or IO
3
or NH
4
+
) was applied in the experiment:
(1) control without N and I, (2) KI application without N, (3) KIO
3
application without N, (4) KI + Ca(NO
3
)
2
,
(5) KIO
3
+ Ca(NO
3
)
2
, (6) KI + (NH
4
)
2
SO
4
and (7) KIO
3
+ (NH
4
)
2
SO
4
. The experiment was arranged in a split-
plot design. Iodine (in both forms) was applied pre-sowing in a dose of 2 kg I ha
1
. Nitrogen in the form of
Ca(NO
3
)
2
and (NH
4
)
2
SO
4
was introduced pre-sowing and as a top dressing, each dose of 100 kg N ha
1
.
Results and discussion: A diverse, statistically signicant inuence of tested factors on the activity of free
radical-scavenging (DPPH) and the content of: dry matter, glucose, fructose, sucrose, total soluble sugars,
soluble solids Brix %, phenolic compounds, phenylpropanoids, avonols, anthocyanins and carotenoids
was noted in carrot roots directly after the harvest as well as at the end of four-month storage. Iodine
applied with relatively high doses of nitrogen decreased the quality of fresh carrot. After storage, opposite
relations were noted for tested combinations (with I and N application) with respect to carrot quality
when compared to results obtained after the harvest. The lowest storage ability was found for carrot trea-
ted with KI without N. Obtained results directly suggest the need for developing individual agronomic
rules for iodine biofortication of carrot for: (a) consumption and/or processing directly after the harvest
and (b) long-term storage.
2014 Elsevier Ltd. All rights reserved.
1. Introduction
Iodine is an important mineral nutrient necessary for the proper
functioning of human body. It is involved in the biosynthesis of
thyroid hormones [thyroxine (T4) and triiodothyronine (T3)] that
regulate numerous metabolic processes in the whole organism.
In medicine, the spectrum of developmental and functional disor-
ders caused by too low an intake of iodine are dened as iodine
deciency disorders (IDD). In the pre- and neonatal period, this
nutrient has a crucial role in neurological and brain development
(Melse-Boonstra & Jaiswal, 2010; Walker et al., 2007).
It is estimated that approximately 3038% of human population
has inadequate iodine intake and is, therefore, at risk of IDD (White
& Broadley, 2009; Winger et al., 2008). Europeans have notably
poor iodine intake with 56.9% children and ca. 60% adults consum-
ing less than the recommended daily intake (Andersson, de
Benoist, Darnton-Hill, & Delange, 2007; de Benoist, Andersson, Egli,
Takkouche, & Allen, 2004) despite numerous public health
programmes, based mainly on salt iodization, in most member
http://dx.doi.org/10.1016/j.foodchem.2014.03.024
0308-8146/ 2014 Elsevier Ltd. All rights reserved.
or IO
3
)
and nitrogen (as NO
3
or NH
4
+
) was applied in the experiment: (1)
control without N and I, (2) KI application without N, (3) KIO
3
application without N, (4) KI + Ca(NO
3
)
2
, (5) KIO
3
+ Ca(NO
3
)
2
, (6)
KI + (NH
4
)
2
SO
4
, (7) KIO
3
+ (NH
4
)
2
SO
4
. The experiment was ar-
ranged in a split-plot design. Each treatment was randomised in
four repetitions on 2.7 5 m (13.5 m
2
) plots. The total area of
the experiment was 378 m
2
.
Iodine (in both forms) was applied pre-sowing in a dose of
2 kg I ha
1
as pure salts (KI POCH Poland, KIO
3
Sigma Aldrich
).
