Article

pubs.acs.org/JAFC

Grain Nutrient Composition of Maize (Zea mays L.) Drought-Tolerant

Populations
Dragana Ignjatovic-Micic,*,† Jelena Vancetovic,† Dejana Trbovic,‡ Zoran Dumanovic,†
Marija Kostadinovic,† and Sofija Bozinovic†
†
Maize Research Institute Zemun Polje, Slobodana Bajica 1, 11185 Belgrade, Serbia
‡
Institute of Meat Hygiene and Technology, Kacanskog 13, 11000 Belgrade, Serbia

ABSTRACT: A total of 13 maize populations from the drought-tolerant mini core collection from Maize Research Institute
gene bank were evaluated for oil, protein, and tryptophan contents, fatty acid (FA) composition, and kernel characteristics. All
accessions are high oil (5.8−7.9%) and protein (10.58−12.45%) genotypes. Most of the accessions showed high contents of
tryptophan (0.070−0.081%) and saturated (12.65−17.91%) and monounsaturated (24.19−45.52%) FAs. Significant positive
correlations were found between oil and protein and between oil and tryptophan contents (p < 0.01). Correlations between oil
and principal FA were non-significant. Several accessions showed multiple nutritional advantages. For example, IP6428 had high
oil (7.3%), tryptophan (0.081%), and saturated FA (17.9%) contents. Moreover, a positive correlation (p < 0.01) between
palmitic (13.68%) and oleic (34.74%) acids enables the use of IP6428 for developing lines high in these FAs. Because drought-
tolerant accessions were selected in both subtropical and temperate zones, they could be used for breeding value-added maize
adapted to both environments.
KEYWORDS: fatty acids, oil, protein, tryptophan, Zea mays L.

■ INTRODUCTION
Maize (Zea mays L.) is one of the leading cereal grains
worldwide, along with wheat and rice.1 Breeding maize has
been primarily aimed at increasing yield potential and stability
under different environmental conditions. However, demands
for nutritionally improved maize that are beneficial to health
have initiated many programs in breeding quality-improved
genotypes.2,3
Typical kernel composition for the commodity yellow dent
corn on a dry matter basis is 71.7% starch, 9.5% protein, 4.3%
oil, 1.4% ash, and 2.6% sugar.4 Nutritional quality of maize oil
and protein can be improved by altering their fatty acid (FA)
and amino acid composition, respectively. Maize kernel
contains an average FA composition of 11% palmitic (16:0),
2% stearic (18:0), 24.1% oleic (18:1), 61.9% linoleic (18:2),
and 0.7% linolenic (18:3) acids.5 Developing maize oil with
different arrangements of altered FA compositions can be
beneficial in various ways. For example, increasing the oleic acid
content would enhance oil oxidative stability6 and provide a
more healthful FA composition that could decrease coronary
heart disease.7 On the other hand, increased saturated (palmitic
and stearic) FA composition would allow for production of
margarines without hydrogenation and the subsequent
formation of undesirable trans FAs.8
Maize is poor in terms of protein content and also has low
biological quality in relation to essential amino acids lysine and
tryptophan. Several natural maize mutants conferring higher
lysine and tryptophan levels were identified, but the recessive
opaque-2 (o2) mutation was found to be the most suitable for
genetic manipulation in breeding programs. The breeding
efforts led to the development of quality protein maize (QPM),
maize high in lysine and tryptophan content with good
agronomical performances.9 QPM was developed primarily for
use in the regions where maize is the main staple food (sub-
Saharan Africa and Asia) and where protein-rich foods to
supplement diets cannot be afforded. The biological value of
QPM is doubled in comparison to normal maize (80 versus
45%, respectively) and similar to the biological value (90%) of
milk. In addition, QPM has other nutritional advantages over
normal maize, such as higher concentration of niacin (B3)
because of higher tryptophan levels and better absorption of
potassium and carotene by the body.10 Also, it is rich in iron
and zinc contents.11
The narrow genetic base of commercial genotypes has
limited variability for value-added traits. One of the ways for
improving these traits is gene introgression from landraces or
wild relatives.12,3 It was shown that teosinte and landraces
harbor phenotypic variation that can facilitate genetic dissection
of kernel traits and grain quality, ultimately leading to
improvement via traditional plant breeding and/or genetic
engineering.13 Maize Research Institute Zemun Polje (MRI)
gene bank has one of the largest maize collections in the world,
with approximately 5500 accessions, including 2217 landraces
from Western Balkan and 3258 introduced populations and
inbred lines from 40 countries. A survey for drought-tolerant
accessions resulted in the establishment of a drought-tolerant
mini core collection, which comprises 15 inbred lines, 13
landraces, and 12 introduced populations.14 These accessions
were selected under a water-controlled regime in Egypt in 2007
as well as in two temperate regions (Serbia and Macedonia,
drought test location) and Egypt in 2008. In the subsequent
Received: September 12, 2014
Revised: November 28, 2014
Accepted: January 9, 2015
Published: January 9, 2015

