Volume 67
PLANT BREEDING AND SEED SCIENCE
2013
DOI 10.2478/v10129-011-0066-2
Ayda Hosseinzadeh-Mahootchi, Kazem Ghassemi-Golezani*
Department of Plant Eco-Physiology, Faculty of Agriculture, University of Tabriz, Tabriz, Iran
(*Corresponding author e-mail: golezani@gmail.com)
THE IMPACT OF SEED PRIMING AND AGING ON PHYSIOLOGICAL PERFORMANCE OF CHICKPEA UNDER DIFFERENT IRRIGATION TREATMENTS
ABSTRACT
A sub-sample of chickpea (Cicer arietinum L. cv.ILC482) seeds was kept as control and two other subsamples were aged at 40 °C for 3 and 5 days. Consequently, three seed lots with different levels of vigor were
provided. These seed lots were soaked in distilled water at 15°C for 12 and 18 hours and then dried back to
initial moisture content at a room temperature of 20-22°C. Then seeds were sown in the field as split plot
factorial based on RCB design. Hydro-priming improved leaf chlorophyll content index of plants from
different seed lots. Hydro-priming also enhanced stomatal conductance of plants from all seed lots
under all irrigation levels, but this advantage for plants from low vigor seed lots particularly under
limited irrigations was higher than that for other treatments. Plants from high vigor seed lot under
different irrigation treatments had higher relative water content, compared with those from low vigor seed
lots. Hydro-priming improved relative water content, membrane stability and grain yield of chickpea plants
from different seed lots under various irrigation treatments. It was concluded that hydro-priming to some
extent can repair aged seeds and improve their performance under different irrigation treatments.
Key words: chickpea, hydro-priming, membrane stability, relative water content, stomatal conductance
INTRODUCTION
Chickpea (Cicer arietinum L.) is an important food legume crop which is
grown in semi-arid regions (Labidi et al., 2009). Drought is perhaps the major
factor negatively affects plant growth and development and causes a sharp
decrease of plants productivity (Pan et al., 2002). Water deficit affects
many physiological processes associated with plant growth and development
(Toker and Cagirgan, 1998). In
drought stress conditions, plants
Communicated by Andrzej Anioł
14
Ayda Hosseinzadeh-Mahootchi, Kazem Ghassemi-Golezani
close their stomata to avoid further water loss (Dulai et al., 2006). This limits
CO2 assimilation which may promote an imbalance between photochemical
activity at photosystem II (PSII) and electron requirement for photosynthesis,
leading to a photoinhibitory damage of PSII reaction centers (Long et al.,
1994). Drought stress affects photosystem efficiency (Fv/Fm) and decreases quantum yield of photosystem II (Ahmed et al., 2002). Investigations based on assessments of chlorophyll a fluorescence have shown that
PSII is quite resistant to water deficits, being either unaffected (Shangguan et
al., 2000) or affected only under very severe drought conditions (Saccardy et
al., 1998). In fact, photo damage may be prevented through processes of
thermal deactivation, down regulating PSII photochemistry, in a so-called
dynamic photo inhibition (Osmond, 1994), that brings the electron transport
capacity into balance with carbon metabolism (Epron et al., 1992).
Another plant response to drought stress is change in photosynthetic pigment
content. Leaf chlorophyll content play important role in harvesting light.
The content of both chlorophylls a and b changes under drought stress
(Farooq et al., 2009). The effects of drought stress on membrane stability
index (MSI) and relative water content (RWC) have also been decreased
under water deficit (Bayoumi et al., 2008). RWC is a reliable parameter for
quantifying the plant drought stress response (Bayoumi et al., 2008). The
deleterious effects of water deficit on plants may be somewhat reduced by
sowing high vigor seed lots or by priming of seeds before sowing
(Ghassemi-Golezani et al., 2012).
Maximum seed vigor is achieved at or slightly after mass maturity (end
of seed filling period), which is previously termed physiological maturity
(Shaw and Loomis, 1950). Thereafter seeds begin to deteriorate on the
mother plant (Ghassemi-Golezani and Mazloomi-Oskooyi, 2008; GhassemiGolezani and Hossinzadeh-Mahootchy, 2009). When deterioration is advanced,
rate and uniformity of seed germination and seedling emergence and tolerance
to environmental stresses decreases (Khan et al., 2003). The slower rate of
emergence frequently associated with low-vigor seeds resulting in smaller
plants, compared with high-vigor seeds (Ellis and Roberts, 1981; Ghassemi
Golezani et al., 2010b).
