Evaluation of stem borer resistant maize genotypes for resistance to fall armyworm (Spodoptera frugiperda J.E. SMITH) infestation

Abstract

Fall armyworm (FAW) is currently the most destructive insect pest of maize in sub-Saharan Africa (SSA). Varieties that combine high grain yield (GY) with tolerance to FAW would enhance and stabilize maize productivity in SSA. Genotypes resistant to lepidopteran pests like stem borer (SB) could serve as potential sources of alleles for development of FAW resistant varieties. This study was conducted to assess some SB-resistant maize genotypes for FAW tolerance, and to identify genotypes that combined high GY with tolerance to FAW. Twenty-nine white maize genotypes with varying levels of SB resistance were evaluated under artificial FAW-infested and FAW-protected conditions using randomized complete block design with three replicates. Genotypic differences were significant for all the traits under both FAW-infested and FAW-protected conditions. Under FAW-infested condition, GY ranged from 3.44 (FAWTH-8) to 5.81 t ha-1 (FAWTH-1) (mean = 4.61 t ha-1), and from 3.42 (FAWTH-25) to 6.85 t ha-1 (FAWTH-18) (mean = 4 .86 t ha-1) under FAW-protected condition. Across genotypes, FAW infestation reduced GY by 5.1% suggesting that SB resistance could confer tolerance to FAW. Association of GY under FAW-infested condition with FAW Leaf Damage (FAWLD; r=-0.45) and FAW Ear Damage (FAWED; r=-0.65) were significant. Base index (BI) was significantly correlated with GY (r=0.93), ear aspect (r=-0.84), FAWLD (r=-0.66) and FAWED (r=-0.78). Six moderately resistant genotypes (FAWTH-1, FAWTH-13, FAWTH-4, FAWTH-10, FAWTH-23 and FAWTH-6) with GY ≥ 5.13 t ha-1 and positive BI ≥ 4.0 were identified. The genotypes varied for FAW tolerance. Base index and low FAW damage scores could serve as selection criteria for combined tolerance to FAW and high GY. The identified genotypes are recommended for further development as FAW tolerant varieties.

Full text

Vol. 15(3), pp. 99-109, July-September 2023
DOI: 10.5897/JPBCS2023.1023
Article Number: E06FE7F71286
ISSN 2006-9758
Copyright ©2023
Author(s) retain the copyright of this article
http://www.academicjournals.org/JPBCS
Journal of Plant Breeding and Crop
Science
Full Length Research Paper
Evaluation of stem borer resistant maize genotypes for
resistance to fall armyworm (Spodoptera frugiperda
J.E. SMITH) infestation
Nesma Alaaeldin Zakaria Moussa1, Ayodeji Abe2*, Anthony Oluwatoyosi Job2,3,
Yinka Odunayo Kolawole2 and Amudalat Bolanle Olaniyan2
1Pan African University Life and Earth Sciences Institute (Including Health and Agriculture), Pan African University,
Ibadan 200284, Nigeria.
2Department of Crop and Horticultural Sciences, Faculty of Agriculture, University of Ibadan, Ibadan 200284, Nigeria.
3Value Seeds Limited, Kaduna State, Nigeria.
Received 6 June, 2023; Accepted 13 September, 2023
Fall armyworm (FAW) is currently the most destructive insect pest of maize in sub-Saharan Africa
(SSA). Varieties that combine high grain yield (GY) with tolerance to FAW would enhance and stabilize
maize productivity in SSA. Genotypes resistant to lepidopteran pests like stem borer (SB) could serve
as potential sources of alleles for development of FAW resistant varieties. This study was conducted to
assess some SB-resistant maize genotypes for FAW tolerance, and to identify genotypes that combined
high GY with tolerance to FAW. Twenty-nine white maize genotypes with varying levels of SB resistance
were evaluated under artificial FAW-infested and FAW-protected conditions using randomized complete
block design with three replicates. Genotypic differences were significant for all the traits under both
FAW-infested and FAW-protected conditions. Under FAW-infested condition, GY ranged from 3.44
(FAWTH-8) to 5.81 t ha-1 (FAWTH-1) (mean = 4.61 t ha-1), and from 3.42 (FAWTH-25) to 6.85 t ha-1
(FAWTH-18) (mean = 4 .86 t ha-1) under FAW-protected condition. Across genotypes, FAW infestation
reduced GY by 5.1% suggesting that SB resistance could confer tolerance to FAW. Association of GY
under FAW-infested condition with FAW Leaf Damage (FAWLD; r=-0.45) and FAW Ear Damage (FAWED;
r=-0.65) were significant. Base index (BI) was significantly correlated with GY (r=0.93), ear aspect (r=-
0.84), FAWLD (r=-0.66) and FAWED (r=-0.78). Six moderately resistant genotypes (FAWTH-1, FAWTH-13,
FAWTH-4, FAWTH-10, FAWTH-23 and FAWTH-6) with GY ≥ 5.13 t ha-1 and positive BI ≥ 4.0 were
identified. The genotypes varied for FAW tolerance. Base index and low FAW damage scores could
serve as selection criteria for combined tolerance to FAW and high GY. The identified genotypes are
recommended for further development as FAW tolerant varieties.
Key words: Base index, fall armyworm ear damage, fall armyworm leaf damage, maize grain yield, stem borer
resistance.
