BI O D I V ER S I T AS ISSN: 1412-033X

Volume 23, Number 8, August 2022 E-ISSN: 2085-4722

Pages: 4169-4179 DOI: 10.13057/biodiv/d230839

Secondary trait and index selection determination for maize genotype

selection in an acidic tidal swamp environment

SUWARTI1,3,, MUNIF GHULAMAHDI2,♥♥, DIDY SOPANDIE2, TRIKOESOEMANINGTYAS2,

EKO SULISTYONO2, MUHAMMAD AZRAI3
1
Graduate School of Institut Pertanian Bogor. Jl. Raya Dramaga Kampus IPB Dramaga Bogor 16680, West Java, Indonesia. Tel./fax.: +62-251-8622642,

email: suwarti_agh@apps.ipb.ac.id
2
Department of Agronomy and Horticulture, Faculty of Agriculture, Institut Pertanian Bogor. Jl. Meranti Kampus IPB Darmaga, Bogor 16680, West
Java, Indonesia. Tel.: +62-251-8629354, email: mghulamahdi@yahoo.com
3
Indonesian Cereals Research Institute. Jl. Dr. Ratulangi No. 274, Maros 90514, South Sulawesi, Indonesia

Manuscript received: 20 June 2022. Revision accepted: 28 July 2022.

Abstract. Suwarti, Ghulamahdi M, Sopandie D, Trikoesoemaningtyas, Sulistyono E, Azrai M. 2022. Secondary trait and index selection
determination for maize genotype selection in an acidic tidal swamp environment. Biodiversitas 23: 4169-4179. Determining secondary
traits as complementary in maize genotypes selection for the stressed abiotic environment is compulsory since employing the only main
character in the form of yield usually provides a biased result. This study aimed to obtain secondary traits for the maize selection in an
abiotic multi-stresses of Fe, Al, and a temporary flooding environment of acid sulfate tidal swamp land using multivariate analysis. 150
(un-repeated) maize lines and five check genotypes (five replicates) were arranged in a randomized augmented plot design on three
environmental treatments to generate 525 single plots. Three types of environments based on irrigation regulation were applied on acid
soil tidal swamp land to study the agronomic characteristics of maize genotypes in each treatment. Saturated Cultivation Technology
(SSC) was considered an optimal environment. Dryland treatment was the condition to let the irrigation only depend on rainwater.
Temporary flooding treatment was carried out at five stages at different growth periods. Several steps of analysis were applied in this
study by taking variance, and heritability was used as the basis for selections. The characters affected by treatments with medium to high
heritability were then analyzed with correlation and pathway analysis. Seven selection indexes, SSI, GMP, YSI, YI, TOL, STI, and MP,
that have a high correlation to yield by both optimum and stress environments, were applied simultaneity with the selected trait character
to screen the genotypes utilizing PCA biplot analysis. Venn analysis was used to group the genotypes based on the result of the analysis.
Indices of STI, MP, GMP, and traits of rows number per ear and the weight of 6 ears were selected as secondary characters for the
maize genotypes selection in Saturated Soil culture with temporary flooding stress. Meanwhile, STI, MP, and ear stand height were the
selected characters in the dryland of acid sulfate tidal swamp land. Based on the combination of all secondary character selection,
genotypes were grouped into five types of stress tolerance in SSC+TF and DL environments. Eight genotypes were tolerant to both
SSC+TF and DL stress, and two genotypes were susceptible to both environments.

Keywords: Acid soil, tidal swamp, heritability, pathway analysis, pyrite, temporary flooding

Abbreviations: SSC: Saturated soil culture, TF: Temporary flooding, SSC+TF: Saturated soil culture+temporary flooding, DL: Dry
land, SSI: Stress susceptibility index, GMP: Geometric mean productivity, YSI: Yield stability index, YI: Yield index, TOL:
tolerance index, STI: Stress tolerance index, MP: Mean productivity

INTRODUCTION
Improvement of maize production through extending
crop area has strict obstacles along with expanding a wide
area by converting arable soil for non-agricultural purposes
(Maulana et al. 2019). The utilization of suboptimal land
such as sulfate acid soil tidal swamp areas to increase
national maize production in Indonesia is an opportunity
since the land is widely available, about 4.3 million
hectares of tidal swampy areas can be managed as crop
cultivation area (Surahman et al. 2018). Pyrite oxidation in
the soil layer was the main restriction to developing maize
crops on the sulfate acid soil tidal swamp to manage the
land. Pyrite oxidation would trigger Fe3+ release in the soil
layer leading to a decreased pH level (Toyip et al. 2019;
Annisa et al. 2020). A low pH level in the soil triggers
aluminum ion solubility and increases acidity (Pujiwati et
al. 2015; 2016). Insoluble iron in the form of Fe3+ is found
at the aerated soil layer and alkaline pH. In contrast, when
soil is flooded and has an anaerobic condition or soil pH is
decreased, there is a reduction of Fe3+ to Fe2+, which is
soluble and dangerous to the plant (Vahedian et al. 2014;
dos Santos et al. 2017).
The toxicity of aluminum and iron to plant tissue has
been studied previously in several experiments. In addition
to damaging the plant tissues, the main risk of heavy metal
stresses of Al and Fe is decreased crop yields. Low pH soil
in acid soils increases the concentrations of Fe and Al in
the solution, leading to the precipitation of inorganic
phosphorus as iron and aluminium phosphates. The excess
condition of Fe becomes toxic to the plant as a highly
reactive Fenton catalyst. The formation of ROS such as
hydroxyl radicals (OH) and superoxide anion radicals (O2-)
is easily generated under Fe toxicity in the plant tissue
4170 B I O D I V ER S I T AS 23 (8): 4169-4179, August 2022
(Nikolic and Pavlovic 2018). Easy observed Fe excess
stress symptoms in plant tissue were the appearance of
bronzing color, starting from tip to basal of the leaf
(Nugraha et al. 2016). There are two types of Fe uptake in
the plant; strategy I for dicot and strategy II for graminea.
Maize plant Fe uptake followed chelation-based Fe uptake
by the so-called phytosiderophores strategy. Those plant
hormones are secreted to the rhizosphere to bind Fe to
become the Fe-phytosiderophores complex that enters the
cell via the root ZmYS1 (Yellow stripe1) and YSL (Yellow
Stripe Like) transporter. Chelating compounds
phytosiderophores belong to the mugeneic acids family,
released as Fe deficiency response (Bartucca et al. 2018).
Maize crop productivity is negatively affected by
aluminum presence in acid soil. The toxicity of aluminium
primarily affects plant root growth, thus, reducing the
ability to exploit the soil water and nutrients (Kopittke et
al. 2015; Matonyei et al. 2020). Furthermore, the condition
of low pH soil releases toxic forms of Al, such as
Al(OH)2+, Al(OH)2+, and Al3+ into the soil solution (Xu et
al. 2017). Plant mechanisms of aluminum tolerance could
be summarized into two types: Al exclusion and
detoxification. The exclusion mechanism is carried out to
limit Al absorbance or reduce the root’s uptake. The
detoxification mechanisms were usually conducted by Al
complexation, followed by transfer and storage of these
complexes in vacuoles (Bian et al. 2018).
Development of maize variety tolerant to several abiotic
stress troughs a breeding program for sulfate acid tidal
swamp area environment is necessary along with the
development of proper agronomic management (Sabagh et
al. 2018). The breeding program for abiotic stress tolerant
genotype usually starts with selecting tolerant genotypes
for the target environment. Grain yield is usually chosen as
the main character in the selection of maize tolerant to
abiotic stress. However, using grain yield as solely
character usually generate a biased result. Secondary traits
and index selection could complement the yielding
character as the plant tolerance to environmental stress
measurement (Fadhli et al. 2020). Multivariate analysis to
obtain the selection index value will simplify the characters
in selecting genotypes tolerant to specific environments.
Furthermore, a selection index based on the calculation of
several formulas simultaneity can be used to strengthen the
applied characters used in a selection in the abiotic stress
environment (Anshori et al. 2019). This study aimed to
obtain secondary characteristics for maize selection
genotypes in acid sulfate tidal soil under Fe, Al, and
temporary flooding stress on the Saturated Soil Culture
crops technique and dryland environment.
MATERIALS AND METHODS
Study area
The study was carried out in the sulfate acid tidal
swamp area of Karya Bhakti Village, Rantau Rasau
District, Tanjung Jabung Timur Jambi Province Indonesia,
from May to September 2019. The coordinates location of
SSC treatment was -1°10’50”S 104°09’44”E, SSC+TF was
-1°10’51”S 104°09’44”E, and DL was -1°10’50”S
104°09’37”E. The results of soil analysis at SSC, SSC+TF,
and DL locations, respectively, showed pyrite levels of 400
ppm; 100 ppm, and 100 ppm, Fe levels were 45200 ppm;
35600 ppm, and 37300 ppm (very high), Sulphur levels of
200 ppm, 500 ppm and 100 ppm, Al3+ levels 52900 ppm,
47600 ppm and 46200 ppm (very high), soil pH H2O 5.10,
5.20 and 5.2 (acidic), pH KCl 3.6, 3.7 and 3.6. Organic
matter C was 7.39 and 11.23 (very high), and 1.28 (low).
Total N 0.21%, 0.31% (moderate), and 0.09%. C/N ratio
35, 36 (very high) and 14 (medium). Available P2O was 12
(medium), 8, and 8 (low). Cation exchange capacities were
17.2 cmol.g-1 and 23.72 cmol.g-1 (medium) and 8.97
cmol.g-1 (low). Climatic data from January to June 2019
obtained from the Meteorology, Climatology and
Geophysics Agency, Muaro Jambi Climatology Station,
shows an average monthly temperature was 22.71°C -
33.65°C, relative humidity 77.22-85.77%, rainfall 42.70-
191.70 mm per month.
Procedures
Design of field experiment
The experiment was arranged in an augmented
randomized block design. Genetic materials consisted of
150 maize lines collected by the Indonesian Cereals
Research Institute and five check genotypes. Check
genotypes consisted of one commercial composite variety
(Sukmaraga) as check number 1, two elite lines (MR-14
and Nei 9008) as check number 2 and check number 3, one
commercial hybrid variety (P27) as check number 4, and
one commercial three-way cross-hybrid variety (Bima 20)
as check number 5. Maize seeds were planted at a depth of
±5 cm, the distance between rows was 70 cm, and between
plants spacing was 20 cm; to gain a population per hectare
of 71428 plants. Each plot is comprised of three rows of
plants, with the row length being 4.2 m. Each block
consisted of 30 test plots of genotypes and five plots of
check genotypes.
Arrangement of soil irrigation in each treatment
There were three soil irrigation system treatments
applied in this experiment. The treatments were Saturated
Soil Culture (SSC), SSC+Temporary Flooding, and Dry
Land (DL) environment. The saturated Soil Culture
technique manages the water level to adjust 5 cm above the
base of the 30 cm wide and 25 cm depth trench, the length
of the trench following the length of the land perpendicular
to the primary trench. This SSC technic purpose is to
manage soil consistently in redox conditions. Floodgate
was a tube applied to flow and retain water in the field at
low tide and discharge water out of the field during high
tides so that it also functions to clean the pyrite layer at the
bottom of the trench.
The SSC+Temporary Flooding (SSC+TF) treatment
was the SSC technique. Still, it allowed the water to
overflow until it reached 10 cm above the root area,
arranged periodically to allow roots in the anoxia
condition. Temporary flooding in SSC+TF period occurred
after planting (0 to 4 DAP); TF at 14-17 DAP; TF at 28-31
DAP; TF at 43-46 DAP (anthesis and tassel periods), and
 SUWARTI et al. – Secondary trait for maize selection in the tidal swamp 4171

TF at 58-61 DAP (R1/grain filling periods) (Figure 1). The
water pump was used conditionally to regulate water flow
if swamp water overflow was inadequate artificially. In
addition, a 50 cm high embankment covered with tarpaulin
was made around the land to prevent water leakage inside
and outside the experimental area. The third environment
treatment was dry land (DL), where soil watering in this
experiment was dependent on rainwater.
Tolerance index formulas
The estimated heritability value was calculated
using the heritability equation in the broad sense, which is
derived from the variance in the broad sense
. In a broad sense, heritability values are
expressed by decimal numbers ranging from 0 to 1 or
percentages; the criteria for heritability values are classified
by category; high , medium
( and low . Finally, stepwise
regression was applied to determine the most important
character with a high correlation with the yield.
Observed yield variables were used to calculate the
tolerance indices. The tolerance index used in this
experiment was; Stress susceptibility index
, with tolerant criteria if S<0,5; moderate
if the value is 0.5<S<1; Sensitive if the value of S>1,
Geometric mean productivity
stability index ; Yield index ; Tolerance
Index ; Stress Tolerance Index
; Mean Productivity .
Description: Yp: Value of certain variables in varieties
under stress; Yo: The value of a certain variable in the
varieties that were not stressed; Xp: the average value of
certain variables in all varieties under stress; Xo: The
average value of a certain variable in all varieties that are
not stressed.
Data analysis
All the collected data were analyzed over several
stages. First, data were analyzed to obtain variance with a
standard error of 5% at the initial process. Characters
significantly affected by genetic variance and have high
heritability values (>50%) were then reselected through
correlation analysis and path analysis to determine the most
important characters on the yielding character and directly
affect the yield. Seven tolerance indices were connected to
yield character by correlation analysis. The selected
characters were then analyzed using PCA (Principal
Component Analysis). Finally, Venn Diagram by Heberle
et al. (2015) was applied to separate all the tested
genotypes into tolerance groups in environmental stress
trials.
RESULTS AND DISCUSSION
Genotypic variance, phenotypic variance, and
heritability
The mean square of agronomic character data is
presented in Table 1. Genotype factor was highly affected
by yield character both in SSC+TF and SSC environments,
while the DL environment showed significantly affected.
Due to high variance, some data were being transformed at
log (x+1) or at (0.5+x)0.5. Most of the characters on SSC
treatment were highly affected by genotypic factors.
; Yield Senescence, leaf length, leaf width, anthesis days, silking
days, and weight of six stems were the characteristics that
were not affected by genotypic factors among all 23
observed traits. In the SSC+TF environment, genotypic
factors did not affect the number of leaves and leaf length.
The dry land treatment generated most characters that were
not affected by genotypic factors compared to the other
treatments. Those characters in dry land were a weight of 6
cobs, the weight of 1 cob, number of harvested plants,
number of harvested cobs, plant aspect, senescence, ear
diameter, stem diameter, and weight of 6 dry stems.

