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Scientia Horticulturae
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Article history:
Received 25 August 2014
Received in revised form 18 March 2015
Accepted 19 March 2015
Keywords:
Allium sativum
Conditioning
Fructan
Sugar
Gene
1-SST
a b s t r a c t
Bulb size and regularity of shape and absence of defects determine garlic commercial quality, with these
factors being related to variety, environment and cultural practices. Low temperatures and short photoperiods at the beginning of development, followed by high temperatures and long photoperiods at the
end of the crop cycle are important for good bulb formation. Conditioning seed cloves at low temperatures before they are sown ahead harvesting and also decreases the incidence of undifferentiated bulbs;
however, its effects on growth pattern, fructan accumulation and gene expression of enzymes associated
with their synthesis are unknown. During 4 different seasons, sets of two hundred garlic bulbs Coreano
were conditioned at 5 C (5C) for 5 weeks and others sets were stored at room temperature (RT), the
cloves were separated and selected by weight (58 g). Three treated replicates, two hundred cloves per
replicate, were sown in commercial elds at Cosio Aguascalientes Mxico. Representative samples of
plants were analyzed for growth parameters, sugar and fructan contents, and 1-SST expression at different developmental stages. During the conditioning at 5C, the buds of different cloves were used to
1-SST expression. The 5C plants had shortened crop cycle (3242 days), and their growth rate was higher
(0.23 cm d1 ) compared to RT plants (0.013 cm d1 ). The fructan accumulation was also accelerated, and
this process could be associated with leaf senescence, apparently modulated by the high glucose content
in leaves of 5C plants (7.7 mg 100 g1 ) in comparison with the RT plants (1.5 mg 100 g1 ). However, the
5C plants did not reach either the yield or quality features of the RT plants. 1-SST expression was higher in
the garlic buds conditioned at 5C than in those of RT plants; however, with respect to plant development,
the expression was higher in all organs of RT plants. The conditioning of garlic bulbs at 5 C was effective
in shortening the harvest time and in eliminating the incidence of defects, allowing them to be harvested
when the garlic supply is low. The changes in fructan and sugar metabolism inuence growth pattern.
Published by Elsevier B.V.
1. Introduction
Corresponding author. Tel.: +52 442 192 1307x5579; fax: +52 4421921304.
E-mail addresses: lorenzogo@yahoo.com (L. Guevara-Olvera),
ramonggg66@gmail.com (R.G. Guevara Gonzlez), gene torres@hotmail.com
(H. Torres-Robles), mercado501120@gmail.com (E.M. Mercado-Silva).
http://dx.doi.org/10.1016/j.scienta.2015.03.030
0304-4238/Published by Elsevier B.V.
151
season, before the sowing, one set of 200 bulbs was stored at
room temperature (RT), 19 C and 77% RH average, and another
set was conditioned for ve weeks at 5 C (5C) and 98% RH average. The cloves from each bulb were separated, and selected those
with commercial size and weight (58 g), treated with fungicides,
nematicides, and bactericides to avoid pests and diseases in the
eld. The cloves of each treatment were divided in three replicates
(two hundred cloves per replicate) and sown in different commercial elds at Cosio, Aguascalientes. The cloves were sown in double
furrows at a distance of 7 cm from each other. All cultural practices
used were those commercially managed and recommended by the
Garlic Producer Association of Aguascalientes.
2.2. Indicative parameters of plant and bulb growth of garlic
The number of leaves, plant height, leaf width, bulb and
pseudostem diameter, plant weight, and bulbing index (bulb
diameter/pseudostem diameter) were used to evaluate coldconditioning effect on growth. Five different sampling periods were
conducted in each crop cycle. For each sampling period, ve plants
from each treatment and replicate were taken, transported on ice,
and washed before measuring the growth parameters.
The medium part of the leaves, pseudostem and bulb were separated and frozen in liquid nitrogen and stored at 70 C until their
analysis. Total yield and harvest quality were evaluated at the end of
each season. The harvest time was determined by the leaves senescence and a clear and complete differentiation of the cloves in the
bulb. The plants from a section of 5 m of the furrow were separated and curing in eld for two weeks (whole plants were left
on the eld until the foliage was dry), after which the bulbs were
weighed and sorted by defects and size accordance to commercial
garlic standards. The data were then extrapolated to one hectare.