Nitrogen in the form of Ca(NO
3
)
2
[Yara International ASA (Hydro)]
and (NH
4
)
2
SO
4
[Zakady Azotowe in Tarnw, Poland] was intro-
duced in two 100 kg N ha
1
doses: pre-sowing and as a top-dress-
ing. Pre-sowing application of nitrogen and iodine was conducted
before ridge formation, whereas the second dose of N at canopy
closure (27 June 08, 26 June 09 and 07 July 10). Carrots were cul-
tivated in one row on 40 cm wide and 30 cm high ridges at a seed-
ing rate of 37 seeds m
1
(approximately 550 000 seeds per
hectare). The seeds were sown on 24 April 08, 24 April 09 and 23
April 10. The carrot roots were harvested on 30 September 08, 23
September 09 and 30 September 10. At harvest, approximately
5 kg samples of carrot storage roots were chosen from each of
the four plots (replications) for laboratory analysis. Additionally,
for each of the experimental treatments, a mixed sample of 25 kg
of roots was collected for storage. During harvest, mean carrot
length was also determined. Only roots qualifying as marketable
yield were taken for further analyses. Marketable yield consisted
of storage roots of cylindrical or close-to-cylindrical shape with
head diameter of P3 cm, undamaged by pests, not infected by fun-
gi or bacteria, with no fractures and heads greened to a maximum
of 0.5 cm. The length of a storage root was 15 cm minimum.
2.2. Storage conditions
Stored carrots from each year of the study were placed in plastic
boxes (60 40 25 cm) and kept at normal atmosphere, 1 C, and
9598% humidity. After four-months, healthy roots (not affected
by diseases) were weighed and taken for analysis.
2.3. Plant analysis after harvest and long-term storage
Stored roots were washed and juiced or diced using a household
processor immediately before analysis. The content of total soluble
solids (Brix) was measured using a Atago Palette PR-32 a digital
refractometer. Dry matter was assessed at 105 C. Levels of glu-
cose, fructose, sucrose and total sugars (calculated) were deter-
mined using RP-HPLC. Determination of these compounds was
conducted in room temperature using a Knauer system (Germany).
Samples (10 ll) were injected on an amine LiChrospher RP 100-10
NH
2
250 4 mm column. The eluent was a mixture of acetonitryl/
water (87:13 v/v) and the ow rate 1.3 ml/min. Measurement was
conducted using Smartline RI refractometric detector 2300 (Bogdanov,
2002). Total carotenoid content was assayed after sample extrac-
tion with acetone/n-hexane (4:6) using a b-carotene standard
curve.
To estimate phenolic constituents and free radical scavenging,
carrot extracts were prepared in 80% methanol. Total phenols, phe-
nylpropanoids, avonols and anthocyanins concentrations were
determined using the spectrophotometric method described by
Fakumoto and Mazza (2000). Free radical scavenging activity was
evaluated on the basis of 30-min plant tissue reaction with diph-
enylpicrylhydrazyl (DPPH) (Pekkarinen, Stockmann, Schwarz,
Heinnonen, & Hopia, 1999).
2.4. Statistical analysis
Results were statistically veried using ANOVA module of Stat-
istica 9.0 PL programme for signicance level P < 0.05. Signicance
of changes was assessed with the use of variance analysis F test.
In case of signicant changes, homogenous groups were distin-
guished on the basis of Duncan test.
All data were presented as means for the years 20082010 as in
each year comparable results of treatments on quality of carrot
were obtained.
3. Results and discussion
3.1. Root weight and dry matter content
Results of numerous studies indicate that IO
3
introduced into
the soil or nutrient medium is less harmful for plants than I
when
applied in the same concentrations (Mackowiak & Grossl, 1999;
Blasco, Rios, Cervilla et al., 2011; Blasco, Rios, Leyva et al., 2011;
Caffagni et al., 2011; Hong et al., 2012). In our research no signi-
cant effect of iodine and nitrogen fertilization was found with re-
spect to root weight at harvest and after long-term storage
S. Smolen et al. / Food Chemistry 159 (2014) 316322 317
(Table 1). In our study, the lack of impact on carrot root weight
suggests that soil application of iodine at 2 kg I ha
1
as either I
or IO
3
, rather thanIO
3
without
nitrogenelicited the greatest water loss fromcarrots during storage.
This observation was reected in higher dry matter content at its
highest after 4 months. Most likelytheserelationships aretheconse-
quence of processes occurring during carrot cultivation and storage.
