Field Crops Research 169 (2014) 123131

Contents lists available at ScienceDirect

Field Crops Research

journal homepage: www.elsevier.com/locate/fcr

Inuence of boron nutrition on the rice productivity, kernel quality

and biofortication in different production systems
Atique-ur-Rehman a,b , Muhammad Farooq a,c,d, , Ahmad Nawaz a , Riaz Ahmad a
a

Department of Agronomy, University of Agriculture, Faisalabad 38040, Pakistan

Department of Agronomy, Bahauddin Zakariya University, Multan, Pakistan
The UWA Institute of Agriculture, The University of Western Australia, 35 Stirling Highway, Crawley 6009, WA, Australia
d
College of Food and Agricultural Sciences, King Saud University, Riyadh 11451, Saudi Arabia
b
c

a r t i c l e

i n f o

Article history:
Received 11 August 2014
Received in revised form
14 September 2014
Accepted 15 September 2014
Available online 17 October 2014
Keywords:
Boron
Application methods
Panicle sterility
Water relations
Kernel quality

a b s t r a c t
Boron (B) deciency is becoming a common problem in water-saving rice production systems of South
Asia. Boron nutrition can potentially improve the crop performance. This two-year eld study was conducted to evaluate the potential of pre-optimized B application through various methods in improving the
performance of rice grown in aerobic culture, alternate wetting and drying and ooded systems. Boron
was delivered as seed priming (0.1 mM B), foliar spray (200 mM B), and soil application (1 kg B ha1 ); no
B application and hydropriming were taken as control. Boron nutrition by either method improved the
plant water relations, total chlorophyll contents, morphological and yield related traits in different rice
production systems. In addition to improvement in kernel yield, boron application also improved the
kernel quality. Foliar application of boron in ooded rice and alternate wetting and drying gave maximum net returns than all other treatment combinations. In crux, B application through foliar spary or
seed priming may be an economically viable option to reduce panicle sterility, improve kernel quality,
rice growth and yield. Improvement in rice yield by B application is attributed to increase in kernel size
and decrease in panicle sterility.
2014 Elsevier B.V. All rights reserved.

1. Introduction
Rice is an important cereal crop being consumed as staple food
by millions across the globe. Conventionally, rice nursery seedlings
are transplanted in the paddy elds, which need more water and
labor inputs. On the eve of decreasing water resources and increasing labor cost, water and labor-saving rice production systems
may replace the conventional production system. Several watersaving rice production systems, such as alternate wetting and
drying (AWD), and aerobic rice are being propagated worldwide

Abbreviations: AWD, alternate wetting and drying; B, boron; Chl, chlorophyll;

cm, centimeter; dS m1 , desi siemen per meter; E, East; FAO, Food and Agriculture
Organization; FW, fresh weight; h, hour; kg m3 , kilogram per cubic meter; LSD,
least signicant difference; m, meter; m3 , cubic meter; masl, meter above see level;
mM, milli molar; N, north; nm, nanometer; ppm, parts per million; RWC, relative
water contents; USA, United State of America; USDA, United State Department of
Agriculture; Wd , dry weight; Wf , fresh weight; Ws , saturated weight; WUE, water
use efciency; Zn, zinc.
Corresponding author at: University of Agriculture, Department of Agronomy,
Faisalabad, Punjab 38040, Pakistan. Tel.: +92 41 920 1098; fax: +92 41 920 0605.
E-mail address: farooqcp@gmail.com (M. Farooq).
http://dx.doi.org/10.1016/j.fcr.2014.09.010
0378-4290/ 2014 Elsevier B.V. All rights reserved.

(Castaneda
et al., 2002), however certain factors like weed and
pest pressure, panicle sterility and lodging are hindering the wide
scale adoption and potential benets of these systems (Farooq et al.,
2011). Among these factors, panicle sterility is of vital importance
(Ikehashi and Araki, 1986; Maheswari et al., 2007), which reduces
the grains per panicle (Nieuwenhuis et al., 2002) leading towards
less rice yields (Gowri, 2005). Any moisture decit during owering and grain lling increases the panicle sterility (Ekanayake et al.,
1989; Bouman and Tuong, 2001; Mazid et al., 2002), which severely
affects the rice yields.
Like other nutrients, boron (B) is an essential nutrient for plant
growth and development. In plants, B takes part in nucleic acid,
carbohydrate, protein, phenol and indole acetic acid metabolisms,
cell wall synthesis, membrane integrity and function (Goldbach
et al., 2001). Boron is also considered to be associated with one
or more of the processes of calcium utilization, cell division,
owering/fruiting, carbohydrate and nitrogen metabolism, disease resistance, water relations, and catalyst for certain reactions
(Sprague, 1951). However, B deciency is spreading in most of rice
growing soils. Although considered tolerant, rice suffers with B deciency resulting in substantial yield loss (Cakmak and Rmheld,
1997; Rashid et al., 2004, 2009). Several factors including drought,

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Atique-ur-Rehman et al. / Field Crops Research 169 (2014) 123131

Table 1
Weather data at the experimental station during course of experimentation.
Months

June
July
August
September
October
November

Rainfall (mm)

Relative humidity (%)

Temperature ( C)

Sunshine (h)