© 2015 American Chemical Society 1251 DOI: 10.1021/jf504301u

J. Agric. Food Chem. 2015, 63, 1251−1260
Journal of Agricultural and Food Chemistry
experiments, they were chosen because of their good
combining ability with elite inbred testers. The landraces and
introduced populations are further being analyzed for macro-
nutrient content15 and composition, with the aim to identify
accessions with drought tolerance and high oil and/or protein
quality. The existence of drought tolerance and grain quality in
a single accession should enable more effective improvement of
commercial genotypes designed to meet the issues of the
ongoing climatic changes and increased food demands because
of the population growth.
The objectives of the present study were to (1) explore the
variability within the drought-tolerant populations previously
identified as potentially high oil accessions by near-infrared
spectroscopy (NIR) for (a) oil and protein contents
determined by standard laboratory methods, (b) FA
composition, and (c) tryptophan content, (2) evaluate kernel
physical parameters, (3) determine correlations between the
traits, and (4) identify genotypes with nutritional value.
Because the drought-tolerant mini core collection was selected
in field trials performed in both subtropical and temperate
zones, the identified genotypes could be used for breeding
value-added maize adapted to both environments.
■
MATERIALS AND METHODS
Plant Material and Field Trials. A total of 25 accessions from the
elite drought-tolerant mini core were increased (via pair crossing, i.e.,
full-sibbing) in two replications, using a completely randomized block
design (RCBD), in Zemun Polje, Serbia, during 2010 and 2011
growing seasons. Walter climate diagrams (Figure 1) illustrate
temperature and precipitation conditions during the growing season
in these 2 years. Standard agronomic practices were used to provide
Figure 1. Walter climate diagrams for the years (A) 2010 and (B)
2011 in Belgrade, Serbia. Tavg, average temperature (°C); Sprec, sum of
precipitation (mm).
1252
Article
adequate nutrition and keep the fields disease-free. At least 80 ears per
population were collected, dried, and shelled, and samples of kernels
from the center of each ear were prepared for the grain quality analysis.
NIR analysis (Infratec 1241 grain analyzer, FOSS, Sweden) with
artificial neural network (ANN) calibrations was used for oil content
determination, on non-destructed kernels (300 g/sample) from each
replicate (spectra were collected between 800 and 1100 nm). The
analyses were performed at the Faculty of Agronomy, Skopje,
Macedonia. A total of 13 potentially high oil accessions, with oil
content over 4.5% (average content for maize), were selected for
further analysis, with 4 of them from Western Balkan, 2 from Iran, 2
from Italy, and 1 each from Bulgaria, France, Pakistan, and the U.S.A.
For one introduced population, there are no available data of origin.
Information on the 13 analyzed accessions, including agro-ecological
group or country of origin, NIR data, kernel type, and color, are given
in Table 1. Kernel types were determined according to the CIMMYT/
IBPGR descriptor.16
Determination of the Oil Content and FA Composition. Each
accession consisted of 50 g of maize kernels. The oil content was
determined by the standard Soxhlet method. Dried kernels were milled
in a Cyclotec 1093 lab mill (FOSS Tecator, Sweden) with a particle
size of <500 μm, and oil was extracted with hexane in a Soxhlet
extractor (INKOLAB, Croatia).
FA composition of the oil samples was determined after
transesterification using 0.25 M trimethylsulfonium hydroxide
(TMSH, Fluka, Switzerland) in methanol according to EN ISO
5509:2000. Prior to transesterification, 0.05 mL (10 mg mL−1) of the
nonadecanoic acid methyl ester (≥99%, Fluka, Switzerland) was added
as an internal standard.
The fatty acid methyl esters (FAMEs) were determined by capillary
gas chromatography on a GC Shimadzu 2010 (Kyoto, Japan)
equipped with a split/splitless injector, fused silica cyanopropyl HP-
88 column (length, 100 m; inner diameter, 0.25 mm; film thickness,
0.20 μm; J&W Scientific, Santa Clara, CA), and flame ionization
detector. The temperature of the injector was 250 °C, with a split ratio
of 1:50, and the detector temperature was 280 °C. Nitrogen was used
as the carrier gas at a flow rate of 1.33 mL min−1. The column oven
temperature was programmed from 125 to 230 °C. The total analysis
time was 50.5 min. FAs were identified by comparing the relative
retention time of peaks in the samples to FAME peaks in the Supelco
37 Component FAME mix standard (Supelco, Bellefonte, PA). The
chromatographic peak areas were corrected by response factors
calculated from the ratios between the peak area of each FA and the
internal standard.
Determination of the Protein Content, Tryptophan Content,
and Quality Index (QI). Each accession was represented by 60
randomly chosen kernels, divided into two samples of 30 kernels each.
Kernels were dried in a controlled oven at 65 °C overnight (16−18 h)
and milled in a Cyclotec 1093 lab mill (FOSS Tecator, Sweden) with a
particle size of <500 μm. The flour was defatted by hexane treatment
for 4 h in a Soxhlet extractor (INKOLAB, Croatia).
The protein content was determined by the standard Kjedahl
method using a 2200 Kjeltec auto distillation unit (FOSS, Tecator,
Sweden) based on nitrogen determination as explained by Vivek et al.9
It was estimated from the nitrogen value as percent protein = percent
nitrogen × 6.25 (conversion factor for maize).
The tryptophan content was determined using the colorimetric
method by Nurit et al.17 The color was developed in the reaction of
flour hydrolysate (obtained by overnight digestion with papain
solution at 65 °C) with 2 mL of reagent containing 56 mg of Fe3+
dissolved in 1 L of glacial acetic acid and 2 mL of 15 M H2SO4. After
incubation at 65 °C for 15 min, absorbance was read at 560 nm using
an UV-1601 Shimadzu spectrophotometer. The tryptophan content
was calculated using a standard calibration curve, developed with
known amounts of tryptophan, ranging from 0 to 30 μg mL−1.
QI, defined as the tryptophan/protein ratio in the sample, was
calculated as QI = 100(tryptophan content in the sample/protein
content in the sample).
Kernel Physical Characteristics. Thousand kernel weight was
measured by weighing 5 × 200 unbroken kernels per sample. Flotation
DOI: 10.1021/jf504301u
J. Agric. Food Chem. 2015, 63, 1251−1260
Journal of Agricultural and Food Chemistry Article

Table 1. High Oil Maize Populations from the Drought-Tolerant Mini Core Collection
accessiona agro-ecological group or country of origin oil percentage (NIR) (%) kernel typeb kernel color
1 L87 white flinty dents Moravac 5.40 semi-dent white
2 L244 denty type of U.S.A. Corn Belt dents 4.80 semi-flint dark yellow
3 L632 denty type of U.S.A. Corn Belt dents 4.75 semi-flint yellow, red stripes
4 L2033 Mediterranean flints 4.75 flint yellow
5 IP4347 Iran 4.65 semi-flint yellow
6 IP4353 Iran 5.05 semi-flint yellow
7 IP4842 France 5.25 semi-dent yellow
8 IP6233 Bulgaria 4.90 flint yellow
9 IP6389 no data 4.65 semi-dent orange
10 IP6427 Italy 4.85 semi-flint yellow/white
11 IP6428 Italy 5.35 semi-dent yellow
12 IP6750 Pakistan 4.65 semi-dent yellow
13 IP7001 U.S.A. 4.60 semi-flint yellow
a
1−4, landraces (L) from Western Balkan; 5−13, introduced populations (IPs). bAccording to the CIMMYT/IBPGR descriptor for maize, semi-
dent kernels are intermediate between dent and flint but closer to dent, while semi-flint kernels are flint kernels with soft cup.