3.2. Quality parameters of carrot at harvest
Fertilizer with iodine and nitrogen had a statistically signicant
inuence on quality parameters determined directly after harvest
including the content of: glucose, fructose, sucrose, total sugars,
Brix (Fig. 1AE), phenolic compounds, phenylpropanoids, avo-
nols, anthocyanins (Fig. 2AD), carotenoids (Fig. 3B) as well as free
radical scavenging activity (Fig. 3A). A general trend of increasing
glucose, fructose, total sugars and soluble solids (Brix) together
with a lower content of phenolic compounds and phenylpropa-
noids was noted for KI and KIO
3
application (without N) when
compared to the control. Additionally, it should be underlined that
the highest levels of total sugars and total soluble solids were
noted in carrot roots from these combinations.
At harvest soil fertilization with KIO
3
+ Ca(NO
3
)
2
and KIO
3
+
(NH
4
)
2
SO
4
signicantly decreased levels of: glucose, sucrose, total
sugars, total soluble solids (Fig. 1A, C, D and E) and avonols in
comparison to the control (Fig. 2C). In carrot roots treated with
KIO
3
+ (NH
4
)
2
SO
4
, a signicantly higher content of sucrose, total
soluble solids (Fig. 1C and D) and carotenoids (Fig. 3B) while lower
lever of fructose was noted after application of KIO
3
+ Ca(NO
3
)
2
. In
addition, the content of fructose in carrot grown using this combi-
nation was the lowest in the whole study (Fig. 1B). The lowest con-
centration of phenylpropanoids was observed in carrots treated
with KIO
3
+ (NH
4
)
2
SO
4
while of carotenoids in roots grown with
KIO
3
+ Ca(NO
3
)
2
.
Results of this study indicate the inuence of both forms of io-
dine on carrot quality assessed directly after harvest was depen-
dent on the chemical form of nitrogen including other elements
(Ca
2+
and SO
4
2
), i.e. type of nitrogen fertilizer. Soil fertilization with
ammonium sulphate did not signicantly decrease the content of
sugar compounds in carrot in comparison to the application of cal-
cium nitrate. This is particularly interesting as there is common
agreement that the process of SO
4
2
reduction and its incorporation
into sulphur-containing amino acids require metabolic energy
(Cram, 1983; Leustek & Saito, 1999). In addition, carrot requires
low sulphur, which may explain its low uptake data presented
previously (Smolen , Sady, Ro_ zek, Ledwo_ zyw, & Strzetelski, 2011).
It is striking, however, that application of KIO
3
together with either
Ca(NO
3
)
2
or (NH
4
)
2
SO
4
contributed to a decrease in carbohydrate
content (glucose, fructose, sucrose and total sugars) and total
soluble solids (Fig. 1AE) when compared to KI application with
nitrogen fertilizers. This can be explained based on the theory
IO
3
to I
reduction
(Hung, Wong, & Dunstan, 2005; Wong & Hung, 2001). In the case
of plants fertilised with both forms of nitrogen, the observed ef-
fects could be explained by increased energy consumption by the
reduction of NO
3
1
0
0
g
-
1
f
.
w
.
)
a a
b
bc cd
e f
f
g
h
i
f
d cd
A
1 2 3 4 5 6 7
0
500
1000
1500
2000
F
r
u
c
t
o
s
e
(
m
g
1
0
0
g
-
1
f
.
w
.
)
a b b
c c
d e
f
f
g
h
B
h
c c
1 2 3 4 5 6 7
0
500
1000
1500
2000
2500
3000
3500
S
u
c
r
o
s
e
(
m
g
1
0
0
g
-
1
f
.
w
.
)
C
a
a
b
c
d e
f
f
g g
g
b
b
b
1 2 3 4 5 6 7
0
1000
2000
3000
4000
5000
6000
7000
T
o
t
a
l
s
u
g
(
m
g
1
0
0
g
-
1
f
.
w
.