Daily maximum

Daily minimum

Daily mean

2010

2011

2010

2011

2010

2011

2010

2011

2010

2011

2010

2011

1.0
277.8
22.6
86.5
0.0
0.0

78.3
118.1
92.6
155.1
0.4
0.0

40.0
63.6
74.6
66.8
59.6
62.3

55
70.3
74.7
75.8
61.0
61.2

40.1
36.0
34.9
33.9
32.9
27.1

38.6
34.7
34.1
32.9
32.2
27.6

27.7
27.9
26.1
23.3
19.7
10.5

26.0
26.0
25.5
23.6
17.2
13.3

33.9
31.9
30.5
28.6
26.3
18.8

32.3
30.4
29.8
28.3
17.2
20.4

9.4
9.0
6.0
7.9
7.6
8.5

9.4
9.0
5.4
6.9
7.6
8.5

Source: Agro-meteorology Cell, Department of Crop Physiology, University of Agriculture, Faisalabad, Pakistan

low soil pH, calcareous nature of soil, and B leaching and xation have been considered as the possible reasons of B deciency
(Goldberg, 1997; Shorrocks, 1997; Rashid et al., 2009). Boron application to rice elds increased rice growth and grain yields in soils
low in B (Rashid et al., 2004, 2009; Dunn et al., 2005; Hussain et al.,
2012); however, some problems with uptake of B in conditions of
low moisture in soil and/or application of B only upon evident B
deciency symptoms have been found (Rashid et al., 2004; Dunn
et al., 2005).
Boron availability is often associated with soil moisture conditions and limited soil water often limits B availability because its
release from organic complexes is restricted and ability of plants to
extract it from the soil is impaired (Tisdale et al., 1985). Rashid et al.
(2002) reported that B deciency not only causes severe reduction
in paddy yield, but also deteriorates its kernel quality (Rashid et al.,
2004). Thus B application may substantially increase the paddy
yield (Rashid et al., 2002).
Boron deciency has been identied as one of the most important factors causing sterility in cereals because of poor development
of anthers and pollen and failure of pollen germination (Cheng and
Rerkasem, 1993), especially in low moisture conditions. Improved
pollen vitality of rice with application of B has been known since
long (Garg et al., 1979).
For efcient uptake of B by the plant, application method plays
a vital role. Among different practices, soil application is the most
prevalent method of B addition in the developed world. However,
seed priming can be used as an attractive and easy physiological
strategy (Farooq et al., 2007, 2012), for applying micronutrients.
For instance in oats (Avena sativa L.), seed priming with B had
no marked effect on seed germination, but tillering was increased
substantially (Saric and Saciragic, 1969). Moreover, nutrient application through foliar sprays helps to cope with deciency rapidly.
In a study, Mohsen and Magda (2004) found that application of
zinc (Zn) through foliar application in wheat increased the yield
signicantly.
As B is important for humans as well, grain biofortication is a
pragmatic and easy option to enhance the level of micronutrients
in grains. Biofortication may be achieved through conventional breeding (genetic biofortication), transgenic approaches
and agronomic means (Cakmak, 2008). Agronomic biofortication is safe, cost-effective and short term approach of grain
nutrient enrichment (Mayer et al., 2008). Agronomic biofortication includes micronutrient delivery as soil, foliar fertilizers and
micronutrient enriched NPK fertilizers (Cakmak, 2008) and seed
priming.
Previously, we found that pre-optimized B application in rice
through seed priming, seed coating, foliar and soil application in
lab and pot experiments signicantly improved germination, early
seedling growth, tillering, leaf expansion, water relations, chlorophyll contents and kernel yield in different rice cultivars (Farooq
et al., 2011; Rehman et al., 2011; Rehman and Farooq, 2012).

However, these B application strategies were not evaluated under

eld conditions. So this study was conducted to investigate the
inuence of B nutrition on the rice productivity, kernel quality and
biofortication in different production systems.
2. Materials and methods
This two-year study was conducted at Agronomic Research
Area, University of Agriculture, Faisalabad (31.25 N, 73.06 E and
183 masl), Pakistan during kharif season of 2010 and 2011. The
experiment was laid out in randomized complete block design
in split-plot arrangement randomizing rice production systems in
main plots and B application methods in sub plots with net plot
size of 8 m 2.2 m. Before sowing of nursery and direct seeding of
aerobic rice, soil samples with the help of auger were collected
from 0 to 30 cm depth at different locations of the experimental site. Composite samples were air dried, grinded and passed
through 2 mm sieve to remove clods and material other than soil.
Soil was sandy loam having pH (8.2, 8.2), electrical conductivity (0.35, 0.36 dS m1 ), organic matter (0.92, 0.99%), total nitrogen
(0.06, 0.04%), available phosphorous (4.88, 4.80 ppm), exchangeable potassium (167, 177 ppm), zinc (0.80, 0.78 ppm) and boron
(0.56, 0.48 ppm) during both years, respectively. The experimental soil belongs to Lyallpur soil series (aridisol-ne-silty, mixed,
hyperthermic Ustalc), Haplarged in USDA classication and Haplic Yermosols in FAO classication. Meterological data during the
experimental period are given in Table 1.
The experiment consisted of three different rice production systems (aerobic rice, alternate wetting and drying (AWD) system and
ooded rice) and three boron application methods viz. seed priming
with 0. 1 mM B, soil application at 1 kg B ha1 and foliar application of 200 mM B. Hydropriming and no B application were taken
as control. Soil application of B was done as basal dose; while B
was foliage applied one week after sowing/transplanting. Boric acid
(10.65% B) was used as the source of B. In aerobic rice, land was prepared by four cultivations followed by two plankings. The detail of
land preparation, sowing time, seed rate, and other crop husbandry
practices is given in Table 2. The whole quantity of phosphorous,
potassium and zinc was applied as basal dose while half quantity
of nitrogen was applied as basal and other half was applied in two
splits at tillering and panicle initiation stage. Super Basmati was
used as test cultivar in all the rice production systems.
In aerobic rice, aerobic conditions were maintained throughout the crop growth period. After the seedling emergence, rst
irrigation was applied one and a half week after sowing. After that
supplementary irrigations were given as per requirement of the
crop. In total, 16 irrigations (52 acre inches) were given to reach
the crop at maturity stage. In AWD, nursery seedlings were transplanted in standing water (57 cm). Water was remained standing
for 30 days after transplanting; afterwards eld was allowed to
dry for some days. The irrigation was applied according to the

