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Corresponding author:Vlado Kovačević, Faculty of Agriculture, Kralja P. Svačića 1d,
31000 Osijek, Croatia; Phone: ++385 31 554 932, e-mail: vkovacevic@pfos.hr
UDC 575.633.15
DOI: 10.2298/GENSR1601297A
Original scientific paper
GENOTYPIC EFFECTS ON BORON CONCENTRATIONS AND RESPONSE ON
BORON FERTILIZATION IN MAIZE INBRED LINES
Luka ANDRIĆ1, Vlado KOVAČEVIĆ2, Imre KADAR3, Antun JAMBROVIĆ1,
Hrvoje PLAVŠIĆ1and Domagoj ŠIMIĆ1
1Agricultural Institute Osijek, Osijek, Croatia
2Faculty of Agriculture of University J. J. Strossmayer in Osijek,
Osijek, Croatia
3Research Institute for Soil Science and Agricultural Chemistry (RISSAC),
Budapest, Hungary
Andrić L., V. Kovačević, I.Kadar A. Jambrović, H. Plavšić and D. Šimić (2015):
Genotypic effects on boron concentrations and response on boron fertilization in maize
inbred lines- Genetika, Vol 48, No. 1,297 - 305.
Boron (B) deficiency in maize can result in barren cobs attributed to silks being non-
receptive which is particularly important for the female parent in seed production. The
objectives of this study were 1) to investigate genotypic differences among nine female
inbred lines used in seed production for B concentration in ear-leaf and grain, as well as
for grain yield and moisture in a three-year experiment (2006-2008) and 2) to determine
response and relations among the traits when four of the female inbred lines are treated by
foliar boron fertilization - three times in 10-days interval with 0.5% Solubor solution
(17.5% B) during one growing season (2008). The investigations were performed on
Experimental field of Agricultural Institute Osijek, (soil type: eutrical cambisol). Highly
significant differences among the nine female inbred lines were detected for B
concentration in ear-leaf (from 14.7 to 46.7 mg B kg-1) and grain (from 1.20 to 2.06 mg B
kg-1) as well as for grain yield (from 3.33 to 4.83 t ha-1) and grain moisture (from 14.7%
to 26.6%). However, there were also significant effects of growing season and the
genotype by environment interaction for all four traits. Positive and moderate correlations
were found between the boron status in plant and grain yield. Although B concentrations
were considerably increased by foliar boron fertilization (averages 41.7 and 125.3 mg B
kg-1 in leaves, 1.79 and 2.80 mg B kg-1 in grain, for control and fertilization,
298 GENETIKA, Vol. 48, No.1, 297-305, 2016
respectively), in general grain yield differences among treatments were non-significant.
(averages 5.21 and 5.15 t ha-1, respectively).
Key words: boron, female parent inbred lines, foliar fertilization, grain, leaves,
maize, yield
INTRODUCTION
Inbred lines of maize are important parent components for producing hybrid seed and
subsequent growing single-cross hybrids for commercial production. Even though the vegetative
growth and thus the canopy of inbred maize vary among genotypes, it is generally decreased
compared with that of hybrid, mainly due to poor rooting ability of inbred genotypes making them
more vulnerable to nutrient deficiencies and imbalances. Although maize is relatively insensitive
to boron (B) deficiency, poor grain-setting can result in barren cobs, and this was attributed to silks
being non-receptive (GUNES et al., 2011; LORDKAEW et al., 2011) which is particularly important
for the female parent in seed production.
Generally, concerning the susceptibility of plants to B deficiency, many studies are
published relating not only to "boron-intensive species of crops", but also to genotypic differences
within a species (BERGMANN, 1992; MENGEL and KIRKBY, 2001). However, response of maize
genotypes to B nutrition is not well documented in literature. The objectives of this study were 1)
to investigate genotypic differences among nine female inbred lines used in seed production for
boron concentration in ear-leaf and grain, as well as for grain yield and moisture in a three-year
experiment and 2) to determine response and relations among the traits when four of the female
inbred lines are treated by foliar boron fertilization.
MATERIALS AND METHODS
Field experiment
Nine inbred lines – female parents of maize hybrids developed by the Agricultural
Institute Osijek were grown during three growing seasons (2006-2008) on experimental field of
Agriculture Institute Osijek. Maize was planted at the end of April /beginning May by planters on
interrow spacing 70 cm and distance in row 22 cm. Two seeds were sown on each sowing place.
At 3-5 leaf stages maize crop was thinned and one plant in each sowing place was leaved (plant
density = 64936 plant ha-1). The experiment was conducted in four replicates (basic plot 28 m2or
four 10-m long rows).
