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Response of provitamin-A maize germplasm to storage weevil Sitophilus

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Article · November 2016

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International Journal of Agronomy and Agricultural Research (IJAAR)

ISSN: 2223-7054 (Print) 2225-3610 (Online)
http://www.innspub.net
Vol. 9, No. 5, p. 1-13, 2016
RESEARCH PAPER OPEN ACCESS

Response of provitamin-A maize germplasm to storage weevil

Sitophilus zeamais (Motschulsky)

Frejus A. K. Sodedji*1,2, Daniel B. Kwemoi3, Godfrey Asea3, Samuel Kyamanywa1

School of Agricultural Science, Makerere University, Kampala, Uganda

1

Faculty of Agronomic Sciences, University of Abomey-Calavi, Cotonou, Benin

2

National Crop Resources Research Institute, Kampala, Uganda

3

Article published on November 8, 2016

Key words: Biofortified maize, Storage pests, Maize weevils

Abstract

Development and dissemination of biofortified maize is essential for communities at risk of malnutrition. This
further requires that the improved maize varieties retain all their nutritional attributes up to the end users.
Weevils (Sitophilus zeamais) are among the most economically important storage pests of maize in the tropics
and present a permanent threat to the availability of maize nutrients for human consumption. This study aimed
at evaluating resistance to maize weevils in 24 provitamin-A maize inbred lines and their single cross hybrids.
Evaluation was done by artificial infestation and the Dobie Index of Susceptibility was used to group the
genotypes. Results showed that Provitamin-A maize genotypes evaluated were in general susceptible to S.
zeamais. Two inbred lines CLHP0014, and CLHP0005; and five single crosses; CLHP00434/CLHP0020,
CLHP00306/CLHP0005, CLHP00434/CLHP0003, CML486/CLHP0005, CLHP0331/CLHP0020 were
moderately resistant to S. zeamais. Grain damage (GD) was strongly positively correlated with number of F1
weevils which emerged (R2= 0.74, P<0.001). The index of susceptibility (IS) was positively and strongly
correlated with number of weevils which emerged (R 2= 0.86, P<0.001) and grain damage (R2= 0.69, P<0.01).
Susceptibility of the maize genotypes was not correlated with differences in kernel colours. Lines with moderate
resistance to weevil were identified and could be improved for the use in developing nutritional qualities maize
varieties to minimize loss due to storage pests.
* Corresponding Author: Frejus A. K. Sodedji  [email protected]

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 Int. J. Agron. Agri. R.

Introduction In many tropical countries, S. zeamais can cause 30

Maize (Zea mays L.) is an important source of daily to 80% seed weight loss in storage (Ajayi and Soyelu,
calories, especially in areas of severe malnutrition and 2013). In the absence of proper post-harvest insect
food insecurity (Nelson et al., 2010; WHES, 2015). It control measures, seed loss can be high as 90% (Giga
is a dominant food crop in much of sub-Saharan et al., 1998; Derera et al., 2014). This results in loss,
African and the American countries, where 17 to 30 %
reduction in nutritional value of maize; reduction in
of children under 5 years age are vitamin A deficient
market value of the crop as well as loss of seed vigor
(Harjes et al., 2008). Although maize is a staple
(Abebe et al., 2009, Tongjura et al., 2010). Hence,
cereal crop, the white kernels maize varieties which
damage due to storage weevils must be controlled to
are very popular among the African population, have
minimize the losses incurred by maize producers,
low nutritional value, mainly with respect to protein
since cares and expenditures for pest control in field
and essential micronutrients such as carotenoids
(FAO, 1992; Sands et al., 2009; Chandler et al., crops would be of no use, if the product was attacked