INTRODUCTION
Maize (Zea mays L.) is one of the most important staple
food and industrial crop in sub-Saharan Africa (SSA)
where it contributes to the livelihoods and food security of
smallholder farmers (Erenstein et al., 2022). In SSA
human consumption accounts for about 63% of maize
produced (Santpoort, 2020) which supplies about 30% of
the food calorie requirement of more than 300 million
people (Shiferaw et al., 2011; Smale et al., 2013; Beyene

100 J. Plant Breed. Crop Sci.
et al., 2016). Although maize has a high yield potential in
SSA (IITA, 2017), average maize grain yield is very low
(1.5 t ha-1) compared to global average of 4.9 t ha-1
(OECD, 2018; FAOSTAT, 2021; Grote et al., 2021).
The low maize productivity and production in SSA is a
function of several biotic (e.g. Striga spp., foliar diseases,
insect pests including stem borers and fall armyworms)
and abiotic (e.g. drought, flood. heat and low soil fertility)
stress factors, as well as socio-economic restrictions
which included fragmented pieces of land, unaffordable
input costs (OECD, 2018), wars and terrorism among
others.
Of all the biotic constraints to maize productivity, insect
pests alone cause an estimated 60% of yield losses in
SSA (Mugo et al., 2018). No insect pests of economic
importance to maize production include Busseola fusca
(African stem borer), Eldana saccharina (African
sugarcane borer), Sesamia calamistis (African pink stem
borer), Chilo partellus (Spotted stem borer), Cicadulina
mbila (Maize leafhopper), some termite species
(Macrotermes and Microtermes species), and more
recently Spodoptera frugiperda (fall armyworm) (Assefa
and Ayalew, 2019). However, stem borers and fall
armyworm (FAW) are the two most important insect pests
of maize in SSA (Ajala et al., 2008; Nagoshi et al., 2017;
Job et al., 2022). The FAW is a highly polyphagous,
invasive pest of global economic importance (Kasoma et
al., 2021a; Matova et al., 2020; Overton et al., 2021) with
a wide host range cutting across over 80 species and
more than 353 plants (Prasanna, 2018; Wan et al., 2021).
Although native to tropical and subtropical regions of
America, FAW was first reported in West Africa in 2016
and has spread rapidly to other regions of the continent
(Goergen et al., 2016; Cock et al., 2017; Tepa-Yotto et
al., 2021). Currently, FAW has assumed the status of the
most destructive, yield-limiting insect pest of maize in
SSA where it causes severe grain yield losses, thereby
becoming a grave threat to food and livelihood security
(Day et al., 2017; Abrahams et al., 2017; Prasanna et al.,
2018; Kumela et al., 2019; Matova et al., 2022).
Depending on the plant’s genetic make-up, extent of
infestation and in the absence of appropriate control
measures, maize grain yield losses due to FAW can be
up to 100% (Prasanna et al., 2018). It is therefore
necessary to design an effective management strategy to
control FAW infestation in farmers’ fields to avert high
grain yield losses on farmers’ fields in SSA.
Common control strategies to FAW attack includes the
use of insecticides, biological control agents, cultural
practices and host plant resistance (Prasanna et al.,
2018, 2021). However, several factors including costs
and legislative barriers hinder availability and use of
these FAW control measures by most smallholder African
farmers. Furthermore, independent deployment of each
of the control strategies in SSA is not neither sustainable
nor effective. Therefore, a multifaceted approach which
includes the use tolerant/resistant varieties is required to
provide a durable and sustainable FAW management in
SSA (Prasanna et al., 2022).
Host-plant resistance is economic, sustainable,
environmentally friendly and compatible with other pest
management strategies (Abrahams et al., 2017; Kumela
et al., 2019; Job et al., 2022; Prasanna et al., 2022).
However, only few commercial maize cultivars with
resistance to FAW are available in Africa. Since the mode
of action of FAW and that of stem borers are very similar,
and significant correlations have been reported between
the resistance indices of both pests (Williams et al., 1998;
Abel et al., 2000; Prasanna et al., 2018), it could be
considered that stem borer resistance would confer
resistance to FAW. Furthermore, resistance to insect
pests in maize has been shown to be genetically broad-
based suggesting that resistance of some maize
genotypes to a given insect pest could influence their
resistance to another insect pest (Brooks et al., 2005).
Hence, evaluating maize genotypes developed for stem
borer resistance or tolerance under FAW infestation will
provide a basis for selecting best performing ones.
Therefore, the present study was carried out to: evaluate
some stem borer resistant tropical white maize genotypes
for their grain yield performance under fall armyworm
infested and protected conditions and to identify and
select maize genotypes that combined high grain yield
with tolerance to fall armyworm infestation.
MATERIALS AND METHODS
Description of experimental site
The experiment was carried out at the experimental field of the
Department of Crop and Horticultural Sciences, along Parry Road,
University of Ibadan (N07.45164°, E003.8906; 208 masl), Oyo
State, Nigeria in two cropping seasons. The location of the
experimental site is characterized by high incidence of fall
armyworm (FAW) infestations both on and off season. The soil at
the experimental site was sandy-loam with a pH (H2O) of 5.5. It was
low in total nitrogen (1.30 g kg-1), available P (0.75 mg kg-1) and K
(0.28 cmol kg-1), while the organic carbon (12.90 g kg-1) was
moderate.
Genetic materials used in the experiment
Twenty-nine white maize genotypes which comprised 19 top cross
hybrids, three single cross hybrids, one population cross hybrid and
six open pollinated varieties of similar maturity and varying levels of
resistance to stem borers were used for the study (Table 1). The
genetic materials were sourced from Value Seeds Ltd, Kaduna,
*Corresponding author. E-mail: a.abe@ui.edu.ng; ayodabe@yahoo.com.