One of the simple and suitable methods which can improve seedling
vigor and establishment and consequently crop performance in the field is
seed priming (McDonald, 2000). Priming appears to reverse the detrimental
effects of seed deterioration (McDonald, 2000). The early improvements
may increase the rate and uniformity of seed germination and seedling
emergence (Farooq et al., 2005, Ghassemi-Golezani et al., 2010a), especially
under stressful conditions (Ghassemi-Golezani et al., 2012). Some of the
deleterious effects of low-vigor seed lots and environmental stresses such
as water limitation on crop performance may be also overcome by seed
priming (Demir et al., 2006), via improving seedling vigor (Ghassemi-
Relationship between root and yield related morphological characters in pea (P.sativum L.)… 15
Golezani, 1992) and stand establishment (Finch-Savage, 2000). Thus, this
research was carried out to investigate the effects of seed vigor on some
physiological characteristics and yield of chickpea under different irrigation
treatments.
MATERIALS AND METHODS
Seeds of chickpea (Cicer arietinum L. cv.ILC482) were obtained from Dry
-land Agricultural Research Institute of Maragheh, Iran. These seeds were
divided into three sub-samples. A sub-sample was kept as control with
100% germination (V1). The other sub-samples with about 20% moisture
content were artificially aged at 40°C for 3 and 5 days, reducing seed germination
to %98 and 89% (V2 and V3, respectively). Consequently, three seed lots
with different levels of vigor were provided. Then, each seed lot was
divided into three sub-samples, one of which was kept as control
(unprimed, P 1) and two other samples were soaked in distilled water at 15°C
for 12 (P2) and 18 (P3) hours and then dried back to initial moisture content at
a room temperature of 20-22°C for 24 hours.
The field experiment was conducted at the Research Farm of the University
of Tabriz (Latitude 38˚05’ N, Lon-gitude 46˚17’ E, Altitude 1360 m above
sea level) in 2011. All the seeds were treated with benomyl at a rate of
2 g × kg-1 before sowing. Seeds were hand sown in about 4 cm depth with
a density of 60 seeds × m-2. Each plot consisted of 8 rows with 4 m length,
spaced 25 cm apart. The experiment was arranged as split plot factorial,
based on RCB design with three replications. All plots were irrigated
immediately after sowing and subsequent irrigations were carried out after
70 (I1), 120 (I2) and 170 (I3) mm evaporation from class A pan. Weeds were
controlled by hand during crop growth and development. Plants were
protected from heliothis caterpillar attack by spraying Diazinon at a rate of
2 ml × l-1 before flowering.
Photochemical efficiency of photosystem II (Fv/Fm) was measured using
a portable chlorophyll fluorometer. Measurements were made after 20 min
dark adaptation (Maxwell et al., 2000) from 3 plants. Chlorophyll content
index of leaves was measured every week by a chlorophyll meter (CCM200). Relative water content was determined according to Barr and
Weatherley (1962). Fresh weight of the youngest fully expanded leaf was
recorded within 24 h after excision. Turgid weight was obtained after soaking the leaf for 24 h in distilled water. After that, the leaves were quickly
and carefully dried with tissue paper prior to determination of turgid
weight. Leaf dry weight was obtained after drying the sample for 48 h at
75°C. Relative water content was calculated from the following equation:
16
Ayda Hosseinzadeh-Mahootchi, Kazem Ghassemi-Golezani
Leaf samples (0.1 g) were taken in 10 ml double-distilled water in glass
vials and kept at 40ºC for 10 min. Initial conductivity (C1) was recorded
with a conductivity meter after transferring the sample to 25ºC. The samples
were kept at 100ºC for 30 min and cooled at 25ºC. Final conductivity (C2)
was measured according to Sairam (1994). The membrane stability index
(MSI) was calculated as:
Stomatal conductance of leaves was determined using a portable prometer
(Delta-T AP4, Cambridge, UK). The measurements were taken on the surface of the leaf at the flowering stage. Finally, plants of 1 m2 in the middle
part of each plot were harvested and grain yield was recorded. Analysis of
variance of the data appropriate to the experimental design and comparison
of means at p≤0.05 were carried out, using MSTATC software.