November 07
NPKZn at
143886825 kg ha1
NPKZn at
15982 g m2

July 22

July 22

40 g m2

40 g m2

Butachlor
(600 g a.i. ha1 )

6579 m3

November 07
NPKZn at
143886825 kg ha1
NPKZn at
15982 g m2
5962 m3

November 04
NPKZn at
143886825 kg ha1

5345 m3

65 kg ha1

Butachlor
(600 g a.i. ha1 )
+2 hoeings
Butachlor
(600 g a.i. ha1 )

Fertilizer applied
Fertilizer for
nursery
Transplanting
Seed rate

June 22
Flooded rice

June 22
Alternate wetting and
drying (AWD)

June 22

Soaking irrigation + 4
cultivations + 1 rotavator + 2
plankings + Direct drilling
Soaking irrigation + 4
cultivations + 2
plankings + Puddling + Nursery
transplanting
Soaking irrigation + 4
cultivations + 2
plankings + Puddling + Nursery
transplanting
Aerobic rice

Direct
seeding
Nursery
sowing
Seedbed preparation
Rice production
systems

Table 2
Detail of crop husbandry practices in different rice production systems during both experimental years.

Weed control

Irrigation

Harvesting

Atique-ur-Rehman et al. / Field Crops Research 169 (2014) 123131

125

crop requirement. A total of 18 irrigations (58 acre inches) were

applied till maturity. In ooded rice, nursery seedlings were transplanted in standing water (57 cm) and water was then kept
standing till physiological maturity. In total, 20 irrigations (64 acre
inches = 6578.57 m3 ), were applied to the crop in this system.
Penultimate leaves were harvested at tillering (55 days after
sowing) to determine different parameters of water relations. Leaf
water potential ( w ) was determined with pressure chamber. The
leaf tissues after determining w were frozen for 48 h, thawed, sap
was expressed, centrifuged at 5000 g and osmotic potential ( s )
was determined with an osmometer (Digital Osmometer, Wescor,
Logan, UT, USA). Relative leaf water contents (RWC) were calculated
following Barrs and Weatherly (1962). Fresh leaves were weighed
to get fresh weight (Wf ) and these weighed leaves were oated on
water for 4 h and weighed to record saturated weight (Ws ). These
leaves were dried at 85 C till constant weight to determine dry
weight (Wd ). Relative water contents were then calculated using
the following formulae,
RWC =

(Wf Wd )
100
(Ws Wd )

For leaf chlorophyll (chl) contents, penultimate leaves were collected at tillering (55 days after sowing). Leaf chl contents were
determined following Arnon (1949). According to this method, pigments were extracted in aqueous 80% acetone and determined in
the same solvent. The concentration of each Chl was determined by
measuring the extinction of the extract at the major red absorption
(QY) maxima of Chl a (664 nm) and b (647 nm). Net assimilation
rate (NAR) was estimated following Hunt (1978).
At maturity, plant height was measured with the help of meter
rod from the base of plant to the tip of ag leaf. Number of tillers
and productive tillers were counted from a unit area selected randomly from each plot. Number of kernels per panicle was counted
from 20 randomly selected panicles from each plot. Kernels were
separated from each panicle and counted separately. After harvesting and threshing of crop, samples were randomly taken from each
treatment, 1000 kernels were counted and weighed with electronic
balance. Whole plots were harvested and threshed separately and
clean rough rice was air-dried. Yield of each plot was weighed with
the help of screw weighing balance; values were adjusted to 12%
moisture and expressed in t ha1 . For panicle sterility estimation,
bulk samples of panicle from each plot were taken randomly at
maturity and then 10% panicles of the bulk samples were taken
as a sub-sample. From the sub-sample the number of competent
spikelets per panicle (a) and the total number of kernels per panicle (b) were counted. The percentage sterility was calculated by the
following Subedi et al. (1997):
Panicle sterility (%) =

(a b)
100
a

Water use efciency (WUE) was calculated as a ratio of kernel

yield and total water applied during the crop season.
Kernels developed into full length, size and were translucent, showed normal starch compaction and light passed through
them were taken as normal kernels. Random samples were taken
from the bulk of kernel yield and normal and abnormal kernels
were counted. Micro-Jheldahl digestion and ammonia distillation
process was used to determine the protein contents of the kernels. Then chlorimetric ammonia assay of the digest was used to
determine nitrogen concentration of the kernels, which was then
converted to protein contents by multiplying with the factor 5.95.
Milled rice kernels were grinded in a grinding mill for determination of kernel amylase contents (Juliano, 1971). The intensity of
blue color was read in a Spectrophotometer at 620 nm. For kernel
length, small samples of 100 kernels were taken from each treatment. Kernel length was measured with the help of digital vernier

