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Food Chemistry
journal homepage: www.elsevier.com/locate/foodchem
Rapid Communication
a r t i c l e
i n f o
Article history:
Available online 8 February 2012
Keywords:
Free-Air Carbon dioxide Enrichment (FACE)
Iron
Grain mineral composition
Protein
Zinc
a b s t r a c t
Wheat (Triticum aestivum L. cv. Yitpi) was grown in the Australian Grains Free-Air Carbon dioxide Enrichment (AGFACE) facility under current ambient [CO2] (384 lmol mol1) and elevated [CO2]
(550 lmol mol1) in combination with two different times of sowing (TOS) to investigate the interactive
effect of [CO2] and grain lling conditions on wheat grain quality. Grains were sampled at harvest maturity and their protein and mineral nutrient (Ca, S, Zn and Fe) concentrations were measured. Protein concentration of the grain was decreased by 12.7% at elevated [CO2] and the largest reduction in grain
protein was observed at the later TOS (TOS2). Concentration of grain S, Ca, Fe and Zn were also signicantly decreased at elevated [CO2]. Most of the grain mineral nutrient concentrations were signicantly
increased at the TOS2 suggesting that rising temperature together with increased water stress are likely
to offset some of the negative effects of elevated [CO2] on grain mineral concentrations.
2012 Elsevier Ltd. All rights reserved.
1. Introduction
Atmospheric carbon dioxide concentration [CO2] is expected to
reach 550 lmol mol1 by the middle of this century (Carter, Jones,
& Lu, 2007). Simultaneously, global temperature is predicted to rise
by an average of 1.54.5 C with more frequent occurrences of extreme climatic events such as heat waves and/or droughts (Carter
et al., 2007). Changes in these environmental variables directly or
indirectly affect plant growth, development, grain yield and quality
(Ainsworth & Rogers, 2007; Loladze, 2002; Seneweera, Makino,
Mae, & Basra, 2005).
Grain yield for C3 crops is expected to increase over the next
100 years due to increasing carbon dioxide concentration [CO2]
(Kimball et al., 2001), but this may occur at the cost of reducing
grain quality traits such as protein, mineral nutrient and starch
properties (Hgy et al., 2009; Rogers et al., 1998; Seneweera
et al., 1996). In particular, reduction of N concentration in grains
has been well documented under elevated carbon dioxide concentration [eCO2] (Erbs et al., 2010; Hgy et al., 2009; Kimball et al.,
2001). Other climatic factors such as high temperatures and soil
water stress are likely to have a negative impact on wheat grain
Corresponding author. Tel.: +61 3 5362 2164, mobile: +61 40187 9853; fax: +61
3 5362 0782.
E-mail address: samans@unimelb.edu.au (S. Seneweera).
0308-8146/$ - see front matter 2012 Elsevier Ltd. All rights reserved.
doi:10.1016/j.foodchem.2012.01.105
1308
conducted to investigate the impact of [eCO2] and high temperature during grain lling under low rainfall conditions. In this
experiment, we tested the hypothesis that [eCO2] further modies
the wheat grain quality in drier and warmer grain lling conditions. Delayed sowing treatment was introduced to mimic high
temperature, but other stress factors such as drought and vapour
pressure decit are also more common. For the rst time, we report the effects of [eCO2] and delayed sowing on the grain protein
and mineral concentration of wheat, using the FACE facility. The
ndings of this experiment will provide much needed information
on how climate change impacts on wheat grain quality.
2. Materials and methods
2.1. CO2 exposure and experiment conditions
The experiment was conducted during the growing seasons of
2008 and 2009 at the Department of Primary Industries Research
Station, Walpeup, Victoria, Australia (35120 S, 142000 E).
The experimental design was a completely randomized block
design. Wheat (Triticum aestivum L. cv. Yitpi) was grown in eight
[aCO2] rings (384 lmol CO2 mol1, 4 m diameter) and eight
[eCO2] rings (550 lmol CO2 mol1, 4 m diameter), completely randomized with two different sowing times: normal or TOS1 (late
May or early June) and late or TOS2 (mid or late June). The four
combinations of two levels of [CO2] and two sowing times were
allocated to the 16 rings with four replicates. Each ring consisted
of two plots (4 m long 1.8 m width) and the plots were sown
in eight rows of 0.21 m space between rows and 0.1 m space within rows.
[CO2] in the FACE rings (4 m diameter) was maintained at
550 lmol CO2 mol1 by injecting pure CO2 into the air from an
octagonal FACE ring (Mollah, Norton, & Huzzey, 2009). The ambient plot had no CO2 injection, where seasonal long average [CO2]
was 384 lmol CO2 mol1. The main soil type dominating in the
study site was Calcarosol according to the Australian Soil Classication Standards. Some of the growth conditions of TOS1 and
TOS2 in both growing seasons are described in Table 1. All agronomic practices in this experiment were similar to normal farming
practices of the region and fertilizer nitrogen supply was at a rate
of 50 kg/ha which is considered to be adequate for prevailing
growing conditions. Accumulated growing degree days (GDD)
were calculated by summing daily degree days according to Darroch and Baker (1990). Daily degree days were calculated as
Tn = (Tmax + Tmin)/2 Tb, where Tmax and Tmin are the maximum
and minimum daily temperatures, respectively, and Tb is the base
temperature (5 C).