Author(s) agree that this article remain permanently open access under the terms of the Creative Commons Attribution
License 4.0 International License

Moussa et al. 101
Table 1. Genetic materials used in the experiment.
S/N
Code
Pedigree
Type
1
FAWTH-1
AWRSYN-W2 × 1393
Top cross
2
FAWTH-2
AWRSYN-W2 × CML 331
Top cross
3
FAWTH-3
AWRSYN-W2 × CKSBL 10060
Top cross
4
FAWTH-4
AMATZBR-WC3 × 1393
Top cross
5
FAWTH-5
AMATZBR-WC3 × CML 331
Top cross
6
FAWTH-6
AMATZBR-WC3 × CKSBL 10060
Top cross
7
FAWTH-7
TZBR-ELd4WC2 × 1393
Top cross
8
FAWTH-8
TZBR-EL4WC2 × CML 331
Top cross
9
FAWTH-9
TZBR-EL4WC2 × CKSBL 10060
Top cross
10
FAWTH-10
TZBSR X 1393
Top cross
11
FAWTH-11
TZBSR × CML 331
Top cross
12
FAWTH-12
TZBSR × CKSBL 10060
Top cross
13
FAWTH-13
TZBR Comp 1- WC2 × 1393
Top cross
14
FAWTH-14
TZBR Comp 1-WC2 × CML 331
Top cross
15
FAWTH-15
TZBR Comp 1-WC2 × CKSBL 10060
Top cross
16
FAWTH-16
TZBR Comp 2-WC2 × 1393
Top cross
17
FAWTH-17
TZBR Comp 2-WC2 × CML 331
Top cross
18
FAWTH-18
TZBR Comp 2-WC2 × CKSBL 10060
Top cross
19
FAWTH-19
1393 × AbSL50
Single cross
20
FAWTH-20
1393 × CML 331
Single cross
21
FAWTH-21
1393 × CKSBL 10060
Single cross
22
FAWTH-22
AWRSYN-W2
OPV
23
FAWTH-23
TZBR Comp 1-WC2
OPV
24
FAWTH-24
TZBR Comp 2-WC2
OPV
25
FAWTH-25
TZBR ELd4-WC2
OPV
26
FAWTH-26
AMATZBR-WC3
OPV
27
FAWTH-27
Sammaz 15
OPV
28
FAWTH-28
SC 651
Top cross
29
FAWTH-29
TZBR Comp-1 WC2 × TZBR Comp-2 WC2
Population cross
OPV = Open pollinated variety.
Nigeria.
Experimental design, crop establishment and management
The experiment was laid out in a randomized complete block
design with three replicates. The experimental field was divided into
two blocks, namely: FAW-infested and FAW-protected. The FAW-
infested block was artificially infested with FAW larvae, while the
FAW-protected block served as control. The two blocks were
separated by a 10.0 m alley to which seeds of a maize population
of similar maturity with the test genotypes were densely sown to
trap insecticide spray drifts from the protected block. Plots
consisted of single rows, each 3.0 m long and spaced 0.75 m.
Seeds were sown 0.25 m apart within rows. Two seeds were sown
per hole and the seedlings later thinned to one plant at two weeks
after sowing (WAS) to achieve a plant population of 53,333 plants
per hectare. The plants were grown under rain-fed conditions, with
supplemental drip irrigation when necessary to prevent drought
stress. Weeds were controlled using a combination of atrazine (250
g L-1) and S-metolachlor (250 g L-1 SC) at the rate of 4.0 L ha-1 as
pre-emergence herbicide, and this was complemented with one
round of hand weeding. At 2WAS, NPK 15:15:15 fertilizer was
applied at the rate of 40 kg N ha-1. An additional 30 kg N ha-1 was
applied at 5WAS using urea. The FAW larvae were raised at the
International Institute for Tropical Agriculture (IITA), Ibadan, Nigeria.
Artificial infestation of the FAW-infested block was done two WAS
by using a camel brush to transfer ten second instar FAW larvae
into the inner whorl of each maize seedling. To ensure uniformity,
infestation of plots was done on the same day. Control of FAW on
the FAW-protected block was achieved by spraying the plants
weekly for five weeks, starting from the second week after sowing,
with the insecticide emamectin benzoate (5% WDG) at the rate of
0.38 g L-1 following manufacturer’s instructions. The densely sown
10 m strip separating the FAW-infested and FAW-protected blocks
was not sprayed and served to harbor FAW larvae and moth.
Data collection
Data were collected on plot basis for all traits under both FAW-
infested and FAW-protected conditions. Under each condition, data
was recorded for days to anthesis (DA) as the number of days from
sowing to the date when 50% of the plants in a plot shed pollen,
while days to silking (DS) was recorded as the number of days from
sowing to the date when 50% of the plants in a plot have emerged

102 J. Plant Breed. Crop Sci.
silks. Anthesis-silking interval (ASI) was expressed in days as the
difference between DS and DA. Plant height (PH) and ear height
(EH) in cm were measured at physiological maturity as the distance
from soil level to the collar of the uppermost leaf and upper ear leaf,
respectively, of five competitive plants. Plant aspect (PASP) was
scored on a scale of 1 to 9 based on uniformity in plant and ear
heights, lodging characteristics, reaction to pests and diseases,
general appeal etc, where 1 = excellent, and 9 = poor. Husk cover
was scored using a scale of 1 to 9, where 1 = husk tightly covers
ear tip and extends beyond it, and 9 = poor husk cover with ear tip
clearly exposed). Ear aspect (EASP) was also scored on a scale of
1 to 9, where 1 = excellent, clean uniform and well filled ears and 9
= ears with poor phenotypic appearance after harvest. The plants
were also scored for streak disease on a scale of 1 to 9, where 1 =
all plants excellent, clean with no streak infection and 9 = all plants
severely streak infected. At harvest, the number of plants per plot
was recorded.