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Atique-ur-Rehman et al. / Field Crops Research 169 (2014) 123131

Table 3
Inuence of boron nutrition on the water relations of rice grown in different production systems.
B application method

2010

2011

AR

AWD

FR

Mean

AR

AWD

FR

Mean

Water potential (MPa)

Control
Hydropriming
Seed priming with B
Foliar application of B
Soil application of B
Mean

2.06
1.89
1.73
1.12
1.17
1.59

2.02
1.88
1.65
1.04
1.21
1.56

2.15
1.84
1.57
1.11
1.22
1.57

2.08A
1.87B
1.65C
1.09D
1.19D

2.13
1.95
1.80
1.18
1.26
1.66

2.09
1.95
1.74
1.12
1.29
1.64

2.23
1.91
1.88
1.17
1.29
1.69

2.15A
1.94B
1.81C
1.16E
1.28D

Osmotic potential (MPa)

Control
Hydropriming
Seed priming with B
Foliar application of B
Soil application of B
Mean

2.39
2.04
1.89
1.70
1.91
1.99

2.64
2.04
1.91
1.77
1.83
2.04

2.47
2.05
2.07
1.68
1.89
2.03

2.51A
2.04B
1.96BC
1.72D
1.87C

2.47
2.09
1.97
1.68
1.99
2.04

2.72
2.07
1.99
1.85
1.91
2.11

2.62
2.08
2.14
1.69
1.97
2.11

2.60A
2.08B
2.04B
1.75C
1.96B

44.35
51.03
56.93
63.00
62.80
55.63

44.73
51.85
62.06
64.87
64.93
57.69

45.42
53.23
62.88
65.33
67.10
58.79

42.73
49.41
55.80
60.75
61.91
54.12

41.77
49.58
57.79
62.53
61.73
54.68

42.42
51.23
59.54
63.69
66.28
56.64

42.31D
50.07C
57.71B
62.32A
63.31A

Relative water contents (%)

Control
Hydropriming
Seed priming with B
Foliar application of B
Soil application of B
Mean

44.84D
52.04C
60.63B
64.40A
64.94A

Interaction and main effects not sharing the same letter for a parameter during an experimental year do not differ signicantly at p 0.05; ARaerobic rice; AWDalternate
wetting and drying; FRooded rice; Bboron.

caliper. Boron contents in kernels were determined according to

John et al. (1975), using a spectrophotometer (Perkin Elmer, CA,
USA) at 420 nm.
Data were analyzed statistically by employing the Fisher analysis of variance technique (Steel et al., 1996) and treatment means
were compared by using Least Signicance Difference (LSD) test at
5% probability level.
3. Results
Boron application methods signicantly affected the water relations of rice during both years (Table 3), but effect of different rice
production systems on water relations of rice was non-signicant
in both years. Foliar B application and soil application of boron
signicantly improved water potential during rst year while during second year foliar application was signicantly better than
soil application. These were followed by B priming during both
years (Table 3). Similarly, foliar application signicantly improved

osmotic potential than other treatments followed by soil application while for pressure potential and relative water contents both
foliar application and soil application were at par with each other
and these were followed by B priming during both years (Table 3).
Likewise, B application signicantly affected chlorophyll
contents of rice in the second year; results being non-signicant
for the rst year (Table 4). Total chlorophyll contents were highest in soil application followed by B priming, foliar application and
hydropriming, respectively (Table 4). Net assimilation rate was also
signicantly affected by rice production systems and boron application methods. Among rice production systems, net assimilation rate
was highest in ooded rice and lowest in aerobic rice in both years
(Table 4). Among B application methods, maximum net assimilation rate was recorded with soil application of B which was similar
with control. However, during second year, net assimilation rate
was highest in foliar application of B (Table 4).
Different B application methods and rice production systems
signicantly affected plant height during both years (Table 5).

Table 4
Inuence of boron nutrition on the chlorophyll contents and seasonal net assimilation rate of rice grown in different production systems.
B application method

2010

2011

AR

AWD

FR

Mean

AR

AWD

FR

Mean

Total chlorophyll (mg g1 FW)

Control
Hydropriming
Seed priming with B
Foliar application of B
Soil application of B
Mean

4.84
5.23
5.13
4.81
4.83
4.96

5.18
4.45
4.67
4.63
4.38
4.66

4.97
4.75
4.47
4.33
4.91
4.68

4.99
4.81
4.76
4.58
4.71

4.47
4.94
5.13
4.92
4.98
4.88

4.35
4.23
4.78
4.46
4.84
4.53

4.47
4.81
4.87
4.63
5.02
4.76

4.43B
4.66AB
4.93A
4.67AB
4.94A

Net assimilation rate (g m2 day1 )

Control
Hydropriming
Seed priming with B
Foliar application of B
Soil application of B
Mean

3.05
2.78
3.01
2.55
3.10
2.90C

2.65
2.97
2.88
2.82
4.89
3.24B

5.74
5.41
3.16
3.26
3.46
4.21A

3.82A
3.72B
3.02C
2.88D
3.82A

1.99
2.30
3.25
3.31
3.24
2.82C

2.45
2.63
3.71
3.80
3.83
3.28B

2.75
3.01
3.85
3.99
3.78
3.48A

2.40D
2.65C
3.60B
3.70A
3.62B

Interaction and main effects not sharing the same letter for a parameter during an experimental year do not differ signicantly at p 0.05; ARaerobic rice; AWDalternate
wetting and drying; FRooded rice; Bboron.