2.2. Grain sample preparation
Ears harvested at maturity were dried at room temperature and
grains were separated and aspirated (Vacuum separator, Kimseed,
Australia) to remove the remaining husk and dust, and stored at
Table 1
Summary of climatic conditions. TOS1: sowing date according to local best practice, TOS2: late sowing date.
2008-TOS1
2008-TOS2
2009-TOS1
2009-TOS2
Days temperature 30
35 C during grain lling
Days
temperature > 35 C
during grain lling
TMAX during
grain lling (C)
TMAX during
growing season
(C)
Grain lling
duration
(days)
127
107
213
110
1372
1300
896
1755
4
4
2
3
3
4
3
12
25.5
26.6
26.2
35.5
19.0
20.2
20.3
22.8
43
37
67
19
1309
In both years, grain protein concentration was signicantly increased at TOS2 (17%, 16%) than at TOS1 (12%, 13%). On an average,
protein concentration increased by 40% and 17% at TOS2 in 2008
and 2009, respectively (Fig. 1). Such increases in protein concentration in TOS2 could be partly due to a lower grain yield which was
mainly due to higher temperatures and water stress experienced
in the later stages of plant development (Table 1). Increased grain
protein synthesis at high temperature (3035 C) has been documented elsewhere (Balla, Karsai, & Veisz, 2009; Panozzo & Eagles,
2000). In both years, TOS2 treatments plants experienced higher
temperatures (above 30 C) for 34 days during anthesis, which
may have accelerated grain protein synthesis. However, a smaller
increase of grain protein concentration in the 2009-TOS2 compared
to 2008-TOS2 could be related to the high temperature induced heat
shock (>40 C) experienced during grain lling (Balla et al., 2009). In
2009 TOS2 crops experienced > 35 C maximum temperature on the
3 days after anthesis including 7 days of >40 C maximum temperature during grain lling. In contrast, in the 2008-TOS2 growing
Table 2
Grain mineral concentration of Fe, Zn, S, Ca, grain yield and grain N uptake of Triticum aestivum L. cv. Yitpi grown under two [CO2] (ambient, 384 and 550 lmol mol1), two times
of sowing (normal, TOS1 and late, TOS2) in two growing seasons. Means and summary of two-way ANOVA results are shown.
Main effects
[aCO2]
[eCO2]
Percentage change
TOS1
TOS2
Percentage change
Interactive effects
TOS1
[aCO2]
[eCO2]
Percentage change
TOS2
[aCO2]
[eCO2]
Percentage change
ANOVA results
[CO2]
TOS
[CO2] TOS
S (g kg1)
Ca (g kg1)
35.00
31.50
(10%)
26.63
39.88
(49%)
31.13
24.25
(22%)
24.38
31.00
(27%)
1.75
1.66
(5%)
1.50
1.90
(27%)
0.33
0.29
(15%)
0.27
0.35
(28%)
102
164
(61%)
152
114
(25%)
2.7
3.7
(37%)
3.1
3.2
(3%)
27.50
25.75
(6%)
27.25
21.50
(21%)
1.52
1.49
(2%)
0.29
0.26
(11%)
120
186
(55%)
2.6
3.7
(42%)
42.50
37.25
(12%)
35.00
27.00
(23%)
1.97
1.84
(7%)
0.38
0.32
(17%)
89
141
(58%)
2.7
3.8
(40%)
ns
ns
**
***
***
***
***
ns
ns (p = 0.057)
ns
ns
ns
ns
ns
ns
ns
41.10
36.80
(11%)
38.90
39.00
(0.3%)
26.00
22.90
(12%)
22.0
26.88
(22%)
1.83
1.65
(10%)
1.59
1.88
(19%)
0.37
0.34
(9%)
0.39
0.32
(16%)
150
223
(48%)
203
170
(16%)
4.0
5.2
(30%)
4.7
4.5
(4%)
40.00
37.75
(6%)
22.75
21.25
(7%)
1.83
1.65
(7%)
0.40
0.38
(5%)
178
227
(28%)
4.4
5.0
(14%)
42.75
35.75
(15%)
29.25
24.50
(16%)
1.59
1.88
(12%)
0.35
0.30
(13%)
122
218
(79%)
3.6
5.5
(52%)
***
***
**
ns
ns
**
***
***
ns
ns
ns
ns
ns
ns (p = 0.09)
1310
Fig. 1. (a) Grain protein concentration at elevated and ambient [CO2], (b) grain protein concentration at TOS1 and TOS2, (c) grain protein yield/m2 ground area at elevated and
ambient [CO2], and (d) grain protein yield/m2 at TOS1 and TOS2 in both growing seasons.
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