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Tools for Improving Sweet Corn Yield

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 ISSN: 2456-7515
International Journal of Advances in Agriculture Sciences
(A Monthly Scientific Journal of Kiban Research Publications)
www.kibanresearchpublications.com

RESEARCH ARTICLE

Tools for Improving Sweet Corn Yield

J Rattin1, J Molinari2, E Giardina2, Adalberto Di Benedetto1-2*
1Faculty of Agricultural Sciences, National University of Mar del Plata, Route 226, km. 73.5 (B7620ZAA),
Balcarce, Province of Buenos Aires, Argentina.
2Facultyof Agronomy, University of Buenos Aires, San Martín 4453 avenue (C1417DSE), Buenos Aires,
Argentina.

*Corresponding Author: Email: dibenede@agro.uba.ar

Abstract: Future demand for high quality agricultural foods will increase over the coming decades,
driven by population growth and changing dietary habits. The two basic options for agricultural
production are expansion of the land area under agricultural production, and agricultural land-use
intensification. Because land expansion is limited, intensification has been and will increasingly become
more important in the future. It has been shown that the use of transplant is the most reliable method to
ensure adequate crop establishment of commercial plantings of various high-value vegetable crops,
including sweet corn. On the other hand, the exogenous hormonal management of crops has emerged as
a promised tool for increasing yield. The aim of this work was to determine the combined effects of a
transplant routine and a single benzyl aminopurine (BAP) spray in pre- and post-transplant on
commercial yield for two different super sweet maize hybrids to test the hypothesis that an increase in
commercial sweet maize yield is possible and related to endogenous hormonal signaling. Our results
showed an increase sweet maize yield over an area basis related to transplanting plants as a result of
higher photo assimilate fixation and partitioning to stems. On the other hand, a single early BAP-spray
additionally increases yield and growth responses through the involvement of cytokine’s as an
endogenous signaling and a correct vascular development.
Keywords: cytokinin; Endogenous signaling; Photo assimilate production; Photo assimilate partition;
Transplant.
Article Received: 01 Sept. 2018 Revised: 11 Sept. 2018 Accepted: 22 Sept. 2018

Abbreviations Used
su (sugary maize mutant hybrid); sh2
(shrunken maize mutant hybrid); PAR
(photosynthetically active radiation); BAP
(benzyl aminopurine); RGR (relative growth
rate); RLAE (rate of leaf area expansion);
NAR (net assimilation rate); LAR (leaf area
ratio); SLA (specific leaf area); LAI (leaf area
index); CGR (crop growth rate); RUE
(radiation use efficiency); HI (harvest index).
Introduction
Sweet corn (Zea mays), considered as a
vegetable, is a special type of corn with sweet
taste, thin pericarp and endosperm both with
a high nutritional value. It is destined
exclusively for human consumption, in fresh
form or in processed foods [1]. Sweet corn
quality has been improved with the use of
different mutants (sugary, su; shrunken, sh2)
[2-3]. In countries with short-seasons such as
those found in the Mar del Plata (Argentine),
planting window exposes the crop to various
stresses and weather risks. It has been
suggested that sh2 corn genotypes should be
planted shallower and at soil temperatures
higher than required for the normal (su)
genotype [4-5]. Although these super sweet
corn phenotypes are favored among
consumers, sweet corn growers have found
that cold soils show slow emergence, and
reduce stands and profitability [6].
Future demand for agricultural products will
increase over the coming decades, driven by
population growth and changing dietary
habits, which means that agricultural
production will have to increase [7]. On the
other hand, the threat of global climate
change is causing concern in agriculture,
given that climatic factors essential for crop
development will be severely affected,
reducing the production and quality of food