Scoring for FAW leaf damage (FAWLD) and FAW ear damage
(FAWED) were done only on the FAW-infested plots at 6WAS and
at harvest, respectively. The FAWLD and FAWED were rated on a
scale of 1 to 9, where 1 = no visible damage, and 9 = severe
damage (Davis et al., 1989, 1992; Prasanna et al., 2018).
Grain yield (GY) was estimated by harvesting all the ears in a
plot and shelled. The fresh weight adjusted for number of plants at
harvest and percent moisture content of shelled grains were used
to estimate GY and reported in kg ha-1 adjusted to 15% moisture
content using the formula below:
Grain yield ( )
=
Data analyses
All data analyses were conducted separately for the FAW-infested
and FAW-protected conditions. Analyses of variance were done
using the PROC MIXED procedure in SAS version 9.3 (SAS
Institute, 2011). Seasons of evaluation were considered as
separate environments. In the model, genotype and genotype ×
environment were random, while environments and replications
within environments were fixed. Significance was declared at 5%
level of probability.
Base index (BI) approach by Badu-Apraku et al. (2015) and
Oloyede-Kamiyo (2019) was used, with slight modifications to
identify FAW tolerant and FAW susceptible genotypes. Traits
included in the estimation of BI were GY, FAWLD, FAWED, EASP,
and PASP as earlier described. To reduce the effect of differences
in scales of measurement of the traits under FAW infestation, all
data were standardized prior to integration into the BI equation. The
BI values were calculated as:
A genotype with a positive BI was considered FAW tolerant,
whereas negative BI indicated susceptibility to FAW. Correlation
analyses were carried out to establish the relationships among
measured traits, and between BI and the traits included in the
selection criteria using PROC CORR in SAS version 9.3 (SAS
Institute, 2011).
RESULTS
Trait variability among genotypes
The analysis of variance revealed highly significant (p ≤
0.01) differences among the genotypes for all traits
measured under both FAW-infested and FAW-protected
conditions (Table 2). The effects of environment and
genotype × environment interaction were not significant
for all traits under both conditions.
Genotype responses under FAW-infested and FAW-
protected conditions
Agronomic and fall armyworm damage traits
Under FAW-infested condition, days to anthesis (DA)
ranged from 3.7 (FAWTH-3) to 61.9 days (FAWTH-24)
with a mean of 57.3 days, while days to silking (DS)
ranged from 57.2 (FAWTH-19) to 63.7 days (FAWTH- 20)
with a mean of 59.5 days (Table 3). Across genotypes,
ASI was 2.3 days. Plant and ear heights ranged from
133.8 cm (FAWTH-8) to 185.1 cm (FAWTH-12) and from
69.4 cm (FAWTH-28) to 105.8 cm (FAWTH-23),
respectively. In general, the plant and ear aspects of all
the genotypes were good, while the husk cover score
ranged from very good to moderate. The highest streak
disease score of 6.7 was recorded for genotypes
FAWTH-3, FAWTH-9 and FAWTH-14, while genotype
FAWTH-4 had the least (3.1). The FAWLD was highest in
genotypes FAWTH-5 and FAWTH-8, but least in
genotypes FAWTH-13, FAWTH-19 and FAWTH-27.
Genotypes FAWTH-6, FAWTH-12 and FAWTH-13 had
the least FAWED, while genotypes FAWTH-15 and
FAWTH-26 had the highest FAWED (Table 3).
Under FAW-protected condition, DA and DS ranged
from 53.6 (FAWTH-12) to 61.7 days (FAWTH-17) with a
mean of 54.6 days and from 54.6 (FAWTH-12) to 64.7
days (FAWTH-17) with a mean of 59.7 days. Averaged
across genotypes, ASI was 2.2 days (Table 4). Genotype
FAWTH-2 (155.1 cm) was the shortest, while genotype
FAWTH-24 (194.7 cm) was tallest. Ear height ranged
from 75.2 cm for FAWTH-2 to 101.1 cm for FAWTH-18.
Plant and ear aspect ranged from 2.0 (FAWTH-12) to 4.0
(FAWTH-25) and from 1.7 (FAWTH-18) to 4.3 (FAWTH-
23, FAWTH-25), respectively. Husk cover score among
the genotypes ranged from very good to moderate.
Genotypes FAWTH-1 had the least streak disease score
(2.3), while genotypes FAWTH-12, FAWTH-16 and
FAWTH-20 had the highest score of 6.0 (Table 4).
Grain yield performance
Genotypic differences were observed for grain yield (GY)
under FAW-infested and FAW-protected conditions (5).
Under FAW-infested condition, GY varied from 3.44
(FAWTH-8) to 5.81 t ha-1 (FAWTH-1), whereas under
FAW-protected condition, GY ranged from 3.42 (FAWTH-
25) to 6.85 t ha-1 (FAWTH-18). Across genotypes, mean
GY under FAW-protected condition (4.86 t ha-1) and

Moussa et al. 103
Table 2. Mean squares from analysis of variance for some agronomic traits and fall armyworm damage parameters of 29 white maize genotypes evaluated under fall armyworm infested
and protected conditions for two seasons in Ibadan, Nigeria.