2.3. Total fructan content
2.3.1. Fructan extraction
A pool from each plant tissue (bulb basal region, pseudo stem
and leaves) was made from ve plants, frozen with liquid nitrogen and freeze-dried before extraction; 0.1 g of freeze-dried sample
powder (0.5 mm particle size) of each organ was mixed in 80 C
water for 15 min then cooled. 50 mL of the mix was ltered (Whatman #. 1) to obtain the fructan raw extract. Total fructan content
was determined using the Fructan HK Assay test kit (Megazyme
International Ireland, Ltd., Wicklow, Ireland), following method
999.03 from the AOAC (McCleary et al., 2000) and the instructions
of the manufacturer.
2.4. Simple carbohydrates, fructooligosaccharides and fructan
prole determination
The carbohydrates were extracted from 100 mg of freeze-dried
tissue placed in 8 mL of water at 80 C, stirred for 15 min, then
cooled and ltered (Whatman #. 1) and made up to 10 mL. The
suspension was diluted to 0.5 mg/mL, and then ltered through a
Millipore nylon membrane (0.2 m). The solution was sonicated
and chromatographically analyzed.
2.4.1. High-performance anion-exchange chromatography with
pulsed amperometric detector (HPAECPAD)
A 25 L sample was injected into an ion chromatograph
Dionex ICS-3000 (Sunnyvale, CA, USA) equipped with a CarboPac
PA-100 (4 mm 50 mm) guard column and a CarboPac-PA100
(4 mm 250 mm) column. Fructan separation was achieved using a
gradient of sodium acetate, in 0.15 M NaOH, at a ow of 0.8 mL/min,
as follows: 05 min, 45 mM NaOH; 560 min, 0375 mM sodium
acetate; 6065 min, 500 mM sodium acetate; and 6575 min,
152
45 mM NaOH, at a column temperature of 25 C. Applied potentials, for detection by the amperometric pulse, were as follows: E1
(400 ms), E2 (20 ms), E3 (20 ms), and E4 (60 ms) of +0.1, 2.0, +0.6,
and 0.1 V, respectively (Martens and Frankenberger, 1990).
data were analyzed by ANOVA and the Tukey test (p < 0.05) for
means comparison, using the statistical software, JMP v.5.9. The
Gompertz equation was used to t the plant height data over
time or DAS. The model includes three constants of physiological
importance:
y = Aee
(BCt)
3. Results
3.1. Plant growth
Between 44 and 56 DAS no signicant differences were observed
on plant height; nevertheless, in the 87103 DAS period, 5C plants
showed greater height (4265 cm) than RT plants (3647 cm).
These differences were more evident at 122 and 137 DAS, 5C
plants were taller (68.778.2 cm), and had greater bulb diameter (36.447.4 mm), greater bulbing index (2.64.4), and a greater
weight gain (64133 g) than RT plants, although RT plants had more
leaves (8.59.8) with a greater width (2136.6 mm).
5C plants completed the crop cycle between 156 and 158 DAS
while RT plants completed it between 191 and 201 DAS, resulting in an earlier harvest of 3242 days in 5C plants. The statistical
comparison at the end of the crop development of both treatments
(157 and 191 DAS for 5C and RT plants respectively) showed different results than those described above (Table 1). After 191201
DAS, RT plants exhibited more and wider leaves, greater height,
greater pseudostem and bulb diameter, and more weight than were
observed in 5C plants. These data indicate that the growth of RT
plants continued after 157 DAS and they reached larger sizes than
did 5C plants by the end of their cycle.
Using the height data for all seasons and the conditioning treatment of the seed cloves into Gompertz model, the constants
values and the t goodness of model was calculated (Table 2).
The model adequately predicted the plant growth, noting that the
plants from 5C cloves had a limit of maximum growth and a turning
point (inection point on growth curve) signicantly lower than
plants from RT cloves. However, the growth rate of plants from 5C
cloves was almost double than shown by plants from RT cloves.
Early harvest represents an economic advantage when the supply of the product in the market is low. For that reason, the
protability of storing seed cloves at low temperatures should
be calculated taking into account the total yield and bulb quality.
The yield per hectare and the undifferentiated and defective bulbs,
as well as the production of each commercial size obtained in each
Table 1
Growth variables values at the end of the crop cycle of Garlic Coreano, obtained from seed cloves stored at room temperature (RT) or conditioning at 5 C (5C) and planted
on different seasons. Data are means standard deviation of 15 determinations.