In third year of testing (2008), based on differences in boron uptake , four inbred lines
were selected for testing of their response to foliar fertilization with boron (Table 3). Foliar
spraying was made with 0.5% solution of Solubor (17.5% B) in three terms as follows: - June 18
(stage of 8-12 leaves), June 27 (stage of 10-14 leaves) and July 3 (before anthesis). Boron
application was made by hand using back sprayer at evening hours.
Sampling,chemical and statistical analysis
Soil sampling of surface layer until 30 cm of depth (one average sample in level of
experiment) was made by auger in October each year after harvesting. Plant available phosphorus
and potassium were determined by ammonium-lactate extraction.
The ear-leaf at anthesis (middle of July: about twenty leaves in mean sample) and grain at
maturity (ten cobs in mean sample) was taken for chemical analysis from each sub-plot. The total
L.ANDRIC et al: GENOTYPIC AND FERTILIZATION EFFECT ON BORON STATUS IN MAIZE 299
amount of boron in the leaf- and grain samples, after microwave digestion using concentrated
HNO3+H2O2, was measured by the ICPAES technique by Jobin-Yvon Ultrace 238 ICP-OES
spectrometer in the laboratory of the Research Institute for Soil Science and Agricultural
Chemistry (RISSAC) of Hungarian Academy of Science and Arts in Budapest, Hungary.
The data were statistically analyzed by ANOVA and treatment means were compared
using t-test and LSD at 0.05 and 0.01probability levels.
Description of the maternal parents of maize hybrids
B1 line belongs to the Iodent heterotic group of maize (JAMBROVIĆ et al. 2014), FAO
group 310, currently used as the female parent of the hybrid Os2983. It is tolerant to water deficit
and high plant density. Our previous data indicated that the line yielded about 2 t ha-1 in seed
production with no substantial deviations. B2 is the female parent of the flint hybrid Tvrtko 303
(KOVAČEVIĆ et al., 2013b) tracing back from a single cross, FAO group 510. It has unique pale
green leaves and white silk indicating putative micronutrient deficit. The line B3 belongs to the
BSSS heterotic group, B73 subgroup (JAMBROVIĆ et al. 2014), FAO group 450. The line is the
female parent of the hybrid Os 499 (KOVAČEVIĆ et al., 2013b) having average grain yield about 2-
3 t ha-1. B4 is the female parent of the hybrid OS 444 (KOVAČEVIĆ et al., 2013b), FAO group 450,
the line of the Lancaster heterotic group (JAMBROVIĆ et al., 2014) yielding about 1.5-2.5 t/ha. B5
belongs to the BSSS heterotic group, the female parent of the hybrid Os 494. It is high yielding in
seed production but with considerable deviations. The line B6 is related to the line B3 and it is the
female parent of the hybrid OSSK 552 (KOVAČEVIĆ et al., 2013b). The dark green inbred line B7
belongs to the BSSS heterotic group and it is the female parent of the hybrid OSSK 596
(KOVAČEVIĆ et al., 2013b), FAO 620. The lines B8 and B9 belong also to the BSSS heterotic
group and they are the female parents of the hybrid OSSK 602, (FAO 620), and OSSK 644 (FAO
650), respectively, with low to average grain yield in seed production.
Soil characteristics
Experimental field of Osijek Agricultural Institute is classified as soil of A-C profile
(eutric cambisol) favorable physical and chemical properties. Reaction of surface layer is neutral
/slightly acid, low in organic matter and normal supplied with plant available phosphorus and
potassium (Table 1).
Table 1. Soil characteristics
Surface soil layer until 30 cm of depth after maize harvesting
Year
pH
mg 100g-1 (AL-method)
%
H2O
KCl
P2O5
K2O
Org. matter
2006
7.35
6.68
15.4
26.7
1.65
2007
6.23
5.28
15.9
25.6
1.78
2008
6.74
5.92
23.9
33.5
2.04
Weather characteristics
Weather characteristics, particularly quantity and distribution of precipitation and air-
temperatures are considerable factors of maize yield and yield variations among years. In general,
lower precipitation and the higher air-temperatures in summer, especially in July and August, are
300 GENETIKA, Vol. 48, No.1, 297-305, 2016
in close connection with the lower yields of maize (MAKLENOVIĆ et al., 2009; MARKULJ et al.,
2010; RASTIJA et al., 2012; MAJDANČIĆ et al., 2015). With that regard, recent climatic change has
mainly negative effects on global food production (PARRY et al., 2005; LOBELL and FIELD, 2007;
SVEČNJAK et al., 2007; SIPOS et al., 2009; VIDENOVIĆ et al., 2013; KOVAČEVIĆ et al.. 2013a;
RENGEL, 2011, 2015).