Atique-ur-Rehman et al. / Field Crops Research 169 (2014) 123131

127

Table 5
Inuence of boron nutrition on morphological, yield-related traits, kernel yield, panicle sterility and water use efciency of rice grown in different production systems.
B application method

2010
AR

Plant height (cm)

Control
Hydropriming
Seed priming with B
Foliar application of B
Soil application of B
Mean
Number of tillers (m2 )
Control
Hydropriming
Seed priming with B
Foliar application of B
Soil application of B
Mean
Number of productive tillers (m2 )
Control
Hydropriming
Seed priming with B
Foliar application of B
Soil application of B
Mean
Number of kernels per panicle
Control
Hydropriming
Seed priming with B
Foliar application of B
Soil application of B
Mean
1000-kernel weight (g)
Control
Hydropriming
Seed priming with B
Foliar application of B
Soil application of B
Mean
Kernel yield (t ha1 )
Control
Hydropriming
Seed priming with B
Foliar application of B
Soil application of B
Mean

2011
AWD

100.50h
106.50g
113.50de
109.50efg
108.00fg
107.60B

705e
675g
720d
780a
750b
726A

604f
573h
616e
674a
6478b
623A

111.00ef
116.50d
121.00bc
116.50d
123.00b
117.60A

FR

Mean

112.00ef
123.00b
122.00b
117.00cd
128.50a
120.50A

107.83C
115.33B
118.83A
114.33B
119.83A

AR

AWD

84.35
92.13
99.66
98.17
96.04
94.07B

105.18
105.05
112.29
108.29
108.31
107.83A

FR

Mean

99.70
103.02
109.36
110.17
108.67
106.18A

96.42B
100.07B
107.11A
105.55A
104.34A

630i
690f
705e
720d
660h
681B

465k
720d
600j
735c
750b
654C

600E
695C
675D
745A
720B

559g
570ef
584ab
585a
576c-f
574.92A

483j
548h
573def
577b-e
558g
548C

493i
558g
583abc
581a-d
568f
557B

512D
559C
580A
581A
567B

528j
589g
603f
618de
558i
579B

365l
620d
500k
635c
650b
555C

499E
594C
573D
642A
618B

386g
405f
439ab
434bc
423d
417.

340h
413e
438ab
440ab
425d
411

346h
427cd
446a
445a
429cd
419

358D
415C
441A
440A
426B

90.67
92.83
107.13
107.80
107.27
101.14B

105.67
114.33
118.33
118.56
118.56
115.09A

108.00
111.78
127.55
110.78
113.12
114.25A

19.06
20.24
20.91
21.55
21.98
20.75

19.15
20.38
21.97
22.71
20.91
21.03

18.01
19.63
20.23
22.70
21.13
20.34

101.44C
106.32BC
117.67A
112.38AB
112.98AB

18.74C
20.08BC
21.04AB
22.32A
21.34AB

68.00
67.10
99.87
97.73
97.50
86.04B

20.04
20.45
21.08
20.73
21.05
20.67

83.73
89.67
110.03
111.53
114.13
101.82A

86.67
86.00
121.77
115.37
112.33
104.43A

79.47B
80.92B
110.56A
108.21A
107.99A

19.72
20.95
21.97
20.76
20.74
20.83

18.69
20.63
21.34
21.43
20.73
20.57

19.48
20.67
21.08
20.97
20.84

3.17e
3.31de
3.68c
3.62c
3.65c
3.48B

3.61cd
3.99b
4.37a
4.44a
3.99b
4.08A

3.53cd
3.48cde
4.25ab
4.37a
4.37a
4.01A

3.44B
3.59B
4.10A
4.15A
4.01A

2.47
2.84
3.11
3.30
3.34
3.01B

3.02
3.36
3.79
3.89
3.62
3.54AB

3.58
3.49
4.03
4.27
4.23
3.92A

3.02B
3.23B
3.64A
3.82A
3.73A

Panicle sterility (%)

Control
Hydropriming
Seed priming with B
Foliar application of B
Soil application of B
Mean

3.16d
3.54cd
3.39cd
3.19d
3.40cd
3.33B

3.64bcd
3.47cd
3.65bcd
3.67bcd
3.75bc
3.64A

4.94a
3.17d
4.05b
3.58bcd
3.25cd
3.79A

3.92A
3.39C
3.69AB
3.48BC
3.47BC

6.99
6.70
6.22
6.23
6.53
6.54

8.21
6.69
6.49
6.12
6.37
6.78

7.61
6.31
6.17
6.21
6.31
6.53

7.61A
6.57B
6.29B
6.18B
6.41B

Water use efciency (kg m3 )

Control
Hydropriming
Seed priming with B
Foliar application of B
Soil application of B
Mean

0.59
0.62
0.69
0.68
0.68
0.65B

0.60
0.67
0.73
0.75
0.67
0.68A

0.54
0.53
0.65
0.66
0.67
0.61C

0.58D
0.61C
0.69AB
0.70A
0.67B

0.37
0.44
0.49
0.50
0.50B
0.46B

0.44
0.46
0.57
0.62
0.52
0.52A

0.39
0.42
0.54
0.63
0.59
0.51A

0.40D
0.44C
0.53B
0.59A
0.54B

Interaction and main effects not sharing the same case letter for a parameter during an experimental year do not differ signicantly at p 0.05; ARaerobic rice; AWDalternate
wetting and drying; FRooded rice; Bboron.