A Di Benedetto et. al.| Oct. 2018 | Vol.3| Issue 10 |01-14 1

 Available online at: http://ijaas.kibanresearchpublications.com/index.php/IJAAS
[8-9]. The two basic options for this are
expansion of the land area under agricultural
production, and agricultural land-use
intensification. Because land expansion is
limited, intensification has been and will
increasingly become more important in the
future [10].
The first strategy for sweet maize yield
increase which included changes in plant
population [11] as an approach to improve a
fast and early PAR interception has been
limited by self-shading [12]. As a result,
sweet maize yield become steady around 15
tons ha-1 and did not change during the last
decade [13-14-15]. A change in cultural
practices is the second option to improve
yield. In this way, mulch practices [16] and a
transplant routine [17-18] appear as
promising tools for improving sweet maize
productivity.
Finally, one of the last alternatives is a
change in hormonal balance through
exogenous applications of cytokinins [15-19].
In the course of development of a maize crop,
the architecture of the stand may differ,
leading to differences in the distribution of
radiation within the stand, which in turn
may be responsible for differences in
productivity indices per unit area [20].
Usually, it has been assumed that corn
biomass accumulation between sowing and
harvest are directly related to incident
photosynthetically active radiation (PAR)
intercepted by the canopy and the efficiency
to convert PAR into dry matter [21].
However, Rattin et al. [15] working with four
super sweet maize, have recently suggested
that yield would be the result of both PAR
and ground stimulus, which determine photo
assimilate production and partition. This
novelty approach would indicate that root
growth could be considered as a limiting
factor to shoot growth and yield in mutants
maize crops. The aim of this work was to
determine the combined effects of a
transplant routine and a single benzyl
aminopurine (BAP) spray in pre- and post-
transplant on commercial yield for two
different super sweet maize hybrids.
The proposed experiments test the
hypothesis that an increase in commercial
sweet maize yield is possible and related to a
change in cropping systems and endogenous
hormonal signaling.
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Materials and Methods
Plant Material and Experiments
To reach the proposed objectives and validate
proposed hypothesis, two experiments were
performed:
Experiment 1
A greenhouse experiment was carried out at
the Faculty of Agronomy, University of
Buenos Aires, Argentina (34° 35’ 59’’S, 58°
22’ 23’’W) during December 2013. The corn
mutant hybrids ‘Butter Sweet’ (sh2) provided
by Semillería Basso (Argentina) was sown in
plastic plug trays (128 cells tray-1; 17.37 cm3
cell-1) using a commercial growing media
(Klasmann 411® medium, Klasmann-
Deilmann, GmbH, Germany) or direct-seeded
in 5-litre pots filled with a Sphagnum
maguellanicum-river waste-perlite (40-40-20,
v/v/v) medium). The plants were arranged at
a density of four plants m-2, which avoided
mutual shading and both direct-seeded and
transplants were at the same growth state at
the beginning of the experiment. Transplants
were grown under greenhouse facilities from
sowing to transplant.
Transplants were conducted by hands.
Seedlings were sprayed with different BAP
(6-benzylaminopurine) (SIGMA EC 214-927-
5) (Sigma-Aldrich Co., St. Louis, MO, USA)
solutions (0, 50, 100 or 200 mg L-1) when first
true leaf pairs appeared (pre-transplant
application or 7 days from direct-seeded
plants) or one week after transplant (post-
transplant application or 15 days from direct-
seeded plants). BAP was previously diluted in
alcohol 80% and plants were run-off sprayed.
At the beginning of the experiment total
porosity (%), air-filled porosity (%), container
capacity (%) and bulk density (g cm-3) were
63.50, 17.06, 40.06 and 0.35 respectively.
Organic matter (%), pH, electric conductivity
(dS m-1) and cation exchange capacity were
45.3, 5.2, 0.71 and 58.9 respectively.
A weekly ferti-irrigation of 1.0: 0.5: 1.0: 0.5
(v/v/v/v) N: P: K: Ca (nitric acid, phosphorus
acid, potassium nitrate, and calcium nitrate)
(150 mg l-1 N) was included through to the
overhead irrigation water. Half hourly
averages of the air temperature were
measured using a HOBO H08-001-02 data
logger (Onset Computer Corporation, MA,
USA) protected from direct radiation by
aluminum foil shades. Minimum
temperature, maximum temperature and
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mean global solar radiation during the
experiment were 25.11°C, 28.31°C and 25.60
MJ m-2 day-1 respectively.
Experiment 2
The experiment was conducted at the INTA
Balcarce Experimental Station, Argentina
(37°45’S, 58°18’W) from November 22th 2012
to February 10th 2013 on a Typic Argiudoll
soil with an organic matter of 5.6% in the
first 25 cm depth. The experiment was
repeated twice from November 26th 2013 to
February 15th 2014 and from November 19th
2014 to February 16th 2015. Water and
nutrients were at non-limiting levels using
an irrigation system, which kept soil water
above 50% of maximum soil available water
in the first meter of depth.
The corn mutant hybrids ‘Canner’ (su1) and
‘Butter Sweet’ (sh2) provided by Semillería
Basso (Argentina) were sown in plastic plug
trays (128 cells tray-1; 17.37 cm3 cell-1) using
a commercial growing media (Klasmann
411® medium, Klasmann-Deilmann, GmbH,
Germany) or direct-seeded. Transplanted
plants were grown under greenhouse
facilities from sowing to transplant and were
conducted by hands. Both direct-seeded and
transplants were at the same growth state at
the beginning of the experiment. Final
population densities were 80,000 plants ha-1.
The experimental field was fertilized with
150 kg N ha-1 (18-46-0) at the beginning of
experiments. Seedlings were sprayed with
different BAP (6-benzylaminopurine)
(SIGMA EC 214-927-5) (Sigma-Aldrich Co.,
St. Louis, MO, USA) solutions (0, 50, 100 or
200 mg L-1) when first true leaf pairs
appeared (pre-transplant application or 7
days from direct seeded plants) or one week
after transplant (post-transplant application
or 15 days from direct-seeded plants). BAP
was previously diluted in alcohol 80% and
plants were run-off sprayed.
Weather records (daily maximum-minimum
air temperature and global solar radiation)
were recorded from a meteorological station
500 meters from the experimental site. The
mean air temperatures were 13.04, 12.06,
14.24°C (minimum) and 27.33, 27.16, 28.82°C
(maximum) during the 2012-2013, 2013-2014
and 2014-2015 experiments respectively.
Mean light were 21.82, 22.57 and 23.02 MJ
m-2 day-1 during the 2012-2013, 2013-2014
and 2014-2015 experiments respectively.
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Growth Evaluations
Experiment 1
Plants for destructive measurements were
harvested (ten plants per treatment) at
emergence and at 7-day intervals during the
experiment. Roots were washed and root,
stem and leaf fresh weights (FW) were
recorded. Dry weights (DW) were recorded
after drying roots, stems and leaves to
constant weight at 80°C for 96 hours.
The number of leaves was recorded and each
leaf area was determined using a leaf area
meter LI-COR FL16 (LI-COR Inc., Lincoln,
NE, USA) the relative growth rate (RGR) was
calculated as the slope of the regression of
the natural logarithm of the whole plant on a
DW basis versus time (in days). The rate of
leaf area expansion (RLAE) was calculated as
the slope of the regression of the natural
logarithm of total leaf area versus time (in
days). The mean net assimilation rate (NAR),
and the leaf area ratio (LAR) were calculated
as follows:
kwW0ekwt
NAR =
A0ekat
Aa ekat
LAR  ka /
kwW0ekwt
Where kW: RGR (g g-1 days-1); W0:
extrapolated value of total dry weight at time
zero (g); A0: extrapolated value of leaf area at
time zero (cm2); ka: RLAE (cm2 cm-2 days-1); t:
time (in days) at the midpoint of the
experimental period and e: base of natural
logarithms. The specific leaf area on a FW
basis (SLA) was calculated as the ratio
between the area of the new individual leaf
and leaf FW. The allometric coefficients
between root and shoot and between leaf
blades and the stem fraction were calculated
as the slope (β) of the straight-line regression
of the natural logarithm of the root DW
versus the natural logarithm of the shoot DW
and between the natural logarithm of the leaf
blade DW versus the natural logarithm of the
stem DW respectively.
Experiment 2
For field experiments, five plants per
treatment at E-T (emergence-transplant) and
V4, V7, V9, VT and R3 stages were harvest.
Roots were washed and root, stem and leaf
FW were recorded. On the other hand, DW
were recorded after drying roots, stems and
leaves to constant weight at 80°C for 96
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hours. Individual leaf area was determined
using a leaf area meter LI-COR FL16 (LI-
COR Inc., Lincoln, NE, USA). The sample
size was five plants per block. Leaf area
index (LAI) was calculated using the total
leaf area per unit sample soil. The crop
growth rate (CGR) related the total DW with
time (in days) and the unit sample soil (m2).
Radiation use efficiency (RUE) was
calculated as the DM accumulated divided by
the intercepted PAR accumulated from E/T to
VT.
The harvest index (HI) was calculated as the
FW of the harvested ears as a percentage of
the total shoot FW of the plants. Samples for
examination of leaf anatomy were collected
and tissue from the middle region of the
lamina was fixed in formalin-acetic-alcohol.
Leaf thickness and phloem/xylem ratio, were
determined from leaf lamina tissues
embedded in paraffin and sectioned at 20 µm
on a rotary microtome. Both materials were
stained with safranin-crystal violet-fast
green. Data are the mean of ten leaf cross-
sections per leaf from three leaves per
treatment. An image analysis system (Image
Pro Express v 6.0, Media Cybernetics, USA)
facilitated quantitative anatomical
measurements.
Statistical Analysis
We used a complete randomized design for
Experiment 1 and a randomized design with
three blocks of five plants for experiment 2.
Four rows of 10 m (0.70 m apart) for each
treatment for Experiment 2 were used. Since
we found no significant differences in
Experiment 2 data between three successive
years, we considered them together (n = 9).
Data were subjected to one-way analysis of
variance and means were separated by
Tukey’s test (P < 0.05); STATISTICA 8 (Stat
Soft) software was used. Slopes from
straight-line regressions of RLA, RLAE,
RGR, NAR, LAR and allometric values were
tested using the SMATR package [22].
Result
Pot Experiment
Dry Weight Accumulation
Figure 1A shows there is no significant DW
change related to control due to a BAP spray
during the first 35 days from sowing in
direct-seeded plants. However, the same
hormonal treatments in transplanted ones
showed a significant DW accumulation
increase, especially when a 200 mg L-1 pre-
transplant BAP spray was applied (Figure
1B). A positive relationship between shoot
DW and root DW for both direct-seeded and
transplanted plants (r2 = 0.966 and 0.965
respectively) was found (Figure 2).