Season treatment
2009/2010
RT
5C
2010/2011
RT
5C
2011/2012
RT
5C
2012/2013
RT
5C
DAS
Number of leaves
Plant height
(cm)
Leaf width
(mm)
Pseudostem diameter
(mm)
Bulb diameter
(mm)
BI
Weight
(g)
194
157
11.4 1.4a
5.3 0.2b
81.1 7.8a
63.4 7.5b
27.1 5.0a
18.9 3.3b
23.9 4.7a
11.9 3.6b
51.0 7.4a
45.4 6.6b
3.4 1.0b
7.8 2.9a
175.5 51.9a
80.0 29.2b
198
158
12.7 1.0a
9.2 0.8b
81.9 8.6a
68.8 6.1b
28.1 4.1a
21.1 3.6b
16.6 4.5a
7.6 2.3b
65.8 5.9a
53.5 5.1b
4.2 1.2b
7.7 2.8a
153.1 41.2a
96.2 24.1b
201
155
7.7 1.4a
6.6 3.0b
95.9 8.4a
77.7 11.0b
26.9 5.3a
22.2 4.5b
17.4 4.8a
16.0 4.9b
59.6 11.1a
56.9 9.4b
4.1 0.9a
3.8 0.9a
237.1 81.5a
154.3 79.5b
191
156
7.5 1.3a
5.1 1.2b
71.4 5.9a
77.3 6.7a
31.0 4.6a
27.6 3.7b
17.7 4.0a
13.6 3.0b
64.4 10.5a
57.0 5.4b
3.7 0.6b
4.3 0.7a
168.4 62.5a
129.6 37.5b
Different letters between the same column and factor, indicate signicant differences (Tukey P < 0.05). Days after sowing (DAS), bulbing index (BI).
153
Table 2
Parameters estimating garlic development by the Gompertz Model using the plant height (cm) and days after sowing (DAS) data.
Season
treatment
r2
RMSE
Parameters
Estimated
All seasons
5C
0.884
8.824
Asymptote
83.841
2.090
79.746
87.937
Growth rate
Inection point
Asymptote
0.023
48.956
96.587
0.002
1.664
2.717
0.020
45.693
91.261
0.026
52.219
101.912
Growth rate
Inection point
Asymptote
Growth rate
Inection point
Asymptote
Growth rate
Inection point
Asymptote
Growth rate
Inection point
Asymptote
Growth rate
Inection point
Asymptote
Growth rate
Inection point
Asymptote
Growth rate
Inection point
Asymptote
Growth rate
Inection point
Asymptote
Growth rate
Inection point
0.013
68.996
65.747
0.0461
24.605
90.908
0.012
58.332
79.282
0.023
50.524
105.326
0.014
82.284
82.580
0.026
40.535
111.051
0.011
79.737
101.081
0.019
66.107
87.291
0.016
61.167
0.0007
68.996
1.334
0.005
2.867
5.540
0.002
6.811
3.567
0.003
5.309
7.131
0.002
6.042
3.601
0.004
3.477
9.032
0.002
9.231
5.557
0.002
3.485
2.951
0.001
2.634
0.012
68.996
63.132
0.036
18.986
80.051
0.009
44.982
72.289
0.018
44.643
93.044
0.011
70.877
75.523
0.018
33.720
93.350
0.008
61.645
90.190
0.015
59.278
81.508
0.014
57.004
0.015
74.224
68.362
0.056
30.224
101.766
0.015
71.682
86.275
0.028
56.405
117.608
0.017
92.690
89.637
0.033
47.350
127.011
0.014
96.049
111.972
0.022
72.937
93.074
0.019
67.329
All Seasons
RT
20092010 5C
0.920
7.38
20092010 RT
0.878
8.93
20102011 5C
0.946
5.95
20102011 RT
0.933
7.73
20112012 5C
0.934
7.67
20112012 RT
0.883
10.36
20122013 5C
0.926
6.77
20122013 RT
0.881
7.56
Error STD
Lower
Upper
Conditioning at 5 C (5C), stored at room temperature (RT), regression coefcient (R2), root mean square error (RMSE).