SSC
SSC + TF
DL
5-13 DAP 18-28 DAP 32-42 DAP 47-57 DAP
14-17 DAP

28-31 DAP

4 DA R1
0-4 DAP

4 DAT

= temporary flooding periode

= saturated soil culture technic periode
= watering depend on rain water

Figure 1. Irrigation treatment scheme on SSC technique and the temporary flooding period in the experimental environment
 4172 B I O D I V ER S I T AS 23 (8): 4169-4179, August 2022

Table 1. Mean square of genotype factor at each character on environment treatments of SSC, SSC+TF, and DL

Source of variance MSg SSC CV MSg SSC+TF CV MSg DL CV

Yield 0.15** 30.59t2 0.03** 15.68t2 0.10* 22.01t2
Weight 6 cobs 0.07** 8.88t2 0.99* 11.73t2 0.01ns 12.71t2
Weight of 1 cob 0.10** 22.85 0.12** 22.19 0.02ns 2.71t2
Number of harvested plants 0.07** 10.59t1 0.99* 32.78t2 0.01ns 11.95t1
Number of harvested cobs 0.10** 13.55t1 0.12** 17.76t1 0.02ns 16.29t1
Kernels/Cob weight Ratio 7.18** 13.07 21.06** 7.89 0.04** 12.80t2
Weight of 1000 seed 2920.57* 23.37 0.02* 5.36t1 0.01** 3.97t1
Plant aspect 0.43** 16.38 0.02** 14.96t1 0.00ns 14.85t1
Senescence 0.02ns 22.98 0.03** 11.75t1 0.01ns 12.86t1
Ear length 9.03** 16.82 6.37** 21.22 0.01* 8.81t1
Ear diameter 0.43** 10.69 0.40* 21.06 0.25ns 17.92
Number of rows per cob 5.03** 10.69 6.14** 21.06 2.74** 17.92
Number of seeds per row 40.86** 23.07 0.07** 8.07t1 0.02** 8.77t1
Number of leaves 0.01* 11.93t1 1.82ns 24.68 2.57** 13.92
Height of cob 125.10** 19.59 0.03* 11.62t1 237.59** 20.01
Plant height 524.82** 14.47 468.22** 19.93 600.18** 13.16
Leaf length 101.09ns 21.31 0.01ns 6.86 84.13* 11.04
Leaf width 0.01ns 11.36t1 0.01* 12.50t1 1.11* 13.18
Stem diameter 6.51* 18.21 6.28* 22.87 0.01ns 9.91t1
SPAD 39.80** 12.29 43.49* 20.14 23.20* 11.04
Anthesis day 11.54ns 7.21 13.76** 4.91 9.21** 2.50
Silking day 16.03ns 6.58 19.12** 4.99 9.88** 2.54
Weight of 6 dry stem 0.01ns 10.91t2 0.01** 5.65t2 0.01ns 10.18t2
Note: **significant effect on 1% level, *significant effect on 5% level, ns non-significant, t1 transformed at ; t2 transformed at
; CV: Coefficient of Variance; MS: Mean square of genotypes

Table 2. Agronomic characters of 150 maize lines under three environments types on acid soil tidal swamp, Jambi 2019

SSC SSC+TF DL
Agronomic Traits
σ2p σ2g h2 σ2p σ2g h2 σ2p σ2g h2
Yield 0.21 0.35 59.81 -0.12 0.61 -19.23 0.10 0.49 20.40
Weight 6 cobs 14.64 14.65 99.95 8.51 8.52 99.96 2.52 2.54 99.18
Weight of 1 cob 17.62 17.62 100.00 6.98 6.98 100.00 5.57 5.57 99.99
Number of harvested plants 9.32 35.95 25.92 3.86 27.14 14.21 0.19 11.85 1.58
Number of harvested cobs 10.57 45.83 23.06 4.92 15.26 32.22 -0.92 31.52 -2.92
Kernels/Cob Weight Ratio -34.04 150.50 -22.62 4.21 4.23 99.52 0.07 0.12 61.57
Weight of 1000 seed 218.23 2168.13 10.07 603.25 3458.62 17.44 228.26 2074.85 11.00
Plant aspect 0.05 0.21 24.80 0.07 0.40 18.33 0.02 0.22 8.57
Senescence 0.45 2.51 17.95 0.55 2.07 26.39 0.01 2.30 0.43
Ear length 0.87 5.20 16.76 0.96 2.54 37.80 0.41 3.37 12.03
Ear diameter 0.07 0.13 56.83 0.05 0.20 23.86 -0.48 2.40 -20.04
Number of Rows per ear 0.67 2.37 28.37 1.12 1.64 68.55 0.59 2.33 25.26
Seeds number per Row 4.64 22.43 20.70 6.34 14.14 44.83 2.68 12.87 20.83
Number of Leaves 0.46 2.03 22.83 0.18 1.11 16.14 0.37 1.15 31.94
Height of Cobs 20.86 41.66 50.08 10.23 52.65 19.42 0.22 0.41 52.49
Plant height 86.56 178.59 48.46 50.78 130.67 38.87 93.22 227.32 41.01
Leaf length 6.71 74.26 9.03 0.00 0.01 4.12 9.11 53.10 17.16
Leaf width 0.11 1.28 8.74 0.10 0.85 11.27 0.14 0.57 24.13
Stem diameter 0.79 3.34 23.80 0.70 3.47 20.28 -0.22 9.21 -2.41
SPAD 5.62 17.31 32.49 5.02 23.39 21.48 2.56 12.88 19.87
Anthesis day 0.54 9.39 5.73 1.85 6.37 28.98 1.65 2.61 63.08
Silking day 1.60 9.61 16.69 2.78 8.02 34.62 1.76 2.84 62.09
Weight of 6 dry stems 0.00 0.00 33.50 0.00 0.01 47.01 0.00 0.01 7.81
Notes: σ2p: Phenotypic variance, σ2g: Genotypic variance, h2: Heritability, SSC: Saturated soil culture, DL: Dry land, SSC+TF:
Saturated soil culture+temporary flooding

Other characters that were highly significantly affected the SSC+TF environment were the weight of 6 ears, the
by genotype and had moderate to high heritability values in weight of 1 ear, number of harvested cobs, the ratio of
 SUWARTI et al. – Secondary trait for maize selection in the tidal swamp
shelled grain to cob, senescence score, ear diameter, ear
length, rows number of kernels, number of seeds per row,
plant height at 10 WAP, stem diameter at 8 WAP, SPAD
value, anthesis day, silking days, and weight of 6 dry stems
(Table 1 and Table 2).
A summary of the phenotypic variance, genotypic variance,
and heritability observed characters are shown in Table 2.
Characters with high heritability values can be used as
selection criteria to increase the effectiveness of selection.
Maize plants have different responses to the environment,
so the characters utilized to increase the effectiveness of
selection also vary, depending on the selection environment.
The highest heritability on yield character was obtained in
the SSC environment (scored 59.81%), followed by the DL
environment, which scored 20.40% and was categorized as
a medium category, and at the lowest was the SSC+TF
environment, it had a negative heritability value (-19.23)
which means there was no genetic progress and can be
considered as 0 (zero). Low heritability indicates that the
character is more affected by the environment rather than
genotypic nature (Wening et al. 2020).
A high broad sense heritability value in the SSC
environment (>50%) was obtained on the weight of 6 ears,
the weight of 1 ear, ear diameter, and ear height position.
Medium heritability values (between 20%-50%) were
obtained on the number of harvested plants, the number of
harvested cobs, plant aspect, seeds number per row, seeds
rows number, plant height, number of leaves, stem
diameter, SPAD value, and weight of 6 dry stems. High
heritability in the SSC+TF environment was shown at
characters of the weight of 6 cobs, the weight of 1 ear
kernels/cob ratio, and the number of rows per ear. The
number of harvested cobs, senescence, ear length, ear
diameter, seeds number per row, plant height, stem
diameter, SPAD value, anthesis day, silking day, and
weight of 6 dry stems were categorized as a medium
heritability. The high-category heritability values of
agronomic characters in DL environment treatment were
obtained on the weight of 6 cobs, the ratio of the kernel to
Table 3. The most important trait correlation based on stepwise regression analysis of SSC tidal swamp treatment
Characters
YieldSSC
Weight of 6 ears (X6e)
Weight of 1 ear (X1e)
Height of ear stand (XeSt)
Notes: **significant at 1% level, *significant at 5% level
Table 4. The most important trait correlation based on stepwise regression analysis on SSC+GS tidal swamp treatment
Characters
YieldSSC+GS
Weight of 6 ears (X6E)
Weight of 1 ear (X1E)
Kernels/cob weight Ratio (XKpC)
Number of rows per ear (XNR)
Notes: **significant at 1% level, *significant at 5% level
4173
cob, anthesis days, and silking days. Medium heritability
values were obtained on the number of rows per ear, seeds
number per row, number of leaves, plant height, and leaf
width. The heritability of each maize character is
influenced by additive or non-additive gene action and
expressed at the phenotype performance (Sesay et al.
2016).
Correlation of important character to the yield
A stepwise regression analysis was conducted to
determine the essential/dominant characters related to the
main character (Ayvat and Omeroglu 2022). The generated
model by stepwise analysis was considered more precise in
predicting the results based on the resulting model than the
correlation involving all observation variables (Andayani et
al. 2016). This study found that, in the SSC environment
weight of 6 ears (X6e), the weight of 1 ear (X1e), and the
height of the ear stand (XeSt) were the dominant
characteristics that affect maize grain yield (Table 3).
Therefore, the SSC treatment in acid soil tidal swamp is
considered an optimum crop technic in the tidal swamp.
The stepwise regression correlation analysis on all
maize characters observed in the SSC+TF environment
showed that four traits had a significant effect, with an R2
value of 70.00% and a model F value of 87.52 significantly
different. These characters were the weight of 6 ears
(X6E), the weight of 1 ear (X1E), kernels per cob weight
ratio (XKpC), and the number of rows per ear (XNR)
(Table 4). The equation model formed from this analysis
were Y(SSC+TF)= -0.09 + 1.63 X6E + 0.71 X1E - 0.01
XKpC + 0.01 XNR. Meanwhile, the stepwise regression
correlation results on dry land treatment showed three
characters (weight of 6 ears, the height of ear stand, and
anthesis day) that were dominant to affected yield.
Equation model formed in dry land treatment was Yield
DL= 4.26 - 0.30 XKpC + 0.02 XESt - 0.07 XAD. The R2
value explained from this model is 40.81%, with a model F
value was 34.7, which is highly significant (Table 5).
YieldSSC (X6e) (X1e)
1
0.75** 1
0.44** 0.34** 1
0.51** 0.53** 0.26**
YieldSSC+GS (X6e) (X1e) (XKpC)
1
0.82** 1
0.1ns 0.05ns 1
-0.23** -0.17* 0.31** 1
0.52** 0.47** 0.17* -0.2*
 4174 B I O D I V ER S I T AS 23 (8): 4169-4179, August 2022
Table 5. The most important trait correlation based on stepwise regression analysis on DL tidal swamp treatment
Characters
YieldDL
Kernels/cob weight Ratio (XKpC)
Height of ear stand (XeSt)
Anthesis day (XAD)
Notes: **significant at 1% level, *significant at 5% level
Each environment shows a different important selected
character with a high correlation to yield. Two characters
selected in optimum treatment (SSC) were similar to those
found in SSC+TF. The result was understandable though
both environments lay in the same area. Height ear stands
as the dominant character that affected grain yield in stress
DL environment was found in SSC optimum environment
likewise. Kernels per cob ratio in SSC+TF is also shown as
the dominant character in the DL environment. The high
correlation between variables shows a close relationship
and can be an indirect selection character for quantitative
traits (Hamawaki et al. 2017).
Direct and indirect effects of agronomic traits on the
yield characters
The pathway analysis results on the Saturated Water
Cultivation (SSC) treatment showed a residual value of
0.39, meaning that the path model could explain 61% of the
variance. The most considerable direct effect was obtained
on the weight of 6 ears, which has a direct effect coefficient
of 0.61, weight of 1 ear get 0.20, and the height of ear stand
get 0.13 score the direct effect. The pathway analysis of
characters in the SSC+TF environment showed a residue of
0.30, meaning that 70% of the yield characters could be
explained by the four characters used in the test. The
character weight of 6 ears (0.73), the weight of 1 ear (0.7),
and the number of rows per ear (0.14) showed a positive
direct effect coefficient on the yield. However, the kernels
per cob ratio have a negative direct effect (-10). Characters
of kernels per cob ratio and anthesis day had a negative
direct effect on grain yield, with values of -0.25 and -0.26,
respectively.
In contrast, the height of the ear stand has a positive
direct effect (0.47) in the DL environment. The residual
effect in this environment stress treatment was 0.59,
meaning that 41% of the path model could explain the yield
character. The positive direct characters to yield in path
analysis indicated that increasing maize grain yield was
supported by increasing the character value in those
environments. A negative direct effect demonstrates that
the yield response is more influenced by the indirect effect
(Putri and Ashari 2019). Path analysis enables breeders to
Table 6. Agronomic characters with direct effect and indirect effect on maize grain yield in SSC treatment in acid soil tidal swamp
Direct effect
Weight of 6 ears (X6e) 0.61
Weight of 1 ear (X1e) 0.20
Height of ear stand (XESt) 0.13
YieldDL (XKpC) (XeSt)
1
-0.31** 1
0.53** -0.12ns 1
-0.32** 0.03ns -0.11Ns
improve understanding of the relationships between several
traits by revealing the correlation coefficients into direct
and indirect effects (Machado et al. 2017).
Correlation of selection index to yield characters
A total of seven stress tolerance indices (GMP, YSI,
SSI, YI, TOL, STI, and MP) were used to determine the
appropriate index for maize genotype selection in tidal
areas under the stress of Al, Fe, and temporary flooding
(Table 9). The correlation coefficient of tolerance indices
with productivity in SSC+TF treatment shows a highly
significant and high value based on the Pearson correlation
on the GMP (r=0.73), STI (r=0.63), and MP (r=0.86)
indices; all those indices had an exact high correlation to
yield on the optimum treatment (SSC) that has correlation
indices of r=0.89, r=0.85, and r=0.82, respectively.
Index selection that has a high correlation with grain
yield on DL environment stress was STI (r=0.81) and MP
(r=0.77), the correlation of optimum SSC treatment on
those selection indices was STI (r=0.76) and MP (r=0.98).
The correlation coefficient score is useful to explain the
relationship within traits in terms of degree, characteristic,
and direction of selection to apply (Sayo et al. 2017).
Appropriate secondary trait and index selection for
genotype selection in tidal swamp land using PCA
analysis
PCA biplot analysis was employed to explain the
relationship between tolerance index and productivity of
stress environment in this experiment. PCA analysis is
widely used as it simplifies large amounts of data into easy-
to-understand visualizations (Metsalu and Vilo 2015).
Furthermore, the data analyzed was standardized
beforehand—data closer to the biplot axis indicates more
stable characters than those more distant from the axis. In
the previous study, Principal Component Analysis is useful
to predict the relationship between germplasm to the
environment and cassava sample according to several of its
enzymatic activity. The analysis work with the approach of
multiple correlations (Wang et al. 2014; Mursyidin and
Khairullah 2020).
Indirect effect
(X6e) (X1e) (XeSt) Residual
0.07 0.07 0.39
0.21 0.03 0.39
0.32 0.05 0.39
 SUWARTI et al. – Secondary trait for maize selection in the tidal swamp 4175