RESULTS
Chlorophyll Content Index (CCI)
Fig.1. Changes in leaf chlorophyll content index (CCI) of chickpea at different stages of growth
and development affected by seed vigor
Chlorophyll content index of chickpea leaves diminished with progressing
plant development (Fig. 1). At the most stages of development, leaf chloro-
Relationship between root and yield related morphological characters in pea (P.sativum L.)… 17
phyll content index of plants from low vigor seed lots (V2 and V3) was
lower than that for plants from high vigor seed lot (V1). CCI of plants from
all seed lots decreased with increasing plant senescence at later stages of
development. Reduction in CCI of V3 plants was started earlier than that of
plants from other seed lots. The rate of reduction was also much higher for
V3 than for V2 and V1 plants (Fig. 1).
The analysis of variance of data showed significant effects of irrigation
level and seed vigor on maximum chlorophyll content index. Interaction of
seed vigor × hydro-priming duration for this trait was also significant
(Table 1). Maximum chlorophyll content of chickpea was decreased with
increasing water limitation (Table 2). Hydro-priming improved leaf CCI of
plants from different seed lots (Table 2), but this beneficial effect of hydropriming for plants from high vigor seed lot was higher than that for plants
from other seed lots.
Table 1
Analysis of variance of the data of chickpea plants from different seed lots
under different irrigation treatments
MS
S.O.V
D.F.
RWC
MSI
Replication
2
0.219
33.309
11.751
Irrigation
level
2
11.392*
880.805*
467.195*
Error
4
0.666
0.066
212.846
48.925
32.534
Vigor
2
8.217**
0.002ns
6189.123**
107.531*
I×V
4
0.629ns
0.001ns
2081.420
Priming
Duration
2
0.790ns
0.009ns
I×P
4
1.210ns
V×P
4
I×V×P
8
Error
C.V [%]
48
CCI
Fv/Fm
Stomatal conductance
0.080
703.494
0. 207ns 216294.827**
Grain yield
853.758
285003.114**
788.110
10.814ns
117851.113**
60.095ns
23.202ns
20394.316**
4688.346**
97.379*
26.933ns
50978.050**
0.055ns
2802.531*
14.784ns
8.746ns
1100.462ns
2.119*
0.032ns
3455.605**
74.978*
33.598*
604.905ns
0.991ns
0.023ns
3288.040**
67.162*
15.210ns
1321.139ns
0.836
0.034
856.284
27.251
11.254
2051.446
8.44
3.98
14.47
27.61
9.04
ns, *,**: No significant and significant at p≤0.05 and p≤0.01, respectively
14.72
Ayda Hosseinzadeh-Mahootchi, Kazem Ghassemi-Golezani
18
Table 2
Means of maximum chlorophyll content index, stomatal conductance, relative water content,
membrane stability and grain yield for different irrigation treatments
Treatments
CCI
Fv/Fm
Stomatal Conductance
[mmol × m-2 × s-1]
RWC
[%]
MSI
[%]
Grain yield
[g × m-2]
Irrigation
I1
6.975a
0.7624a
410.9a
67.73a
89.02a
418.387a
I2
6.307b
0.6350a
327.9b
61.53b
82.55b
289.290b
I3
5.676c
0.5946a
232.0c
56.32b
81.25b
215.392c
Different letters at each column indicate significant difference at p≤ 0.05
I1, I2 and I3: Irrigation after 70,120 and 170 mm evaporation from class A pan, respectively
Table 3
Means of maximum chlorophyll content index and membrane stability for different unprimed
and primed seed lots
Traits
CCI
MSI (%)
Treatment
P1
P2
P3
v1
6.448bc
6.804ab
7.570a
v2
5.864bc
6.134bc
6.394bc
v3
6.340bc
5.734c
5.587c
v1
81.90b
87.46a
86.78a
v2
83.22b
85.42ab
85.14ab
v3
82.06b
85.05ab
85.44ab
Different letters at each column indicate significant difference at p≤ 0.05
V1, V2and V3: Seed lots with 100, 85 and 74% viability, respectively
P1, P2 and P3: non-primed, primed for 12 and 18 hours, respectively
Fv/Fm
Efficiency of photosystem II (Fv/Fm) was not significantly affected by
water limitation, seed vigor and hydro-priming duration.