Table 2. Oil and Protein Parameters of High Oil Maize Populations from the Drought-Tolerant Mini Core Collection
oil
total FA protein
accession OCa,b SFAa,c MUFAa,d PUFAa,e ω6:3a PCa,f TCa,g QIa,h
L87 7.9 a 13.32 h 28.02 g 58.65 b 86.86 b 12.25 b 0.070 bcde 0.58 de
L244 7.7 a 16.40 b 35.00 c 48.60 f 74.52 de 12.25 b 0.075 abcd 0.62 bcd
L632 7.0 bc 12.76 i 45.52 a 41.65 h 60.51 g 11.11 ef 0.077 ab 0.70 a
L2033 6.9 c 15.46 d 31.90 e 52.90 d 67.40 f 11.68 d 0.078 ab 0.67 abc
IP4347 6.6 de 12.65 i 24.19 h 63.15 a 79.40 cd 11.69 d 0.079 ab 0.68 abc
IP4353 7.1 bc 13.55 g 32.21 e 54.22 c 67.47 f 11.02 f 0.076 abc 0.69 a
IP4842 6.5 ef 15.15 e 33.65 d 51.20 e 81.90 bc 11.15 ef 0.066 de 0.60 de
IP6233 6.3 f 16.16 c 32.44 e 51.40 e 98.48 a 11.24 e 0.076 abcd 0.67 abc
IP6389 6.0 g 15.26 de 36.76 b 48.20 f 58.05 g 12.26 b 0.067 cde 0.55 e
IP6427 6.6 ef 16.54 b 32.28 e 51.17 e 72.45 ef 12.37 ab 0.074 abcd 0.61 cde
IP6428 7.3 b 17.91 a 35.00 c 47.10 g 61.66 g 11.93 c 0.081 a 0.68 ab
IP6750 6.8 cd 16.02 c 32.76 de 51.22 e 67.65 f 12.45 a 0.076 abc 0.62 bcde
IP7001 5.8 g 14.30 f 30.82 f 54.89 c 70.19 ef 10.58 g 0.063 e 0.59 de
mean 6.8 15.04 33.12 51.87 73.15 11.69 0.074 0.63
SDi 0.87 1.63 4.86 5.32 14.35 1.18 0.01 0.07
CVj 2.42 0.91 1.92 1.25 4.92 0.58 5.97 5.92
LSD0.05k 0.25 0.21 0.98 0.99 5.54 0.15 0.01 0.07
a
Means followed by the same letter(s) within the same columns are not significantly different (p < 0.05). bOC = oil content. cSFA = saturated fatty
acid. dMUFA = monounsaturated fatty acid. ePUFA = polyunsaturated fatty acid. fPC = protein content. gTC = tryptophan content. hQI = quality
index. iSD = standard deviation. jCV = coefficient of variation. kLSD0.05 = least significant difference at p < 0.05.

index (FI) was measured by soaking 100 randomly selected kernels Fisher’s least significant difference (LSD). All statistical analyses were
into an aqueous sodium nitrate solution with a specific weight of 1.25 g performed in MSTAT-C software. Pearson’s correlation coefficient
cm−3 at 35 °C, and values were expressed as a percentage of floating was used for determining correlations between the estimated traits.

kernels. Kernel dimensions were obtained by measuring thickness,
width, and length of a 100 kernels per sample using a digital
micrometer. The hull, endosperm, and germ were manually dissected
after soaking 50 g of kernels for 2 min in 100 mL of water. After the
water was drained, the wet kernels were first manually separated into
hull, endosperm, and germ, with the resulting anatomical parts dried in
an oven set at 60 °C and weighed to calculate relative percentages. The
kernel hardness was measured by a modified Stenvert−Pomeranz
method, by milling 20 g of kernels in the micro hammer mill (3600
rpm, 2 mm sieve). Results were expressed as milling response and hard
fraction portion (%). The milling response index presents the time (s)
necessary for grain grinding until the top level of the material collected
in a glass cylinder (125 × 25 mm) reaches the level of 17 mL.
Statistical Methods. All chemical analyses were performed in two
replicates, and the results were statistically analyzed. A factorial analysis
of variance (ANOVA) for trials was conducted using randomized
complete block (RCB) design. Treatment means were tested using
1253
■
RESULTS AND DISCUSSION
Biochemical Analysis. Laboratory analysis of the oil
content in the samples (Table 2) revealed higher values
compared to previously performed NIR analysis (Table 1).
However, correlation between the two methods was significant
at p < 0.05 (data not shown). This could be expected, because
it was shown that NIR estimation and reference analysis for the
oil content in maize kernels were different and that more
studies are needed before obtaining accurate measurements
using NIR methods.18 In this context, the oil content
determined by the standard Soxhlet method is more accurate.
Populations from the drought-tolerant mini core collection
showed significant variability in oil and protein contents as well
as in FA composition and tryptophan content (Tables 2 and 3).
DOI: 10.1021/jf504301u
J. Agric. Food Chem. 2015, 63, 1251−1260
Journal of Agricultural and Food Chemistry Article