Table 7. Agronomic characters with direct effect and indirect effect to maize grain yield of SSC+TF treatment in acid soil tidal swamp

Indirect effect
Direct effect
(X6E) (X1E) (XKpC) (XNR) Residual
Weight of 6 ears (X6e) 0.73 0.00 0.02 0.07 0.30
Weight of 1 ear (X1e) 0.07 0.04 -0.03 0.02 0.30
Kernels/cob weight Ratio (XKpC) -0.10 -0.12 0.02 -0.03 0.30
Number of rows per ear (XNR) 0.14 0.34 0.01 0.02 0.30

Table 8. Agronomic characters with direct effect and indirect effect on maize grain yield of DL treatment in acid soil tidal swamp

Indirect effect
Direct effect
(XKpC) (XESt) (XAD) Residual
Kernels/cob weight ratio (XKpC) -0.25 -0.06 -0.01 0.59
Height of ear stand (XESt) 0.47 0.03 0.03 0.59
Anthesis day (XAD) -0.26 -0.01 -0.05 0.59

Table 9. Pearson’s correlation of tolerance indices and grain yield characters on the SSC + TF treatment acid soil tidal swamp

Characters Yp Ys_gs GMP YSI SSI YI TOL STI MP

Yp 1
Ys_dl 0.42 ** 1
GMP 0.89 ** 0.73 ** 1
YSI -0.2 ** 0.07 -0.2 ** 1
SSI -0.2 ** 0.07 -0.2 ** 1 ** 1
YI 0.42 ** 1 ** 0.73 ** 0.07 0.07 1
TOL 0.46 ** -0.6 ** 0.06 -0.3 ** -0.3 ** -0.6 ** 1
STI 0.85 ** 0.63 ** 0.9 ** -0.1 -0.1 0.63 ** 0.13 1
MP 0.82 ** 0.86 ** 0.96 ** -0.1 -0.1 0.86 ** -0.1 0.87 ** 1
Notes: ** significant at 1% level, Yp: The yield on SSC treatment, Ys_gs: The yield on SSC+GS treatment; GMP: Geometric mean
productivity; YSI: Yield stability index; SSI: Susceptibility index; YI: Yield index; TOL: Tolerance index; STI: Stress tolerance index;
MP: Mean productivity

Table 10. Pearson’s correlation of tolerance indices and grain yield characters on the DL treatment acid soil tidal swamp

Characters Yp Ys_dl GMP YSI SSI YI TOL STI MP

Yp 1
Ys_gs 0.62** 1
GMP 0.87** 0.9** 1
YSI -0.2* 0.16 -0.1 1
SSI 0.18* -0.2* 0.07 -1** 1
YI 0.62** 1** 0.9** 0.16 -0.2 1
TOL 0.95** 0.33** 0.67** -0.3** 0.28** 0.33** 1
STI 0.76** 0.81** 0.89** -0 0.04 0.81** 0.58** 1
MP 0.98** 0.77** 0.95** -0.1 0.1 0.77** 0.85** 0.84** 1
Notes: ** significant at 1% level, Yp: The yield on SSC treatment, Ys_dl: Yield in dry land treatment; GMP: Geometric mean
productivity; YSI: Yield stability index; SSI: Susceptibility index; YI: Yield index; TOL: Tolerance index; STI: Stress tolerance index;
MP: Mean productivity.

The results of the biplot analysis show that the STI,
MP, and XESt indices vectors coincide with the yield
character in the SSC+TF environments (Figure 2). In the
DL environment, the STI, GMP, MP, XNR, and X6E were
at PC 1. The value of PC1 on SSC+TF was able to explain
52.40% of the variance and on PC2 explained 16.30% of
the variance. PC1 in the DL environment explained 57.10%
variance on PC1 and 16.60% variance on PC2. Data in
Table 9 show that the highest PC coefficient was obtained
on PC 1 in SSC+TF and DL environment treatments. As
the main character, yield is given 3 points in the model to
optimize the selection. PC1 was the fit model to explain the
character factor based on the eigenvalue. Hence models to
explain index selection in both SSC and DL environments
were:
ISSC+TF = 3*0.94 Yield+0.98GMP+0.89STI+0.90MP+
0.83X6E+ 0.59XNR
IDL = 3*0.90Yield+0.84STI+0.96MP+0.65XESt
4176 B I O D I V ER S I T AS 23 (8): 4169-4179, August 2022

Figure 1. Biplot principal components 1 and 2 for all tolerance indices with yield in dryland (DL) and saturated water+temporary
flooding (SS+TF) cultivation treatments

Table 11. PCA eigenvalues for each dimension in SSC + TF treatment and treatment (DL) dry land in tidal land

PC 1 PC 2 PC 3 PC 4 PC 5
SSC+TF treatment
Yield SSC+TF 0.94 -0.05 -0.02 0.07 -0.13
Geometric Mean Productivity (GMP) 0.98 0.01 -0.11 -0.04 0.12
Stress Tolerance Index (STI) 0.89 0.10 -0.29 -0.13 0.07
Mean Productivity (MP) 0.90 0.08 -0.19 -0.12 0.25
Weight of 6 Ear (X6E) 0.83 -0.07 0.02 0.29 -0.43
Number of Rows Per Ear (XNR) 0.59 -0.03 0.73 0.25 0.23
Variance percent 57.07 16.57 10.44 7.35 4.80
Cumulative variance percent 57.07 73.64 84.08 91.44 96.24
Eigenvalue 4.57 1.33 0.84 0.59 0.38
DL treatment
Yield DL 0.90 -0.02 -0.02 -0.03 0.42
Stress Tolerance Index (STI) 0.84 0.10 0.13 0.43 -0.28
Mean Productivity (MP) 0.96 -0.02 0.09 0.19 0.03
Height of Ear Stand (XESt) 0.65 -0.05 0.41 -0.62 -0.16
Variance percent 52.35 16.33 14.50 11.07 4.98
Cumulative variance percent 52.35 68.69 83.19 94.26 99.24
Eigenvalue 3.14 0.98 0.87 0.66 0.30

Tolerance grouping based on multivariate analysis
The grouping of genotype tolerances was based on the
index value obtained from the calculation following the
index selection model. The genotypes were then classified
into three categories, i.e., tolerant, moderately tolerant, and
susceptible. The tolerant category score was Xi>X +
SE*ttable; the susceptible category score was Xi<X -
SE*ttable, while the moderately-tolerant category score was
between these two values. Finally, a Venn diagram was
applied to describe the tolerance category in each
environment. The results of the tolerance grouping in the
two experimental environments classified the genotypes
into five categories (Figure 3A). Eight genotypes were
tolerant in both SSC+TF and DL stress environments, and
47 genotypes were classified tolerant in the DL
environment and moderately tolerant in SSC+TF
environments. Forty-seven genotypes were moderately
tolerant in both SSC+TF and DL environments, and 51
genotypes were moderately tolerant at SSC+TF and
susceptible in the DL environment. Two genotypes were
susceptible to SSC+TF and DL environments (Figure 3B).
 SUWARTI et al. – Secondary trait for maize selection in the tidal swamp 4177

A B
Figure 2. A. Venn diagram summarizing adaptation of maize lines on SSC+TF and B. DL of acid soil tidal swamp environments based
on selected indices

Table 12. Venn diagram of maize genotype grouping based on the tolerance in SSC+TF and DL stress acid soil tidal swamp environment

Mean yield Number of

Group of tolerance Genotypes
SSC+TF DL genotypes
[Tolerant DL] and Sukmaraga, P27, BIMA20URI, M-35-1-B, MPop11_01, MPop18_02, 0.93 2.14 8
[Tolerant TF] MPop27_08, MPop28_01
[Tolerant DL] and MPop27_01, MPop28_02, MPop28_03, C44-3-1, C27-4-B, 0.20 1.90 47
[Moderate TF] MPop28_04, E57-B, MGOLD, MPop28_05, DKL9001, M1-3-B, M-
10-1-B, DYW7-B, DYW34-B, MPop03_06, MPop02_02, MPop02_04,
MPop02_06, MPop11_06, MPop18_01, MPop26_01, MPop26_03,
MPop27_04, MPop05_01, MPop05_02, MPop05_04, MPop05_06,
MPop24_01, MPop24_02, MPop24_03, MPop27_05, MPop24_06,
MPop24_08, MPop24_09, MPop24_10, MPop21_03, MPop21_04,
MPop21_08, MPop21_09, MPop27_06, PAC2244-2-3-1-3-9-B-B,
PAC2244-2-3-1-4-11B-B, PAC9996-2-4-1-2-11B-B, PAC2247-1-1-1-
1-8B-B, PAC2244-1-6-1-1-9B-B, PAC2245-3-1-1-2-8B-B, C17-1-B

[Medium DL] and NEI9008, C39-1-B, C31-1-B, C25-1-B, E31B, E90B, M42-1-B, 0.14 1.11 47
[Moderate TF] DYW15-B, DYW24-B, CLYN231, AL46, MAL03, MR15, B11209,
G1026-12, G1044-14, MPop03_01, CY-7, MPop03_09, MPop03_10,
MPop10_01, MPop10_07, MPop10_08, MPop02_03, MPop15_01,
MPop11_02, MPop11_03, MPop11_08, MPop05_03, MPop24_04,
MPop24_05, MPop24_07, PAC99912-2-4-1-4-10-B-B, PAC2247-1-1-
1-4-12B-B, PAC2247-1-1-1-2-9BB, MPop27_07, PAC2244-1-6-1-3-
7B-B, PAC2244-2-3-1-2-9B-B, PAC2246-2-4-1-2-10B-B, PAC2245-
3-1-1-4-5B-B, PAC9999-1-2-1-4-9B-B, PAC2247-1-1-1-2-8B-B, C40-
2-B, C61-1-B, C32-5-B, C58-1-B, C19-4-B

[Moderate TF] and MR-14, C93-1-B, C63-1-B, C48-5-B, C46-2-B, C72-1-B, C47-3-B, 0.08 0.71 51
[Susceptible DL] C73-2-B, C6-3-B, C62-2-B, C37-1-B, C11-1-B, C60B-B, C16B, C41-
1-B, E100B, E93B, E52B, E102B, WYW27B, WYW18B, WYW11B,
M-6-1-B, DYW-17-B, MPop28-06, MPop03_03, MPop27_02,
MPop10_02, Pop10_03, MPop27_03, MPop26_02, MPop05_05,
MPop21_10, G104414, G2013631, PAC2245-3-1-1-4-4B-B,
PAC9999-1-2-1-1-10BB, PAC2243-1-5-1-1-9B-B, PAC99912-3-1-1-
3-9B-B, PAC9997-1-1-1-3-8B-B, PAC99912-2-4-1-3-8B-B,
PAC99912-2-4-1-3-7B-B, PAC2241-2-3-1-3-4B-B, PAC2246-2-4-1-1-
7B-B, PAC99912-2-4-1-1-7B-B, C54-1-B, C43-1-B, C56-1-B, C12-1-
B, C42-3-B, C10-2-B