Table 3
Total yield and harvest quality of garlic bulbs harvested at different seasons from seed cloves stored at room temperature (RT) or conditioning at 5 C (5C) for ve weeks.
Data are expressed as ton/ha. Undifferentiated bulbs not included into total yield.
Season treatment
Undifferentiated bulbs
Defects
C<6
C6
C7
C8
C9
C10
C11
C12
C > 12
20092010 RT
20102011 RT
20112012 RT
20092010 5C
20102011 5C
20112012 5C
16.3
15.8
20.1
12.3
9.7
10.0
4.6
0.3
3.3
0.0
0.0
0.0
0.8
0.7
1.1
2.4
0.4
0.5
0.3
0.1
1.4
0.4
0.4
0.4
0.5
0.2
0.2
1.1
0.8
0.6
2.8
0.6
0.4
1.8
2.7
1.8
2.5
1.3
0.6
3.1
3.7
2.5
0.6
2.9
3.9
2.5
1.7
1.7
3.7
3.6
3.7
0.9
0.0
1.9
3.2
3.8
4.8
0.1
0.0
0.3
1.0
2.6
2.9
0.0
0.0
0.3
0.9
0.0
1.1
0.0
0.0
0.0
Garlic size; diameter in cm: C6, 4.55; C7, 55.5; C8, 5.56; C9, 66.5; C10, 6.57.0; C11, 77.5; and C12, 7.58.
season were analyzed (Table 3). The overall yield was lower for
5C (9.712.3 ton per ha) than for RT plants (15.820.1 ton per ha).
Additionally, bulbs from RT plants showed higher calibers (C8 and
C9) compared with those from the 5C plants (C7C9). Nevertheless
bulbs from 5C plants did not show undifferentiated bulb production
whereas for RT plants, this production was high (0.34.6 ton per
ha). However, the successive conditioning at 5 C during consecutive seasons decreased more the overall yield (data not show) being
important for growers to maintain RT plants for seed production.
3.1.1. Changes in total fructan content
The conditioning only had a signicant effect on total fructan
content at 53 DAS (Table 4). However, as expected, the fructan
content in the bulbs was signicantly higher than in pseudostems.
The interaction temperature*organ conrmed these results; the
bulbs of both treatments presented higher values than those found
in pseudostems with no signicant differences between the two
treatments. These results conrmed that bulbs have a high capacity for storing these carbohydrates, and that pseudostems function
as temporary storage sites for these compounds, which are then
transported to the bulbs as the growing cycle takes place.
Table 4
Means comparison of fructan content (%DW) of bulb and pseudostem during development of garlic plants generated from seeds cloves conditioning at 5 C for 5
weeks or stored at room temperature (RT). Data from 2011 to 2012 season, each
data set represents the mean of 30 samples.
Factor/DAS
Temperature
5 C
RT
Organ
Bulb
Pseudostem
Temperature*organ
5 C*Bulb
RT*Bulb
5 C*Pseudostem
RT*Pseudostem
53
102
137
157
36.4a
28.9b
31.9a
18.1a
28.2a
26.3a
30.5a
25.0a
37.9
27.4b
35.0
24.9b
40.9
13.6b
45.2
10.4b
48.4
39.0a
36.8a
33.8a
20.9b
35.4a
34.7a
20.7a
29.2a
46.1a
35.6ab
6.4b
20.8ab
56.0a
34.4b
5.03c
15.7bc
54.2a
36.6ab
194
27.9
19.2b
Different letters between the same column and factor, indicate signicant differences (Tukey P < 0.05). Room temperature (RT). Days after sowing (DAS).
154
Fig. 1. Sugar (glucose, black circles and solid lines; fructose, white circles with dotted lines; sucrose, black triangles and short dashes) and fructooligosaccharides, FOS
(kestose, light gray bars; neo-kestose, white bars; nistose, gray bars; DP5, black bars) content during crop development of different garlic organs: leaves, pseudostems and
bulbs of RT plants (A, B, and C, respectively), and 5C plants (D, E and F, respectively). All data is presented by means of three samples with standard deviation bars. Data from
2011 to 2012 season.
a high sucrose content (Fig. 1D), but this increased until reaching
its highest values at 137 DAS (62.7 mg 100 g1 ) and then fell drastically at harvest time (5.8 mg 100 g1 ). Fructose content showed
a moderate increment (4.212.9 mg 100 g1 ) respect of the values observed for RT plants whereas glucose content showed wider
variation (0.37.7 mg 100 g1 ) in a shorter period of time, (53137
DAS). These data indicated a greater capacity for sucrose synthesis
or accumulation in 5C plants leaves up to 137 DAS, but this capacity drastically decreased in association with the bulb formation and
clove differentiation.