Figure 1: Changes in total dry weight for ‘Butter Sweet’ (sh2) super sweet maize hybrid under direct-seeded (A) or
transplanting (B) from experiment 1. Plants were sprayed with zero (control plants) or different BAP concentrations
(50, 100 or 200 mg L-1) in pre-transplant (or 7 days from emergence in direct-seeded plants) (empty symbols) or post-
transplant (or 15 days from emergence in direct-seeded plants) (full symbols). Vertical lines indicate least significant
differences (LSD) (n = 3)

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Figure 2: Shoot-root dry weight relationships during the experiment 1 in ‘Butter Sweet’ (sh2) super sweet maize
hybrid under direct-seeded (full symbols) or transplanting (empty symbols). Plants were sprayed with different BAP
concentrations (0, 50, 100 or 200 mg L-1) in pre-transplant (or 7 days from emergence in direct-seeded plants) or post-
transplant (or 15 days from emergence in direct-seeded plants). The linear regression equations are Shoot dry
weight-direct seeded = 3.01 Root dry weight - 0.04 (r2 = 0.966, P < 0.001) and Shoot dry weight-transplanted = 3.25 Root
dry weight - 0.12 (r2 = 0.965, P < 0.001). The probability of the slope being zero was P < 0.001 for both implantation
routines
Leaf Area and Biomass Accumulation treatments showed higher RLA than
controls. Transplants showed lower RLA and
In direct-seeded plants, only 50 and 100 mg
higher SLA values than direct-seeding plants
L-1 BAP increased RLA over the rest of the
so in controls as in all of the BAP spray
treatments, including controls. In
tested. No significant differences in RLAE
transplanted plants, all BAP spray
were found (Table 1).