2013). Carotenoids molecules in maize are precursors and destroyed when stored (Alleoni and Ferreira,

of vitamin A (Provitamin-A) in human diet, and 2006). Development of resistant maize varieties
represent an important source of vitamin A offers an effective way and long term solution to the
particularly for populations of developing countries, damage due to S. zeamais (Gudrups et al., 2001;
whose poor economic conditions do not allow an easy Mwololo et al., 2012b). A number of studies have
diet balance of plants and animal foods (Sivaranjani been conducted to screen maize varieties against S.
et al., 2013). zeamais and have identified genotypes having
resistance genes that can therefore be used to
Food biofortification seek thus to enhance the
introgressing genes of resistance into available
concentration of various micronutrients in maize,
especially the provitamin-A carotenoids, so as to germplasm (Kim and Kossou, 2003; Abebe et al.,
address the problem of micronutrient deficiencies in 2009; Mwololo et al., 2012a). However, the genotypes
developing countries (Hotz and McClafferty, 2007; used in most of these studies were not known as
Stein, 2010; Bouis and Welch, 2010). Consequently, provitamin-A maize and to date there is a need to
provitamin-A maize cultivars are introduced in identify genotypes with good performance for
regions of severe malnutrition. However, success in
provitamin A trait and resistant to storage weevils.
biofortification of maize for areas at risk of
Study was thus conducted to assess the response of
malnutrition requires that the improved maize
varieties retain all their nutritional attributes up to provitamin-A inbred lines and their single crosses to
the end users. Even if significant progress have storage weevils under controlled storage conditions.
recently been made to increase provitamin-A content
in maize, there is a danger that if the losses caused by Materials and methods
storage pests are not well managed, the improved Experimental materials
provitamin-A maize will not be available in sufficient A sub set of 24 elite provitamin-A maize inbred lines
quantity and quality to meet the overwhelming need (Table 1) were selected from introduction from
of provitamin-A maize varieties in areas at risk of CYMMYT and used to develop 72 single crosses
vitamin A deficiency (VAD). hybrids by crossing four of the inbred lines (CML300,
CLHP0020, CLHP0005, CLHP0003) used as males
Maize weevils (Sitophilus zeamais) represent a threat to the other 20 inbred lines (females) at the National
to the sustainable storage of maize grains in tropics. Crop Resources Research Institute (NaCRRI) of
They are economically important field-to-store pests Uganda in 2014 (September to December).
of maize that start to infest the ripening maize crop in Concurrently, seeds of the inbred lines were
the field as early as when the grain moisture content multiplied during that season and used for a first
is still 50-55% (Ojo and Omoloye, 2012). screening of the parental lines.

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At the onset of first rainy season in 2015 (April to The F1 insects at 1 to 10 days old were sieved out with
August), seeds of the provitamin-A inbred lines and mesh sieve (Endecotts Ltd, UK) and used in the
their single crosses were self-pollinated at NaCRRI in kernel screening.
Namulonge and the National Semi-Agricultural
Hybrids kernel colour scoring
Research Institute (NaSAARI) in Serere. The inbred
Kernel colour of the hybrids was visually scored prior
lines were selfed to produce enough seeds for
to the grain resistance test as described by Chandler
screening while F1s seeds were advanced to produce
et al.(2013). This involves grouping a total of 100
F2 grains. F2 grains are the generation of grains randomly selected seeds of each genotype into colour
normally stored by farmers and therefore the most classes using an ordinal standardized colour scale
vulnerable to weevils damage (Siwale et al., 2009). ranging from 1 (lightest yellow) to 12 (darkest
Seeds of yellow conversions of popular single cross orange). The score of a specific genotype was
testers CML202/CML395, CML312/CML442 and computed as an average of the different colour scores
Longe10H a popular three ways maize variety in observed in the seeds bulk.
Uganda were multiplied at the same time. Longe 5, an
open pollinated maize variety highly susceptible to Preparation and infestation of grains samples with
weevils was included as a susceptible check. weevils
After a proper cleaning operation, four subsamples
Experimental sites (replicates) each of 50 grams seeds were weighed for
Seeds multiplication was done at the National Crop each of the 100 genotypes (24 inbred lines and 76
Resources Research Institute (NaCRRI), Namulonge hybrids) using a compact balance electronic weighing
and the National Semi-Agricultural Research Institute scale brand type. The weighed samples were