The highest glucose content observed in 5C plants leaves (137
DAS), could indicate a physiological change related to the sugar
signaling process associated with senescence of leaves.
On the other hand, the pseudostem of RT plants (Fig. 1B) showed
important differences in sugar content compared with that of 5C
plants (Fig. 1E). RT pseudostems showed a high sucrose content
at 53 and 102 DAS (24.6 and 44.6 mg 100 g1 ), which quickly
decreased at 137 and 157 DAS, and then increased again at the
harvest time, 194 DAS (41.9 mg 100 g1 ). Exhibiting a similar tendency, fructose content showed high values (11 mg 100 g1 ) at 53
DAS, which signicantly decreased at 157 DAS (0.6 mg 100 g1 )
and increased again at 194 DAS (37.8 mg 100 g1 ). Glucose content was very low during the rst 157 DAS and high at harvest time
(36.0 mg 100 g1 ). 5C plants pseudostems showed lower sucrose
content from 53 to 137 DAS (4.55.7 mg 100 g1 ), which also
increased at harvest time or 157 DAS (28.8 mg 100 g1 ), while
fructose and glucose were not detected within the 53137 DAS
period, and only high contents were registered at harvest time, 157
DAS (49.5 mg 100 g1 and 9.8 mg 100 g1 for fructose and glucose,
respectively).
As with the leaves and pseudostems, the RT plants bulbs also
showed important differences in sugars content (Fig. 1C) when
compared with the 5C plants bulbs (Fig. 1F). RT plants bulbs
showed increases in sucrose during the period from 53 to 137 DAS
(7.443.7 mg 100 g1 ), after which these levels reduced drastically
at 157 DAS and increased again at harvest time, 194 DAS (18.2 mg
100 g1 ). Fructose and glucose were not detected in RT plant bulbs.
The sucrose content of 5C plant bulbs was only determined at 53
DAS (11.1 mg 100 g1 ) and no content at 102 and 137 DAS was
recorded. It was determined again at harvest time, 157 DAS reaching its highest value (19.2 mg 100 g1 ). Fructose was determined
at 53 DAS (12.1 mg 100 g1 ) and at harvest time, 157 DAS showing
a lower value (0.6 mg 100 g1 , Fig. 1F). Similarly, to RT plants, 5C
plants did not show glucose content.
In a general way, the presence of these sugars in the three organs
could be related to the fructan synthesis and restructuring process
in those organs, which seems to be modulated by the length of
the crop cycle. This means that RT plants, with a longer crop cycle,
sucrose uctuations also occurred for longer periods; while for 5C
plants, that uctuation took place in a shorter time.
Fructooligosaccharides (FOS) content also showed important
differences between the two treatments. Leaves of RT plants
(Fig. 1A) showed lower levels of kestose (0.43.7 mg 100 g1 ),
neo-kestose (0.96.5 mg 100 g1 ), and nistose (0.32 mg 100 g1 )
whereas in 5C plant leaves, the FOS content was higher (kestose,
4.411.4 mg 100 g1 ; neo-kestose, 48.9 mg 100 g1 ). This data
showed that conditioning at 5 C induced a high temporary accumulation of FOS in leaves of 5C plants; this effect could be due to
an increased induction of 1-SST and 6G-FFT activities, key enzymes
of 1-kestose, and neo-kestose synthesis, respectively.
1-kestose content in RT plant pseudostems (Fig. 1B) decreased
as the crop cycle proceeded (6.73.4 mg 100 g1 ). Neo-kestose
showed values between 7 and 8 mg 100 g1 at 53 and 102 DAS
respectively, and after that, it decreased at 137 DAS and then
increased from six to 11 mg 100 g1 at 157 and 194 DAS. Nistose remained relatively constant while the FOS DP5 (degree of
155
156
Fig. 2. Changes in high molecular weight fructan content during crop cycle expressed as sum of areas under chromatograms HPAEC-PAD. RT plants (A) and 5C plants (B).