Table 1: Changes in the rate of leaf appearance (RLA), the relative leaf area expansion rate (RLAE) and the specific
leaf area (SLA) for ‘Butter Sweet’ (sh2) super sweet corn hybrid under direct-seeded or transplant. Different
lowercase letters indicate significant differences (P < 0.05) between control and BAP-sprayed plants, while different
capital letters indicate significant differences (P < 0.05) among sowing routines (n = 3)
Sowing routine BAP RLA RLAE SLA
(mg L-1) (leaves week-1) (cm2 cm-2 day-1) (cm2 g-1)
Direct-seeded 0 0.261bA 0.146aA 304.29aB
7-50 0.306aA 0.149aA 294.03aA
7-100 0.285aA 0.146aA 280.98bA
7-200 0.267bA 0.153aA 271.41bA
15-50 0.265bA 0.144aA 282.94bB
15-100 0.265bA 0.143aA 295.20aB
15-200 0.265bA 0.145aA 301.58aB

Transplant 0 0.219bB 0.132aB 351.27aA

Pre-50 0.239aB 0.137aB 305.23bA
Pre-100 0.231aB 0.138aA 298.42bA
Pre-200 0.232aB 0.129aB 275.29cA
Post-50 0.231aB 0.129aB 314.06bA
Post-100 0.239aB 0.131aB 328.11bA
Post-200 0.242aB 0.135aB 327.20bA

There were small but no significant sprayed with 100 or 200 mg L-1 pre-
differences in RGR and LAR between BAP transplant or 50, 100 and 200 mg L-1 BAP
treatments or implantation routines. sprayed-plants. BAP increased NAR in
However, NAR values were significantly direct-seeded plants as well (Table 2).
higher in transplanted plants and in those

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Table 2: Changes in the relative growth rate (RGR), the net assimilation rate (NAR) and the leaf area ratio (LAR) for
‘Butter Sweet’ (sh2) super sweet corn hybrid under direct-seeded or transplant. Different lowercase letters indicate
significant differences (P < 0.05) between control and BAP-sprayed plants, while different capital letters indicate
significant differences (P < 0.05) among sowing routines (n = 3)
Sowing routine BAP RGR NAR LAR
(mg L-1) (g g-1 day-1) (g cm-2 day-1) (cm2 g-1)
(10-3)
Direct-seeded 0 0.154bA 0.818bB 188.73bA
7-50 0.161bA 0.850bB 189.22bA
7-100 0.162bA 0.925aB 175.21bA
7-200 0.160bA 0.894aB 179.04bA
15-50 0.157bA 0.890aB 176.80bA
15-100 0.172aA 0.740cB 232.32aA
15-200 0.163bA 0.913aA 178.39bA

Transplant 0 0,167aA 0.892bA 186.89aA

Pre-50 0,171aA 0.950aA 180.30aA
Pre-100 0,164aA 0.953aA 171.95aA
Pre-200 0,176aA 0.978aA 179.99aA
Post-50 0,161aA 0.921bA 175.18aA
Post-100 0,161aA 0.893bA 179.76aB
Post-200 0,160aA 0.890bA 179.63aA

Direct seeding control plants showed higher A single BAP spray at different doses and
root: shoot and lower stems: leaves slope application time decreased root: shoot β
straight-line coefficients (β) than coefficient in both direct-seeded plants and
transplanted plants from plants allometries. transplanted ones (Table 3).

Table 3: Changes in allometric relationships between roots and shoots and between stem and leaves for ‘Butter
Sweet’ (sh2) super sweet corn hybrid under direct-seeded or transplant. The slope straight-line (β) are indicated. The
probability of the slope being zero was P < 0.001. Different lowercase letters indicate significant differences (P < 0.05)
between control and BAP-sprayed plants, while different capital letters indicate significant differences (P < 0.05)
among sowing routines (n = 3)
Sowing routine BAP Root: Shoot Stem: Leaves
(mg L-1)
β β
Direct-seeded 0 0.854aA 1.147aB
7-50 0.731bA 1.064bA
7-100 0.701bA 1.095bA
7-200 0.702bA 1.129aA
15-50 0.674cA 1.140aA
15-100 0.701bA 1.108bA
15-200 0.716bA 1.084bA

Transplant 0 0.790aB 1.174aA

Pre-50 0.707bB 1.081bA
Pre-100 0.620cB 1.084bA
Pre-200 0.661cB 1.107bA
Post-50 0.616dB 1.117bA
Post-100 0.628dB 1.098bA
Post-200 0.624cB 1.095bA

Figure 3: Relationship between RGR (A), NAR (B) and the root dry weight (RFW) in plants of ‘Butter Sweet’ (sh2)
super sweet corn hybrid under direct-seeded (empty symbols) or transplanting (full symbols) from experiment 1.
Plants were sprayed with different BAP concentrations (0, 50, 100 or 200 mg L -1) in pre- or post-transplant. The
straight-line regressions were RGR = 0.010 root DW + 0.13 (r 2 = 0.712 P < 0.001), NAR = 0.106 root DW + 0.55 (r2 = 0.605
P < 0.001) (n = 3)

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Field Experiments the implantation routine and genotype.