During rst year of study, maximum plant height was recorded

with soil application of Boron in ooded rice (Table 5), followed
by soil application in AWD and B priming and hydropriming in
ooded rice. In aerobic rice, none of the treatment gave as much
height as obtained in ooded rice or AWD. During second year,
all B application methods improved plant height than control and
hydropriming and these were at par with each other (Table 5).

Regarding number of tillers, more tillers were recorded with foliar

application in aerobic rice followed by soil application in ooded
rice during rst year (Table 5). In second year, foliar application in
aerobic rice was also at the top followed by B priming in aerobic rice
and foliar application and B priming in ooded rice (Table 5). The
same trend was recorded for productive tillers during rst year.
However, during second year maximum productive tillers were

128

Atique-ur-Rehman et al. / Field Crops Research 169 (2014) 123131

Table 6
Inuence of boron nutrition on kernel quality and grain boron biofortication of rice grown in different production systems.
B application method

Normal Kernels (%)

Control
Hydropriming
Seed priming with B
Foliar application of B
Soil application of B
Mean

2010

2011

AR

AWD

FR

Mean

AR

AWD

FR

60.00
61.67
74.23
74.03
73.27
68.64

60.33
60.97
75.40
74.57
73.93
69.04

59.73
60.80
75.03
74.80
74.80
69.03

60.02C
61.14B
74.89A
74.47A
74.00A

59.10
60.67
73.83
73.53
72.53
67.93AB

59.07
59.77
75.07
74.23
73.67
68.36A

58.17
59.27
73.53
73.23
73.67
67.57B

7.24
7.24
7.13
7.12
7.15
7.17

7.25A
7.15A
7.12B
7.13B
7.13B

7.29
7.28
7.11
7.13
7.11
7.18

7.29
7.28
7.11
7.13
7.12
7.18

7.28
7.28
7.09
7.12
7.12
7.18

7.29A
7.28B
7.11E
7.13C
7.12D

27.26A
27.12A
24.94C
25.83B
25.26C

27.63
27.17
25.37
26.53
25.67
26.47

27.93
27.73
25.67
26.23
26.37
26.78

27.83
27.80
25.97
25.73
25.10
26.48

27.80A
27.57A
25.67B
26317B
25.71B

Kernel protein contents (%)

Control
Hydropriming
Seed priming with B
Foliar application of B
Soil application of B
Mean

7.26
7.27
7.09
7.09
7.09
7.16

Kernel amylose contents (%)

Control
Hydropriming
Seed priming with B
Foliar application of B
Soil application of B
Mean

27.16ab
26.49bc
24.47f
25.93cd
25.40de
25.89

7.26
7.23
7.14
7.17
7.15
7.18

27.23ab
27.40a
24.73ef
25.70cd
25.73cd
26.16

27.40a
27.47a
25.63d
25.87cd
24.63ef
26.20

Mean
58.78C
59.90B
74.14A
73.67A
73.29A

Kernel length (mm)

Control
Hydropriming
Seed priming with B
Foliar application of B
Soil application of B
Mean

6.18
6.24
6.28
6.29
6.28
6.25

6.21
6.23
6.26
6.27
6.27
6.25

6.18
6.27
6.29
6.32
6.31
6.27

6.19C
6.25B
6.28AB
6.29A
6.29AB

6.08
6.14
6.16
6.22
6.22
6.17

6.12
6.11
6.12
6.15
6.18
6.14

6.09
6.14
6.15
6.17
6.16
6.14

6.09C
6.13BC
6.14B
6.18A
6.19A

Kernel B contents (mg g1 )

Control
Hydropriming
Seed priming with B
Foliar application of B
Soil application of B
Mean

1.55
1.53
2.00
2.46
2.01
1.91A

1.52
1.47
2.01
2.36
2.03
1.87B

1.51
1.56
1.92
1.96
1.87
1.76C

1.52C
1.52C
1.97B
2.26A
1.97B

1.43
1.41
1.88
2.34
1.89
1.79A

1.40
1.35
1.89
2.24
1.91
1.76B

1.39
1.44
1.80
1.84
1.75
1.64C

1.41C
1.40C
1.86B
2.14 A
1.85B

Interaction and main effects not sharing the same letter for a parameter during an experimental year do not differ signicantly at p 0.05; ARaerobic rice; AWDalternate
wetting and drying; FRooded rice; Bboron.