[Susceptible DL] and AMB30, AMB20 0.00 0.78 2

[Susceptible TF]
4178 B I O D I V ER S I T AS 23 (8): 4169-4179, August 2022
Sukmaraga, P27, and BIMA20URI varieties belong to
the tolerant group in two stress treatment environments of
SSC+TF and DL. Those check varieties belonged to
heterozygous genotypes of open-pollinated, hybrid, and
three-way cross variety, respectively. Maize tolerance to
abiotic stress was commonly controlled by additive genes,
which is why heterozygotes are more desirable to survival
in an abiotic stress environment. Genotypes of M-35-1-B,
MPop11_01, MPop18_02, MPop27_08, MPop28_01.
Genotypes of AMB30 and AMB20 were susceptible in
both SSC+TF and DL treatment environments.
In conclusion, based on multivariate analysis, secondary
characters/selection indices employed for selecting maize
genotype in acid tidal swamp land using saturated soil
culture technique with temporary flooding (SSC+TF) were
STI, MP, GMP, rows number per ear, and weight of 6 ears.
STI, MP, and ear stand height characters were the most
appropriate for selecting maize genotype on the acidic tidal
swamp land on dryland stress.
ACKNOWLEDGEMENTS
The authors thank the Research and Development
Agency of the Ministry of Agriculture through the DIPA
Learning Officer for the financial support for this research
activity. We also thank PT FKS Multi Agro for allowing
this research to be carried out on plant development land in
the tidal land of Tanjung Jabung Timur, Jambi.
REFERENCES
Andayani NN, Aqil M, Syuryawati. 2016. The application of stepwise
regression model in the determination of white corn yields.
Informatika Pertanian 25 (1): 21-28. DOI:
10.21082/ip.v25n1.2016.p21-28. [Indonesian]
Annisa W, Husnain, Djufry F. 2020. Effect of reactive phosphate rock to
corn on acid sulphate soil in South Kalimantan. IOP Conf Ser Earth
Environ Sci 484: 012093. DOI: 10.1088/1755-1315/484/1/012093.
Anshori MF, Purwoko BS, Dewi IS, Ardie SW, Suwarno WB. 2019.
Selection index based on multivariate analysis for selecting doubled-
haploid rice lines in lowland saline prone area. Sabrao J 51 (2): 161-
174.
Ayvat P, Omeroglu SK. 2022. Mortality estimation using APACHE and
CT scores with stepwise linear regression method in COVID-19
intensive care unit: A retrospective study. Clin Imaging 8: 4-8. DOI:
10.1016/j.clinimag.2022.04.017.
Bartucca ML, Di Michele A, Del Buono D. 2018. Interference of three
herbicides on iron acquisition in maize plants. Chemosphere 206:
424-431. DOI: 10.1016/j.chemosphere.2018.05.040.
Bian H, Pang J, Huang C, Zhang W. 2018. The response of transitional
pedogenic characteristics of loess in the Yunxian Basin to abrupt
climatic events in the northern subtropics since the Last Glacial
Maximum. Catena 171: 166-175. DOI: 10.1016/j.catena.2018.06.016.
Fadhli N, Farid M, Rafiuddin, Efendi R, Azrai M, Anshori MF. 2020.
Multivariate analysis to determine secondary characters in selecting
adaptive hybrid corn lines under drought stress. Biodiversitas 21 (8):
3617-3624. DOI: 10.13057/biodiv/d210826.
Heberle H, Meirelles VG, da Silva FR, Telles GP, Minghim R. 2015.
InteractiVenn: A web-based tool for the analysis of sets through Venn
diagrams. BMC Bioinformatics 16: 169. DOI: 10.1186/s12859-015-
0611-3.
Kopittke PM, Moore KL, Lombi E, Gianoncelli A, Ferguson BJ, Blamey
FPC, Menzies NW, Nicholson TM, McKenna BA, Wang P,
Gresshoff PM, Kourousias G, Webb RI, Green K, Tollenaere A.
2015. Identification of the primary lesion of toxic aluminum in plant
roots. Plant Physiol 167 (4): 1402-1411. DOI: 10.1104/pp.114.253229.
Machado BQV, Nogueira APO, Hamawaki OT, Rezende GF, Jorge GL,
Silveira IC, Medeiros LA, Hamawaki RL, Hamawaki CDL. 2017.
Phenotypic and genotypic correlations between soybean agronomic
traits and path analysis. Genet Mol Res 16 (2): 2-11. DOI:
10.4238/gmr16029696.
Matonyei TK, Barros BA, Guimaraes RGN, Ouma EO, Cheprot RK,
Apolinário LC, Ligeyo DO, Costa MBR, Were BA, Kisinyo PO,
Onkware AO, Noda RW. Gudu SO, Magalhaes JV, Guimaraes CT.
2020. Aluminum tolerance mechanisms in Kenyan maize germplasm
are independent from the citrate transporter ZmMATE1. Sci Rep 10
(1): 1-10. DOI: 10.1038/s41598-020-64107-z.
Maulana AI, Ghulamahdi M, Lubis I. 2019. Response of corn varieties
under saturated soil culture and temporary flooding on tidal swamp. J
Trop Crop Sci 6 (1): 41-49. DOI: 10.1016/j.proenv.2016.03.060.
Metsalu T, Vilo J. 2015. ClustVis: A web tool for visualizing clustering of
multivariate data using Principal Component Analysis and heatmap.
Nucleic Acids Res 43 (W1): W566-W570. DOI: 10.1093/nar/gkv468.
Mursyidin DH, Khairullah I. 2020. Genetic evaluation of tidal swamp rice
from South Kalimantan, Indonesia based on the agro-morphological
markers. Biodiversitas 21 (10): 4795-4803. DOI:
10.13057/biodiv/d211045.
Nikolic M, and Pavlovic J. 2018. Plant responses to iron deficiency and
toxicity and iron use efficiency in plants. Elsevier Inc. Amsterdam
Nugraha Y, Ardie SW, Rumanti IA, Suwarno S, Ghulammahdi M,
Aswidinnoor H. 2016. Responses of selected Indonesian rice varieties
under excess iron condition in media culture at seedling stage.
Penelitian Pertanian Tanaman Pangan 35(3): 181. DOI:
10.21082/jpptp.v35n3.2016.p181-190.
Pujiwati H, Ghulamahdi M, Yahya S, Aziz SA, Haridjaja O. 2015. The
application of peaty mineral soil water in improving the adaptability
of black soybean toward aluminium stress on tidal mineral soil with
saturated water culture. Agrivita 37 (3): 284-289. DOI:
10.17503/Agrivita-2015-37-3-p284-289.
Pujiwati H, Ghulamahdi M, Yahya S, Aziz SA, and Haridjaja O. 2016.
Productivity of three soybean genotypes with different water and
water depth on various land conditions in tidal swamp. Jurnal
Agronomi Indonesia 44 (3): 248-254. DOI: 10.24831/jai.v44i3.12926.
[Indonesian]
Sabagh, A El, Hossain A, Barutçular C, Khaled AA, Fahad S, Anjorin FB,
Islam MS, Ratnasekera D, Kizilgeçi F, Yadav GS, Yıldırım M,
Konuskan O. 2018. Sustainable maize (Zea Mays L.) production
under drought stress by understanding its adverse effect, survival
mechanism and drought tolerance indices. J Exp Biol Agric Sci 6 (2):
282-295. DOI: 10.18006/2018.6(2).282.295.
dos Santos RS, De Araujo AT, Pegoraro C, de Oliveira AC. 2017. Dealing
with iron metabolism in rice: from breeding for stress tolerance to
biofortification. Genet Mol Biol 40 (1): 312-325. DOI: 10.1590/1678-
4685-gmb-2016-0036.
Sayo S, David KO, Omolayo JA, Silvestro M, Lawrence SF, Ayoola OO,
Adebiyi, OO. 2017. Correlation and path coefficient analysis of top-
cross and three-way cross hybrid maize populations. Afr J Agric Res
12 (10): 780-789. DOI: 10.5897/ajar2016.11997.
Sesay S, Ojo DK, Ariyo OJ, Meseka S. 2016. Genetic variability,
heritability and genetic advance studies in top-cross and three-way
cross maize (Zea mays L.) hybrids. Maydica 61 (2): 1-7.
Surahman M., Ghulamahdi M, Murdianto, Prastowo, Sutrisno, Sapei A,
Purwanto YJ, Suharnoto Y, Wijaya H, Suwarto, Sehabudin U,
Budiman C, Nindita A, Furqoni H, Ritonga A W, Zamzani A,
Amarilis A, Rau, MI. 2018. Five steps toward the Indonesian soybean
self-sufficiency. IOP Conf Ser Earth Environ Sci 196: 012044. DOI:
10.1088/1755-1315/196/1/012044.
Toyip, Ghulamahdi M, Sopandie D, Aziz SA, Sutandi A, Purwanto MYJ.
2019. Physiological responses of four soybean varieties and their
effect to the yield in several saturated soil culture modification.
Biodiversitas 20 (8): 2266-2272. DOI: 10.13057/biodiv/d200822.
Vahedian A, Aghdaei SA, Mahini S. 2014. Acid sulphate soil interaction
with groundwater: A remediation case study in East Trinity. APCBEE
Procedia 9: 274-279. DOI: 10.1016/j.apcbee.2014.01.049.
Wang N, Huang Q, Sun J, Yan S, Ding C, Mei X, Li D, Zeng X, Su X,
Shen Y. 2014. Shade tolerance plays an important role in biomass
production of different poplar genotypes in a high-density plantation.
For Ecol Manage 331: 40-49. DOI: 10.1016/j.foreco.2014.06.046.
Wening RH, Purwoko BS, Suwarno WB, Rumanti IA, Khumaida DN.
2020. Simultaneous selection of leaf drying and yield traits on rice
 SUWARTI et al. – Secondary trait for maize selection in the tidal swamp 4179

lines. Jurnal Agronomi Indonesia 47 (3): 232-239. DOI: Xu L, Liu W, Cui B, Wang N, Ding J, Liu C, Gao S, Zhang S. 2017.
10.24831/jai.v47i3.26076. [Indonesian] Aluminium tolerance assessment of 141 maize germplasms in a solution
culture. Univ J Agric Res 5 (1): 1-9. DOI: 10.13189/ujar.2017.050101.
 SURAT PERNYATAAN
KONTRIBUTORSHIP

Yang bertanda tangan di bawah ini, kami penulis artikel berjudul: Secondary trait and
index selection determination for maize genotype selection in an acidic tidal swamp
environment yang diterbitkan pada Jurnal BIODIVERSITAS Volume 23, Number 8, Agustus
2022, menyatakan bahwa kontributor penulisan dari Karya Tulis Ilmiah tersebut adalah sebagai
berikut:

No Nama Penulis Status Kontributor

1 Suwarti Kontributor Utama
2 Munif Ghulamahdi Kontributor Utama
3 Muhammad Azrai Kontributor Utama
4 Didy Sopandie Kontributor Utama
5 Eko Sulistyono Kontributor Utama
6 Trikoesoemaningtyas Kontributor Utama

Demikian surat pernyataan ini dibuat dengan sebenar-benarnya untuk dapat

dipergunakan sebagaimana mestinya.

Bogor, 30 November 2022

Kami yang membuat pernyataan,

No Nama Penulis Tanda Tangan

1 Suwarti 1.

2 Munif Ghulamahdi 2.

3 Muhammad Azrai 3.

4 Didy Sopandie 4.

5 Eko Sulistyono 5.

6 Trikoesoemaningtyas 6.
Biodiversitas Journal of Biological Diversity Tasks 0  English  View Site  warti

Notifications 
11449 / SUWARTI et al. / Secondary trait and index selection determinati Library

[biodiv] Editor Decision

2022-06-21 09:24 AM
Submissions Workflow Publication

Submission Review Copyediting Production

Suwarti Mujiharto, Ghulamahdi:

We have reached a decision regarding your submission to Biodiversitas Journal of Biological Diversity,
Round 1 Determination
"Secondary Trait and Index Selection Round 2 for Maize
Round 3
Genotype Selection on Tidal Swamp Acid
Environment".

Our decision is: Revisions Required

Round 3 Status
Submission accepted.

Notifications
------------------------------------------------------
Reviewer A:
[biodiv] Editor Decision 2022-06-21 09:24 AM

-This manuscript has outdated references. At least, you need to compose a minimum 80% of scientific
[biodiv] Editor Decision 2022-07-19 06:54 AM
journals published in the last 10 years (2012-2022)
[biodiv] Editor Decision 2022-07-20 12:12 PM
-Maximum 10% of reference in Indonesian
[biodiv] Editor Decision 2022-08-09 05:29 AM
-Please write the references based on the author's guidelines, include DOI. Kindly check the author's
guidelines here https://smujo.id/biodiv/guidance-for-author
[biodiv] Editor Decision 2022-08-12 03:48 AM

Recommendation: Revisions Required

[biodiv] Editor Decision 2022-08-12 03:57 AM

------------------------------------------------------

Reviewer's Attachments  Search

No Files

________________________________________________________________________
Biodiversitas Journal of Biological Diversity

Revisions
 e so s
Biodiversitas Journal of Biological Diversity Tasks 0  English  View Site  warti
 Search Upload File
Notifications
No Files

[biodiv] Editor Decision
Review Discussions Add discussion

2022-06-21 09:24 AM Name From Last Reply Replies Closed

— warti warti 0
2022-07-14 2022-07-14

Suwarti Mujiharto, Ghulamahdi: 01:48 AM 03:43 AM

— warti - 0
We have reached a decision regarding your submission to Biodiversitas Journal of Biological Diversity,
2022-07-14
"Secondary Trait and Index Selection Determination for Maize Genotype Selection on Tidal Swamp Acid
03:43 AM
Environment".
— warti - 0
Our decision is: Revisions Required 2022-07-29
06:43 AM

Uncorrected dewinurpratiwi warti 1

Proof 2022-08-04 2022-08-06
05:18 AM 11:51 PM
------------------------------------------------------
Reviewer A: BILLING dewinurpratiwi dewinurpratiwi 2
2022-08-04 2022-08-09
-This manuscript has outdated references. At least, you need to
05:20 AM compose a minimum
12:08 AM 80% of scientific
journals published in the last 10 years (2012-2022)

-Maximum 10% of reference in Indonesian

-Please write the references based on the author's guidelines, include DOI. Kindly check the author's
guidelines here https://smujo.id/biodiv/guidance-for-author

Recommendation: Revisions Required

------------------------------------------------------

________________________________________________________________________
Biodiversitas Journal of Biological Diversity
Biodiversitas Journal of Biological Diversity Tasks 0  English  View Site  warti

Notifications 
11449 / SUWARTI et al. / Secondary trait and index selection determinati Library

[biodiv] Editor Decision

2022-07-19 06:54 AM
Submissions Workflow Publication

Submission Review Copyediting Production

Suwarti Mujiharto, Ghulamahdi:

We have reached a decision regarding your submission to Biodiversitas Journal of Biological Diversity,
Round 1 Determination
"Secondary Trait and Index Selection Round 2 for Maize
Round 3
Genotype Selection on Tidal Swamp Acid
Environment".

Our decision is: Revisions Required

Round 3 Status
Submission accepted.

Notifications
------------------------------------------------------
Reviewer A:
[biodiv] Editor Decision 2022-06-21 09:24 AM

This article contains comprehensive data sufficient for publication as a research article in Biodiversitas.
[biodiv] Editor Decision 2022-07-19 06:54 AM
The article has been well written, the data are solid and well presented, the discussion is comprehensive,
and the results contain novelty[biodiv]
in the field.
EditorHowever,
Decisionthere is still inappropriate use of terms in the text,
2022-07-20 12:12 PM
which is unusual in the plant breeding subject. Please see the corrections for clarification and revision.
[biodiv] Editor Decision 2022-08-09
need 05:29
AM
Additional comments and editorial corrections are also provided for revision. All the references to
be rewritten following the journal Guidance for Author.
[biodiv] Editor Decision 2022-08-12 03:48 AM

Recommendation: Revisions Required

[biodiv] Editor Decision 2022-08-12 03:57 AM

------------------------------------------------------

Reviewer's Attachments  Search

No Files

________________________________________________________________________
Biodiversitas Journal of Biological Diversity

Revisions
 e so s
Biodiversitas Journal of Biological Diversity Tasks 0  English  View Site  warti
 Search Upload File
Notifications
No Files

[biodiv] Editor Decision
Review Discussions Add discussion

2022-07-19 06:54 AM Name From Last Reply Replies Closed

— warti warti 0
2022-07-14 2022-07-14

Suwarti Mujiharto, Ghulamahdi: 01:48 AM 03:43 AM

— warti - 0
We have reached a decision regarding your submission to Biodiversitas Journal of Biological Diversity,
2022-07-14
"Secondary Trait and Index Selection Determination for Maize Genotype Selection on Tidal Swamp Acid
03:43 AM
Environment".
— warti - 0
Our decision is: Revisions Required 2022-07-29
06:43 AM

Uncorrected dewinurpratiwi warti 1

Proof 2022-08-04 2022-08-06
05:18 AM 11:51 PM
------------------------------------------------------
Reviewer A: BILLING dewinurpratiwi dewinurpratiwi 2
2022-08-04 2022-08-09
This article contains comprehensive data sufficient for05:20
publication
AM as a research article in Biodiversitas.
12:08 AM
The article has been well written, the data are solid and well presented, the discussion is comprehensive,
and the results contain novelty in the field. However, there is still inappropriate use of terms in the text,
which is unusual in the plant breeding subject. Please see the corrections for clarification and revision.
Additional comments and editorial corrections are also provided for revision. All the references need to
be rewritten following the journal Guidance for Author.

Recommendation: Revisions Required

------------------------------------------------------

________________________________________________________________________
Biodiversitas Journal of Biological Diversity
 1 Secondary trait and index selection determination for maize genotype
2 selection on acidic tidal swamp acid environment

3 SUWARTI 1,3♥, MUNIF GHULAMAHDI 2♥♥, DIDY SOPANDIE2, TRIKOESOEMANINGTYAS2, EKO

4 SULISTYONO2, MUHAMMAD AZRAI3
5 1
Graduate School of IPB University, Bogor 16680, Indonesia
6 
email: suwarti_agh@apps.ipb.ac.id
7 2
Department of Agronomy and Horticulture, Faculty of Agriculture, IPB University, Bogor 16680, Indonesia.
8 
email: mghulamahdi@yahoo.com
9 3
Indonesian Cereals Research Institute. Jl. Dr. Ratulangi No. 274, Maros 90514, South Sulawesi, Indonesia.