Since we found no significant differences
between three successive years, mean data
are presented.
Leaf Area Expansion
Direct-seeded sweet maize plants showed
higher leaf area than transplants at both
flowering (Vt) and ear harvest (R3). A single
BAP spray increased leaf area although the
doses for highest response are related to both
Anyway, a higher proportion of the leaf area
expanded remained at the ear harvest stage
in transplanted plants (data not shown).
Although there were no significant RLA and
RLAE differences between control and BAP-
sprayed plants in both sweet maize
genotypes tested, direct-seeded plants
showed higher values than transplanted ones
(Table 4).

Table 4: Changes in the total leaf area at both flowering (Vt) and ear harvest (R3), the leaf area appearance and the
relative leaf expansion area (RLAE) for two sweet corn hybrids (‘Canner’ su1 and ‘Butter Sweet’ sh2) under direct-
seeded or transplanting. Plants were sprayed with different BAP concentrations (0, 50, 100 or 200 mg L -1) in the pre-
transplant stage or 7 days from emergence in direct-seeded plants. Different lower case letters indicate significant
differences (P < 0.05) between control and BAP-sprayed plants, while different capital letters indicate significant
differences (P < 0.05) among sowing routines (n = 9)
BAP Leaf area RLA RLAE
(mg L-1) (cm2 plant-1) (leaves week-1) (cm2 cm-2 day-1)

‘Canner’
Direct-seeded 0 4,208.75bA 1,448.93bA 0.142aA 0.212aA
5 4,674.82aA 2,319.12aA 0.143aA 0.213aA
50 4,576.54aA 2,503.01aA 0.144aA 0.213aA
100 4,674.82aA 2,490.84aA 0.143aA 0.214aA
200 4,573.37aA 2,471.30aA 0.145aA 0.213aA

Transplanted 0 2,182.47bB 1,208.40bB 0.122aB 0.183aB

5 2,675.48aB 1,613.37aB 0.122aB 0.184aB
50 2,661.21aB 1,759.22aB 0.123aB 0.185aB
100 2,201.50bB 1,518.26aB 0.128aB 0.183aB
200 2,066.75bB 1,309.14bB 0.125aB 0.182aB

‘Butter Sweet’
Direct-seeded 0 3,821.95bA 2,052.79bA 0.140aA 0.211aA
5 4,586.05aA 2,702.75aA 0.140aA 0.212aA
50 4,500.44aA 2,490.33aA 0.140aA 0.212aA
100 4,649.46aA 2,385.70aA 0.138aA 0.213aA
200 4,484.59aA 2,452.28aA 0.139aA 0.213aA

Transplanted 0 2,219.22bB 1,570.57bB 0.112aB 0.185aB

5 3,338.11aB 1,700.56aB 0.117aB 0.186aB
50 3,090.81aB 1,697.39aB 0.117aB 0.185aB
100 2,814.98aB 1,740.19aB 0.112aB 0.184aB
200 2,550.25bB 1,347.06bB 0.116aB 0.183aB

Biomass Accumulation transplanted plants, a single BAP spray

increased NAR. NAR values were always
Table 5 showed no significant differences for
significantly higher for transplanted plants
RGR between controls and BAP-sprayed
than those direct-seeded ones. On the other
plants in both sowing routine and for both
hand, CGR was higher in BAP-sprayed
sweet corn genotype. ‘Canner’ plants showed
plants and in those transplanted ones.
no NAR differences, but in ‘Butter Sweet’

Table 5: Changes in the relative growth rate (RGR), the net assimilation rate (NAR) and crop growth rate (CGR) for
two sweet corn hybrids (‘Canner’ su1 and ‘Butter Sweet’ sh2) under direct-seeded or transplanting. Plants were
sprayed with different BAP concentrations (0, 50, 100 or 200 mg L -1) in the pre-transplant stage or 7 days from
emergence in direct-seeded plants. Different lower case letters indicate significant differences (P < 0.05) between
control and BAP-sprayed plants, while different capital letters indicate significant differences (P < 0.05) among
sowing routines (n = 9)
BAP RGR NAR CGR
(mg L-1) (g g-1 day-1) (g cm-2 day-1) (g m-2 day-1)

‘Canner’
Direct-seeded 0 0.072aA 0.066aB 709.38bB
5 0.071aA 0.055aB 1,016.29aB
50 0.069aA 0.049aB 971.67aB
100 0.072aA 0.059aB 979.49aB
200 0.071aA 0.055aB 1,091.45aB

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Transplanted 0 0.078aA 1.496aA 1,544.56bA

5 0.079aA 1.448aA 1,865.00aA
50 0.079aA 1.397aA 1,969.39aA
100 0.079aA 1.544aA 1,873.31aA
200 0.080aA 1.511aA 2,082.29aA

‘Butter Sweet’
Direct-seeded 0 0.069aA 0.050aB 827.42cB
5 0.071aA 0.053aB 1,210.68aA
50 0.071aA 0.066aB 1,325.64aB
100 0.070aA 0.046aB 884.65cB
200 0.070aA 0.052aB 1,026.52bB

Transplanted 0 0.069aA 0.802cA 1,012.98cA

5 0.072aA 0.920bA 1,251.01bA
50 0.074aA 1.113aA 1,513.34aA
100 0.074aA 1.113aA 1,555.98aA
200 0.075aA 1.216aA 1,302.44bA