(NaSAARI), Serere in Uganda. NaCRRI, is located in thereafter labelled and wrapped in polythene bags

Wakiso District, central Uganda at an elevation 1200 and then frozen at -20°C for 14 days to kill any
insect/egg that could have attacked the grains in the
masl. It is within the bimodal rainfall region and has a
field (Siwale et al., 2009). After freezing, the seed
tropical wet and mild dry climate with slightly humid
samples were transferred to 250 cm3 evaluation jars
conditions(NARO, 2005). NaSSARI is located in
and left to achieve uniform temperature and moisture
Eastern Uganda at an elevation of 1080 masl;
content before the infestation. A total of 32 unsexed
between the bimodal rainfall regions and the
insects was used to infest the maize kernels in each
unimodal rainfall regions (Mubiru et al., 2012). The
glass jar (Derera, Pixley, & Giga, 2010). The jars were
trials for on grains resistance test were conducted in sealed as described earlier. The inbred lines were laid
the entomology laboratory unit at NaCRRI. in 4 x 6 alpha Lattice Design while single crosses were
set in 4 x 19 Alpha Lattice design within the
Mass rearing of weevils for the kernel screening laboratory (Dhliwayo et al., 2005). A Thermo-
The weevils used in this study were mass reared in Hygrometer was used to monitor relative humidity
entomology laboratory mass rearing unit at NaCRRI. (70±5%) and temperature (28±2°C) status during the
A total of 200 unsexed adult insects were sampled experimentation period (Mwololo et al., 2012a)
and used to infest 1000 grams of Longe 5 (highly
susceptible to weevils). The grains were put in 3000 Data collection
cm3 plastic jars and covered with lids which were first Samples were given 10-days oviposition period after
perforated and stuck with wire mesh to allow which all adult insects, dead and living were removed
maximum aeration without the insects escaping from and counted. The samples were then incubated for 25
the jars. The unsexed adult weevils were given 10 days days to allow a development period of 35 days which
to oviposit after which they were removed and the is the average period for the weevils to complete one
samples were left for 35 days of incubation to ensure cycle, under optimum conditions (Gwinner et al.,
that enough F1 insects were produced. 1996; Derera et al., 2001a).

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 Int. J. Agron. Agri. R.

F1 progeny insects were then counted and removed Statistical analysis

from the jars at 2 days interval until no more insect Data were subjected to an analysis of variance across
emerged from the jars. This interval of counts locations using the Restricted Maximum Likelihood
prevents any chance of F1 progeny laying eggs in the (ReML) in GenStat (12th Editon). The genotypes were
maize samples to produce F2 generation weevils. This considered as a fixed effect while environment and
is based on the fact that individuals of S. zeamais do replications were random. The linear model for the
not mate before they are three days old. Five across environments was as follows:
parameters were assessed for each genotype,
including adult insect mortality, number of F1 insects
emerged during the storage period, median
Where 𝑦𝑖𝑗𝑘 = observed value from each experimental
development period, Dobie’s index of susceptibility
unit, u = grand mean, 𝑙𝑖 = effect of the ith
and grain damage as described below.
environment,𝑟/𝑙𝑖𝑗 = effect of the jth replication nested
Adult mortality to the ith environment,𝑔𝑘 = effect of kth genotype, 𝑔𝑙𝑖𝑘
Adult mortality was estimated as percentage of the = interaction effect of kth genotype by the ith
number of dead adult insects after ten 10 days of environment and eijk is the experimental error.
oviposition period.
Means were separated using Least Significant
Number of F1 insects emerged (N° F1 insects) Differences (LSD) at a 5% probability level. The index
It represents the cumulative sum of F1 progeny of susceptibility scale, ranging from 0 to 11, was used
insects counted and removed from the jars at 2 days to classify the response of the maize genotypes into
interval until no more insects emerged from the jars. resistance groups. The resistance classes were 0 - 3 =
resistant, 4 - 7 = moderately resistant, 8 - 10 =
Median development period (MDP)
susceptible and ≥11 = highly susceptible (Issa et al.,
The median development period was computed as the
2011). kernel colour scale of 1 to 12 (Chandler et al.,
number of days from the middle period of oviposition
2013) was used to categorize the hybrids maize
(5 days) to the middle emergence of progeny (50%
genotypes into four groups (1-4= yellow, 4.1-6 =
emergence of F1 insects), (Dobie, 1974; Siwale et al.,
orange pale, 6.1-8=orange and 8.1 -12= deep orange).
2009).
Correlation analyses were performed between
Index of susceptibility (IS) measured traits. The kernel colour scores of the
The index of susceptibility was calculated using the hybrids were also regressed against their index of
method proposed by Dobie (1977). This involves the susceptibility to check if there is a relationship
number of F1 progeny and the median development between these two traits.
period.
Results
Response of provitamin-A inbred lines to maize
weevil infestation
There were significant differences among the inbred
Grain damage (GD)
lines (Table 2) for adult mortality (P < 0.01), Number
Once all adult insects which emerged were removed
of F1 insects emerged (P <0.001), Median
from the infested jars, 100 grains were randomly
development period (P < 0.01), Index of susceptibility
taken from each jar and the number of damaged
(P < 0.001), and grain damage (P < 0.001).
grains (holed grains) by weevil feeding was
Genotypes by Environment Interaction was also
determined as a proportion of the total number of
significant for all the five grain susceptibility
grain sampled (Caneppele et al., 2003; Abebe et al.,
parameters (Table 2).
2009).