Table 3. FA Composition (Percentage of Total Oil) of High Oil Maize Populations from the Drought-Tolerant Mini Core
Collection
FAs
a,b a,c a,d a,e a,f a,g
accession 16:0 16:1 17:0 18:0 18:1 18:2 18:3a,h 20:0a,i 20:1a,j 20:3a,k 22:0a,l 24:0a,m
L87 11.25 g 0.110 a 0.030 a 1.47 l 27.70 h 57.92 b 0.675 ef 0.320 g 0.205 b 0.055 i 0.115 d 0.140 g
L244 13.49 a 0.065 c 0.030 a 2.20 h 34.72 c 47.86 f 0.645 ef 0.435 f 0.210 b 0.095 fg 0.110 d 0.140 g
L632 9.71 j 0.040 d 0.025 a 2.20 h 45.24 a 40.86 h 0.680 def 0.525 cd 0.235 a 0.110 ef 0.135 c 0.160 def
L2033 12.18 de 0.090 b 0.060 a 2.37 f 31.36 fg 52.03 d 0.790 ab 0.510 cde 0.200 b 0.075 h 0.150 bc 0.190 ab
IP4347 10.25 i 0.085 b 0.060 a 1.69 k 23.96 i 62.26 a 0.785 ab 0.330 g 0.145 d 0.105 fg 0.150 bc 0.170 cde
IP4353 10.62 h 0.085 b 0.025 a 2.12 i 31.95 ef 53.33 c 0.795 ab 0.465 ef 0.170 c 0.095 fg 0.140 bc 0.175 bcd
IP4842 11.64 f 0.035 d 0.030 a 2.59 e 33.44 d 50.49 e 0.620 f 0.585 ab 0.170 c 0.090 gh 0.153 b 0.165 cdef
IP6233 12.05 e 0.065 c 0.065 a 3.18 b 32.29 ef 50.72 e 0.515 g 0.540 bcd 0.085 e 0.155 ab 0.155 b 0.170 cde
IP6389 12.34 cd 0.095 ab 0.030 a 1.89 j 36.24 b 47.22 f 0.820 a 0.585 ab 0.200 b 0.165 a 0.210 a 0.205 a
IP6427 12.88 b 0.040 d 0.025a 2.81 c 32.07 ef 50.35 e 0.700 cde 0.505 cde 0.170 c 0.125 de 0.155 b 0.165 cdef
IP6428 13.68 a 0.085 b 0.065 a 3.25 a 34.74 c 46.19 g 0.750 bc 0.610 a 0.170 c 0.155 ab 0.155 b 0.150 fg
IP6750 12.48 c 0.095 ab 0.070 a 2.63 d 32.52de 50.33 e 0.745 bcd 0.545 bc 0.150 d 0.135 cd 0.145 bc 0.155 efg
IP7001 11.07 g 0.085 b 0.060 a 2.35 g 30.58 g 53.97 c 0.770 ab 0.495 de 0.150 d 0.145 bc 0.140 bc 0.180 bc
mean 11.82 0.075 0.044 2.36 32.84 51.04 0.715 0.496 0.174 0.116 0.147 0.167
SDn 1.22 0.19 0.03 0.58 4.85 5.32 0.11 0.1 0.05 0.07 0.03 0.02
CVo 1.07 13.25 20.95 6.33 1.99 1.28 6.05 7.41 10.69 9.33 9.8 7.09
LSD0.05p 0.19 0.015 0.049 0.015 1.01 1.01 0.069 0.049 0.015 0.015 0.015 0.015
a
Means followed by the same letter(s) within the same columns are not significantly different (p < 0.05). b16:0 = palmitic acid. c16:1 = palmitoleic
acid. d17:0 = heptadecanoic acid. e18:0 = stearic acid. f18:1 = oleic acid. g18:2 = linoleic acid. h18:3 = linolenic acid. i20:0 = arachidic acid. j20:1 =
eicosenoic acid. k20:3 = eicosatrieonic acid. l22:0 = behenic acid. m24:0 = lignoceric acid. nSD = standard deviation. oCV = coefficient of variation.
p
LSD0.05 = least significant difference at p < 0.05.