A single BAP spray significantly increased were always for transplanted plants.
LAI with higher values for direct-seeded Anyway, different maize genotypes showed
plants. On the contrary, a similar response different absolute LAI and RUE values
pattern was found for RUE and HI but, in (Table 6).
these growth parameters, the higher values
Table 6: Changes in the leaf area index (LAI), the radiation use efficiency (RUE) and the harvest index (HI) for two
corn hybrids (‘Canner’ su1 and ‘Butter Sweet’ sh2) under direct-seeded or transplanting. Plants were sprayed with
different BAP concentrations (0, 50, 100 or 200 mg L-1) in the pre-transplant stage or 7 days from emergence in direct-
seeded plants. Different lower case letters indicate significant differences (P < 0.05) between control and BAP-
sprayed plants, while different capital letters indicate significant differences (P < 0.05) among sowing routines (n = 9)
BAP LAI RUE HI
(mg L-1) (m-2 m-2) (g MJ-1 m-2 day-1)
‘Canner’
Direct-seeded 0 1.079bA 0.0120bB 0.400cB
5 1.855aA 0.0183aA 0.429aB
50 2.002aA 0.0208aA 0.414bB
100 1.673aA 0.0204aB 0.418bA
200 1.977aA 0.0192aA 0.413bA

Transplanted 0 1.031bA 0.0169bA 0.472aA

5 1.291aB 0.0194aA 0.453aA
50 1.407aB 0.0210aA 0.455aA
100 1.215aB 0.0190aA 0.406bA
200 0.967bB 0.0185aA 0.418bA

‘Butter Sweet’
Direct-seeded 0 1.642bA 0.0209bB 0.407cB
5 2.162aA 0.0257aA 0.413bB
50 1.992aA 0.0268aA 0.447aB
100 1.909aA 0.0256aA 0.451aA
200 1.962aA 0.0236aA 0.423bA

Transplanted 0 1.056bB 0.0230bA 0.434cA

5 1.360aB 0.0245aA 0.459bA
50 1.358aB 0.0252aB 0.504aA
100 1.392aB 0.0231bB 0.438cA
200 1.078bB 0.0231bA 0.434cA

Yield Components decrease yield in direct-seeded ‘Canner’

plants but even more increased yield in
When yield was analyzed on a plant basis,
transplanted ones. On the contrary, BAP-
control ‘Canner’ showed higher values in
sprayed ´Butter Sweet’ plants increased yield
transplanted plants but no significant
related to controls in both implantation
differences for ‘Butter Sweet’. A BAP spray
routines. Anyway, direct-seeded losses at the
did not change ‘Canner’ yield but increased it
germination-emergence stage (14% and 1-2%
in ‘Butter Sweet’ plants for both direct-
for direct-seeded and transplanted
seeded and transplanted ones. When yield is
respectively) (data not shown) could explain
estimated on an area basis, transplanted
higher yield in transplanted plots (Table 7).
plants began the higher values. A BAP spray

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Table 7: Changes in yield for two sweet corn hybrids (‘Canner’ su1 and ‘Butter Sweet’ sh2) under direct-seeded or
transplanting. Plants were sprayed with different BAP concentrations (0, 50, 100 or 200 mg L -1) in the pre-transplant
stage or 7 days from emergence in direct-seeded plants. Different lower case letters indicate significant differences
(P < 0.05) between control and BAP-sprayed plants, while different capital letters indicate significant differences (P
< 0.05) among sowing routines (n = 9). Ear number on an area basis (ha -1) calculated yield (ton fresh weight ha-1) as
the product of ear FW weight by plant number. Plant numbers in direct-seeded plants were decreased by
germination-emergence losses (mean 14% for both corn mutant hybrids). Mean post-transplant losses were 1% and 2%
for ‘Canner’ and ‘Butter Sweet’ respectively
BAP Yield Yield
(mg L-1) (g plant-1) (ton ha-1)
‘Canner’
Direct-seeded 0 246.17aB 16.94aB
5 212.94cB 14.65bB
50 226.24bB 15.57bB
100 208.60cB 14.35bB
200 204.50cB 14.07bB

Transplanted 0 264.23cA 20.93cA

5 394.38aA 31.03aA
50 314.07bA 24.87bA
100 274.24cA 21.72cA
200 264.73cA 20.97cA
‘Butter Sweet’
Direct-seeded 0 242.98bA 16.72bB
5 285.12aA 19.62aA
50 294.74aA 20.28aB
100 284.72aA 19.59aA
200 264.30bA 18.18bA

Transplanted 0 243.38bA 19.28cA

5 290.83aA 23.03aA
50 301.41aA 23.87aA
100 275.78aA 21.84bA
200 228.75bB 18.12cA

showed higher vascular bundles and

Anatomical Measurements
Leaf thicknesses were not significantly
different between controls direct-seeded
plants and transplanted ones. However, a
single BAP spray increased leaf thickness.
On the other hand, transplanted plants
phloem/xylem ratio than direct-seeded ones
so for controls as BAP-sprayed plants. A
single BAP spray increased phloem/xylem
ratio for the most treatments tested (Table
8).