Stomatal conductance
Irrigation treatments, seed vigor and hydro-priming duration had significant effects on stomatal conductance (SC) of chickpea leaves. Interactions of irrigation ×
hydro-priming, seed vigor × hydro-priming and irrigation ×vigor× hydro-priming
for SC were also statistically significant (Table 1). Stomatal conductance of plants
from all seed lots significantly decreased with decreasing water availability (Table
2). Hydro-priming improved stomatal conductance of plants from all seed lots un-
Relationship between root and yield related morphological characters in pea (P.sativum L.)… 19
der all irrigation levels, but this advantage for plants from low vigor seed lots particularly under limited irrigations was higher than that for other treatments (Fig. 2).
Fig. 2. Stomatal conductance of chickpea plants from various seed lots under different irrigation treatments
I1, I2 and I3: Irrigation after 70,120 and 170 mm evaporation from class A pan, respectively
V1, V2and V3: Seed lots with 100, 85 and 74% viability, respectively
P1, P2 and P3: non-primed, primed for 12 and 18 hours, respectively
Relative Water Content (RWC)
Irrigation treatments, seed vigor and hydro-priming duration had significant
effects on relative water content (RWC) of chickpea leaves (Table 1). Interactions of seed vigor × hydro-priming and irrigation × vigor × hydropriming for RWC were also statistically different (Table 1). Leaf relative
water content was decreased as water deficit increased. However, there was
no significant difference between plants under I2 and I3 (Table 2). Plants
from high vigor seed lot (V1) under different irrigation treatments had
higher relative water content, compared with those from low vigor seed
lots. Hydro-priming improved relative water content of chickpea plants from
different seed lots under various irrigation treatments. This improvement for
plants from low vigor seed lot under severe water limitation was greater
than that under other irrigation treatments (Fig.3).
20
Ayda Hosseinzadeh-Mahootchi, Kazem Ghassemi-Golezani
Fig. 3. Relative water content of chickpea plants from various seed lots under different irrigation treatments
I1, I2 and I3: Irrigation after 70,120 and 170 mm evaporation from class A pan, respectively
V1 , V2and V3: Seed lots with 100, 85 and 74% viability, respectively
P1, P2 and P3: non-primed, primed for 12 and 18 hours, respectively
Membrane Stability Index (MSI)
Membrane stability index of chickpea leaves was significantly influenced
by irrigation levels. However, no significant effects of seed vigor and hydro
-priming duration on this trait were found (Table 1). Interaction of seed
vigor × hydro-priming for MSI was also significant. Membrane stability of
chickpea leaves was decreased with increasing water severity (Table 2).
Hydro-priming enhanced membrane stability of chickpea plants from all
seed lots (Table3).
Grain yield
Grain yield was significantly influenced by irrigation treatments, seed
vigor and hydro-priming duration (Table 1). Interaction of irrigation × seed
vigor for grain yield was also significant (Table 1). Grain yield among
plants from various seed lots diminished with increasing water limitation
(Figure 4, Table 2). Plants from high vigor seed lot (V1) under different irrigation treatments had higher grain yield, compared with those from low
vigor seed lots (Fig. 4). Hydro-priming significantly enhanced grain yield
from 258.19 g/m2 to 339.55 g/m2.
Relationship between root and yield related morphological characters in pea (P.sativum L.)… 21
Fig. 4. Mean grain yield of chickpea for different seed lots under different irrigation treatments
Different letters at each column indicate significant difference at p≤ 0.05 I1, I2 and I3
: Irrigation after 70,120 and 170 mm evaporation from class A pan, respectively
V1, V2and V3: Seed lots with 100, 85 and 74% viability, respectively
Relative water content, membrane stability index, chlorophyll content
index and stomatal conductance had significant and positive correlations
with each other and also with grain yield. RWC and stomatal conductance
showed the highest positive correlations with grain yield (Table 4).
Table 4
Correlation coefficients among some physiological parameters of chickpea
Traits
1. Relative water content
1
2
3
4
5
6
1
2. Membrane stability index
0.650**
1
3. Fv/Fm
0.363ns
0.550**
1
4. Chlorophyll content index
0.680**
0.528**
0.348 ns
5. Stomatal conductance
0.790**
0.771**
0.505**
0.670**
6. Grain yield
0.809**
0.696**
0.355 ns
0.747**
ns ,**: no significant and significant at and p≤0.01
1
1
0.808**
1
22
Ayda Hosseinzadeh-Mahootchi, Kazem Ghassemi-Golezani
DISCUSSION
Greater beneficial effect of hydro-priming on chlorophyll content of plants
from V1 and V2 seed lots (Fig.1) could be attributed to early improvements in
rate and uniformity of seed germination and seedling emergence (Farooq et
al., 2005, Ghassemi-Golezani et al., 2010a). Rapid emergence of seedlings
could lead to the production of vigorous plants (Ghassemi-Golezani et al.,
2008a) with high chlorophyll content in their leaves (Ghassemi-Golezani et
al., 2008b) as it was shown for plants from high vigor seed lots of chickpea
(Table 2). Delayed emergence of seedlings from poor vigor seed lots resulted in inefficient use of environmental resources. Early decline in leaf
chlorophyll content of these plants at later stages of growth (Fig. 1) closely
related with poor resistance to stressful condition. The reduction of chlorophyll content was probably related to the enhanced activity of the enzyme
chlorophyllase (Reddy and Vora, 1986) and inducing the destruction of
chloroplast structure and the instability of pigment protein complex (Singh
and Dubey, 1995).