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 Int. J. Agron. Agri. R.

Adult mortality in the inbred lines was generally low Number of F1 insects emerged varied with the mean
with an average value of 17.52% (Table 3). Line value of 83 insects (Table 3). Lines CLHP00328 and
CML451 had the lowest percent of adult mortality the CLHP0301 had the highest number of weevils
while the line CLHP0014 exhibited the highest mean emergency while the line CLHP0014 had the lowest
of adult mortality. number of F1 insects emerged.

Table 1. Provitamin-A inbred lines evaluated for resistance to storage weevils.

Genotype Pedigree
CML304 CML304-B-B-B-B
CML486 CML486-B-B-B
CML451 CML451-B-B-B
CLHP00306 (KUIcarotenoidsyn-FS17-3-1-B-B-B-B//(KUIcarotenoidsyn-FS17-3-1-B-B-B/(CML297-B×KUICarotenoidsyn-
FS17-3-2-B/KUI3×SC55)))-plant19(HH)-3-1-B-B-B-B-B-B
CLHP00478 (CLQRCWQ97-B///((KUIcarotenoidsyn-FS17-3-2-B-B-B/(KU1409/DE3/KU1409)S2-18-2-B)//CarotenoidSyn3-
FS11-4-3-B-B-B))-B-25-6-B-B-B-B-B-B
CLHP00476 (CLQRCWQ97-B///((KUIcarotenoidsyn-FS17-3-2-B-B-B/(KU1409/DE3/KU1409)S2-18-2-B)//CarotenoidSyn3-
FS11-4-3-B-B-B))-B-25-2-B-B-B-B-B-B
CLHP0310 ((CML506//CML506/CML305)//((CML506//CML506/CML305)/KUIcarotenoidsyn-FS11-1-1-B-B-B-B)-B)-6-3-
B-B-B-B-B
CLHP0290 ([DTPYC9-F65-2-3-1-1-B-BxDTPYC9-F65-2-2-1-1-B-B]-3-4-2-B-B-B//([DTPYC9-F65-2-3-1-1-B-BxDTPYC9-
F65-2-2-1-1-B-B]-3-4-2-B-B/(CML297-B×KUICarotenoidsyn-FS17-3-2-B/KUI3×B77)))-plant3(H)-1-3-B-B-B-B-
B-B
CLHP00308 ((CML506//CML506/CML305)//((CML506//CML506/CML305)/(KU1409/DE3/KU1409)S2-18-2-B-B-B)-B)-
11-1-B-B-B-B-B
CLHP0302 (CML486/(CML297-B×KUICarotenoidsyn-FS17-3-2-B/KUI3×B77))-B-11-1-B-B-B-B-B-B-B
CLHP0352 ([DTPYC9-F65-2-3-1-1-B-BxDTPYC9-F65-2-2-1-1-B-B]-3-4-2-B-B/(CML297-B×KUICarotenoidsyn-FS17-3-2-
B/KUI3×B77))-B-21-1-B-B-B-B-B-B-B
CLHP00294 (KUIcarotenoidsyn-FS17-3-1-B-B-B-B//(KUIcarotenoidsyn-FS17-3-1-B-B-B/(CML297-B×KUICarotenoidsyn-