Source of
variation
DF
GY
(t ha-1)
DA
DS
ASI
PH (cm)
EH (cm)
PASP
(1 - 9)
EASP
(1 - 9)
Streak score
(1 - 9)
HCOV
(1 - 9)
FAWLD
(1 – 9)
FAWED
(1 – 9)
Infested
Env
1
0.045
0.000
0.006
0.008
58.279
27.006
0.033
0.011
0.059
0.066
0.695
0.002
Rep(Env)
4
1.857***
25.558***
13.078***
2.143*
1779.279***
621.107***
2.313***
0.288
2.487***
0.564
35.889***
3.156***
Geno
28
1.916***
26.803***
28.319***
5.357***
1465.099***
569.899***
0.846***
1.196***
6.113***
1.102***
2.748***
3.618***
Env*Geno
28
0.016
0.118
0.147
0.077
7.480
3.150
0.017
0.015
0.026
0.019
0.086
0.036
Error
112
0.261
1.334
1.697
0.802
68.279
33.689
0.214
0.285
0.327
0.267
0.801
0.468
Protected
Env
1
0.273
0.266
0.445
0.023
6.829
0.336
0.002
0.006
0.015
0.001
-
-
Rep(Env)
4
2.053***
19.470**
8.624
3.537*
1957.469***
288.048*
0.984**
0.388
0.533
0.162
-
-
Geno
28
3.557***
15.505***
22.230***
4.668***
738.056***
344.360***
1.230***
2.236***
4.924***
1.100***
-
-
Env*Geno
28
0.022
0.244
0.147
0.110
8.895
5.251
0.015
0.022
0.044
0.019
-
-
Error
112
0.233
4.316
4.389
1.284
122.861
83.281
0.200
0.321
0.716
0.244
-
-
*;**;***: significant respectively at 0.05, 0.01, 0.001 probability levels. Env: Environment; Rep: Replication; Geno: Genotype; DF: Degree of freedom; GY: Grain yield; DA: Days to anthesis;
DS: Days to silking; ASI: Anthesis-silking interval; PH: Plant height; EH: Ear height; PASP: Plant aspect; EASP: Ear aspect; HCOV: Husk cover; FAWLD: Fall armyworm leaf damage;
FAWED: Fall armyworm ear damage.
Table 3. Mean performance for some agronomic traits of 29 white maize genotypes evaluated under fall armyworm infested condition for two seasons in Ibadan, Nigeria.
Entry
DA
DS
ASI
PH
(cm)
EH
(cm)
PASP
(1 - 9)
EASP
(1 - 9)
Streak score
(1 - 9)
HCOV
(1 - 9)
Fall army worm damage
FAWLD (1 - 9)
FAWED (1 - 9)
Mean
FAWTH-1
56.0
57.3
1.3
171.9
91.4
3.0
2.6
4.6
2.9
4.2
3.4
3.80
FAWTH-2
57.9
60.8
2.9
148.8
76.6
2.7
3.6
5.6
3.0
5.9
4.9
5.40
FAWTH-3
53.7
56.7
3.0
158.5
86.0
3.0
3.3
6.7
3.6
4.9
4.6
4.75
FAWTH-4
57.0
60.0
3.0
172.7
85.9
2.9
3.0
3.1
4.1
4.3
3.3
3.80
FAWTH-5
56.6
60.6
4.0
134.2
70.3
3.0
3.4
6.0
3.3
6.2
4.7
5.45
FAWTH-6
54.8
57.9
3.1
177.4
90.9
2.4
2.4
5.7
3.5
5.9
3.0
4.45
FAWTH-7
57.6
59.8
2.2
167.0
91.7
2.9
3.0
4.3
2.6
4.3
3.6
3.95
FAWTH-8
59.3
62.6
3.4
133.8
73.4
3.0
3.6
6.3
3.6
6.2
4.1
5.15
FAWTH-9
55.2
57.4
2.2
169.2
84.9
3.1
3.7
6.7
3.4
5.6
4.4
5.00
FAWTH-10
55.2
57.5
2.3
166.4
86.5
3.4
2.7
4.6
4.0
4.6
3.4
4.00
FAWTH-11
58.3
61.4
3.2
149.8
78.4
3.4
3.4
6.0
3.3
5.8
4.1
4.95
FAWTH-12
55.6
57.6
2.0
185.1
97.5
3.0
3.0
6.0
3.4
5.1
3.0
4.05
FAWTH-13
57.4
58.6
1.3
181.9
103.6
2.9
2.4
4.0
4.0
4.0
3.0
3.50

104 J. Plant Breed. Crop Sci.
Table 3. Contd.