Table 8: Changes in leaf thickness and phloem/xylem ratio for two sweet corn hybrid (‘Canner’ su1 and ‘Butter Sweet’
sh2) under direct-seeded or transplanting. Plants were sprayed with different BAP concentrations (0, 50, 100 or 200
mg L-1) in the pre-transplant stage or 7 days from emergence in direct-seeded plants. Different lower case letters
indicate significant differences (P < 0.05) between control and BAP-sprayed plants, while different capital letters
indicate significant differences (P < 0.05) among sowing routines (n = 9)
BAP Leaf thickness Phloem/Xylem
(mg L-1) (µm plant-1) Ratio
(%)
‘Canner’
Direct-seeded 0 186.67bA 253.57dB
5 246.97aA 394.87cB
50 222.73bA 417.07cB
100 200.00bA 502.33bB
200 200.61bA 584.00aA

Transplanted 0 180.30bA 320.93dA

5 195.45aB 422.37cA
50 195.45aA 543.90bA
100 192.42aA 598.21aA
200 212.12aA 603.03aA
‘Butter Sweet’
Direct-seeded 0 183.33bA 287.50cB
5 180.30bA 332.26bA
50 200.00aA 347.73bA
100 178.79bB 415.00aA
200 198.48aB 421.21aA

Transplanted 0 168.18eA 324.00cA

5 184.85dA 350.91bA
50 218.18cA 355.85bA
100 231.06bA 393.33aA
200 253.03aA 432.73aA

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Discussion used. Anyway, a change in hormonal balance

