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J.

Agronomy & Crop Science (2008) ISSN 0931-2250

CHILLING/FREEZING STRESS

Seed Priming Enhances the Performance of Late Sown


Wheat (Triticum aestivum L.) by Improving Chilling
Tolerance
M. Farooq1, S. M. A. Basra2, H. Rehman2 & B. A. Saleem3
1 Department of Agronomy, University of Agriculture, Faisalabad, Pakistan
2 Department of Crop Physiology, University of Agriculture, Faisalabad, Pakistan
3 Institute of Horticultural Sciences, University of Agriculture, Faisalabad, Pakistan

Keywords
growth; late sowing; seed priming; stand
establishment; wheat; yield
Correspondence
Dr M. Farooq
Department of Agronomy, University of
Agriculture, Faisalabad 38040, Pakistan
Tel.: +92 41 9200161-9/2936
Fax: +92 41 9201098
Email: farooqcp@gmail.com
Accepted October 9, 2007
doi:10.1111/j.1439-037X.2007.00287.x

Abstract
In ricewheat systems, late sowing of wheat is the major reason of low yield.
This yield reduction is principally due to lower and erratic germination, and
poor crop establishment because of low temperature prevailing. The present
study was conducted to explore the possibility of improving late sown wheat
performance by seed priming techniques. Seed priming strategies were: on-farm
seed priming, hydropriming for 24 h, seed hardening for 12 h and osmohardening with KCl or CaCl2 for 12 h. Seed priming improved emergence, stand
establishment, tiller numbers, allometry, grain and straw yield, and harvest
index. However, seed priming techniques did not affect plant height, number
of spikelets, number of grains and 1000 grain weight. Osmohardening with
CaCl2 gave more grain and straw yield and harvest index compared with control and other priming treatments, followed by osmohardening with KCl and
on-farm seed priming. Improved yield was attributed principally to better stand
establishment and improved number of fertile tillers. Seed priming techniques
can be effectively used to improve the performance of late sown wheat.

Introduction
Ricewheat systems cover about 32 % of the total rice
area and 42 % of the total wheat area in India, Pakistan,
Bangladesh and Nepal (Iqbal et al. 2002). The farmers in
the ricewheat zone of Punjab in India and Pakistan principally grow basmati varieties, which are late maturing
compared with coarse varieties of rice. Therefore, paddy
harvest is generally delayed at most of the farms in this
area. The late paddy harvest coupled with poor soil structure and loose plant residues creates problems for preparation of a good seedbed, and planting of wheat often
gets late (Byerlee et al. 1984). Moreover, the occurrence
of rain during land preparation operations may cause a
further delay of 23 weeks in wheat sowing (Aslam et al.
1993), and late planting is the principal limitation to
wheat productivity in many areas of South Asia (Hobbs
et al. 1994, Ahmed and Meisner 1996). Studies have
reported that after mid-November a days delay in
planting of wheat results in a yield loss of 1 % ha)1
2007 The Authors
Journal compilation 2007 Blackwell Verlag, 194 (2008) 5560

(Randhawa et al. 1981, Hobbs et al. 1988). Farmers also


have to resort to the broadcast method for wheat sowing
which results in poor and patchy plant stands.
Good crop establishment is one of the major challenges
to crop production and its importance is recognized by
farmers as well as researchers (Chivasa et al. 1998). Constraints to good establishment include seedbed preparation (Joshi 1987), low quality seed, untimely sowing (van
Oosterom et al. 1996), poor sowing technique (Radford
1983), inadequate soil moisture (Harris 1996), adverse
soil conditions (Lee et al. 1998) and high temperatures
(Ougham et al. 1988, Peacock et al. 1993).
Seed priming has been found a doable technology to
enhance rapid and uniform emergence, high vigour, and
better yields in vegetable and flower species (Dearman et al.
1987, Parera and Cantliffe 1994, Bruggink et al. 1999)
and some field crops (Hartz and Caprile 1995, Chiu et al.
2002, Giri and Schillinger 2003, Murungu et al. 2004,
Basra et al. 2005, 2006, Kaur et al. 2005, Farooq et al.
2006a,b, 2007a,b). Seed priming has been commonly
55