No significant effect of water stress on photosystem II efficiency (Fv/Fm)
(Table 1) was a result of higher resistance of photosynthetic machinery to
water limitation (Chaves et al., 2002). Photo damage may be prevented through
processes of thermal deactivation, down regulating PSII photochemistry, in
a so-called dynamic photo inhibition (Osmond, 1994), that brings the electron transport capacity into balance with carbon metabolism (Epron et al.,
1992).
Decreasing water availability decreased stomatal conductance of chickpea
plants (Fig. 2), due to the closure stomata which decrease transpiration rate
and loss of water. During drought, leaves are subjected to both heat and water
deficiency stress (Clarke et al., 1993). As a consequence of the reduction in
transpiration rates of leaves, leaf temperature increases (Kusvuran, 2012).
Improved stomatal conductance of plants from primed seeds under all irrigation
treatments (Fig. 2) was due to rapid emergence of plants and higher resistance
of vigorous plants to unfavorable conditions (Ghassemi-Golezani et al.,
2012).
The decrease in leaf RWC (Fig. 3) could be related with low water availability under stress conditions (Shalhevet, 1993), or to poor root system,
which is not able to compensate for water loss by transpiration (Gadallah,
2000). Higher RWC of plants from high vigor seed lot (V1) and the efficiency of seed hydro-priming for better RWC under stressful condition
were the result of rapid and uniform seedling emergence (GhassemiGolezani et al., 2012). This can lead to the production of vigorous plants
with a potential to use environmental resources efficiently. The resulting
plants better tolerate drought stress, reduce pest damage and increase crop
yield (Harris et al., 1999).
Relationship between root and yield related morphological characters in pea (P.sativum L.)… 23
Membrane lipids peroxidation, membrane damage and ion leakage under
water stress (Katsuhara et al., 2005) led to reduction in membrane stability
index (MSI) of chickpea leaves (Table 2). Beneficial effects of hydropriming on membrane stability of chickpea plants (Table 3) could be attributed
to maintenance of positive leaf turgor, efficient and longer use of plants
from soil resources under water stress by early establishment of seedlings.
Water limitation considerably reduced grain yield of chickpea, due to
reductions in leaf, stomatal conductance, chlorophyll content, relative water
content and membrane stability (Table 2). The earliest response to the leaf
water deficit is stomata closure, which limits CO2 diffusion to chloroplasts
and limits photosynthesis (Cronic and Masacci, 1996). Inhibition of chlorophyll
synthesis due to water stress (Fig. 1, Sayed, 2003) and decrease of relative
water content affected by low water potential (Fig. 2, Krouma, 2010) can influence
plant growth (Ohashi et al., 2000) and field performance (Fig. 4).
Lower grain yield of plants from low vigor seed lots (Fig. 4) related to
slow emergence of seedlings from aged seeds, poor stand establishment and
delayed flowering of plants (Ghassemi-Golezani et al., 2010b). The advantage
of high vigor seeds in improving field performance and enhancement of grain
yield of chickpea due to hydro-priming (Fig. 4) directly related with rapid
seedling emergence, optimal stand establishment, efficient and longer use of
plants from light and soil resources during growth and development and
production of larger grains under all irrigation treatments (GhassemiGolezani et al., 2012). Decreasing the superiority of plants from high vigor
seeds under low water supply (Figure 4) is the result of early emergence
and high density of plants from these seeds which increased competition of
individual plants for water and other resources under limited irrigation conditions
(Ghassemi-Golezani et al., 2012). High positive correlations of stomatal
conductance and relative water content with grain yield (Table 3) indicate
that these parameters can be used to estimate potential field performance of
chickpea under different irrigation conditions.
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