recorded with B priming and foliar application in ooded rice and

AWD and in B priming in aerobic rice (Table 5). Kernels per panicle and 1000-kernels weight was signicantly improved with B
application methods and all B application treatments were at par
with each other during rst year, however, 1000-kernel weight
was non-signicant during second year (Table 5). Boron application treatments signicantly improved kernel yield in AWD and
ooded rice and these were at par with each other, however, none
of the treatment recorded in aerobic rice produced as much yield as
in AWD or ooded rice. During second year, B application methods
were also at par with each other and among different production
systems, kernel yield was highest in ooded rice followed by AWD
(Table 5). Boron application signicantly decreased panicle sterility during both years and all these were at par with each other
(Table 5).
Regarding WUE, signicant differences were observed among
different rice production systems and B application methods in
both years. During rst year, WUE was highest in AWD while it
was lowest in ooded rice. However during second year, WUE was
similar in AWD and ooded rice and lowest in AR. Among boron
application methods, maximum WUE was recorded in foliar application of B in both years (Table 5).
Similarly B application methods signicantly improved the
quality of rice (Table 6). All B application methods improved normal
kernels and kernel length signicantly while protein and amylase
contents were not improved than control or hydropriming in both

years (Table 6). Moreover, maximum kernel boron contents were

recorded in AR during both years while they were minimum in
FR in both years (Table 6). Among the boron application methods,
maximum kernel B contents were found where B was foliar applied
(Table 6).
Economic analysis of different B application methods in various
rice production systems indicated that B application, through either
method, improved net return and benetcost ratio in aerobic rice,
AWD and ooded rice (Table 7). However, among all productions
systems and B application treatments, foliar application in ooded
rice gave maximum net returns and benetcost ratio (Table 7).
4. Discussion
Boron application signicantly improved water relations,
chlorophyll contents, morphology, yield-related traits, kernel quality and biofortication in rice. Improvement in water relations
due to boron application might be due to increased water uptake
through increased number of root tips and mycorrhizas (Mttnen
et al., 2001), which may have been resulted in more water uptake by
the plants helping them to grow vigorously. In our earlier studies,
we found that B application increased the root score in rice (Farooq
et al., 2011; Rehman et al., 2011), which might have enabled plants
to extract water efciently resulting in better water relations as
was observed in this study. Several other studies have reported
improvement in water status due to B application in various eld

CKcontrol; HPhydropriming; SPseed priming with boron; FAfoliar application of boron; SAsoil application of boron; B:C ratiobenet cost ratio; PKRPakistani rupees.

SA

4.30
3.87
1467.7
10.0
9
258.8
1726.4
14.94
1711.5
1.47
4.32
3.89
1474.5
11.1
10
287.5
1762.0
4.02
1758.0
1.54

FA
SP

4.14
3.73
1413.1
10.6
9.5
273.1
1686.2
2.3
1683.9
1.47
3.49
3.14
1189.5
10.1
9.1
261.6
1451.1
1.15
1450.0
1.27

HP
CK

3.56
3.20
1213.4
10.3
9.3
267.4
1480.8

1480.8
1.30
3.81
3.42
1298.7
10.3
9.3
267.4
1566.1
14.94
1551.2
1.34

SA
FA

4.17
3.75
1421.6
10.5
9.5
273.1
1694.7
4.02
1690.7
1.48
4.08
3.67
1392.6
10.3
9.2
264.5
1657.1
2.3
1654.8
1.45

SP
HP

3.68
3.31
1254.4
10.1
9.1
261.6
1516.0
1.15
1514.8
1.33
3.32
2.98
1131.5
10
9
258.8
1390.2

1046.6
0.92

CK

3.50
3.15
1192.9
10.5
9.4
270.3
1463.2
14.94
1448.2
1.25

SA
FA

3.46
3.11
1181.0
10.4
9.4
270.3
1451.2
4.02
1447.2
1.26
3.40
3.06
1158.8
10.3
9.3
267.4
1426.2
2.30
1423.9
1.25

SP
HP

3.08
2.77
1049.6
10.2
9.2
263.9
1313.5
1.15
1312.3
1.15

CK

2.82
2.54
962.5
10.1
9.1
261.6
1224.2

1224.2
1.07

Flooded rice
Alternate wetting and drying
Aerobic rice
Parameter

Table 7
Economics of rice as affected by boron application in different production systems.

Kernel yield
Adjusted yield
Value
Straw yield
Adjusted straw yield
Value
Gross benets
Cost that vary
Net benets
B:C ratio

Remarks

t ha1
10% less than actual
$ 15.17/40 kg
t ha1
10% less than actual
$ 1.15/40 kg
$ ha1
$ ha1
$ ha1
1$ = 87 PKR