)
D
a
b
c
d
e
f
g h i
j k
l
de
a
a
r
s
1 2 3 4 5 6 7
0
2
4
6
8
10
12
14
16
B
r
i
x
(
%
)
E
a
b
c c d
e
f
g h h i
j k
l
At harvest After storage
At harvest After storage
Fig. 1. Content of: glucose (A), fructose (B), sucrose (C), total sugars (D) and total
soluble solids (E) in carrot roots at harvest and after storage depending on iodine
and nitrogen fertilization. Treatments: 1 control without N and I fertilization;
2 KI fertilizationwithout Napplication; 3 KIO
3
fertilization without Napplication;
4 KI+Ca(NO
3
)
2
fertilization; 5 KIO
3
+Ca(NO
3
)
2
fertilization, 6 KI+(NH
4
)
2
SO
4
fertilization, 7 KIO
3
+(NH
4
)
2
SO
4
fertilization. Means followed by the same letters are
not signicantly different for P < 0.05 (n = 12).
1 2 3 4 5 6 7
0
10
20
30
40
50
60
70
80
P
h
e
n
o
l
i
c
c
o
m
p
o
u
n
d
s
(
m
g
1
0
0
g
-
1
f
.
w
.
)
A
a
ab
bc bcd
d
e
ef
fg
gh
hi
i
j
cd
ab
1 2 3 4 5 6 7
0
5
10
15
20
P
h
e
n
y
l
p
r
o
p
a
n
o
i
d
s
B
a ab ab
abc bc
bc
d
d
e
g
h
f
e
c
(
m
g
1
0
0
g
-
1
f
.
w
.
)
1 2 3 4 5 6 7
0
2
4
6
8
10
12
F
l
a
v
o
n
o
l
s
C
a
b
c
d
e
d
c c
c
b
b
ab a a
(
m
g
1
0
0
g
-
1
f
.
w
.
)
1 2 3 4 5 6 7
0.0
0.1
0.2
0.3
0.4
0.5
0.6
0.7
A
n
t
h
o
c
y
a
n
i
n
s
D
a
a
a a
a
a
a
a
ab
b
ab
b
b
b
(
m
g
1
0
0
g
-
1
f
.
w
.
)
At harvest After storage
At harvest After storage
Fig. 2. Content of phenolic compounds (A), phenylpropanoids (B), avonols (C) and
anthocyanins (D) in carrot roots at harvest and after storage depending on iodine
and nitrogen fertilization. Treatments: 1 control without N and I fertilization;
2 KI fertilization without Napplication; 3 KIO
3
fertilizationwithout Napplication;
4 KI+Ca(NO
3
)
2
fertilization; 5 KIO
3
+Ca(NO
3
)
2
fertilization, 6 KI+(NH
4
)
2
SO
4
fertilization, 7 KIO
3
+(NH
4
)
2
SO
4
fertilization. Means followed by the same letters are
not signicantly different for P < 0.05 (n = 12).
S. Smolen et al. / Food Chemistry 159 (2014) 316322 319
in carrot (Fig. 2AD). Particularly of interest is the fact that, in the
pot experiment, no signicant inuence on the content of soluble
sugars was found (Smolen, Ledwozyw, et al., 2009). Additionally,
treatment with KIO
3
together with calcium nitrate and ammonium
sulphate led to a signicant decrease [for Ca(NO
3
)
2
] or increase [for
(NH
4
)
2
SO
4
] in the content of soluble solids, phenolic compounds,
phenylpropanoids and avonols in carrot when compared to KI
application with the same nitrogen fertilizers (Smolen, Ledwozyw,
et al., 2009; Smolen, Sady, et al., 2009). In the case of total sugars
and soluble solids, results obtained by (Smolen, Ledwozyw, et al.,
2009) were different to those presented here (Fig. 1D, E). This dis-
crepancy may be due to different growing conditions, e.g. eld ver-
sus pot cultivation. In reference to the impact of I and N on levels of
total phenolic compounds, phenylpropanoids, and avonols, it is
noteworthy that iodine as I
and IO
3
application
during carrot cultivation stimulates catabolic processes that use
carbohydrates substrates during post-harvest storage. There is a
signicant linkage between these processes and nitrogen fertiliza-
tion of plants. Currently, it is not possible to suggest a reason for
this, as knowledge of iodine inuence on plant metabolism re-
mains insufcient. It is not unlikely that the introduction of iodine
in different oxidation states and, therefore, characterised by vari-
ous levels of toxicity triggers adverse physiological and biochemi-
cal response in plants. The process of iodine methylation followed
by the synthesis and volatilization of CH
3
I may be involved in plant
detoxication fromexcessive iodine (Landini et al., 2012; Muramatsu
& Yoshida, 1995). Generally, the activity of S-adenosylmethionine-
dependent halide/thiol methyltransferase (HTMT) an enzyme
responsible for iodine methylation is greater in leaves than roots