In accordance with mentioned observations, the 2006 and particularly 2007 growing
season were less favorable for maize growth compared to the 2008 growing season. Water deficit
and high air-temperature in July are main adversely factor of maize growth in 2006, while about
40% lower precipitation and for 1.6 oC higher temperature characterized the 2007 growing
season. Water deficit in 2007 was particularly observed in June-August period (about 50% lower
precipitation compared to usual) and it was accompanied with 2.5 oC higher air-temperature. Total
precipitation in the 2008 growing season was similar to 2006 growing season, but their monthly
distribution was more balanced and more favorable for maize. Also, temperature regime in
summer months of 2008 was more close to usual than in the previous two years (Table 2).
Table 2. The meteorological data (SHS, 2008)
Osijek: Precipitation and mean air-temperatures (61-90: averages 1961-1990 )
Year
Monthly precipitation (mm)
Monthly mean air-temperatures (oC)
Apr.
May
June
July
Aug
Sept
Σ
Apr.
May
June
July
Aug
Sept
X
2006
87
79
91
15
134
11
415
12.7
16.2
20.1
23.5
19.3
17.8
18.3
2007
3
56
33
27
45
65
230
13.3
18.2
22.3
23.8
22.2
14.5
19.1
2008
50
67
76
79
46
86
405
12.5
18.1
21.5
21.8
21.8
15.7
18.6
61-90
54
58
88
65
59
45
368
11.3
16.5
19.5
21.1
20.3
16.6
17.5
RESULTS AND DISCUSSION
The B contents of monocotyledons are about 2 to 6 mg kg-1. Ranges of 6 - 15 mg B kg-1
contents in dry matter of maize leaves are adequate for normal maize growth (BERGMANN, 1992).
According to these criteria, leaf-B status from 8.8 to 58.1 mg B kg-1 in our study (Table 3) is
adequate for maize.
Both growing season and genotype affected significantly on leaf-B status in maize. Under
unfavorable weather conditions of the 2007 growing season (Table 2) average leaf-B concentration
in maize was 19.7 mg B kg-1 or about twofold lower compared to values in remaining two growing
seasons. Differences of leaf-B among genotypes (3-year averages) were from 14.7 (the line B2) to
46.9 (the line B4) mg B kg-1 and these differences were considerably higher than among years. In
five genotypes, leaf-B concentrations were in range between 34 and 39 (average 36.9) mg B kg-1,
while in two genotypes between 22 and 29 (average 25.5 ) mg B kg-1 (Table 3). Significant
differences for B status among maize inbred lines and their diallel crosses were also reported by
KOVAČEVIĆ et al. (2001) for B-grain and among maize hybrids for B-leaf and B-grain (KOVAČEVIĆ
et al., 2013b), while BRKIĆ et al. (2015) reported about significant differences among 127 maize
genotypes for B concentration in root.
L.ANDRIC et al: GENOTYPIC AND FERTILIZATION EFFECT ON BORON STATUS IN MAIZE 301
Table 3 . Impacts of growing season and genotype on maize status: leaf-boron, grain-boron, grain yield
and grain moisture
Year (the factor A: A1=2006, A2=2007, A3=2008) and genotype (the factor B) impacts
on boron status in maize and grain yield on Osijek eutric cabisol
Maize genotype
Mean
Year
B1
B2
B3
B4
B5
B6
B7
B8
B9
A
Leaf-boron (mg B kg-1 in dry matter od the ear-leaf at silking stage)
A1
24.1
13.8
38.2
51.7
32.7
34.8
38.9
44.5
37.2
35.1
A2
14.1
8.8
18.6
30.7
15.1
21.1
23.6
24.2
21.3
19.7
A3
29.3
21.6
57.4
58.1
37.3
57.9
41.7
47.6
46.0
44.1
x B
22.5
14.7
38.1
46.9
28.4
37.9
34.7
38.8
34.8
P 0.05
A: 3.1
P 0.05
B: 2.8
P 0.05
AB: 6.0
P 0.01
5.1
P 0.01
3.6
P 0.01
9.0
Grain-boron (mg B kg-1 in dry matter at maturity stage)
A1
1.14
0.73
2.03
2.60
1.76
2.24
1.70
1.48
1.43
1.68
A2
2.20
1.60
2.07
1.69
1.37
1.67
1.34
1.33
1.37
1.63
A3
1.84
1.28
1.63
1.88
1.60
2.16
1.91
1.71
1.53
1.73
x B
1.73
1.20
1.91
2.06
1.58
2.03
1.65
1.51
1.44
P 0.05
A: n.s.