However, they can be all considered as high oil and protein
genotypes when compared to oil and protein contents of the
yellow commodity maize given by Watson;4 oil content of the
analyzed accessions was higher from 35% (IP7001) to 84%
(L87), and protein content of the analyzed accessions was
higher from 11% (IP7001) to 31% (IP6750). Enhanced
nutritional quality of maize with high levels of essential
amino acids tryptophan and lysine, such as o2 and QPM,
requires tryptophan content and QI (tryptophan/protein ratio)
in the whole grain sample to be higher than 0.075 and 0.80%,
respectively.9 Four accessions (L87, IP4842, IP6389, and
IP7001) had a low tryptophan content typical for standard
maize genotypes. One accession (IP6427) had an elevated
tryptophan content of 0.074% (a value close to the threshold of
0.075%) and should be further tested for defining it as a high
tryptophan accession. The remaining eight accessions had a
tryptophan level over 0.075% and can be considered as high
tryptophan genotypes. This also implies that they have high
levels of lysine because tryptophan and lysine values are highly
correlated and the value of lysine is 3 times that of tryptophan.9
QI was low in all accessions, with the average value of 0.63.
FA composition for each accession is given in Table 3.
Typical dent maize oil has 61.9% linoleic, 24.1% oleic, 11%
palmitic, 2% stearic, and 0.7% linolenic acids, while all of the
other acids are present only in traces.12 In our research,
palmitoleic (16:1) and heptadecanoic (17:0) acids exhibited
less than 0.1% and eicosenoic (20:1), eicosatrieonic (20:3),
behenic (22:0), and lignoceric (24:0) acids exhibited less than
0.2% of the total FA content. Arachidic acid (20:0) was the
most abundant trace FA, with the average value of 0.49%.
As expected, linoleic (18:2) and oleic (18:1) acids were the
most abundant FAs present in the oil samples, accounting for
approximately 81−85% of the total FA composition. Their
average values were 51.04 and 32.84%, respectively, showing
low linoleic acid content and high oleic acid content in
comparison to typical dent maize. On average, polyunsaturated
linolenic (18:3) and saturated palmitic (16:0) and stearic
1254
(18:0) acids were slightly higher than in typical dent maize.
Considering saturated fatty acids (SFAs) and unsaturated FAs
(Table 2), the most abundant were polyunsaturated fatty acids
(PUFAs) with 51.87%, followed by monounsaturated fatty
acids (MUFAs) with 33.12% and SFAs with 15.04%. Most of
the accessions can be considered high in oleic acid (MUFA)
and low in linoleic and linolenic acids (PUFA). Breeding for
high oleic acid maize oil can improve the average modern diet
rich in saturated fats from animal sources. It was shown that
oleic acid has positive effects in reducing serum cholesterol and
low-density lipoproteins (LDLs)7 as well as in preventing
degenerative diseases and cancer.7,19 The same authors found
similar positive effects of PUFAs on health, but none of the
high oil accessions from the drought-tolerant mini core showed
a high content of these FAs. The only exception was accession
IP4347, with slightly higher contents of linoleic and linolenic
acids than the contents typical for dent maize (62.26 and
0.785% versus 61.9 and 0.70%, respectively). Also, the ω6:3
(linoleic/linolenic) ratios were high, with the average value of
73.15. A significant increase of linolenic acid of over 4% and,
thus, a significant decrease in ω6:3 ratio (with values below 10)
was achieved in a long-term breeding program through eight
cycles of recurrent selection for agronomic traits and high oil
content in four subtropical white and yellow maize
populations.20 These changes in linolenic acid content and
ω6:3 ratio were achieved from initial populations that had
similar contents of linoleic and linolenic acids as our high oil
drought-tolerant accessions, suggesting the possibility that they
also could be the source for such improvements. This is
important because linolenic acid exerts better health benefits
but is more prone to oxidation and flavor reversion.
Several accessions can be considered as high SFAs, and some
of them had SFAs over 16%. Accession IP6428 proved to be
the most prominent high SFA (17.9%), with 13.68% palmitic,
3.20% stearic, and 0.61% arachidic acids. Maize genotypes with
high total SFAs can improve manufacturing of margarines and
DOI: 10.1021/jf504301u
J. Agric. Food Chem. 2015, 63, 1251−1260
Journal of Agricultural and Food Chemistry Article

Table 4. Kernel Physical Characteristics of High Oil Maize Populations from the Drought-Tolerant Mini Core Collection
kernel dimensionsa kernel proportionsa kernel hardnessa
thousand kernel width length thickness hull endosperm germ hard fraction milling
accession weighta (g) FIa (%) (mm) (mm) (mm) (g/100 g) (g/100 g) (g/100 g) portion (%) response (s)
L87 305.00 fg 20.44 d 8.7 bcd 8.8 h 4.8 a 9.41 a 77.50 d 13.09 a 64.97 bc 13.44 a
L244 320.00 de 22.44 d 8.1 fg 10.4 c 4.4 cde 6.54 b 83.56 c 9.90 b 63.38 de 11.34 ab
L632 340.00 ab 43.09 ab 8.3 ef 10.5 c 4.2 ef 6.97 ab 83.86 bc 9.17 bc 58.02 g 11.10 ab
L2033 308.08 efg 48.20 a 8.5 cde 9.8 ef 4.5 bcde 5.83 b 87.32 a 6.85 cd 60.62 f 11.82 ab
IP4347 298.05 gh 32.30 c 7.8 h 10.8 ab 4.2 ef 5.61 b 87.01 a 7.40 cd 59.93 f 12.18 ab
IP4353 273.00 i 3.10 f 7.1 i 10.9 a 3.9 f 7.05 ab 85.90 abc 7.03 cd 62.64 e 12.21 ab
IP4842 325.00 cd 3.34 f 8.4 de 10.4 cd 4.3 de 6.03 b 87.27 a 6.70 d 64.84 bc 12.39 ab
IP6233 342.50 ab 10.03 ef 8.8 bc 10.3 cd 4.6 abcd 6.45 b 87.55 a 6.00 d 64.52 bcd 11.15 ab
IP6389 333.13 bc 8.27 ef 8.6 bcd 9.7 efg 4.8 ab 6.88 ab 86.45 a 6.66 d 69.22 a 12.11 ab
IP6427 350.00 a 34.52 bc 9.2 a 10.0 de 4.7 abc 5.40 b 88.16 a 6.44 d 60.84 f 11.92 ab
IP6428 312.50 def 25.94 cd 8.8 b 9.6 fg 4.5 abcd 5.45 b 88.10 a 6.45 d 63.98 cd 11.85 ab
IP6750 288.05 h 16.95 de 8.3 efg 9.4 g 4.4 cde 7.24 ab 86.00 ab 6.75 d 65.53 b 11.70 ab
IP7001 315.00 def 3.19 f 8.0 gh 10.5 bc 4.3 de 6.95 ab 86.67 a 6.38 d 64.86 bc 10.72 b
mean 316.20 20.91 8.4 10.1 4.4 6.60 85.80 7.60 63.33 11.84
SDb 2.318 15.401 0.542 0.61 0.299 1.358 2.908 2.138 3.091 1.241
CVc 1.86 21.39 1.6 1.8 3.42 17.99 1.28 14.45 2.19 10.53
LSD0.05d 12.83 9.744 0.3 0.4 0.3 2.59 2.40 2.39 1.17 2.72
a
Means followed by the same letter(s) within the same columns are not significantly different (p < 0.05). bSD = standard deviation. cCV =
coefficient of variation. dLSD0.05 = least significant difference at p < 0.05.