FAWTH-14
57.1
59.4
2.3
142.7
72.6
2.6
3.9
6.7
3.5
5.8
5.3
5.55
FAWTH-15
54.4
57.3
2.9
166.2
90.9
2.6
3.7
5.7
3.7
4.7
5.7
5.20
FAWTH-16
59.0
60.9
1.9
180.3
94.1
3.5
2.7
5.6
3.7
4.8
4.4
4.60
FAWTH-17
57.3
60.0
2.7
156.4
83.1
2.7
3.7
6.6
3.6
5.3
4.7
5.00
FAWTH-18
56.1
58.3
2.2
176.8
87.1
2.6
2.9
6.0
3.4
4.8
3.5
4.15
FAWTH-19
55.2
57.2
2.0
170.1
85.0
3.0
3.3
5.4
4.4
4.0
3.9
3.95
FAWTH-20
60.5
63.7
3.2
138.6
70.6
3.0
3.4
5.6
4.0
5.2
4.5
4.85
FAWTH-21
53.7
55.5
1.7
147.3
82.0
3.4
3.4
6.4
2.9
5.5
4.1
4.80
FAWTH-22
56.0
57.6
1.6
146.7
70.1
3.3
2.6
5.4
4.0
5.0
4.2
4.60
FAWTH-23
59.9
60.3
0.4
180.5
105.6
3.0
3.0
4.4
4.0
4.2
3.6
3.90
FAWTH-24
61.9
62.3
0.4
185.0
90.9
2.6
3.4
4.4
4.3
5.1
3.9
4.50
FAWTH-25
58.2
58.4
0.3
148.8
75.9
2.3
3.7
3.9
3.6
5.2
4.7
4.95
FAWTH-26
58.4
61.6
3.3
148.2
76.3
3.0
4.0
5.7
3.6
5.5
5.7
5.60
FAWTH-27
59.8
62.2
2.4
170.0
81.5
4.0
3.0
3.6
3.0
4.0
3.6
3.80
FAWTH-28
60.0
63.2
3.2
161.7
69.4
2.6
3.4
6.0
3.6
5.4
3.0
4.20
FAWTH-29
58.4
60.1
1.7
173.0
85.9
3.6
3.4
4.6
3.6
4.5
3.7
4.10
Mean
57.3
59.5
2.3
162.4
84.1
3.0
3.2
5.4
3.6
5.0
4.1
4.55
SED
0.67
0.75
0.52
4.77
3.35
0.27
0.31
0.33
0.30
0.52
0.39
CV (%)
2.0
2.2
39.5
5.1
6.9
15.5
16.5
9.0
14.4
17.8
16.8
DA: Days to anthesis; DS: Days to silking; ASI: Anthesis-silking interval; PH: Plant height; EH: Ear height; PASP: Plant aspect; EASP: Ear aspect; HCOV: Husk cover; FAWLD: Fall
armyworm leaf damage; FAWED: Fall armyworm ear damage; BI: Base index; SED: Standard error of the difference; CV: Coefficient of variation.
Table 4. Mean performance for some agronomic traits of 29 white maize genotypes evaluated under fall armyworm protected condition for two seasons in
Ibadan, Nigeria.
Entry
DA
DS
ASI
PH (cm)
EH (cm)
PASP (1 - 9)
EASP (1 - 9)
Streak score (1 - 9)
HCOV (1 - 9)
FAWTH-1
56.5
59.7
3.3
184.9
97.3
3.4
2.4
2.3
2.6
FAWTH-2
57.5
61.1
3.6
155.1
75.2
3.0
3.6
5.5
3.4
FAWTH-3
55.8
58.0
2.2
175.4
92.2
3.0
2.6
4.7
3.0
FAWTH-4
56.4
59.7
3.3
194.5
98.6
3.4
3.0
5.1
3.0
FAWTH-5
57.1
58.4
1.3
175.4
91.1
3.0
2.7
5.4
3.1
FAWTH-6
57.6
60.7
3.2
171.4
86.2
2.6
3.4
5.0
3.0
FAWTH-7
57.3
60.8
3.5
183.2
98.5
3.7
3.0
5.3
4.0
FAWTH-8
58.6
60.5
1.8
165.2
80.2
3.0
3.4
4.7
3.6

Moussa et al. 105
Table 4. Mean performance for some agronomic traits of 29 white maize genotypes evaluated under fall armyworm protected condition for two
seasons in Ibadan, Nigeria.
FAWTH-9
55.4
57.5
2.1
171.5
85.6
3.0
2.6
5.5
3.4
FAWTH-10
56.9
59.4
2.5
183.1
88.9
3.4
2.7
5.1
2.7
FAWTH-11
56.9
58.2
1.3
171.9
92.2
3.4
2.6
5.4
3.0
FAWTH-12
53.6
54.6
1.0
185.5
93.3
2.0
2.6
6.0
3.0
FAWTH-13
56.9
58.5
1.6
182.0
94.7
2.7
2.4
3.0
3.4
FAWTH-14
58.9
62.4
3.5
183.9
93.7
2.6
3.3
5.7
4.0
FAWTH-15
56.8
58.6
1.8
160.4
80.1
3.3
3.9
5.6
3.4
FAWTH-16
57.4
60.3
2.8
184.5
95.8
2.6
2.4
6.0
3.2
FAWTH-17
61.7
64.7
3.0
166.7
76.8
3.1
3.6
5.0
4.0
FAWTH-18
55.4
56.3
0.9
191.1
101.0
3.6
1.7
3.7
3.4
FAWTH-19
58.8
60.9
2.1
179.3
85.5
3.7
3.0
4.6
2.6
FAWTH-20
56.8
59.9
3.1
163.4
85.2
3.7
3.5
6.0
3.4
FAWTH-21
56.2
59.0
2.8
166.6
87.2
3.3
3.0
4.8
3.0
FAWTH-22
59.3
61.0
1.7
166.2
81.0
2.7
3.5
5.1
3.3
FAWTH-23
59.7
60.1
0.4
156.9
83.9
2.7
4.3
4.2
4.4
FAWTH-24
58.0
60.3
2.4
194.7
97.8
2.6
3.3
4.5
3.5
FAWTH-25
57.8
59.2
1.4
164.3
84.8
4.0
4.3
5.6
3.7
FAWTH-26
56.0
57.7
1.7
184.9
95.8
3.3
2.3
3.7
3.4
FAWTH-27
59.6
61.5
1.9
167.3
79.6
3.6
3.3
5.9
3.0
FAWTH-28
58.3
60.0
1.7
164.2
75.3
2.6
2.6
5.9
2.9
FAWTH-29
58.2
61.0
2.8
180.8
93.1
2.9
2.3
4.6
3.0
Mean
57.4
59.7
2.2
175.0
88.6
3.1
3.0
5.0
3.3
SED
1.20
1.21
0.65
6.40
5.27
0.26
0.33
0.49
0.29
CV
3.6
3.5
50.8
6.3
10.3
14.4
18.8
14.2
15.0
DA: Days to anthesis; DS: Days to silking; ASI: Anthesis-silking interval; PH: Plant height; EH: Ear height; PASP: Plant aspect; EASP: Ear aspect; HCOV:
Husk cover; SED: Standard error of the difference; CV: Coefficient of variation.