10 Manuscript received: DD MM 2022 (Date of abstract/manuscript submission). Revision accepted: .................... 2022.

11 Abstract. Determining secondary traits as complementary in maize genotypes selection for the stressed abiotic environment is
12 compulsory since employing the only main character in the form of yield usually provides biased result. This study aimed to obtain
13 secondary traits for the maize selection in an abiotic multi-stresses of Fe, Al, and temporary flooding environment of acid sulphate tidal
14 swamp land using multivariate analysis. A total of 150 (un-repeated) maize lines and five check genotypes (repeated five timesfive
15 replicates) were arranged in a randomized augmented plot design on three environmental treatments to generate 525 single plots. Three
16 types of environment based on irrigation regulation were applied on acid soil tidal swamp land to study the agronomic characteristics of
17 maize genotypes in each treatment. Saturated Cultivation Technology (SSC) was considered an optimal environment. Dryland treatment
18 was the condition to let the irrigation only depending on rainwater. Temporary flooding treatment was carried out at five stages at
19 different growth periods. Several steps of analysis were applied in this study by taking variance, and heritability was used as the basis
20 for selections. Affected characters by treatments on medium to high heritability were then analyzed with correlation and pathway
21 analysis. Seven selection indexes, SSI, GMP, YSI, YI, TOL, STI, and MP, that have a high correlation to yield by both optimum and
22 stress environments were applied simultaneity with the selected trait character to screen theing genotypes with the utilizationing of a
23 PCA biplot analysis. Venn analysis was used to group the genotypes based on the result of the whole analysis. Indexes Indices of STI,
24 MP, GMP, and traits of rows number per ear and the weight of 6 ears were selected as secondary characters for the maize genotypes
25 selection in Saturated Soil culture with temporary flooding stress. Meanwhile, STI, MP, and ear stand height were selected characters in
26 the dryland of acid sulphate tidal swamp land. Based on the combination of all secondary character selection, genotypes were grouped
27 on into five types of stress tolerance on SSC+TF and DL environment. A number of eEight genotypes were tolerant of to both SSC+TF
28 and DL stress, and two genotypes were susceptible in to both environments.

29 Keywords: acid soil tidal swamp, heritability, pathway analysis, pyrite, temporary flooding.

30 Abbreviations : SSC=saturated soil culture, TF=temporary flooding, SSC+TF=saturated soil culture+temporary flooding, DL=Dry
31 land, SSI= Stress susceptibility index, GMP= Geometric mean productivity, YSI=Yield stability index, YI= Yield index, TOL=
32 Tolerance Index, STI= Stress Tolerance Index, and MP= Mean Productivity.

33 Running title: secondary trait for maize selection in tidal swamp.

34 INTRODUCTION

35 Improvement of maize production through extending crop area has strict obstacles along with expanding a wide area by
36 converting arable soil for non-agricultural purposes (Maulana et al., 2019). The utilization of suboptimal land such as
37 sulfate acid soil tidal swamp area to increase national maize production in Indonesia is an opportunity since the land is
38 widely available, about 4.3 hectares out of a total of 20.1 hectares of tidal swampy areas, which can be managed as crops Commented [A1]: Please check the number of hactares:
39 area (Surahman et al., 2018). Pyrite oxidation in the soil layer was the main restriction to developing maize crops on the only 4.3 hectares is considered widely available??????
40 sulfate acid soil tidal swamp to manage the land. Pyrite oxidation would trigger Fe3+ released in the soil layer and leading
41 to a decreased pH level (Toyip et al., 2019; Annisa and Djufry, 2020). A low pH level in the soil triggers aluminum ion
42 solubility and increases acidity (Pujiwati et al., 2015, 2016). Insoluble iron in the form of Fe3+ is found at the aerated soil
43 layer and alkaline pH. In contrast, when soil is flooded and has an anaerobic condition or soil pH is decreased, there is a
44 reduction of Fe3+ to Fe2+, which is soluble and dangerous to the plant (Vahedian et al., 2014; dos Santos et al., 2017).
45 The toxicity of aluminum and iron to plant tissue has been studied previously in several experiments. In addition to
46 damaging the plant tissues, the main risk of heavy metal stresses of Al and Fe is the is decrease ofd the crop yields. Low
47 pH soil in acid soils increases the concentrations of Fe and Al in the solution, leading to the precipitation of inorganic
 48 phosphorus as iron and aluminium phosphates. The excess condition of Fe becomes toxic to the plant as a highly reactive
49 Fenton catalyst. Formation of ROS such as hydroxyl radicals (.OH) and superoxide anion radicals (O2.-) is easily generated
50 under Fe toxicity in the plant tissue (Nikolic and Pavlovic, 2018). Easy observed Fe excess stress symptoms in plant tissue
51 were the appearance of bronzing color, starting from tip to basal of the leaf (Nugraha et al., 2016). There are two types of
52 Fe uptake in plant; strategi strategy I for dicot and strategy II for graminea. Maize plant Fe uptake followed chelation-
53 based Fe uptake by the so-called phytosiderophores strategy. Those plant hormones are secreted to the rhizosphere to bind
54 Fe to become the Fe-phytosiderophore complex that enters the cell via the root ZmYS1 (Yellow stripe1) and YSL (Yellow
55 Stripe Like) transporter. Chelating compounds phytosiderophore belong to mugeneic acids family, released as Fe
56 deficiency response (Bartucca et al., 2018).
57 Maize crop productivity is negatively affected by aluminum presence in acid soil. The toxicity of aluminium primarily
58 affects plant root growth, thus, reducing the ability to exploit the soil water and nutrients (Kopittke et al., 2015; Matonyei
59 et al., 2020). The condition of low pH soil releases toxic forms of Al, such as Al(OH)2+, Al(OH)2+, and Al3+ into the soil
60 solution (Xu et al., 2017). Plant mechanisms of aluminum tolerance could be summarized into two types: Al exclusion and
61 detoxification. The exclusion mechanism is carried out in a way of limiting Al absorbance or reduces reducing the root’s
62 uptake. The detoxification mechanisms were usually conducted by Al complexation, followed by transfer and storage of
63 these complexes in vacuoles (Bian et al., 2018).
64 Development of maize variety that has been tolerant to several abiotic stress by trhough a breeding program for
65 sulphate acid tidal swamp area environment is necessary along with the development of proper agronomic management
66 (Sabagh et al., 2018). The initiation of breeding program for abiotic stress tolerant genotype usually starts with selecting
67 tolerant genotypes for the target environment. Grain yield is usually chosen as the main character in the selection of maize
68 tolerant to abiotic stress. However, using grain yield as solely character usually generate a biased result. Secondary traits
69 and index selection could complement the yield character as the plant tolerance to environmental stress measurement
70 (Fadhli et al., 2020). Multivariate analysis to obtain the selection index value will simplify the characters in selecting
71 genotypes tolerant to specific environments. A selection index based on the calculation of several formulas simultaneity
72 can be used to strengthen the applied characters used in a selection in the abiotic stress environment (Anshori et al., 2019).
73 This study aimed to obtain secondary characteristics for maize selection genotypes in acid sulfate tidal soil under Fe, Al,
74 and temporary flooding stress on Saturated Soil Culture crops technic and Dry Landdryland environment.

75 MATERIALS AND METHODS

76 Study area
77 The study was carried out in the sulphate acid soil tidal swamp area of Karya Bhakti Village, Rantau Rasau District,
78 Tanjung Jabung Timur Jambi Province, from May to September 2019. The coordinates location of SSC treatment was -
79 1°10’50”S 104°09’44”E, SSC+TF was -1°10’51”S 104°09’44”E, and DL was -1°10’50”S 104°09’37”E. The results of soil
80 analysis at SSC, SSC+TF and DL locations, respectively, showed pyrite levels of 400 ppm; 100 ppm, and 100 ppm, Fe
81 levels were 45200 ppm; 35600 ppm and 37300 ppm (very high), sulphur levels of 200 ppm, 500 ppm and 100 ppm, Al3+
82 levels 52900 ppm, 47600 ppm and 46200 ppm (very high), soil pH H2O 5.10, 5.20 and 5.2 (acidic), pH KCl 3.6, 3.7 and
83 3.6. Organic matter C 7.39 and 11.23 (very high), and 1.28 (low). Total N 0.21%, 0.31% (moderate), and 0.09%. C/N ratio
84 35, 36 (very high) and 14 (medium). Available P2O was 12 (medium), 8 and 8 (low). Cation exchange capacities were 17.2
85 cmol.g-1 and 23.72 cmol.g-1 (medium) and 8.97 cmol.g-1 (low). Climatic data from January to June 2019 obtained from the
86 Meteorology, Climatology and Geophysics Agency, Muaro Jambi Climatology Station, shows an average monthly
87 temperature was 22.71°C - 33.65°C, relative humidity 77.22-85.77%, rainfall 42.70-191.70 mm per month
88

89 Procedures
90 Design of field experiment field
91 The experiment was arranged in an augmented randomized block design. Genetic Material materials used genetics
92 consisted of 150 unrepeated maize lines collection by the Indonesian Cereals Research Institute, and five check genotypes
93 repeated five times. Check genetics genotypes consisting consisted of one commercial composite variety (Sukmaraga) as
94 check number 1, two elite lines (MR-14 and Nei 9008) as check number 2 and check number 3, one commercial hybrid
95 variety (P27) as check number 4 and one commercial three-way cross-hybrid variety (Bima 20) as check number 5. Maize
96 seeds were planted at a depth of ± 5 cm, the distance between rows was 70 cm, and the distance between plants was 20 cm;
97 to gain a population per hectare was of 71428 plants. Each plot consisted of three rows of plants, with the row length being
98 4.2 m. Each block consisted of 30 test plots of genotypes and five plots of check genotypes.
99
100 Regulation Arrangemment of soil irrigation in each treatment
101 There was three regulation of soil irrigation system treatments applied in this experiment. The treatments were
102 Saturated Soil Culture (SSC), SSC+Temporary Flooding, and Dry Land (DL) environment. The saturated Soil Culture
103 technic technique manages the water level to adjust 5 cm above the base of the 30 cm wide and 25 cm depth trench, the
104 length of the trench following the length of the land perpendicular to the primary trench. This SSC technic purpose is to
105 manage soil consistently in redox conditions. Floodgates was a tube (polongan) that were applied to flow and retain water
106 in the field at low tide and discharge water out of the field during high tides so that it also functions to clean the pyrite
107 layer at the bottom of the trench.
108 The SSC+Temporary Flooding (SSC+TF) treatment was SSC technic technique but allowed the water to overflow until
109 it reached 10 cm above the root area, arranged periodically to allow roots area in the anoxia condition. Temporary flooding
110 in SSC+TF period is occurred on after planting (0 to 4 DAP); TF at 14-17 DAP; TF at 28-31 DAP; TF at 43-46 DAP
111 (anthesis and tassel periods), and TF at 58-61 DAP (R1/grain filling periods) (Figure 1). The water pump is was used
112 conditionally to regulate water flow if swamp water overflow is inadequate artificially. A 50 cm high embankment covered
113 with tarpaulin is was made around the land to prevent water leakage inside and outside the experimental area. The third
114 environment treatment was dry land (DL), where soil watering in this experiment was dependent on rainwater.

SSC
SSC + TF
DL
5-13 DAP 18-28 DAP 32-42 DAP 47-57 DAP
14-17 DAP

28-31 DAP

4 DA R1
0-4 DAP

4 DAT

= temporary flooding periode

= saturated soil culture technic periode
115 = watering depend on rain water
116
117 Figure 1. Irrigation treatment scheme on SSC technique and the temporary flooding period in the experimental environment

118 Tolenace Index Tolerant Formulas

119 The estimated heritability value is was calculated using the heritability equation in the broad sense, which is
120 derived from the variance in the broad sense . Heritability values, in a broad sense, are expressed by
121 decimal numbers ranging from 0 to 1 or percentages, the criteria for heritability values are classified by category; high
122 , medium ( and low . Stepwise regression was applied to determine the most
123 important character with a high correlation with the yield.
124 The tolerance index is was calculated based on the observed yield variables. The tolerance index used in this
125 experiment was; Stress susceptibility index , with tolerant criteria if S < 0,5; moderate if the value is 0.5
126 < S <1; Sensitive if the value of S > 1, Geometric mean productivity ; Yield stability index ;
127 Yield index ; Tolerance Index ; Stress Tolerance Index ; Mean Productivity
128 . Description: Yp = Value of certain variables in varieties under stress; Yo= The value of a certain variable
129 in the varieties that were not stressed; Xp = the average value of certain variables in all varieties under stress; Xo = The
130 average value of a certain variable in all varieties that are not stressed.

131 Data Analysis

132 All the collected data were analyzed over several stages. Data were analyzed to obtain variance with a standard error of
133 5 % at the initial process. Characters significantly affected by genetic variance and have high heritability values (>50%)
134 are were then reselected through correlation analysis and path analysis to determine the most important characters on the
135 yield character and directly affect the yield. Seven tolerance indexes tolerantices were connected to yield character by
136 correlation analysis. The selected characters were then analyzed with using PCA (Principal Component Analysis). Venn
137 Diagram by (Heberle et al., 2015) was applied to separate all the tested genotypes into tolerance groups in environmental
138 stress trials.
139