Farooq et al.

employed to reduce the time between seed sowing and


seedling emergence and the synchronization of emergence
(Parera and Cantliffe 1994). Priming of wheat seed in
osmoticum or water may improve germination and emergence (Ashraf and Abu-Shakra 1978) and promote
vigorous root growth (Carceller and Soriano 1972) under
low soil water potential compared with untreated seeds.
Harris et al. (1999) promoted a low-cost, low-risk technology called on-farm seed priming that would be
appropriate for all farmers, irrespective of their socioeconomic status. On-farm seed priming involves soaking the
seed in water, surface drying and sowing the same day.
The rationale is that sowing soaked seed decreases the
time needed for germination and may allow the seedling
to escape deteriorating soil physical conditions. Besides
better establishment, farmers have reported that primed
crops grew more vigorously, flowered earlier and yielded
higher. In wheat, researchers have recorded mean yield
increases in six large series of on-farm trials of 5 % (nonprimed yield 4.2 t ha)1) to 36 % (non-primed yield 1.4
t ha)1) (Harris et al. 2001).
Although Kant et al. (2003, 2006) reported very valuable
findings, there is still a lot of potential to improve stand
establishment, growth and yield of late sown wheat in rice
wheat systems. No comprehensive report is available in this
regard. The present study was therefore conducted to
investigate the possibility of improving the germination,
crop stand establishment, allometry and grain yield of late
sown wheat in ricewheat systems by seed priming.

27 3 C for 12 h and redrying to initial moisture


content and this cycle repeated once and (d, e) osmohardening, similar to hardening but in the presence of
CaCl2 or KCl solutions of ws = )1.25 MPa (Farooq et al.
2006c). Seeds receiving no prior treatment served as
controls. Primed seeds were given three washings with
water and re-dried closer to original moisture (ca. 8 %)
under forced air at 27 3 C (except for on-farm priming, where seeds were surface dried). These seeds were
put in polythene bags and stored in a refrigerator at
5 1 C until use (except for on-farm priming, where
freshly primed seeds were used).
In all the plots, four cultivations with a tractor-drawn
cultivator followed by two plankings were given to
achieve the desirable soil structure, and crop was sown
with a seed drill in 22 cm-apart rows on 15 December.
In all treatments, the seeding rate was 100 kg ha)1. Fertilizers P and K were applied basally at the rate of 60 and
25 kg ha)1 respectively. Nitrogen was applied at the rate
of 110 kg ha)1, of which 50 % was applied basally, 25 %
at crown root initiation and 25 % at flowering. Four irrigations were given to all treatments at crown root initiation, tillering, flowering and the dough stage. The crop
was harvested when spikes were fully ripened at the
approximate moisture of 20 %, and each plot was
threshed separately. During the course of the study, mean
minimum and maximum temperatures (DecemberApril)
were 5 and 31 C, respectively, while mean precipitation
was 30 mm.

Materials and Methods

Measurements

Site description
The study was conducted at a farmers plot (31.45N,
73.26E and 193 m) in 20052006. Experimental soil was
sandy, sandy loam with pH 8.1, 0.30 mS cm)1 total
exchangeable salts and 0.75 % organic matter.

Number of emerged seeds was recorded daily according


to the seedling evaluation handbook of the Association of
Official Seed Analysis (AOSA) (1983). Time taken to
50 % emergence of seedlings (E50) was calculated according to the following formula of Coolbear et al. (1984)
modified by Farooq et al. (2005):
N

Experimental details
Seeds of wheat (Triticum aestivum L. cv. Auqab 2000)
used for this study were obtained from the Punjab Seed
Corporation, Faisalabad, Pakistan. Moisture content of
the seed was ca. 8 %. The experiment was laid out in a
randomized block design with three replicates and plot
sizes of 10 m 7 m.
Seed priming treatments used in the study were
selected from previous experience (Basra et al. 2005,
2006, Farooq et al. 2006a,b, 2007a,b). These were: (a)
on-farm priming, soaking in tap water for 14 h, (b) hydropriming, soaking seed in aerated distilled water for
24 h, (c) hardening, soaking seeds in tap water at
56