P 0.05
B: 0.14
P 0.05
AB: 0.34
P 0.01
P 0.01
0.19
P 0.01
0.53
Grain yield (t ha-1 on 14% grain moisture basis)
A1
2.81
2.94
4.41
3.84
3.81
3.35
4.42
4.00
3.47
3.67
A2
3.98
4.72
5.12
3.66
3.80
4.84
3.21
1.80
2.20
3.70
A3
4.64
5.50
4.96
5.76
5.98
6.23
5.22
5.27
4.31
5.32
x B
3.81
4.39
4.83
4.42
4.53
4.81
4.28
3.69
3.33
P 0.05
A: 0.62
P 0.05
B: 0.56
P 0.05
AB: 1.19
P 0.01
0.84
P 0.01
0.73
P 0.01
1.56
Grain moisture (%) at harvesting
A1
14.7
18.9
19.2
20.5
18.4
22.9
26.4
29.9
30.5
22.4
A2
13.0
18.7
17.5
19.1
17.2
17.8
24.9
24.5
23.8
19.6
A3
16.4
22.3
224
23.3
21.0
20.5
23.9
23.7
25.5
22.1
x B
14.7
20.0
19.7
21.0
18.9
20.4
25.1
26.0
26.6
Grain-B was considerably lower compared to leaf-B status in maize (3-year averages 1.68
and 33.0 mg B kg-1, respectively). Differences of average grain-B among years were non-
significant, but differences among genotypes were from 1.20 (B2) to 2.06 (B4) mg B kg-1. In two
302 GENETIKA, Vol. 48, No.1, 297-305, 2016
genotypes (the lines B4 and and B6) grain-B was above 2.0, while in four genotypes it was below
1.6 mg B kg-1 (the lines B5, B8, B9 and B2).
Differences among B concentrations in level of year x genotype interaction were from 8.8
to 58.1 (leaf-B) and from 0.73 to 2.60 (grain-B) mg B kg-1. The lowest values were found in in B2
(both in leaf and grain) in 2007 (leaf) and 2006 (grain). The highest values were found in parent
135-88 in 2008 (leaf) and 2006 (grain).
No strong associations were found between B status in maize with yield because two the
most divergent genotypes had similar yields (3-year averages 4.39 and 4.42 t ha-1, for the B2 and
B.4 lines, respectively), probably because B status in maize was in adequate levels. Across all
three growing seasons, pooled correlations coefficients were moderate between the leaf-B and
grain yield (r=0.41) and between the grain B concentrations and grain yield (r=0.44). No
association was detected between the grain B concentrations and grain moisture (4=0.04).
Table 4. Impact of genotype and foliar spraying with boron solution on maize status
Impact of genotype and foliar fertilization (A1 = the control, A2 = three-times foliar fertilization* ) with Solubor* on
boron status , yield and grain moisture in maize (the 2008 growing season )
Treat-
Maize genotype (the factor B) *
Maize genotype (the factor B)*
ment
B1
B2
B4
B6
Mean A
B1
B2
B4
B6
Mean A
Leaf -B (mg B kg-1 in dry matter)
Grain-B (mg B kg-1 in dry matter)
A1 ( 0 )
29.3
21.6
58.1
57.9
41.7
1.84
1.28
1.88
2.16
1.79
A2 (FF)
100.1
88.0
147.0
166.0
125.3
2.96
2.40
3.16
2.90
2.80
Mean B
64.7
54.8
102.6
112.0
2.40
1.84
2.52
2.53
A
B
AB
A
B
AB
P 0.05 11.6
P 0.05 16.4
P 0.05 ns
P 0.05 0.08
P 0.05 0.15
P 0.05 ns
P 0.01 16.0
P 0.01 22.7
P 0.01 0.11
P 0.01 0.19
Grain yield ( t ha-1)
Grain moisture (%) at harvest
A1 ( 0 )
4.64
5.50
4.96
5.76
5.21
16.4
22.3
23.3
20.5
20.6
A2 (FF)
4.54
5.18
4.67
6.19
5.15
17.0
22.1
23.7
20.1
20.7
Mean B
4.59
5.34
4.82
5.98
16.7
22.2
23.5
20.3
20.7
A
B
AB
A
B
AB
P 0.05 ns
P 0.05 0.38
P 0.05 ns
P 0.05 ns
P 0.05 1.8
P 0.05 ns
P 0.01 0.54
P 0.01 2.5
* foliar spraying (3x) with 0.5 % of Solubor (17,5 % B) solution ( June 18 and 27, July 3)
In experiment with B foliar fertilization performed with four genotypes, the lines B1 and
B2 had considerably lower leaf-B concentrations (average 59.8 mg B kg-1) compared B4 and B6
parents (average 134.4 mg B kg-1). Application of B fertilizer had considerable effect on leaf-B
concentrations (averages 41.7 and 125.3 mg B kg-1, for control and B fertilization, respectively),
but grain yields were independent on B fertilization (averages 5.21 and 5.15 t ha-1, respectively).