140 RESULTS AND DISCUSSION

141 Genotypic Variance, Phenotypic Variance, and Heritability of 150 maize genotypes under three environments on
142 acid soil, tidal swamp land stress
143 The mean square of agronomic character data is presented in Table 1. Genotype factor was highly significantly affected
144 to yield character both in SSC+TF and SSC environments, while DL environment showed slightly affected. Some of the Commented [A2]: What is the criteria for slightly affected?
Please use the standard term.
145 data were being transformed due to high variance. Most of the characters on SSC treatment were highly affected by Commented [A3]: Pelase mention the type of data
146 genotypic factors. Senescence, leaf length, leaf width, anthesis days, silking days, and weight of six stems were the transformation carried out.
147 characteristics that were not affected by genotypic factors among all 23 observed traits. On SSC+TF environment,
148 character the number of leaves and leaf length are were not affected by genotypic factors. The dry land treatment generated
149 most characters that were not affected by genotypic factors compared to the other treatments. Those characters in dry land
150 were a weight of 6 cobs, the weight of 1 cob, number of harvested plants, number of harvested cobs, plant aspect, Commented [A4]: What exactly????
151 senescence, ear diameter, stem diameter, and weight of 6 dry stems.
152
153 Table 1. Mean square of genotype factor at each character on environment treatments of SSC, dan SSC+TF, and DL
Source of Variance MSg SSC CV MSg SSC+TF CV MSg DL CV
Yield 0.15** 30.59t2 0.03** 15.68t2 0.10* 22.01t2
Weight 6 cobs 0.07** 8.88t2 0.99* 11.73t2 0.01ns 12.71t2
Weight of 1 cob 0.10** 22.85 0.12** 22.19 0.02ns 2.71t2
Number of harvested plants 0.07** 10.59t1 0.99* 32.78t2 0.01ns 11.95t1
Number of harvested cobs 0.10** 13.55t1 0.12** 17.76t1 0.02ns 16.29t1
Kernels/Cob weight Ratio 7.18** 13.07 21.06** 7.89 0.04** 12.80t2
Weight of 1000 seed 2920.57* 23.37 0.02* 5.36t1 0.01** 3.97t1
Plant Aspect 0.43** 16.38 0.02** 14.96t1 0.00ns 14.85t1
Senescence 0.02ns 22.98 0.03** 11.75t1 0.01ns 12.86t1
Ear Length 9.03** 16.82 6.37** 21.22 0.01* 8.81t1
** *
Ear Diameter 0.43 10.69 0.40 21.06 0.25ns 17.92
** **
Number of Rows per Cob 5.03 10.69 6.14 21.06 2.74** 17.92
Number of Seeds per Row 40.86** 23.07 0.07** 8.07t1 0.02** 8.77t1
Number of Leaves 0.01* 11.93t1 1.82ns 24.68 2.57** 13.92
Height of Cob 125.10** 19.59 0.03* 11.62t1 237.59** 20.01
Plant Height 524.82** 14.47 468.22** 19.93 600.18** 13.16
Leaf Length 101.09ns 21.31 0.01ns 6.86 84.13* 11.04
Leaf Width 0.01ns 11.36t1 0.01* 12.50t1 1.11* 13.18
Stem Diameter 6.51* 18.21 6.28* 22.87 0.01ns 9.91t1
SPAD 39.80** 12.29 43.49* 20.14 23.20* 11.04
Anthesis Day 11.54ns 7.21 13.76** 4.91 9.21** 2.50
Silking Day 16.03ns 6.58 19.12** 4.99 9.88** 2.54
Weight of 6 dry stem 0.01ns 10.91t2 0.01** 5.65t2 0.01ns 10.18t2
154 Note : **significant effect on 1% level, *significant effect on 5% level, ns non significant, t1 transformed at ; t2 transformed at
155 ; CV=Coefficient of Variance; MS=mean square of genotypes
156
157 Other characters that showed were highly significantly affected by genotype and had moderate to high heritability
158 values in the SSC+TF environment were the weight of 6 ears, the weight of 1 ear, number of harvested cobs, the ratio of
159 shelled grain to cob, senescence, ear diameter, ear length, rows number of kernels, number of seeds per row, plant height Commented [A5]: What is the unit?????
160 at 10 WAP, stem diameter at 8 WAP, SPAD value, anthesis day, silking days, and weight of 6 dry stems (Table 1 and
161 Table 2).
162 A summary of the phenotypic variance, genotypic variance, and heritability observed characters are shown in Table 2.
163 Characters with high heritability values can be used as selection characters criteria to increase the effectiveness of
164 selection. Maize plants have different responses to the environment, so the characters utilized to increase the effectiveness
165 of selection also vary, depending on the selection environment. The highest heritability on yield character was obtained in
166 the SSC environment (scored 59.81%), followed by DL environment, which scored 20.40% and was categorized as a
167 medium category, and at the lowest was SSC+TF environment, it had a negative heritability value (-19.23) which means
168 there was no genetic progress and can be considered as 0 (zero). Low heritability indicates that a the character is more
169 affected by the environment rather than genotypic nature (Wening et al., 2020).
170 A high heritability category value in the broad sense of the agronomic character in the SSC environment (> 50%) was
171 obtained on the weight of 6 ears, the weight of 1 ear, ear diameter, and ear height position. Heritability values in the
172 medium category (between 20% - 50%) were obtained on the number of harvested plants, the number of harvested cobs,
173 plant aspect, seeds number per row, seeds rows number, plant height, number of leaves, stem diameter, SPAD value, and
174 weight of 6 dry stems. High heritability in the SSC+TF environment was shown at characters of the weight of 6 cobs, the
175 weight of 1 ear kernels/cob ratio, and the number of rows per ear. The number of harvested cobs, senescence, ear length,
176 ear diameter, seeds number per row, plant height, stem diameter, SPAD value, anthesis day, silking day, and weight of 6
177 dry stems were categorized as a medium heritability. The high-category heritability values of agronomic characters in DL
178 environment treatment were obtained on the weight of 6 cobs, the ratio of kernel to cob, anthesis days, and silking days.
179 Medium heritability values were obtained on the number of rows per ear, seeds number per row, number of leaves, plant
180 height, and leaf width. Heritability of each maize character is influenced by the action of gene additive or non-additive
181 gene action and express at the phenotype performance (Sesay et al., 2016).
182
183
184
185
186 Table 2. Agronomic characters of 150 maize lines under three environments types kind on acid soil tidal swamp, Jambi 2019
Agronomic Traits SSC SSC+TF DL
σ2p σ2g h2 σ2p σ2g h2 σ2p σ2g h2
Yield 0.21 0.35 59.81 -0.12 0.61 -19.23 0.10 0.49 20.40
Weight 6 cobs 14.64 14.65 99.95 8.51 8.52 99.96 2.52 2.54 99.18
Weight of 1 cob 17.62 17.62 100.00 6.98 6.98 100.00 5.57 5.57 99.99
Number of harvested plants 9.32 35.95 25.92 3.86 27.14 14.21 0.19 11.85 1.58
Number of harvested cobs 10.57 45.83 23.06 4.92 15.26 32.22 -0.92 31.52 -2.92
Kernels/Cob weight Ratio -34.04 150.50 -22.62 4.21 4.23 99.52 0.07 0.12 61.57
Weight of 1000 seed 218.23 2168.13 10.07 603.25 3458.62 17.44 228.26 2074.85 11.00
Plant Aspect 0.05 0.21 24.80 0.07 0.40 18.33 0.02 0.22 8.57
Senescence 0.45 2.51 17.95 0.55 2.07 26.39 0.01 2.30 0.43
Ear Length 0.87 5.20 16.76 0.96 2.54 37.80 0.41 3.37 12.03
Ear Diameter 0.07 0.13 56.83 0.05 0.20 23.86 -0.48 2.40 -20.04
Number of Rows per ear 0.67 2.37 28.37 1.12 1.64 68.55 0.59 2.33 25.26
Seeds number per Row 4.64 22.43 20.70 6.34 14.14 44.83 2.68 12.87 20.83
Number of Leaves 0.46 2.03 22.83 0.18 1.11 16.14 0.37 1.15 31.94
Height of Cobs 20.86 41.66 50.08 10.23 52.65 19.42 0.22 0.41 52.49
Plant Height 86.56 178.59 48.46 50.78 130.67 38.87 93.22 227.32 41.01
Leaf Length 6.71 74.26 9.03 0.00 0.01 4.12 9.11 53.10 17.16
Leaf Width 0.11 1.28 8.74 0.10 0.85 11.27 0.14 0.57 24.13
Stem Diameter 0.79 3.34 23.80 0.70 3.47 20.28 -0.22 9.21 -2.41
SPAD 5.62 17.31 32.49 5.02 23.39 21.48 2.56 12.88 19.87
Anthesis Day 0.54 9.39 5.73 1.85 6.37 28.98 1.65 2.61 63.08
Silking Day 1.60 9.61 16.69 2.78 8.02 34.62 1.76 2.84 62.09
Weight of 6 dry stems 0.00 0.00 33.50 0.00 0.01 47.01 0.00 0.01 7.81
187 2 2 2
Notes: σ p = phenotipic variance, σ g = genotipic variance, h =heritability, SSC=sturated soil culture, DL=dry land, SSC+TF= saturated
188 soil culture+temporary flooding.
189

190 Correlation of important character to the yield based on stepwise regression analysis
191 Stepwise regression analysis was conducted to determine the essential/dominant characters related to the main
192 character (Ayvat and Omeroglu, 2022). The generated model by stepwise analysis was considered to be more precise in
193 predicting the results based on the resulting model than the correlation involving all observation variables (Andayani et al.,
194 2016). This study found that, in SSC environment weight of 6 ears (X6e), the weight of 1 ear (X1e), and the height of ear
195 stand (XeSt) were the dominant characters that affect maize grain yield (Table 3). The SSC treatment in acid soil tidal
196 swamp is considers considered as an optimum crop technic in tidal swamp.
197 The stepwise regression correlation analysis on all maize characters observed in the SSC+TF environment showed that
198 four traits had a significant effect, with an R2 value of 70.00% and a model F value of 87.52 significantly different. These
199 characters were the weight of 6 ears (X6E), the weight of 1 ear (X1E), kernels per cob weight ratio (XKpC), and the
200 number of rows per ear (XNR) (Table 4). The equation model formed from this analysis were Y(SSC+TF)= -0.09 + 1.63
201 X6E + 0.71 X1E - 0.01 XKpC + 0.01 XNR. Meanwhile, the stepwise regression correlation results on dry land treatment
202 showed three characters of the( weight of 6 ears, the height of ear stand, and anthesis day) that were dominant to affected
203 yield. Equation model formed in dry land treatment was Yield DL= 4.26 - 0.30 XKpC + 0.02 XESt - 0.07 XAD. The R2
204 value explained from this model is 40.81%, with a model F value was 34.7, highly significant (Table 5).

205 Table 3. The most important trait correlation based on stepwise regression analysis of SSC tidal swamp treatment
Characters YieldSSC (X6e) (X1e)
YieldSSC 1
Weight of 6 ear (X6e) 0.75** 1
Weight of 1 ear (X1e) 0.44** 0.34** 1
Height of Ear stand (XeSt) 0.51** 0.53** 0.26**
206 Notes: **significant at 1% level, *significant at 5% level

207 Table 4. The most important trait correlation based on stepwise regression analysis on SSC+GS tidal swamp treatment
Characters YieldSSC+GS (X6e) (X1e) (XKpC)
YieldSSC+GS 1
Weight of 6 ear (X6E) 0.82** 1
Weight of 1 ear (X1E) 0.1ns 0.05ns 1
Kernels/Cob weight Ratio (XKpC) -0.23** -0.17* 0.31** 1
Number of Rows per ear (XNR) 0.52** 0.47** 0.17* -0.2*
208 Notes: **significant at 1% level, *significant at 5% level

209

210 Table 5. The most important trait correlation based on stepwise regression analysis on DL tidal swamp treatment
Characters YieldDL (XKpC) (XeSt)
YieldDL 1
Kernels/Cob weight Ratio (XKpC) -0.31** 1
Height of Ear stand (XeSt) 0.53** -0.12ns 1
Anthesis Day (XAD) -0.32** 0.03ns -0.11Ns
211 Notes: **significant at 1% level, *significant at 5% level
212
213 Each environment shows a different important selected character with a high correlation to yield. Two characters are
214 chosen in optimum treatment (SSC) were similar to those found in SSC+TF. The result was understandable though both
215 environments lay in the same area. Height ear stands as the dominant character that affected grain yield in stress DL
216 environment was found in SSC optimum environment likewise. Kernels per cob ratio in SSC+TF is also shown as the
217 dominant character in DL environment. The high correlation between variables shows a close relationship and can be an
218 indirect selection character for quantitative traits (Hamawaki et al., 2017).
219

220 Direct and indirect effects of agronomic traits on the yield characters
221 The pathway analysis results on the Saturated Water Cultivation (SSC) treatment showed a residual value of 0.39,
222 meaning that the path model could explain 61% of the variance. The most considerable direct effect was obtained on the
223 weight of 6 ears, which has a direct effect coefficient of 0.61, weight of 1 ear get 0.20, and the height of ear stand get 0.13
224 score the direct effect. The pathway analysis of characters in SSC+TF environment showed a residue of 0.30, meaning that
225 70% of the yield characters could be explained by four characters used in the test. The character weight of 6 ears (0.73),
226 the weight of 1 ear (0.7), and the number of rows per ear (0.14) showed a positive direct effect coefficient on the yield.
227 However, the kernels per cob ratio have a negative direct effect (-10). Characters of kernels per cob ratio and anthesis day
228 had a negative direct effect on grain yield, with values of -0.25 and -0.26, respectively.
229 In contrast, the height of the ear stand has a positive direct effect (0.47) in DL environment. The residual effect in this
230 environment stress treatment was 0.59, meaning that 41 % of the path model could explain the yield character. The
231 positive direct characters to yield in path analysis indicated that in those environments, increasing maize grain yield was
232 supported by increasing the character value. A negative direct effect demonstrates that the yield response is more
233 influenced by the indirect effect (Putri and Ashari, 2019). Path analysis enable breeders to improve understanding of the
234 relationships between several traits by revealing the correlation coefficients into direct and indirect effects (Hamawaki et
235 al., 2017).
236
237 Table 6. Agronomic characters with direct effect and indirect effect character to maize grain yield of in SSC treatment in acid soil tidal
238 swamp
Indirect Effect
Direct Effect (X6e) (X1e) (XeSt) Residual
Weight of 6 ears (X6e) 0.61 0.07 0.07 0.39
Weight of 1 ear (X1e) 0.20 0.21 0.03 0.39
Height of Ear stand (XESt) 0.13 0.32 0.05 0.39
239
240 Table 7. Agronomic acahracters with direct effect and indirect effect character to maize grain yield of SSC+TF treatment in acid soil
241 tidal swamp
Direct Indirect Effect
Effect (X6E) (X1E) (XKpC) (XNR) Residual
Weight of 6 ears (X6e) 0.73 0.00 0.02 0.07 0.30
Weight of 1 ear (X1e) 0.07 0.04 -0.03 0.02 0.30
Kernels/Cob weight Ratio (XKpC) -0.10 -0.12 0.02 -0.03 0.30
Number of Rows per ear (XNR) 0.14 0.34 0.01 0.02 0.30

242 Table 8. Agronomic characters with direct effect and indirect effect character to maize grain yield of DL treatment in acid soil tidal
243 swamp
Indirect Effect
Direct Effect (XKpC) (XESt) (XAD) Residual
Kernels/Cob weight Ratio (XKpC) -0.25 -0.06 -0.01 0.59
Height of Ear stand (XESt) 0.47 0.03 0.03 0.59
Anthesis Day (XAD) -0.26 -0.01 -0.05 0.59
244
245 Correlation of selection index to yield characters
246 A total of seven stress tolerance indexes indices (GMP, YSI, SSI, YI, TOL, STI, and MP) were used to determine the
247 appropriate index for maize genotype selection in tidal areas under the stress of Al, Fe, and temporary flooding (Table 9).
248 The correlation coefficient of tolerance index indices with productivity in SSC+TF treatment shows a very highly
249 significant and high value based on the Pearson correlation on the GMP (r=0.73), STI (r=0.63), and MP (r=0.86)
250 indexesindices; all those indexes indices had an exact high correlation to yield on the optimum treatment (SSC) that has
251 correlation indexes indices of r=0.89, r=0.85, and r=0.82, respectively.
252 Index selection that has high correlation with grain yield on DL environment stress was STI (r=0.81) and MP
253 (r=0.77), the correlation of optimum SSC treatment on those index selection was STI (r=0.76) and MP (r=0.98).
254 Correlation coefficient score is useful to explain the relationship within traits in term of degree, characteristic, and
255 direction of selection to apply (Sayo et al., 2017).
256