Table 5. ANOVA Mean Squares for Biochemical Components

principal FAs
source of variation dfa OCb SFAc MUFAd PUFAe 16:0f 18:0g 18:1h 18:2i 18:3j PCk TCl
year (Y) 1 8.074m 4.935m 678.204 38.769n 1.765 0.449 17.960 38.873n 0.093n 45.947o 0.001m
genotype (G) 12 1.509o 10.487o 604.735o 113.938o 5.847o 1.125o 95.762o 113.750o 0.030o 1.519o 0.000o
G×Y 12 0.957o 0.329o 158.778o 1.873o 0.176o 0.040 1.172n 1.779m 0.008m 0.625o 0.000o
error 24 0.027 0.019 12.935 0.418 0.016 0.022 0.429 0.427 0.002 0.005 0.000
a
df = degrees of freedom. bOC = oil content. cSFA = saturated fatty acid. dMUFA = monounsaturated fatty acid. ePUFA = polyunsaturated fatty
acid. f16:0 = palmitic acid. g18:0 = stearic acid. h18:1 = oleic acid. i18:2 = linoleic acid. j18:3 = linolenic acid. kPC = protein content. lTC =
tryptophan content. mSignificant at the 0.05 level. nSignificant at the 0.1 level. oSignificant at the 0.01 level.

shortenings without trans fats, which raise LDL−cholesterol
and lower high-density lipoprotein (HDL)−cholesterol.12
Kernel Physical Parameters. Profiles of grain physical
parameters are given in Table 4. These parameters are
important for dry milling and food processing industries.
Because the accessions analyzed in our work are sources of
favorable alleles for breeding genotypes with improved
nutritional quality, the kernel parameters are to be understood
in that context. Kernels with thousand kernel weights greater
than 320 g and FI less than 20% are desirable in food
processing.21 In six accessions, thousand kernel weight was over
the desirable 320 g, while the remaining seven accessions had
lower values from approximately 2% (IP6428 and IP7001) to
15% (IP4353). Great variability was found for FI among the
analyzed accessions. It was below 10% in four accessions and
below 20% in another two accessions. In the remaining seven
accessions, FI was in the range from 20.44% (L87) to 48.2%
(L2033). The hardness classification based on FI proposed by
Salinas et al.22 defined that very hard kernels have values
between 0 and 12%, hard kernels have values between 13 and
37%, intermediary hard kernels have values between 38 and
62%, and soft kernels have values over 63%. Results showed
that none of the accessions had soft kernels, because the highest
detected value was 48.20% (L2033).
Kernel dimensions are important for breakfast cereals and oil
crushing.21 The average values of kernel dimensions as well as
1255
kernel proportions were compared to average values of these
traits given for high oil populations in Preciado-Ortiza et al.20
Comparisons were performed with data for C7 cycle of
selection of four populations, because their oil content (7.5, 7.6,
6, and 6.3%) coincided with the oil content of the accessions.
The average kernel dimensions of the accessions were 8.4, 10.1,
and 4.4 mm for width, length, and thickness, respectively.
Kernel thickness was in good agreement with this trait value
found in high oil populations. However, kernel width and
length were lower for 5 and 14%, respectively. Considering
kernel proportions, the largest part of the kernels was made of
endosperm (85.8% on average), followed by the germ (7.6% on
average) and the hull (6.6% on average). Endosperm
proportions were in good agreement with the same trait by
Preciado-Ortiza et al.,20 but the germ proportions were smaller
for most accessions for approximately 28%, except for
accessions L244, L632, and L87.
The hardness of the maize kernel is the most important
physical characteristic in dry milling and alkaline cooking
industries as well as for feed efficiency. Results obtained for
milling response showed low variability among the accessions,
while higher variability was found for the hard fraction portion.
In comparison to 20 commercial hybrids analyzed by
Radosavljevic et al.,23 the average value of milling response
was lower (11.84 versus 12.80 s) and that of the hard fraction
portion was higher (63.33 versus 58.80%) among the
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Journal of Agricultural and Food Chemistry
Table 6. ANOVA Mean Squares for Kernel Physical Characteristics
kernel dimensions
source of variation dfa 1000KWb FIc KWd KLe
year (Y) 1 5.480 349.718 0.019l 0.534
genotype (G) 12 19.912m 937.855m 1.510m 1.162m
G×Y 12 2.174 13.804 0.022 0.032
error 24 0.347 20.002 0.033 0.018
a
df = degrees of freedom. b1000KW = thousand kernel weight. cFI = flotation index. dKW = kernel width. eKL = kernel length. fKT = kernel
thickness. gHP = hull proportion. hGP = germ proportion. iEP = endosperm proportion. jHFP = hard fraction portion. kMR = milling response.
l
Significant at the 0.1 level. mSignificant at the 0.01 level. nSignificant at the 0.05 level.
accessions. The hybrids analyzed by Radosavljevic et al.23
comprised dent, high oil, waxy, white endosperm, flint, and
popcorn hybrids, which showed great variability for hardness.
Milling response ranged from 9.4 s (in a dent hybrid) to 19.9 s
(in a popcorn hybrid), and the hard fraction portion ranged
from 49.6% (in a dent hybrid) to 75.2% (in a popcorn hybrid).
The two analyzed oil hybrids showed milling response of 14.1
and 11.6 s, while their hard fraction portion was 63.4 and
58.8%. Because these values coincide with the values obtained
for the accessions (Table 4), it can be said that kernel hardness
of the high oil accessions is within the range of kernel hardness
of the commercial oil hybrids.
Statistical Analysis. Results of the ANOVA for oil content,
total FAs, five principal FAs, and protein and tryptophan
contents are given in Table 5. Genotype effect was significant
for all of the traits analyzed (p < 0.01). Year effect was not
significant for MUFAs and palmitic, stearic, and oleic acids,
while the highest effect was noted for the protein content (p <
0.01). Except for stearic acid, all of the other analyzed
biochemical components were affected by the genotype ×
year interaction. Considering trace FAs, the year effect was
significant only for eicosenoic (p < 0.05) and eicosatrienoic (p
< 0.01) acids, while genotype and the genotype × year
interaction affected all of them (data not shown).
ANOVA for kernel physical characteristics (Table 6)
revealed that most of them were not affected by the year,
except kernel length, endosperm hardness, and milling response
(p < 0.1). The genotype effect was significant for all of the traits
analyzed at p < 0.01, except for hull proportion (p < 0.05) and
milling response (non-significant). The genotype × year effect
was not significant for any of the traits measured. The
differences of the values of the traits that were significantly
affected by the year are illustrated in Figure 2.
The years differed in total precipitation and number of days,
with a maximum temperature over 30 °C during the vegetation
period (from April to September), with 2011 having less rainfall
and higher temperatures (Figure 1). Biochemical and kernel
physical characteristics that were significantly influenced by the
year are presented in Figure 2. Although it can be stated that
2010 was optimal and that drought occurred in 2011, oil and
protein contents significantly increased in 2011. A significant
increase in the same year was also noted for tryptophan, linoleic
and linolenic acids, total PUFAs, one monounsaturated trace
FA (eicosenoic acid), and two kernel hardness parameters
(hard fraction portion and milling response). On the contrary,
eicosatrienoic acid, SFAs, and kernel width values were
significantly higher in 2010. Most of these results were not
expected. It has been shown that drought significantly reduces
oil and linoleic acid contents and increases oleic acid content in
maize.24 Similar results were obtained for sunflower25,26 and in
Moringa oliefera.27 However, Petcu et al.28 found different
1256
Article
kernel proportions kernel hardness
KTf HPg GPh EPi HFPj MRk
0.425 0.260 0.775 1.908 32.564l 7.104l
0.285m 4.513n 16.007m 32.932m 32.121m 1.913
0.015 0.464 0.873 0.963 0.553 1.031
0.023 1.411 1.207 1.209 1.921 1.554
trends under drought stress, such as an increase in linoleic acid
and a decrease in oleic acid of sunflower. This is in accordance
with our results, because oleic acid decreased but not
significantly (data not shown). Different data on the protein
content change under drought stress can be found in the
literature. A significant decrease was obtained by Ali et al.,24 and
a significant increase was obtained by Zaidi et al.29 In the later
paper, a significant increase for tryptophan and lysine contents
under drought was also recorded. Our results could possibly be
explained by rainfall distribution throughout the vegetation
period, because total precipitation was not substantially
different during the pollination and grain filling period (from
July to September) in 2010 and 2011 growing seasons (147
versus 164 mm, respectively). Also, the genotypes analyzed
were selected as highly drought tolerant in both subtropical and
temperate regions.14 To precisely determine the effects of
drought on the examined kernel traits, an experiment under
controlled temperature and water regimes should be
performed.
Correlations between biochemical components as well as
between biochemical and physical components are given in
Table 7. Significant positive correlations were found between
oil and protein contents as well as oil and tryptophan contents
(p < 0.01). Several other studies have shown positive
correlations between oil and protein contents in maize
kernels. 30−32 Song and Chen32 reported high positive
correlation between lysine and oil, which is in accordance
with our results, because lysine and tryptophan values are
highly correlated and their ratio is normally 3:1.9 These positive
relationships are important for simultaneous improvement of
different nutritional quality traits, because maize is widely used
in both human and animal nourishment.
All correlations between oil and principal FA contents were
non-significant (p < 0.05), but significance was noted with all
trace FAs (data not shown). Non-significant correlations
between oil and saturated palmitic and stearic acids are in
accordance with the results presented in other papers.33,34
However, in the analysis of elite maize breeding material,
Dunlap et al.35 found small but significant positive correlations
between the oil content and levels of stearic and oleic acids.
Several other studies representing different genetic back-
grounds showed significant positive correlations between oil
and oleic acid contents as well as significant negative
correlations between oil and PUFA (linoleic and linolenic
acids) contents.33,34,36 In our study, non-significant correlations
had the same trend; oleic acid was positively correlated (r =
0.017) while linoleic and linolenic acids were negatively
correlated (r = −0.016 and −0.1363, respectively) with the
oil content (data not shown). The lack of significant positive
correlations between oil and FAs disables simultaneous
selection for increased oil content and FAs, but instead, the
DOI: 10.1021/jf504301u
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Journal of Agricultural and Food Chemistry Article