E50 ti


 ni tj  ti
nj  ni

where N is the final number of emerged seeds, and ni and


nj are the cumulative number of seeds emerged by adjacent counts at times ti and tj when ni < N/2 < nj.
Mean emergence time (MET) was calculated according
to the equation of Ellis and Roberts (1981):
P
Dn
2
MET P
n
where n is the number of seeds which emerged on day D,
and D is the number of days counted from the beginning
of emergence.
2007 The Authors
Journal compilation 2007 Blackwell Verlag, 194 (2008) 5560

Seed Priming Improves the Performance of Late Sown Wheat

Coefficient of uniformity of emergence (CUE) was


calculated using the formula of Bewley and Black
(1985):
P
n
3
CUE P 

t  t2 n
where t is the time in days, starting from day 0, the day
of sowing, n is the number of seeds completing emergence on day t and t is equal to MET.
At harvesting, agronomic traits and yield components
were examined following standard procedures. From the
onset of physiological maturity, growth traits were examined four times at 15-day intervals, and yield components
and spike characteristics were recorded at full maturity.
Crop growth rate (CGR) was calculated using the
formula of Hunt (1978). Crop was harvested when fully
ripened to determine grain and straw yield, and harvest
index.
Statistical analysis
Data were statistically analysed using the software
MSTAT-C. Analysis of variance was used to test the significance of variance sources, while LSD test (P = 0.05)
Table 1 Effect of seed priming on seedling emergence and stand
establishment in late sown wheat

Treatments

Time to start
emergence
(days)

E50
(days)

Control
On-farm priming
Hydropriming
Hardening
Osmohardening KCl
Osmohardening CaCl2

8.82
7.46
7.25
7.46
7.71
7.14

10.13
8.37
8.23
8.43
8.33
8.16

a
b
b
b
b
b

a
b
b
b
b
b

MET
(days)

CUE

15 a
10 c
11 b
10 c
10 c
10 c

0.15
0.67
0.51
0.69
0.81
0.85

e
c
d
c
b
a

Mean values sharing the same letter in a column do not differ significantly at P = 0.05. E50, time to 50 % germination; MET, mean emergence time; CUE, coefficient of uniformity of emergence.

was used to compare the differences among treatment


means.
Results
Seed priming techniques significantly affected seedling
emergence and stand establishment in late sown wheat
(Table 1). All the seed priming techniques reduced the
time to start emergence, time to 50 % emergence (E50)
and MET compared with control. Minimum time to start
emergence, E50 and MET were recorded from osmohardening with CaCl2 that was similar to all the seed priming
treatments in case of time to start emergence and E50 and
with on-farm priming, hardening and osmohardening
with KCl in case of MET (Table 1). All the seed priming
treatments also improved the CUE compared with control; however, maximum CUE was recorded from osmohardening with CaCl2 followed by osmohardening with
KCl (Table 1).
Seed priming techniques also significantly affected tiller
numbers, grain and straw yield and harvest index, while
plant height, number of spikelets, number of grains and
1000 grain weight remained statistically unaffected
(Table 2). The maximum tiller numbers, fertile tillers,
grain yield and harvest index were recorded from osmohardening with CaCl2 that was similar to osmohardening
with KCl in case of tiller numbers, and fertile tillers.
However, maximum straw yield was recorded from osmohardening with KCl that was similar to osmohardening
with CaCl2 and on-farm seed priming (Table 2).
Seed priming treatments also significantly affected
CGR in late sown wheat (Fig. 1). Post-anthesis CGR was
higher than that of pre-anthesis in all the treatments
including control and all the priming techniques
improved CGR at both crop stages (Fig. 1). Maximum
CGR at both pre-anthesis and post-anthesis stages was
recorded from osmohardening with CaCl2 followed by
osmohardening with KCl that was similar to on-farm
priming (Fig. 1).

