xu S, Hu J, Li Y, Ma W, Zheng Y, Zhu S (2011) Chilling tolerance in
Nicotiana tabacum induced by seed priming with putrescine.
Plant Growth Regul 63:279290
Yadav PV, Kumari M, Ahmed Z (2011) Seed priming mediated
germination improvement and tolerance to subsequent exposure
to cold and salt stress in capsicum. Res J Seed Sci 4:125136
Yagmur M, Kaydan D (2008) Alleviation of osmotic strength of water
and salt in germination and seedling growth of triticale with seed
priming treatments. Afr J Biotechnol 7:21562162
Yari L, Sheidaie S (2011) Effect of seed priming on seed germina-
tions behavior of rice (Oryza sativa L.). Int J Agric Sci 1:4551
Yeh YM, Chiu KY, Chen CL, Sung JM (2005) Partial vacuum
extends the longevity of primed bitter gourd seeds by enhancing
their anti-oxidative activities during storage. Sci Hortic
104:101112
Yuan-Yuan S, Yong-Jian S, Ming-Tian W, Xu-Yi LI, Xiang GUO,
Rong HU, Jun MA (2010) Effects of seed priming on germi-
nation and seedling growth under water stress in rice. Acta
Agron Sin 36:19311940
Zhao TJ, Liu Y, Yan YB, Feng F, Liu WQ, Zhou HM (2007)
Identication of the aminoacids crucial for the activities of
drought responsive element binding factors (DREBs) of Brassica
napus. FEBS Lett 581:30443050
Zhou YG, Yang YD, Qi YG, Zhang ZM, Wang XJ, Hu XJ (2002)
Effects of chitosan on some physiological activity in germinating
seed of peanut. J Peanut Sci 31:2225
Zhou ZS, Guo K, Elbaz AA, Yang ZM (2009) Salicylic acid alleviate
mercury toxicity by preventing oxidative stress in roots of
Medicago sativa. Environ Exp Bot 65:2734
Zhu JK, Liu J, Xiong L (1998) Genetic analysis of salt tolerance in
Arabidopsis: evidence for a critical role of potassium nutrition.
Plant Cell 10:11811191
1396 Acta Physiol Plant (2013) 35:13811396
1 3