Figure 2. Biochemical and kernel physical characteristics that were significantly influenced by the year. Green bar, optimal temperature and
precipitation conditions (2010); yellow bar, drought (2011).

oil content could be increased independently, along with
altering FA composition into a desired profile.
Considering correlations between principal FAs, palmitic acid
was positively correlated with oleic acid (p < 0.01) and
1257
negatively correlated with linoleic acid (p < 0.05). Correlations
between 18-carbon FAs were either non-significant (stearic and
oleic, oleic and linolenic, and linoleic and linolenic acids) or
significantly negative at p < 0.01 (stearic and linoleic, stearic
DOI: 10.1021/jf504301u
J. Agric. Food Chem. 2015, 63, 1251−1260
 Table 7. Correlations between Kernel Biochemical and Physical Characteristics
principal FAs
a b c
OC 16:0 18:0 18:1d 18:2e 18:3f SFAg MUFAh PUFAi ω6:3 PCj TCk QIl
16:0 ns
18:0 ns ns
18:1 ns 0.433m ns
18:2 ns −0.317n −0.402m −0.955m
18:3 ns ns −0.453m ns ns
SFA ns 0.927m 0.738m ns ns −0.290n
MUFA ns ns ns 0.999m −0.953m ns ns
PUFA ns −0.317n −0.407m −0.955m 1m ns −0.422m −0.953m
ω6:3 ns ns ns −0.380m 0.350n −0.800m ns −0.393m 0.334n
PC 0.567m ns ns ns ns 0.376m ns ns ns −0.313n
TC 0.437m ns ns ns ns ns ns ns ns ns 0.548m
QI ns ns ns ns ns ns ns ns ns ns ns 0.526m
Journal of Agricultural and Food Chemistry