FAW-infested condition (4.61 t ha-1) indicated a
GY reduction of 5.1% due to FAW infestation.
Under both FAW-infested and FAW-protected
conditions, the top 15 genotypes in each case had
GY higher than the respective means (Table 5).
Using BI as selection criteria, the BI of the
maize genotypes ranged from –6.80 (FAWTH-
8) to 8.07 (FAWTH-13) (Table 5). The top nine
high yielding genotypes had high positive BIs
which ranged from 3.16 (FAWTH-28) to 8.07
(FAWTH-13). The GY of the top nine genotypes
ranged from 4.90 (FAWTH-18) to 5.81 t ha-1
(FAWTH-1) with a mean of 5.27 t ha-1. The bottom
five genotypes had negative BIs ranging from –
6.80 for FAWTH-8 to -3.16 for FAWTH-20.
Associations of base index with selection
indices under FAW-infested condition
Base index had significant positive relationship

106 J. Plant Breed. Crop Sci.
Table 5. Grain yield performance of 29 white maize genotypes evaluated under fall armyworm
infested and protected conditions for two seasons in Ibadan, Nigeria.
Entries
Grain yield (t ha-1)
Yield reduction (%)
BI
Protected
Infested
FAWTH-13
5.26
5.48
-4.2
8.07
FAWTH-1
5.4
5.81
-7.6
7.71
FAWTH-4
4.41
5.46
-23.8
5.82
FAWTH-6
5.24
5.13
2.1
5.34
FAWTH-23
3.97
5.2
-31
4.39
FAWTH-10
4.97
5.28
-6.2
3.97
FAWTH-18
6.85
4.9
28.5
3.85
FAWTH-12
5.67
5.11
9.9
3.52
FAWTH-28
4.73
5.09
-7.6
3.16
FAWTH-7
4.52
4.69
-3.8
2.7
FAWTH-22
4.19
4.77
-13.8
1.02
FAWTH-19
6.71
4.36
35.0
0.65
FAWTH-27
4.16
4.78
-14.9
0.57
FAWTH-16
5.43
4.68
13.8
-0.01
FAWTH-3
5.12
4.69
8.4
-0.42
FAWTH-29
5.02
4.46
11.2
-1.28
FAWTH-24
5.02
3.97
20.9
-1.53
FAWTH-2
3.84
4.78
-24.5
-1.86
FAWTH-17
4.79
4.45
7.1
-2.1
FAWTH-15
4.02
4.43
-10.2
-2.32
FAWTH-25
3.42
3.99
-16.7
-2.53
FAWTH-20
3.78
4.07
-7.7
-3.16
FAWTH-9
4.64
4.43
4.5
-3.28
FAWTH-21
4.88
4.22
13.5
-3.6
FAWTH-11
4.78
4.09
14.4
-4.51
FAWTH-5
5.22
4.1
21.5
-4.79
FAWTH-26
5.14
4.14
19.5
-6.26
FAWTH-14
5.17
3.75
27.5
-6.29
FAWTH-8
4.51
3.44
23.7
-6.8
Mean
4.86
4.61
5.1
SED
0.28
0.29
CV
9.9
11.1
BI: Base index; SED: Standard error of the difference; CV: Coefficient of variation.
with GY and significant negative relationships with FAW
leaf damage (FAWLD), ear aspect (EASP), and FAW ear
damage (FAWED) (Table 6). Negative significant
associations were recorded between GY on the one
hand, and EASP, FAWLD and FAWED. The associations
among FAWLD, FAWED and EASP were positive and
significant. However, PASP did not exhibit significant
relationships with any of the traits.
DISCUSSION
The raging infestation by FAW is overwhelming and has
become a major yield-limiting factor to maize production
in SSA. Depending on extent of infestation, susceptibility
of genotype and in the absence of appropriate control
measures, FAW can cause up to 100% loss in maize
grain yield (Prasanna et al., 2018). Host-plant resistance
is the most sustainable management strategy to FAW
infestation on maize in SSA. Stem borers (SB) are
Lepidopteran pests like FAW, and exhibit similar mode of
infestation on maize. Therefore, a search into maize
germplasm exhibiting SB resistance could be a ready
source of genes for tolerance to FAW infestation. In this
study, 29 white maize genotypes with varying levels of
resistance to SB were evaluated under artificial FAW-

Moussa et al. 107
Table 6. Linear relationships between base index and selection indices in 29 white maize
genotypes evaluated under fall armyworm infested condition.
Genotype
GY
FAWLD
EASP
FAWED
PASP
BI
0.932***
-0.660***
-0.835***
-0.784***
-0.057
GY
-0.583***
-0.747***
-0.644***
0.059
FAWLD
0.493**
0.405*
-0.290
EASP
0.739***
-0.217
FAWED
-0.137
*;**;***: significant respectively at 0.05, 0.01, 0.001 probability levels. BI: Base index; GY: Grain
yield; FAWLD: Fall armyworm leave damage; EASP: Ear aspect; FAWED: Fall armyworm ear
damage; PASP: Plant aspect.
infested and FAW-protected conditions.