L.ANDRIC et al: GENOTYPIC AND FERTILIZATION EFFECT ON BORON STATUS IN MAIZE 303
Application of B had also significant impact on grain-B status in maize inbreeds (averages 1.79
and 2.80 mg B kg-1, respectively.
Regarding grain-B, only B2 had considerably lower B concentrations (average 1.84 mg B
kg-1), because in remaining three genotypes average B concentrations were 2.48 mg B kg-1. (Table
4). However, the results indicate that foliar boron fertilization did not consistently affects grain
yield due to non-significant difference between the two treatments. However, the inbred line B6
did respond on B fertilization having considerably higher yield when treated.
CONCLUSIONS
In the three-year experiment, highly significant differences among the nine female inbred
lines used in seed production were detected for boron concentration in ear-leaf and grain as well as
for grain yield and grain moisture. However, there were also significant effects of growing season
and the genotype by environment interaction for all four traits. Positive and moderate correlations
were found between the boron status in plant and grain yield. Foliar boron fertilization affected
boron status in ear-leaf and grain but there was generally no effect on grain yield with the
exception of only one female inbred line.
Received December 03th, 2015
Accepted February 16th, 2016
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GENETIČKI EFEKTI NA KONCENTRACIJE BORA I REAKCIJA INBRED LINIJA
KUKURUZA NA ĐUBRENJE BOROM
Luka ANDRIĆ1, Vlado KOVAČEVIĆ2, Imre KADAR3; Antun JAMBROVIĆ1,
Hrvoje PLAVŠIĆ1and Domagoj ŠIMIĆ1
1Agricultural Institute Osijek, Osijek, Croatia
2Faculty of Agriculture of University J. J. Strossmayer in Osijek, Osijek, Croatia
3Research Institute for Soil Science and Agricultural Chemistry (RISSAC), Budaimpešta,
Mađarska
Izvod
Nedostatak bora (B) u kukuruzu može biti uzrokom pojavi jalovog klipa, jer svila ne može
prihvatiti polen, a što je veoma značajno za majčinsku komponentu u proizvodnji semena.
Predmeti ovoga rada bili su 1) u trogodišnjim istraživanjima (2006.-2008.) ustanoviti genetske
razlike u koncentracijama bora u listu i zrnu, prinosima zrna i vlažnosti zrna, između devet
majčinskih inbred linija koje se koriste u proizvodnji semena i 2) ustanoviti reakciju četiri
odabrane linije na folijarno đubrenje borom - tri puta u 10-dnevnom u intervalu s 0.5% rastvorom
Solubora (17,5% B) tokom jedne godine (2008.). Istraživanja su provedena na oglednom polju
Poljoprivrednog instituta Osijek, na tipu zemljišta eutrični kambisol. Ustanovljene su visoko
signifikantne razlike između devet genotipova kukuruza u koncentracijama bora u listu ispod klipa
tokom svilanja (od 14.7 do 46.7 mg B kg-1) i u zrnu (od 1.20 do 2.06 mg B kg-1), te u prinosima
zrna (od 3.33 do 4.83 t ha-1) i vlažnosti zrna (od 14.7% do 26.6%). Takođe, postojali su i značajni
uticaji faktora godina, te interakcije genotipa i okoline na sva četiri analizirana svojstva. Pozitivne
i umerene korelacije su ustanovljene između koncentracija bora u biljci i prinosa zrna. Iako su
koncentracije bora značajno povećane folijarnom prihranom (prosečno 41.7 i 125.3 mg B kg-1 u
listu, 1.79 i 2.80 mg B kg-1 u zrnu, za kontrolu odnosno tretman folijarnog đubrenja), razlike
prinosa nisu bile statistički značajne (5,21, odnosno 5,15 t ha-1).
Primljeno 03. XII 2015.
Odobreno 16. II. 2016.