1000KWo ns ns 0.277n 0.407m −0.436m −0.370n ns 0.408m 0.443m ns ns ns ns

FIp ns ns ns ns ns ns ns ns ns ns ns 0.307n 0.345n
KWq ns 0.512m 0.394m ns −0.309n −0.328n 0.548m ns −0.312n ns ns ns ns
KLr ns −0.413m ns ns ns ns −0.302n ns ns ns −0.346n ns 0.278n
KTs ns 0.515m ns ns ns ns 0.420m ns ns ns ns ns ns
HPt ns ns −0.429m ns ns ns −0.346n ns ns ns ns ns ns
EPu −0.430m ns ns ns ns ns 0.426m ns ns ns ns ns ns

1258
GPv 0.467m ns −0.446m ns ns ns −0.360m ns ns ns ns ns ns
HFPw ns 0.317n ns ns ns ns ns ns ns ns 0.321n ns −0.483m
MRx 0.372m ns ns −0.286n 0.306n ns ns −0.370m 0.303n ns 0.381m ns ns
a
OC = oil content. b16:0 = palmitic acid. c18:0 = stearic acid. d18:1 = oleic acid. e18:2 = linoleic acid. f18:3 = linolenic acid. gSFA = saturated fatty acid. hMUFA = monounsaturated fatty acid. iPUFA =
polyunsaturated fatty acid. jPC = protein content. kTC = tryptophan content. lQI = quality index. mSignificant at the 0.01 level. nSignificant at the 0.05 level. o1000KW = thousand kernel weight. pFI =
flotation index. qKW = kernel width. rKL = kernel length. sKT = kernel thickness. tHP = hull proportion. uEP = endosperm proportion. vGP = germ proportion. wHFP = hard fraction portion. xMR =
milling response.
Article

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Journal of Agricultural and Food Chemistry
and linolenic, and oleic and linoleic acids). These results are
only partially in accordance with literature data. Wassom et al.36
found non-significant or weak correlations between palmitic
acid and 18-carbon FAs, while all 18-carbon FAs were
significantly mutually correlated. On the other hand, Orhun
and Korkut34 reported significant correlations between palmitic
acid and 18-carbon FAs and non-significant correlations among
18-carbon FAs. However, a near 1:1 negative correlation
between oleic and linoleic acids found within the analyzed
accessions was also reported in other studies.34,36−39 The
genetic correlation between oleic and linoleic acids, which is
near unity, indicates that most or all of the genes responsible
for variation in the accumulation of these two acids are closely
linked or pleiotropic.36
Correlations between kernel biochemical and physical
characteristics revealed, as expected, that the germ proportion
was positively correlated with the oil content (p < 0.01). This
kernel characteristic has been recognized as the most relevant
characteristic in high oil breeding programs.40−43 Among
principal FAs, only stearic acid was significantly correlated
with germ percent (negatively; p < 0.01). This differs from the
results presented by Preciado-Ortiza et al.,20 where all FAs were
significantly correlated with germ proportion. In our work,
significant correlations with germ were found for all trace FAs
(data not presented).
Thousand kernel weight was positively correlated with stearic
and oleic acids but negatively correlated with linoleic and
linolenic acids. No significant correlation was found between
this trait and the oil content, while in the study by Dorsey-
Redding et al.,44 these two traits were correlated at p < 0.001.
They also noted that the correlation between the protein
content and thousand kernel weight was not significant, and
this is in accordance with our results for the accessions.
Considering other biochemical components analyzed, trypto-
phan was not significantly correlated with this kernel parameter,
while significant positive correlation was found for MUFAs and
PUFAs.
The SFA content was significantly correlated with all kernel
dimensions and kernel proportions, positively with kernel
width, kernel thickness, and endosperm proportion (p < 0.01),
while negatively with kernel length and hull and germ
proportions (p < 0.01, p < 0.01, and p < 0.05, respectively).
The hard fraction portion was in positive correlation with
palmitic acid and protein content (p < 0.05), while milling
response was positively correlated with oil, linoleic acid, total
PUFAs, and protein content (p < 0.01, p < 0.05, p < 0.05, and p
< 0.01, respectively) and negatively correlated with oleic acid
and total MUFAs (p < 0.05 and p < 0.01, respectively). These
two hardness parameters were also positively correlated with oil
and protein contents by Dorsey-Redding et al.44
Oil composition and FA concentrations are regulated by
more than 40 QTL.38,45 It is possible that discrepancies in the
statistical results of different experiments are due to different
genetic backgrounds of the analyzed genotypes and different
environmental effects. Differences between our results and
results obtained in other studies could also be due to
differences in materials analyzed (non-selected populations,
versus elite, commercial material). These accessions could
harbor alleles not present in commercial germplasm, leading to
gene actions involved in oil, FA, and protein metabolic
pathways different from the pathways present in elite
germplasm.
1259
Article
It was shown that maize kernel nutritional value can be
improved by introducing landraces into breeding programs.
Genes from exotic germplasm were used for developing maize
lines with high total SFAs,12 and a single cross hybrid was
developed from an Italian landrace with high carotenoid
content that is being used by an Italian beer industry.3 The
results of biochemical analysis presented in this report indicate
considerable potential of the drought-tolerant accessions for
improving value-added traits of the commercial genotypes.
Moreover, several accessions with multiple nutritional advan-
tages were identified. For example, L632 had high oil and oleic
acid contents as well as high tryptophan content, while IP6428
besides high oil and tryptophan contents was also abundant in
SFA. On the other hand, positive correlation between palmitic
and oleic acids enables the use of IP6428 for improving lines for
high contents of these two FAs. Because the drought-tolerant
accessions were selected in field trials performed in both
subtropical and temperate zones, the identified genotypes could
be used for breeding maize with value-added traits adapted to
both environments.
■
AUTHOR INFORMATION
Corresponding Author
*Telephone: +381-11-3756704. Fax: +381-11-3756707. E-mail:
idragana@mrizp.rs.
Funding
This research was supported by the Ministry of Education,
Science and Technological Development of Republic of Serbia
through Project TR31028 “Exploitation of Maize Diversity To
Improve Grain Quality and Drought Tolerance”.
Notes
The authors declare no competing financial interest.
■
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1260 DOI: 10.1021/jf504301u

J. Agric. Food Chem. 2015, 63, 1251−1260