(Itoh et al., 2009). However, there is no direct evidence suggesting
that CH
3
I synthesis by HTMT can occur during storage. Enzymes
extracted from suspension culture catalysed methylation of caf-
feoyl-CoA to feruloyl-CoA in the presence of S-adenosyl-L-methio-
nine, which determines the activity of HTMT. This process was
described as related to plant defense against Phytophthora mega-
sperma f. sp. Glycinea infection (Khnl, Koch, Heller, & Wellmann,
1989). On the basis of this information, it can be supposed that
during long-term storage carrot carbohydrates are used not only
in catabolic processes (mainly cell respiration), but also in the syn-
1 2 3 4 5 6 7
0
2
4
6
8
10
12
D
P
P
H
(
%
)
A
a
ab
b
c
c
d
e
f
e
de
b
b a b a ab
1 2 3 4 5 6 7
0
5
10
15
20
25
30
C
a
r
o
t
e
n
o
i
(
m
g
1
0
0
g
-
1
f
.
w
.
)
B
a
ab bc bc
bc c
d de de
e
f
f
c
bc
d
s
At harvest After storage
At harvest After storage
Fig. 3. Free radical scavenging activity (A) and content of carotenoids (B) in carrot
roots at harvest and after storage depending on iodine and nitrogen fertilization.
Treatments: 1 control without N and I fertilization; 2 KI fertilization without N
application; 3 KIO
3
fertilization without N application; 4 KI+Ca(NO
3
)
2
fertilization; 5 KIO
3
+Ca(NO
3
)
2
fertilization, 6 KI+(NH
4
)
2
SO
4
fertilization,
7 KIO
3
+(NH
4
)
2
SO
4
fertilization. Means followed by the same letters are not
signicantly different for P < 0.05 (n = 12).
320 S. Smolen et al. / Food Chemistry 159 (2014) 316322
thesis of various compounds, including methyl iodide and second-
ary metabolites important for plant defense against storage
diseases.
4. Conclusion
Based on the results from our study with Kazan F
1
cultivar, it is
possible to assess to some extent the efciency of iodine biofortif-
ication of carrot taking into consideration its inuence on yield
quality and storage potential of this crop. Increasing iodine content
in carrot may be advisable due to a high level of b-carotene a di-
rect precursor of vitamin A that has been shown to have a positive
effect on thyroid function (Zimmermann, 2007). Iodine application
combined with plant fertilizer with relatively high doses of nitro-
gen (100 kg N ha
1
applied pre-sowing and as a top dressing in
the form of calcium nitrate and ammonium sulphate) did, how-
ever, decrease the quality of carrot yield evaluated at harvest. As
far as storage potential is concerned, the lowest value was noted
for carrots grown with KI without N. Analysis of our results sug-
gests there is a need to develop independent methods of iodine
biofortication for carrots intended for (a) consumption and/or
processing directly after harvest or (b) long-term storage. The com-
bination of iodine biofortication programmes with appropriate
crop fertilization with mineral nutrients should also be taken into
account. Furthermore, it seems reasonable to employ plant-breed-
ing techniques to select cultivars characterised by increased iodine
accumulation without decreased yield quality and storage
potential.
Basing on conducted research it can be proposed that for the
cultivation of carrot biofortied with iodine, KI form of this ele-
ment should be applied rather than KIO
3
(also when combined
with nitrogen fertilization) due to a signicantly more deleterious
effect of iodates on carrot quality after harvest. In the production of
carrot for long-term storage of roots high doses of nitrogen fertiliz-
ers need to be avoided (particularly in the form of calcium nitrate)
when applied together with iodine in order to prevent increased
losses of root weight or the decrease in its quality.
Acknowledgement
This work was nanced by research grants from the Ministry of
Science and Higher Education Republic of Poland.
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