The observed genotypic differences, coupled with the
enormous contribution of the sum of square for genotype
to the total sum of squares for all the traits under both
FAW-infested and FAW-protected environments
demonstrated the existence of sufficient genetic variability
among the test genotypes, which could be exploited for
FAW tolerance breeding. Also, the main effects of
environment, and genotype × environment interaction
were non-significant, with very low contributions to the
total sum of squares for all the traits under both FAW-
infested and FAW-protected conditions. This indicated
that the performance of the test genotypes was
essentially due to their genetic make-up and little
influenced by environmental factors. Similar observations
were reported by Kamweru et al. (2023) and could be an
indication of the preponderance of additive gene effects
for the traits. It is pertinent to note that the test genotypes
in this study had varying levels of tolerance to SB. In
studies involving artificial SB infestation, Karaya et al.
(2009), Beyene et al. (2011) and Olayiwola et al. (2021)
reported the preponderance of additive gene effect in the
inheritance of GY and SB damage traits in maize.
Averaged across genotypes, a comparison of the grain
yields under FAW-infested and FAW-protected condition
revealed a 5% reduction, which suggests that SB
resistance could confer tolerance to FAW infestation.
The level of resistance to FAWLD and FAWED
exhibited by most of the test genotypes in this study was
moderate. None of the genotypes was highly resistance
or highly susceptible to both FAW damage parameters.
Cultivation of partially resistant genotypes could serve as
an interim management strategy for farmers as well as
valuable genetic resource for breeding programs targeted
at the development maize genotypes with resistance to
the twin effects of FAW and SB infestation. Other studies
(Ni et al., 2014; Abel et al. 2020; Kasoma et al., 2020,
2021b; Kamweru et al. 2023) have also found maize
genotypes expressing moderate resistance to FAW.
Results from this study revealed that GY was
significantly but negatively correlated with FAWLD and
FAWED, which implied FAW infestation reduced maize
GY. Similar negative relationships between GY and FAW
damage parameters had been reported by previous
studies (Assefa and Ayalew, 2019; Overton et al., 2021;
Job et al. 2022; Kamweru et al., 2023). Grain yield is
directly impacted by FAWLD. The FAW larvae's leaf
feeding and whorl damage causes a reduction in the
plant's capacity to photosynthesize, leading to a
disruption in assimilate translocation and partitioning,
which results in impaired growth, poor grain filling and
yield. The older caterpillars burrow into the maize cob,
damaging the maize ear and kernels, and predisposing
the kernels to secondary infections (Buntin, 1986; Anjorin
et al., 2022). Additionally, FAWED leads to a reduction in
seed and grain quality by predisposing the kernels to
fungal attack, rot and mycotoxin accumulation (Williams
et al., 2018). The positive and significant association
among FAWLD, FAWED and EASP suggests that any
one of the traits could be used to predict the other two.
Matova et al. (2022) also reported similar positive
correlations among FAW damage parameters.
In the present study, a BI which included five traits (GY,
EASP, FAWLD, FAWED and PASP) was used as a
selection criterion. The highly significant correlations
between BI and GY (r=0.93), EASP (r=-0.84), FAWLD
(r=-0.66) and FAWED (r=-0.78) indicated that high BI
could be effectively used to select genotypes that
combined high grain yield with FAW tolerance/resistance.
Oloyede-Kamiyo (2019) has shown the effectiveness of
base index in the selection of desirable maize genotypes
under stem borer infestation.
Conclusion
Fall armyworm has assumed the status of the most
destructive yield-limiting insect pest of maize in sub-
Saharan Africa. Cultivation of varieties with resistance to
fall armyworm is most economical, sustainable and
compatible with other management options targeted at
enhancing maize grain yields in sub-Saharan Africa. In
this study, some white maize genotypes with varying
levels of tolerance to stem borer were evaluated for their

108 J. Plant Breed. Crop Sci.
agronomic performance under artificial fall armyworm
infestation.
The genotypes evaluated varied widely for grain yield,
agronomic traits and fall armyworm damage traits. Our
study revealed the utility of stem borer resistant
germplasm as reservoir for fall armyworm tolerant genes.
Averaged across genotypes, grain yield reduction under
fall armyworm infestation was low (5.1%) suggesting that
resistance to stem borer could also confer tolerance to
fall armyworm damage. Grain yield was negatively and
significantly related with fall armyworm damage
parameters, indicating that low fall armyworm damage
scores can be used to identify tolerant and high yielding
genotypes. Highly significant correlations were also found
between base index and grain yield, ear aspect, fall
armyworm leaf damage and fall armyworm ear damage
indicating it could be effective as selection criteria for
combined fall armyworm tolerance and high grain yield.
Genotypes FAWTH-1, FAWTH-13, FAWTH-4, FAWTH-
10, FAWTH-23 and FAWTH-6 with positive base index ≥
4.0 and grain yield ≥ 5.13 t ha-1 under fall armyworm
infested condition were identified as promising candidates
that combined tolerance to fall armyworm with high grain
yields.
FUNDING
This study received funding from Value Seeds Limited,
Kaduna State, Nigeria and M.Sc. scholarship award by
the African Union through the Pan African University Life
and Earth Sciences Institute (Including Health and
Agriculture), University of Ibadan, Nigeria to N.A.Z.M.
CONFLICT OF INTERESTS
The authors have not declared any conflict of interests.
ACKNOWLEDGMENTS
The authors express their appreciation to the
management of Value Seeds Limited, Kaduna State,
Nigeria for the fund and the African Union for the M.Sc.
scholarship grant to N.A.Z.M.
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