Atique-ur-Rehman et al. / Field Crops Research 169 (2014) 123131

129

crops (Sharma and Ramchandra, 1990; Will et al., 2011; Bastias

et al., 2004), which establishes that B has direct role in plant water
relations. In this study, water status was poor in the rice plants
which were not supplied with the B. Few previous reports also
afrm that B decient leaves have poor water status than leaves
with normal B nutrition (Baker et al., 1956; Bastias et al., 2004).
Leaf chlorophyll contents of rice were increased with B seed
priming, foliar application and soil application of B, which might
be due to positive inuence of B on the Hill reaction activity and
net photosynthesis (Sharma and Ramchandra, 1990), which might
have increased the development of chloroplast. On the other hand,
we observed less chlorophyll contents in the rice plants which were
not feeded with B, which might be due to photosynthesis inhibition which might have decreased the amount and development of
chloroplasts (Wang et al., 2007). Boron deciency also results in a
signicant reduction in maximum quantum yield which is thought
to be related to the reduction of chlorophyll content and photochemical quenching of electrons (Plesnicar et al., 1997). Boron
deciency also severely affects the functions of enzymes and other
proteins in the plasma membrane, transport processes across the
membrane and membrane integrity (Cakmak and Rmheld, 1997;
Brown et al., 2002; Goldbach et al., 2002), which may have reduced
the chlorophyll contents in rice plants in which no B was applied.
It was also reported by Garcia-Gonzalez et al. (1990) that B deciency signicantly reduced the photosynthetic rate because of
reduced photosynthetic pigments in Anabaena. In another study,
it was found that B has a positive effect on chlorophyll contents of
the leaves as a result of which efciency and rate of photosynthesis
are increased (Zheng et al., 1989).
Boron application methods improved plant height and tillering intensity in rice, which might be due to active involvement of
B in meristematic growth of plant (Bohnsack and Albert, 1977),
which may have increased the cell division and elongation (Shelp,
1993; Mouhtaridou et al., 2004), resulting in more plant height
and improved tillering. Moreover, improvement in tillering and
plant height due to B application may be due to involvement
of B in regulation of plant hormones, stimulation of metabolic
pathways as well as the involvement of B in cell division and
its essentiality for actively growing regions of plants such as bud
development, root tips and new leaf (Ahmad et al., 2009). Some
recent reports have also indicated the involvement of B in cellular
signaling through interaction with some other transcription factors
(Gonzlez-Fontes et al., 2008) thus regulating the growth cascades.
Moreover, improvement in tillering might be due to increase in the
metabolic activities within the younger seedlings in the presence
of B (Goldbach et al., 2001). In another study, Wongmo et al. (2004)
reported that tillering was increased at lower level of B in some
wheat and barley genotypes. We observed more tillers in aerobic
rice than transplanted rice, which might be due to more number
of rice seeds per unit area. As seeds were directly sown in aerobic rice instead of nursery transplanting which was done in AWD
and ooded rice, more seeds were emerged from a unit area which
resulted in increased plant population, ultimately more number of
tillers and panicles per unit area.
Kernels per panicle and 1000-kernel weight was also improved
by B nutrition, which might be due to role of B in reproduction,
assimilate partitioning (Woodbridge et al., 1971; Dear and Lipsett,
1987; Noppakoonwong et al., 1997), pollen germination and pollen
tube growth (Mozafar, 1993), while B deciency reduces the pollen
germination and pollen tube growth (Bergmann, 1984). Moreover,
improvement in kernel weight by B application might be due to
improvement in source-sink relationship, which may have ensured
better assimilate translocation during grain development. In addition, increase kernel size may be due to increase in chlorophyll
contents which may have ensured the availability of photosynthetic
assimilates in stay green rice plants for longer time.

130

Atique-ur-Rehman et al. / Field Crops Research 169 (2014) 123131

We observed that B application improved the panicle fertility while more panicle sterility was observed in the plants where
no B was applied. Sterility is induced by B deciency in many
species of monocots and dicots, especially in cereals (Dell and
Huang, 1997), which reduces pollen viability (Subedi et al., 1998)
and pollen grains appear empty, misshaped, shrivelled, or may
be normal in shape but lack reserves of storage materials such
as starch (Dell and Huang, 1997). In wheat, severe B deciency
depressed anthers development resulting in small arrow-shaped
structures largely devoid of cells in the anther locules (Dell and
Huang, 1997). In another study in wheat, low B resulted in few
number of pollen grains (<100 per anther vs. normally 10002000
per another) with small size (<half normal size) and misshaped
(Rerkasem and Jamjod, 1997). Rawson (1996) also reported that
the primary cause of sterility is the inadequate B transport to the
owers in the critical 610 days of pollen formation. Moreover,
Zhang et al. (1994) also reported that boron deciency in oilseed
rape reduced the stamen growth resulting in small stamens (Zhang
et al., 1994).
All yield-related parameters were improved by B application
which might be due to role of B in reproductive growth, especially owering, better fruit and seed set resulting in better yields
(Dear and Lipsett, 1987; Noppakoonwong et al., 1997). Improvement in yield contributing parameters due to B application resulted
in improved kernel yield which might be the result of more number
of productive tillers, increased kernel weight and reduced panicle
sterility. Boron application also improved the net assimilation rate
and water use efciency than control. This increase in water use
efciency may be due to increased in yield per unit of water applied
due to boron application.
Application of B to rice not only increased its yield but it
also improved the kernel quality. In another study, Rashid et al.
(2004) also reported that inadequate B fertilization in rice, may
affect rice quality deteriorating its market price. They further
reported that application of B improved cooking quality of rice
which was visible through reduced stickiness, bursting upon cooking and increased elongation of kernel upon cooking. Similarly
kernel milling recovery and head recovery were also increased
with B application (Rashid et al., 2004). In some other studies, it
is well documented that nutritional quality of rice can be improved
with soil or foliar application of B (Rengel et al., 1999; Cakmak,
2002).
Boron application also improved the kernel B contents. In an
earlier study, Mayer et al. (2008) also argued that agronomic biofortication is safe cost-effective and short term approach of grain
nutrient enrichment. Moreover, increased net returns form foliar
application of B in ooded rice may be attributed to the fact that
only small quantity of B is required for foliar spray which reduced
the input cost.

5. Conclusion
B application has tremendous scope in rice production, however, application of B by foliar application or through seed priming
in various rice production systems is quite better and cost-effective
as it improved water relations, chlorophyll contents, growth,
yield, quality characteristics and boron biofortication in rice with
increased net benets. Thus B application in rice may improve rice
yield and quality by decreasing the panicle sterility.

Acknowledgements
This work was fully supported by the grant of the Higher Education Commission of Pakistan.

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