257 Table 9. Pearson’s Correlation of tolerance index indices and grain yield characters on the SSC + TF treatment acid soil tidal swamp
Characters Yp Ys_gs GMP YSI SSI YI TOL STI MP
Yp 1
Ys_dl 0.42** 1
GMP 0.89** 0.73** 1
YSI -0.2** 0.07 -0.2** 1
SSI -0.2** 0.07 -0.2** 1** 1
YI 0.42** 1** 0.73** 0.07 0.07 1
TOL 0.46** -0.6** 0.06 -0.3** -0.3** -0.6** 1
STI 0.85** 0.63** 0.9** -0.1 -0.1 0.63** 0.13 1
MP 0.82** 0.86** 0.96** -0.1 -0.1 0.86** -0.1 0.87** 1
258 Notes: ** significant at 1% level, Yp: the yield on SSC treatment, Ys_gs: the yield on SSC+GS treatment; GMP: geometric mean
259 productivity; YSI: yield stability index; SSI: susceptibility index; YI: yield index; TOL: tolerance index; STI: stress tolerance index; MP:
260 mean productivity.
261

262 Table 10. Pearson’s Correlation of tolerance index indices and grain yield characters on the DL treatment acid soil tidal swamp
Characters Yp Ys_dl GMP YSI SSI YI TOL STI MP
Yp 1
Ys_gs 0.62** 1
GMP 0.87** 0.9** 1
YSI -0.2* 0.16 -0.1 1
SSI 0.18* -0.2* 0.07 -1** 1
YI 0.62** 1** 0.9** 0.16 -0.2 1
TOL 0.95** 0.33** 0.67** -0.3** 0.28** 0.33** 1
STI 0.76** 0.81** 0.89** -0 0.04 0.81** 0.58** 1
MP 0.98** 0.77** 0.95** -0.1 0.1 0.77** 0.85** 0.84** 1
263 Notes: ** significant at 1% level, Yp: the yield on SSC treatment, Ys_dl: yield in dry land treatment; GMP: geometric mean
264 productivity; YSI: yield stability index; SSI: susceptibility index; YI: yield index; TOL: tolerance index; STI: stress tolerance index; MP:
265 mean productivity.
266

267 Appropriate Secondary Trait and Index Selection For Genotype Selection In Tidal Swamp Land Using PCA
268 Analysis
269 PCA biplot analysis was employed to explains the relationship between tolerance index and productivity of stress
270 environment in this experiment. PCA analysis is widely used as it effectively simplifies large amounts of data into easy-to-
271 understand visualizations (Metsalu and Vilo, 2015). The data analyzed was standardized beforehand—data closer to the
272 biplot axis indicates more stable characters compared to those more distant data from axis. In the previous study, Principal
273 Component Analysis is useful to predict the relationship between germplasm to the environment and cassava sample
274 according to several of its enzymatic activity. The analysis work with the approach of multiple correlations (Wang et al.,
275 2014; Mursyidin and Khairullah, 2020).
276 The results of the biplot analysis showed that the STI, MP, and XESt indexes vectors coincide with the yield character
277 in the SSC+TF environments (Figure 2). In the DL environment, the STI, GMP, MP, XNR, and X6E were at PC 1. The
278 value of PC1 on SSC+TF explained 52.40% of the variance, and on PC2, 16.30% of the variance. PC1 in the DL
279 environment explained 57.10% variance, and PC2 explained 16.60% variance.
280
281
282 SSC+TF DL
283 Figure 2. Biplot principal components 1 and 2 for all tolerance indexes indices with yield in dryland (DL) and saturated
284 water+temporary flooding (SS+TF) cultivation treatments
285
286
287 Table 11. PCA eigenvalues for each dimension in SSC + TF treatment and treatment (DL) dry land in tidal land
PC 1 PC 2 PC 3 PC 4 PC 5
SSC+TF Treatment
Yield SSC+TF 0.94 -0.05 -0.02 0.07 -0.13
Geometric Mean Productivity (GMP) 0.98 0.01 -0.11 -0.04 0.12
Stress Tolerance Index (STI) 0.89 0.10 -0.29 -0.13 0.07
Mean Productivity (MP) 0.90 0.08 -0.19 -0.12 0.25
Weight of 6 Ear (X6E) 0.83 -0.07 0.02 0.29 -0.43
Number of Rows Per Ear (XNR) 0.59 -0.03 0.73 0.25 0.23
Variance percent 57.07 16.57 10.44 7.35 4.80
Cumulative Variance percent 57.07 73.64 84.08 91.44 96.24
Eigenvalue 4.57 1.33 0.84 0.59 0.38
DL Treatment
Yield DL 0.90 -0.02 -0.02 -0.03 0.42
Stress Tolerance Index (STI) 0.84 0.10 0.13 0.43 -0.28
Mean Productivity (MP) 0.96 -0.02 0.09 0.19 0.03
Height of Ear Stand (XESt) 0.65 -0.05 0.41 -0.62 -0.16
Variance percent 52.35 16.33 14.50 11.07 4.98
Cumulative Variance percent 52.35 68.69 83.19 94.26 99.24
Eigenvalue 3.14 0.98 0.87 0.66 0.30
288
289 The results of the biplot analysis show that the STI, MP, and XESt indexes indices vectors coincide with the yield
290 character in the SSC+TF environments (Figure 2). In the DL environment, the STI, GMP, MP, XNR, and X6E were at PC
291 1. The value of PC1 on SSC+TF was able to explain 52.40% of the variance and on PC2 explained 16.30% of the
292 variance. PC1 in the DL environment explained 57.10% variance on PC1 and 16.60% variance on PC2. Data in Table 9
293 show that the highest PC coefficient was obtained on PC 1 in SSC+TF and DL environment treatments. As the main
294 character, yield is given 3 points in the model to optimize the selection. Based on the eigenvalue, PC1 was the fit model to
295 explain the characters factor. Hence models to explain index selection in both SSC and DL environments were:
296 ISSC+TF= 3*0.94 Yield+0.98GMP+0.89STI+0.90MP+0.83X6E+0.59XNR,
297 IDL=3*0.90Yield+0.84STI+0.96MP+0.65XESt.
298

299 Tolerance Grouping Based on Multivariate Analysis

300 The grouping of genotype tolerances was based on the index value obtained from the calculation following the index
301 selection model. The genotypes are were then parted classified into three categories: , i.e., tolerant, mediummoderately-
302 tolerant and susceptible. Tolerant category score was Xi > X + SE*ttable, the susceptible category score was Xi < X -
303 SE*ttable, while the mediummoderately-tolerant category score was between these two values. A Venn diagram was applied
304 to describe the tolerance category in each environment. The results of the tolerance grouping in the two experimental
305 environments classified the, genotype was divided sinto five categories (Figure 3a). Eight genotypes were tolerant in both
306 SSC+TF and DL stress environments, 47 genotypes were classified tolerant in the DL environment and medium
307 moderately tolerant in SSC+TF environments. Forty-seven genotypes were medium moderately tolerant in both SSC+TF
308 and DL environments, and 51 genotypes were medium moderately tolerant at SSC+TF and susceptible in the DL
309 environment. Two genotypes were susceptible to SSC+TF and DL environments (Figure 3b).

310
311 (a) (b)
312 Figure 3. Venn diagram summarizing adaptation of maize lines on SSC+TF and DL of acid soil tidal swamp environments
313 based on selected indexes indices
314
315
316 Table 12. Venn diagram of maize genotype grouping based on the tolerance in SSC+TF and DL stress acid soil tidal swamp
317 environment
Mean Yield
Group of Average Number of
Genotypes
Tolerance Genotypes
SSC+TF DL
[Tolerant DL] and Sukmaraga, P27, BIMA20URI, M-35-1-B, MPop11_01, MPop18_02, 0.93 2.14 8
[Tolerant TF] MPop27_08, MPop28_01
[Tolerant DL] and MPop27_01, MPop28_02, MPop28_03, C44-3-1, C27-4-B, MPop28_04, 0.20 1.90 47
[Medium E57-B, MGOLD, MPop28_05, DKL9001, M1-3-B, M-10-1-B, DYW7-B,
Moderate TF] DYW34-B, MPop03_06, MPop02_02, MPop02_04, MPop02_06,
MPop11_06, MPop18_01, MPop26_01, MPop26_03, MPop27_04,
MPop05_01, MPop05_02, MPop05_04, MPop05_06, MPop24_01,
MPop24_02, MPop24_03, MPop27_05, MPop24_06, MPop24_08,
MPop24_09, MPop24_10, MPop21_03, MPop21_04, MPop21_08,
MPop21_09, MPop27_06, PAC2244-2-3-1-3-9-B-B, PAC2244-2-3-1-4-
11B-B, PAC9996-2-4-1-2-11B-B, PAC2247-1-1-1-1-8B-B, PAC2244-1-6-
1-1-9B-B, PAC2245-3-1-1-2-8B-B, C17-1-B
[Medium DL] and NEI9008, C39-1-B, C31-1-B, C25-1-B, E31B, E90B, M42-1-B, DYW15- 0.14 1.11 47
[Medium B, DYW24-B, CLYN231, AL46, MAL03, MR15, B11209, G1026-12,
Moderate TF] G1044-14, MPop03_01, CY-7, MPop03_09, MPop03_10, MPop10_01,
MPop10_07, MPop10_08, MPop02_03, MPop15_01, MPop11_02,
MPop11_03, MPop11_08, MPop05_03, MPop24_04, MPop24_05,
MPop24_07, PAC99912-2-4-1-4-10-B-B, PAC2247-1-1-1-4-12B-B,
PAC2247-1-1-1-2-9BB, MPop27_07, PAC2244-1-6-1-3-7B-B, PAC2244-
2-3-1-2-9B-B, PAC2246-2-4-1-2-10B-B, PAC2245-3-1-1-4-5B-B,
PAC9999-1-2-1-4-9B-B, PAC2247-1-1-1-2-8B-B, C40-2-B, C61-1-B,
C32-5-B, C58-1-B, C19-4-B
[Medium MR-14, C93-1-B, C63-1-B, C48-5-B, C46-2-B, C72-1-B, C47-3-B, C73-2- 0.08 0.71 51
Moderate TF] and B, C6-3-B, C62-2-B, C37-1-B, C11-1-B, C60B-B, C16B, C41-1-B,
[Susceptible DL] E100B, E93B, E52B, E102B, WYW27B, WYW18B, WYW11B, M-6-1-B,
DYW-17-B, MPop28-06, MPop03_03, MPop27_02, MPop10_02,
Pop10_03, MPop27_03, MPop26_02, MPop05_05, MPop21_10,
G104414, G2013631, PAC2245-3-1-1-4-4B-B, PAC9999-1-2-1-1-10BB,
PAC2243-1-5-1-1-9B-B, PAC99912-3-1-1-3-9B-B, PAC9997-1-1-1-3-8B-
B, PAC99912-2-4-1-3-8B-B, PAC99912-2-4-1-3-7B-B, PAC2241-2-3-1-
3-4B-B, PAC2246-2-4-1-1-7B-B, PAC99912-2-4-1-1-7B-B, C54-1-B,
C43-1-B, C56-1-B, C12-1-B, C42-3-B, C10-2-B
[Susceptible DL] AMB30, AMB20 0.00 0.78 2
 and [Susceptible
TF]
318 Sukmaraga, P27, and BIMA20URI varieties belong to the tolerant group in two stress treatment environments of
319 SSC+TF and DL. Those check varieties belonged to heterozygous genotypes of open-pollinated variety, hybrid variety,
320 and three-way cross variety, respectively. Maize tolerance to abiotic stress was commonly controlledencouraged by
321 additive genes, which is why heterozygotes are more desirable to survival in an abiotic stress environment. Genotypes of
322 M-35-1-B, MPop11_01, MPop18_02, MPop27_08, MPop28_01. Line gGenotypes of AMB30 and AMB20 were
323 susceptible in both SSC+TF and DL treatment environments and had lower index scores than all check genotypes.
324

325 Conclusion

326 Based on multivariate analysis, secondary characters/selection indices that can be to eemploy on thefor selection of maize
327 genotype selection iin acid soil tidal swamp land by using saturated soil culture technic technique with temporary flooding
328 (SSC+TF) stress were STI, MP, GMP, rows number per ear, and weight of 6 ears. STI, MP, and ear stand height was were
329 the most appropriate characters to apply to thefor selection of maize genotype selection on the acidic soil tidal swamp land
330 on dry soil land stressstress.

331 ACKNOWLEDGEMENTS

332 Acknowledgments are conveyedThe authors thank to the Research and Development Agency of the Ministry of
333 Agriculture through the DIPA Learning Officer , who hasfor the financed financial supports all offor this research activity.
334 We also thank and to PT FKS Multi Agro for allowing this research to be carried out on plant development land in the
335 tidal land of Tanjung Jabung Timur, Jambi.
336