Table 2 Effect of seed priming on agronomic traits and yield components in late sown wheat

Treatments

Plant
height (cm)

Tillers
(m)2)

Fertile
tillers (m)2)

No. spikelets/
spike

No. grains/
spike

1000 grain
weight (g)

Straw
yield (t ha)1)

Grain
yield (t ha)1)

HI (%)

Control
On-farm priming
Hydropriming
Hardening
Osmohardening KCl
Osmohardening CaCl2

79.04
81.88
81.45
79.47
82.33
81.78

508 e
562 c
538 d
533 d
585 a
588 a

411 c
433 b
420 b
425 b
448 a
451 a

15.43
15.33
16.21
15.66
15.29
16.01

47.33
49.33
47.00
47.00
48.33
48.66

40.11
40.67
41.33
41.00
40.66
40.33

9.85
11.23
10.60
10.14
11.35
11.24

3.11
3.75
3.46
3.48
3.86
4.16

23.99
25.03
24.60
25.55
25.38
27.01

a
a
a
a
a
a

a
a
a
a
a
a

a
a
a
a
a
a

a
a
a
a
a
a

c
a
b
b
a
a

d
b
c
c
b
a

d
b
c
b
b
a

Means sharing the same letters in a column do not differ significantly at P = 0.05. HI, harvest index.
2007 The Authors
Journal compilation 2007 Blackwell Verlag, 194 (2008) 5560

57

Farooq et al.

Crop growth rate (g m2 d1))

18

Control

On-farm seed priming

Hydropriming

Hardening

Osmohardening KCl

Osmohardening CaCl2

15

12

theless, further investigations are imperative on plant


growth changes taking place in time and space. The
greater efficiency of osmohardening with CaCl2 and KCl
is possibly related to the osmotic advantage that both K+
and Ca2+ have in improving cell water status, and also in
that they act as cofactors in the activities of numerous
enzymes (Taiz and Zeiger 2006), most of which are active
when reserve mobilization and radical protrusion are in
progress (Farooq et al. 2006c).
In crux, seed priming techniques can be employed to
improve the performance of late sown wheat. Osmohardening performed better than all other treatments including control; nonetheless, CaCl2 was the most effective.

0
Post-anthesis

Pre-anthesis

Crop sage
Fig. 1 Effect of seed priming on crop growth rate (CGR) in late sown
wheat (S.E.).

Discussion
This study suggests that performance of late sown wheat
can be improved by employing seed priming techniques.
Prevalence of low temperature at sowing results in poor
seed germination, seedling establishment and vigour if
wheat is sown late (Table 1). Poor stand establishment
then results in less tillers and ultimately reduced grain
yield (Table 2). Seed priming improves the germination
rate, speed and uniformity even under less than optimum
field conditions (Lee et al. 1998, Kant et al. 2006) thus
enabling the establishment of uniform and good crop
stand establishment. Due to readily available food during
germination (Kaur et al. 2005, Farooq et al. 2006a,b),
primed seeds are better able to complete the process of
germination in a short time and cope with environmental
stresses including low temperature (Kant et al. 2006).
This early and unformed stand establishment enables the
crop to complete other phonological events in the crop
ontogeny well in time (Kant et al. 2003, 2006). Zheng
et al. (2002) also reported significantly higher and more
rapid germination of osmoprimed rice seeds at low temperature (5 C). Another manifestation of seed priming
was the substantial increase in the number of total and
fertile tillers (Table 1), stemming from the events taking
place during earlier stages of crop growth such as faster
production of more vigorous seedlings. Previous studies
showed that seed priming led to more uniform, vigorous
and faster emergence of seedlings, bestowing wide-ranging
phonological and yield-related benefits (Kaur et al. 2005,
Farooq et al. 2006a,b, 2007a,b). Higher post-anthesis
CGR than that of pre-anthesis seems to be the result of
better weather conditions prevailing at that time. None-

58

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