Bars represent means of three samples with standard deviation bars. Data from 2011 to 2012 season.
Fig. 3. 1-SST gene expression level in garlic sprouts during conditioning at 5 C or room temperature (A), and in different organs during crop development (B). Room
temperature (RT). Conditioning at 5 C (5C). Bulb (b). Pseudostem (Ps). Leaves (L). Days after sowing (DAS). ACTIN was used as positive control to normalize the expression
levels. Data from 2011 to 2012 season.
4. Discussion
The early harvest time of Coreano garlic resulting from conditioning the seed cloves at 5 C coincides with the accelerated
bulb formation period previously reported by Del Pozo et al. (1997)
and Braz et al. (1997) in garlic varieties from Chile and Brazil.
Other studies (Rahim and Fordham, 1988, 2001) had observed an
advancement of 15 days in harvest time when they conditioned
seed cloves for 30 days at 5 C; their scanning electron microscopy
observations indicated that clove primordia appeared two weeks
before in 5C plants compared with that of the control plants. Additionally, they observed that bulb development and differentiation
were completed at 87 and 117 DAS in both treatments, respectively.
Nevertheless, they also indicated that the bulbs of 5C plants had
fewer but larger cells; these data conrm our results of the early
harvest, and seem to explain the low yield and bulb sizes observed.
Khokhar et al. (2007) working with onions, also observed a
shortening of the harvest time, with fewer leaves, lower bulb fresh
weight, and lower yields. These authors also observed high induction of ower stalk bolting in 5C plants and they hypothesized
that the metabolism of plants conditioned at low temperature was
strongly inuenced by this structure, which produced plants with
fewer leaves. In this work, the ower stalk incidence was over
90% higher in 5C plants than in RT plants, which could modify the
metabolism, accelerate carbohydrate transport, and the senescence
of leaves and pseudostems. In contrast, Ade-Ademilua et al. (2009)
conditioned seed cloves at 8 C for 15 days, and after 70 DAS in
open eld observed a higher number of leaves, greater leaf area,
larger size, higher fresh weight and dry weight when compared
with plants with no conditioning. However, they did not report the
treatment effects on yield and bulb quality at the end of the crop
cycle. Thus, our results only match partially with their results.
The data indicate that the conditioning at 5 C induced a fast
emergence and growth of garlic 5C plants during the early stages of
the crop cycle; however, this was reversed during the intermediate
and nal stages by the RT plants.
The Gompertz model conrmed that the conditioning of seed
cloves at 5 C for ve weeks induced a faster plant growth (high
growth rate), but did not reach the sizes achieved by the RT plants,
although these constants did not explain why 5C plants had a
smaller size and lower yields. Reis et al. (2014) used different
mathematical models (logistic, von Bertalanfy, and Gompertz) to
describe similarities and differences between garlic varieties in the
dry matter accumulation during growth. They indicated a good t
of the experimental data with the logistic model; however, other
models also showed high values of R2 .
The ndings of Rahim and Fordham (1988) with respect to
the early formation of clove primordia in 5C plants allowed us to
hypothesize that early clove differentiation induced early leaf and
pseudostem senescence that ended the crop cycle.
The bulbing index gives an indication of the physiological and
metabolic changes during plant development, which could indicate
the stage at which the photosynthates are directed toward bulbs,
and that the plant is ready to generate cloves as a propagule. Fig. 4
shows the change of this index during different seasons for RT and
5C plants and shows at least three stages. The rst stage (4050
DAS) corresponds to initial plant development when nutrients are
transferred from the seed clove to the new plant, the initial
Fig. 4. Bulbing index, BI (bulb diameter/pseudo stem diameter) for all seasons
(20092013). Solid circles represent means for room temperature (RT) conditioning
plants. White circles, means for 5 C (5C) conditioning plants. Solid line, Gompertz
regression for RT plants; and dotted line, Gompertz regression for 5C plants. Solid
arrows represent the different growth stages of RT plants, and white arrows represent the different growth stages for 5C plants.
157
158
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related genes and anthocyanin content in Coreano garlic (Allium sativum)
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download/Q/QC/E (consulted July/2014).
Khokhar, K.M., Hardley, P., Pearson, S., 2007. Effect of cold temperature durations
of onion sets in store on the incidence of bolting, bulbing and seed yield. Sci.
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