337 REFERENCES Commented [A6]: Please rewrite the References according

the Biodiversitas Journal Guide for Authors. Please visit the
338 Andayani, N.N., M. Aqil, and Syuryawati. 2016. The application of stepwise regression model in the determination of white corn yields. Inform. Pertan. journal website for more details.
339 25(1): 21–28.
340 Annisa, W., and F. Djufry. 2020. Effect of reactive phosphate rock to corn on acid sulphate soil in South Kalimantan. ICFST 2019. p. 1–7 Formatted: Highlight
341 Anshori, M.F., B.S. Purwoko, I.S. Dewi, S.W. Ardie, and W.B. Suwarno. 2019. Selection index based on multivariate analysis for selecting doubled-
342 haploid rice lines in lowland saline prone area. Sabrao J. 51(2): 161–174.
343 Ayvat, P., and S.K. Omeroglu. 2022. Mortality estimation using APACHE and CT scores with stepwise linear regression method in COVID-19 intensive
344 care unit: A retrospective study. Clin. Imaging 88(March): 4–8. doi: 10.1016/j.clinimag.2022.04.017.
345 Bartucca, M.L., A. Di Michele, and D. Del Buono. 2018. Interference of three herbicides on iron acquisition in maize plants. Chemosphere 206: 424–
346 431. doi: 10.1016/j.chemosphere.2018.05.040.
347 Bian, H., J. Pang, C. Huang, and W. Zhang. 2018. Catena The response of transitional pedogenic characteristics of loess in the Yunxian Basin to abrupt
348 climatic events in the northern subtropics since the Last Glacial Maximum ( C ). Catena 171(January): 166–175. doi:
349 10.1016/j.catena.2018.06.016.
350 Fadhli, N., M. Farid, Rafiuddin, R. Efendi, M. Azrai, et al. 2020. Multivariate analysis to determine secondary characters in selecting adaptive hybrid
351 corn lines under drought stress. Biodiversitas 21(8): 3617–3624. doi: 10.13057/biodiv/d210826.
352 Hamawaki, O.T., G.F. Rezende, G.L. Jorge, I.C. Silveira, L.A. Medeiros, et al. 2017. Phenotypic and genotypic correlations between soybean agronomic
353 traits and path analysis. Genet. Mol. Res. 16(2): 2–11.
354 Heberle, H., V.G. Meirelles, F.R. da Silva, G.P. Telles, and R. Minghim. 2015. InteractiVenn: A web-based tool for the analysis of sets through Venn
355 diagrams. BMC Bioinformatics 16(1): 1–7. doi: 10.1186/s12859-015-0611-3.
356 Kopittke, P.M., K.L. Moore, E. Lombi, A. Gianoncelli, B.J. Ferguson, et al. 2015. Identification of the primary lesion of toxic aluminum in plant roots.
357 Plant Physiol. 167(4): 1402–1411. doi: 10.1104/pp.114.253229.
358 Matonyei, T.K., B.A. Barros, R.G.N. Guimaraes, E.O. Ouma, R.K. Cheprot, et al. 2020. Aluminum tolerance mechanisms in Kenyan maize germplasm
359 are independent from the citrate transporter ZmMATE1. Sci. Rep. 10(1): 1–10. doi: 10.1038/s41598-020-64107-z.
360 Maulana, A.I., M. Ghulamahdi, and I. Lubis. 2019. Response of corn varieties under saturated soil culture and temporary flooding on tidal swamp. J.
361 Trop. Crop Sci. 6(1): 41–49. doi: 10.1016/j.proenv.2016.03.060.
362 Metsalu, T., and J. Vilo. 2015. ClustVis: A web tool for visualizing clustering of multivariate data using Principal Component Analysis and heatmap.
363 Nucleic Acids Res. 43(W1): W566–W570. doi: 10.1093/nar/gkv468.
364 Mursyidin, D.H., and I. Khairullah. 2020. Genetic evaluation of tidal swamp rice from south kalimantan, indonesia based on the agro-morphological
365 markers. Biodiversitas 21(10): 4795–4803. doi: 10.13057/biodiv/d211045.
366 Nikolic, M., and J. Pavlovic. 2018. Plant responses to iron Deficiency and toxicity and iron use efficiency in plants. Elsevier Inc.
367 Nugraha, Y., S.W. Ardie, I.A. Rumanti, S. Suwarno, M. Ghulammahdi, et al. 2016. Responses of selected Indonesian rice varieties under excess iron
368 condition in media culture at seedling stage. J. Penelit. Pertan. Tanam. Pangan 35(3): 181. doi: 10.21082/jpptp.v35n3.2016.p181-190.
369 Pujiwati, H., M. Ghulamahdi, S. Yahya, S.A. Aziz, and O. Haridjaja. 2015. The application of peaty mineral soil water in improving the adaptability of
370 black soybean toward aluminium stress on tidal mineral soil with saturated water culture. Agrivita 37(3): 284–289. doi: 10.17503/Agrivita-2015-
371 37-3-p284-289.
372 Pujiwati, H., M. Ghulamahdi, S. Yahya, S.A. Aziz, and O. Haridjaja. 2016. Productivity of three soybean genotypes with different water and water depth
373 on various land conditions in tidal swamp. (Produktivitas tiga genotipe kedelai dengan air berbeda dan kedalaman muka air pada berbagai kondisi
374 tanah di pasang surut). J. Agron Indones. 44(3): 248–254.
375 Sabagh, A. El, A. Hossain, C. Barutçular, A.A. Khaled, S. Fahad, et al. 2018. Sustainable maize (Zea Mays L.) production under drought stress by
376 understanding its adverse effect, survival mechanism and drought tolerance indices. J. Exp. Biol. Agric. Sci. 6(2). doi:
377 10.18006/2018.6(2).xxx.xxx.
378 dos Santos, R.S., A.T. De Araujo, C. Pegoraro, and A.C. de Oliveira. 2017. Dealing with iron metabolism in rice: From breeding for stress tolerance to
379 biofortification. Genet. Mol. Biol. 40(1): 312–325. doi: 10.1590/1678-4685-gmb-2016-0036.
380 Sayo, S., K.O. David, J.A. Omolayo, M. Silvestro, S.F. Lawrence, et al. 2017. Correlation and path coefficient analysis of top-cross and three-way cross
381 hybrid maize populations. African J. Agric. Res. 12(10): 780–789. doi: 10.5897/ajar2016.11997.
382 Sesay, S., D.K. Ojo, O.J. Ariyo, and S. Meseka. 2016. Genetic variability, heritability and genetic advance studies in top-cross and three-way cross maize
383 (Zea Mays L) hybrids. Maydica 61(2): 1–7.
384 Surahman, M., M. Ghulamahdi, Murdianto, Prastowo, Sutrisno, et al. 2018. Five steps toward the Indonesian soybean self-sufficiency. IOP Conference
385 Series: Earth and Environmental Science 196 (Forum IPIMA). p. 1–7
386 Toyip, M. Ghulamahdi, D. Sopandie, S.A. Aziz, A. Sutandi, et al. 2019. Physiological responses of four soybean varieties and their effect to the yield in
387 several saturated soil culture modification. Biodiversitas 20(8): 2266–2272. doi: 10.13057/biodiv/d200822.
388 Vahedian, A., S.A. Aghdaei, and S. Mahini. 2014. Acid Sulphate Soil Interaction with Groundwater: A Remediation Case Study in East Trinity.
389 APCBEE Procedia 9(Icbee 2013): 274–279. doi: 10.1016/j.apcbee.2014.01.049.
390 Wang, N., Q. Huang, J. Sun, S. Yan, C. Ding, et al. 2014. Shade tolerance plays an important role in biomass production of different poplar genotypes in
391 a high-density plantation. For. Ecol. Manage. 331: 40–49. doi: 10.1016/j.foreco.2014.06.046.
392 Wening, R.H., B.S. Purwoko, W.B. Suwarno, I.A. Rumanti, and D.N. Khumaida. 2020. Simultaneous selection of leaf drying and yield traits on rice lines
393 (Seleksi simultan karakter daun mengering dan produktivitas pada galur-galur padi). J. Agron. Indones. (Indonesian J. Agron. 47(3): 232–239.
394 doi: 10.24831/jai.v47i3.26076.
395 Xu, L., W. Liu, B. Cui, N. Wang, J. Ding, et al. 2017. Aluminium tolerance assessment of 141 maize germplasms in a solution culture. Univers. J. Agric.
396 Res. 5(1): 1–9. doi: 10.13189/ujar.2017.050101.
397
398
Biodiversitas Journal of Biological Diversity Tasks 0  English  View Site  warti

Notifications 
11449 / SUWARTI et al. / Secondary trait and index selection determinati Library

[biodiv] Editor Decision

2022-07-20 12:12 PM
Submissions Workflow Publication

Submission Review Copyediting Production

Suwarti Mujiharto, Ghulamahdi:

We have reached a decision regarding your submission to Biodiversitas Journal of Biological Diversity,
Round 1 Determination
"Secondary Trait and Index Selection Round 2 for Maize
Round 3
Genotype Selection on Tidal Swamp Acid
Environment".

Our decision is: Revisions Required

Round 3 Status
Submission accepted.

Notifications
------------------------------------------------------
Reviewer A:
[biodiv] Editor Decision 2022-06-21 09:24 AM

This article contains comprehensive data sufficient for publication as a research article in Biodiversitas.
[biodiv] Editor Decision 2022-07-19 06:54 AM
The article has been well written, the data are solid and well presented, the discussion is comprehensive,
and the results contain novelty[biodiv]
in the field.
EditorHowever,
Decisionthere is still inappropriate use of terms in the text,
2022-07-20 12:12 PM
which is unusual in the plant breeding subject. Please see the corrections for clarification and revision.
[biodiv] Editor Decision 2022-08-09
need 05:29
AM
Additional comments and editorial corrections are also provided for revision. All the references to
be rewritten following the journal Guidance for Author.
[biodiv] Editor Decision 2022-08-12 03:48 AM

Recommendation: Revisions Required

[biodiv] Editor Decision 2022-08-12 03:57 AM

------------------------------------------------------

Reviewer's Attachments  Search

No Files

________________________________________________________________________
Biodiversitas Journal of Biological Diversity

Revisions
 e so s
Biodiversitas Journal of Biological Diversity Tasks 0  English  View Site  warti
 Search Upload File
Notifications
No Files

[biodiv] Editor Decision
Review Discussions Add discussion

2022-07-20 12:12 PM Name From Last Reply Replies Closed

— warti warti 0
2022-07-14 2022-07-14

Suwarti Mujiharto, Ghulamahdi: 01:48 AM 03:43 AM

— warti - 0
We have reached a decision regarding your submission to Biodiversitas Journal of Biological Diversity,
2022-07-14
"Secondary Trait and Index Selection Determination for Maize Genotype Selection on Tidal Swamp Acid
03:43 AM
Environment".
— warti - 0
Our decision is: Revisions Required 2022-07-29
06:43 AM

Uncorrected dewinurpratiwi warti 1

Proof 2022-08-04 2022-08-06
05:18 AM 11:51 PM
------------------------------------------------------
Reviewer A: BILLING dewinurpratiwi dewinurpratiwi 2
2022-08-04 2022-08-09
This article contains comprehensive data sufficient for05:20
publication
AM as a research article in Biodiversitas.
12:08 AM
The article has been well written, the data are solid and well presented, the discussion is comprehensive,
and the results contain novelty in the field. However, there is still inappropriate use of terms in the text,
which is unusual in the plant breeding subject. Please see the corrections for clarification and revision.
Additional comments and editorial corrections are also provided for revision. All the references need to
be rewritten following the journal Guidance for Author.

Recommendation: Revisions Required

------------------------------------------------------

________________________________________________________________________
Biodiversitas Journal of Biological Diversity
Biodiversitas Journal of Biological Diversity Tasks 0  English  View Site  warti

Notifications 
11449 / SUWARTI et al. / Secondary trait and index selection determinati Library

[biodiv] Editor Decision

2022-08-09 05:29 AM
Submissions Workflow Publication

Submission Review Copyediting Production

SUWARTI, MUNIF GHULAMAHDI, DIDY SOPANDIE, TRIKOESOEMANINGTYAS, EKO SULISTYONO,
MUHAMMAD AZRAI:

We have reached a decisionRound 1 yourRound

regarding 2
submission to Round 3
Biodiversitas Journal of Biological Diversity,
"Secondary trait and index selection determination for maize genotype selection in an acidic tidal swamp
environment".

Round 3 Status
Our decision is to: Accept Submission
Submission accepted.

Notifications

[biodiv] Editor Decision

________________________________________________________________________ 2022-06-21 09:24 AM

Biodiversitas Journal of Biological Diversity

[biodiv] Editor Decision 2022-07-19 06:54 AM

[biodiv] Editor Decision 2022-07-20 12:12 PM

[biodiv] Editor Decision 2022-08-09 05:29 AM

[biodiv] Editor Decision 2022-08-12 03:48 AM

[biodiv] Editor Decision 2022-08-12 03:57 AM

Reviewer's Attachments  Search

No Files

Revisions
 e so s
Biodiversitas Journal of Biological Diversity Tasks 0  English  View Site  warti
 Search Upload File
Notifications
No Files

[biodiv] Editor Decision
Review Discussions Add discussion

2022-08-09 05:29 AM Name From Last Reply Replies Closed

— warti warti 0
2022-07-14 2022-07-14
01:48 AM 03:43EKO
SUWARTI, MUNIF GHULAMAHDI, DIDY SOPANDIE, TRIKOESOEMANINGTYAS, AMSULISTYONO,

MUHAMMAD AZRAI:
— warti - 0
2022-07-14
We have reached a decision regarding your submission to Biodiversitas Journal of Biological Diversity,
03:43 AM
"Secondary trait and index selection determination for maize genotype selection in an acidic tidal swamp
environment". — warti - 0
2022-07-29
Our decision is to: Accept Submission 06:43 AM

Uncorrected dewinurpratiwi warti 1

Proof 2022-08-04 2022-08-06
05:18 AM 11:51 PM

BILLING dewinurpratiwi dewinurpratiwi 2

________________________________________________________________________
2022-08-04 2022-08-09
Biodiversitas Journal of Biological Diversity 05:20 AM 12:08 AM
Biodiversitas Journal of Biological Diversity Tasks 0  English  View Site  warti

Notifications 
11449 / SUWARTI et al. / Secondary trait and index selection determinati Library

[biodiv] Editor Decision

2022-08-12 03:48 AM
Submissions Workflow Publication

Submission Review Copyediting Production

SUWARTI, MUNIF GHULAMAHDI, DIDY SOPANDIE, TRIKOESOEMANINGTYAS, EKO SULISTYONO,
MUHAMMAD AZRAI:

Round
The editing of your submission, 1 Round
"Secondary 2 indexRound
trait and 3 determination for maize genotype
selection
selection in an acidic tidal swamp environment," is complete. We are now sending it to production.

Submission URL: https://smujo.id/biodiv/authorDashboard/submission/11449

Round 3 Status
Submission accepted.

Notifications
________________________________________________________________________
[biodiv]
Biodiversitas Journal of Biological Editor Decision
Diversity 2022-06-21 09:24 AM

[biodiv] Editor Decision 2022-07-19 06:54 AM

[biodiv] Editor Decision 2022-07-20 12:12 PM

[biodiv] Editor Decision 2022-08-09 05:29 AM

[biodiv] Editor Decision 2022-08-12 03:48 AM

[biodiv] Editor Decision 2022-08-12 03:57 AM

Reviewer's Attachments  Search

No Files

Revisions
 e so s
Biodiversitas Journal of Biological Diversity Tasks 0  English  View Site  warti
 Search Upload File
Notifications
No Files

[biodiv] Editor Decision
Review Discussions Add discussion

2022-08-12 03:48 AM Name From Last Reply Replies Closed

— warti warti 0
2022-07-14 2022-07-14
01:48 AM 03:43EKO
SUWARTI, MUNIF GHULAMAHDI, DIDY SOPANDIE, TRIKOESOEMANINGTYAS, AMSULISTYONO,

MUHAMMAD AZRAI:
— warti - 0
2022-07-14
The editing of your submission, "Secondary trait and index selection determination for maize genotype
03:43 AM
selection in an acidic tidal swamp environment," is complete. We are now sending it to production.
— warti - 0
Submission URL: https://smujo.id/biodiv/authorDashboard/submission/11449
2022-07-29
06:43 AM

Uncorrected dewinurpratiwi warti 1

Proof 2022-08-04 2022-08-06
05:18 AM 11:51 PM

________________________________________________________________________
BILLING dewinurpratiwi dewinurpratiwi 2
Biodiversitas Journal of Biological Diversity 2022-08-04 2022-08-09
05:20 AM 12:08 AM