gave a yield increase between 17.3 to 49.2%
Advances in crop management coupled with
(Table 7). In agreement, Santos et al. [1]
improved stress tolerance and efficiency in
indicated that the exploitation of sweet corn
plant growth accounts for the nearly six-fold
through plant breeding, requiring the
increase in hybrid corn yield over the last 75
evaluation of the response of cultivars to
years. The transplant systems offered best
environmental conditions. The root system of
control of water availability than direct-
a maize seedling has seminal roots that
seeded, which promoted uniform both root
consist of the radicle or primary root and a
and shoot growth and resulted in higher field
variable number of lateral roots that arise
survival [23]. As transplant provides optimal
adventitiously at the base of the first
environmental conditions and water supply
internode of the stem, just above the
for seed germination, sweet corn has been
scutellar node.
transplanted experimentally in an attempt to
improve stands [17].
The seminal root initials are present in the
embryo and are the most important for early
Transplanting sweet corn is feasible because
growth and establishment, which have the
transplant technologies enable large-scale
capacity to form highly branched lateral roots
transplant production with minimal labour
[28]. A limited plug tray volume from sowing
[24]. The growing period could be
gives a vertical root restriction even under a
significantly shortened with transplantation
wide water, nutrient and oxygen supply [29].
of sweet corn plants compared to direct
In agreement with Rattin et al. [15], Figure 2
seeded [19]. Although sweet corn quality has
showed that shoot growth has limited by root
been improved from the use of mutants,
growth. Cytokinins are root factors, which
which preventing the conversion of
are transported via the xylem to the shoots,
carbohydrate to starch, yield showed a high
where they exert a major regulatory
stability around 15 ton ha-1 [25].
influence on growth; photosynthesis and
timing of senescence would be available [30].
Controls tested in our experiments (su and
sh2 mutant F1 hybrids) showed yields near to
Root cytokinins are part of the mechanism by
17 ton ha-1, which are in agreement with
which the shoot/root ratio is regulated. In an
expected yields. However, yield from
intact sweet maize plant, the living cells in
transplanted controls increased until 19.7
the root are capable of producing cytokinins,
(´Butter Sweet’ sh2) and 20.0 (‘Canner’ su)
which is transported by the xylem in the
ton ha-1 (Table7) mainly due to a decrease in
transpiration stream [31]. While El-Hendawy
germination-emergence losses, in agreement
et al. [32] found a positive relationship
with our previous reports [11-14-15-26]. Yield
between a decrease in root growth and
increases in maize have been largely
drought, Alí et al. [33] suggested that this
attributed to genetic gains (50 to 70%) and to
abiotic stress can be overlapped with the a
superior agronomic-management practices
BAP dose (50 mg L-1) similar to those applied
increasingly adopted by growers (30 to 50%)
in our experiments. In the same way, O’Hare
[27].
and Turnbull [34] showed that zeatin
ribosides (the active form of the endogenous
Data from Table 7 showed that transplant
cytokinins) increased with an increase in root
routine would involve an additional 17%
growth. On the other hand, canopy structure
yield increase over controls. On the other
defines both photosynthetic rate and
hand, a first report from our laboratory [15]
endogenous hormonal balance.
showed that a single BAP spray showed a
response related to genotype and the
Boonman and Ponds [35] found a positive
implantation routine. Data from Table 7 are
relationship between PAR and endogenous
in agreement with this previous report and
cytokinin concentrations. Foliar application
showed higher response for sh2 sweet maize
of 50 and 100 mg L-1 BAP significant increase
hybrid. Amin et al. [19] showed that a foliar
growth characters and photosynthetic
application of benzyl adenine (50 and 100 mg
pigments in maize blades at different stages
L-1) increased white maize yield, in
of growth; the higher BAP concentration the
agreement with our present results but, with
higher yield and its components [19]. In
significant differences related to maize
maize, the shoot apical meristem (SAM)
genotype tested and implantation routine
typically initiates a fixed number of leaves
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 Available online at: http://ijaas.kibanresearchpublications.com/index.php/IJAAS
during the vegetative phase of growth [36].
SAM initiates three to five more leaves, each
leaf opposite from the previous one, before
kernel maturation and the onset of
dormancy. Since plant growth is a function of
light interception, the rate of leaf area
establishment after planting is very
important to subsequent crop growth.
The leaf area developmental phase occurs
between seedling emergence and anthesis
and is dependent on leaf number, the rate at
which leaves are initiated and the expansion
rate of individual leaves. Rapid canopy
development may be particularly critical in
relatively cool, short growing season
environments. For these reasons, breeding
select genotypes with high rates of
photosynthesis per unit leaf area and with
high rates of leaf area production during the
pre-silking phase. However, self-shading
early appear at high plant populations [11].
Our results showed that control direct-seeded
plants showed higher RLA than transplanted
ones (Tables 1 and 4), which explain the
differences in total leaf area and RLAE
(Table 4). However, Sánchez-Andonova et al.
[14] showed a lower number of dead leaves
and late senescent leaves at the R3 stage in
transplanted plants, which would improve
light interception during the critical period
bracketing silking. We found a similar result
for both maize hybrids tested in field
experiments (data not shown).
The leaf area increases in some BAP-treated
plants (Table 4) would be associated to the
previously published effects of cytokinins on
SAM [37], however, because BAP increase
LAI (Table 6) for both direct-seeded and
transplants, it is not enough to equal both
total leaf area canopies. Once leaf area is
maximized, biomass accumulation depends
on RUE (at crop level) and RGR (at plant
level) because of photo assimilate
accumulation through the photosynthetic
processes.
On one hand, our results showed higher RUE
from transplants and significant increases in
both implantation routine BAP-sprayed
plants (Table 6). On the other hand, no
significant RGR differences (Tables 2 and 5)
between controls, BAP-spray plants or
implantation routines were found. RGR is the
product of LAR, the so-called ‘morphological
component’ and NAR, the ‘physiological
A Di Benedetto et. al.| Oct. 2018 | Vol.3| Issue 10 |01-14
component’. A change in dry weight
partitioning towards the development of leaf
area would be reflected in an increased LAR,
while an increased efficiency of DW fixation
would be associated with higher NAR values,
since this variable is largely the net result of
DW gain and DW losses [38]. Our results
showed that lack of RGR significant
differences could be explained by increases in
NAR and a slight decrease in LAR (Table 2),
in agreement with our previous reports in
other vegetables [39-40] including sweet
maize hybrids [14-15].
On the other hand, changes in both RGR and
NAR can be explained by root growth, at
least during the first growth weeks from
sowing (Figure 3). Rattin et al. [15] recorded
similar results between sowing and harvest
date. When the mesophyll thickness of the
leaf is increased, the maximum
photosynthetic rate increased as well; this
probably explains the strong relationship
between NAR and mesophyll thickness and
between NAR and the proportion of
intercellular spaces found by Gandolfo et al.
[41].
Light-saturated rates of photosynthesis on
leaf area basis depend not only on
photosynthetic biochemistry but also on
mesophyll structure. Because resistance to
CO2 diffusion from the sub stomatal cavity to
the stroma is substantial, it is likely that
mesophyll structure affects the
photosynthetic rate by affecting CO2 diffusion
in the leaf [42]. Although initiation routine
did not change sweet maize leaf thickness,
BAP-sprayed plants showed an increase in
leaf thickness (Tables 1 and 8) in agreement
with Gandolfo et al. [41]. Under low RGR
differences between direct-seeded and
transplants, significant yield differences only
can be explained through changes in
photosynthetic source-sink balance [43] as
was previously showed and/or by changes in
photo assimilate allocation [44].
Harvest Index (HI), the ratio of grain yield to
total plant mass, has been taken as a
measure of success in partitioning photo
assimilated to harvestable product. Harvest
Index or grain weight divided by total shoot
weight, is considered stable for maize grown
in absence of environmental constraint.
Nevertheless, modifications introduced by
genotype and stages of growth as a ratio of
grain to biomass yield, the HI of cereal crops
11
 Available online at: http://ijaas.kibanresearchpublications.com/index.php/IJAAS
can be affected by any factor which influences
the components of yield to different extents.
As was previously published [11-15], HI was
higher in transplants than in direct-seeded
plants and was increased by an early single
BAP spray (Table 6). A higher available
photo assimilate to grain fill needs
translocation from other plant sources and
involve both an endogenous signaling [45]
and a correct vascular development [46-47-
48], which includes hormone interactions
between auxins and cytokinins [49].
In agreement, it has been indicated that
cytokinins can influence the leaf structure
[41]. At the early stages of leaf development,
treatment with exogenous BAP accelerates
division of mesophyll cells, whereas at the
later stages of development, BAP treatment
activates expansion of growing cells and
those, which have just accomplished their
growth [50]. In agreement with Sánchez-
Andonova et al. [14], our results from Table 8
showed higher phloem/xylem ratio from
transplants and increasing differences in
BAP-sprayed plants.
On the other hand, pot experiment
(experiment 1) allow to collect both DW
shoots and roots and perform plant
allometries, which indicate photo assimilate
partitioning. In this way, our results (Table
3) showed early inverse photo assimilate
allocation in direct-seeded and transplants
and increasing photo assimilate partition to
stems in BAP-sprayed plants. Transplants
give smallest plants than direct-seeded ones,
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