FS17-3-2-B/KUI3×SC55)))-plant19(HH)-4-2-B-B-B-B-B-B
CLHP00328 ([[[NAW5867/P30SR]-43-2/[NAW5867/P30SR]-114-1]-9-3-3-B-1-B/CML395-1]-B-13-1-B-4-#/[BETASYN]BC1-
8-1-1-1-B-B/(CML297-B×KUICarotenoidsyn-FS17-3-2-B/KUI3×B77))-B-1-1-B-B-B-B-B-B-B
CLHP0301 (CML486/(CML297-B×KUICarotenoidsyn-FS17-3-2-B/KUI3×B77))-B-11-1-B-B-B-B-B-B-B
CLHP0331 (CML491-B///((KUIcarotenoidsyn-FS11-1-1-B-B-B/(KU1409/DE3/KU1409)S2-18-2-B)//[[[NAW5867/P30SR]-
40-1/[NAW5867/P30SR]-114-2]-16-2-2-B-2-B/CML395-6]-B-20-1-B-3-#/[BETASYN]BC1-3-1-1-#-B-B-B))-B-3-
1-B-B-B-B-B
CLHP0289 (CML496//(CML496/(CML297-B×KUICarotenoidsyn-FS17-3-2-B/A619×SC55)))-plant11(HF)-1-3-B-B-B-B-B-B
CLHP00434 ((([CLRQ00502xB109]-F2)-04-05-K1/KUIcarotenoidsyn-FS17-3-2-B-B-B-B)//(([CLRQ00502xB109]-F2)-04-
04-K1/CarotenoidSyn3-FS11-4-3-B-B-B-B)//(KUIcarotenoidsyn-FS11-1-1-B-B-B/(KU1409/DE3/KU1409)S2-18-
2-B)-B-2)-12-1-B-B-B-B-B
CLHP0014 CarotenoidSyn3-FS8-4-3-B-B-B-B-B-B-B-B-B-B-B-B
CLHP0002 CML489/[BETASYN]BC1-2-#-B-B-B-B-B-B-B-B-B-B-B
CLHP0006 CML488/[BETASYN]BC1-15-7-1-1-1-B-B-B-B-B-B-B
CML300 CML300-B-B-B
CLHP0005 MAS[206/312]-23-2-1-1-B-B-B/[BETASYN]BC1-11-3-1-#-B-B-B-B-B-B-B-B-B-B-B-B
CLHP0003 CML537/[BETASYN]BC1-10-3-#-B-B-B-B-B-B-B-B-B-B-B-B
CLHP0020 KUIcarotenoidsyn-FS17-3-2-B-B-B-B-B-B-B-B-B-B-B-B

The median development period of weevils on the A classification of the 24 provitamin-A inbred maize
inbred lines maize varied from 43 days to 54 days for lines into four resistance classes based on their index
the lines CLHP0014 and CLHP0005, respectively of susceptibility, revealed lines CLHP0014 and
(Table 3). Percentage of damaged grains due to weevil CLHP0005 as moderately resistant lines while the
infestation was high for the lines CML304, rest of the inbred lines were susceptible or highly
CLHP0006, CLHP00328 and CLHP0301 while the susceptible genotypes. The most susceptible
line CLHP0014 had the lowest percent of damaged genotypes were the lines CLHP0352 and CLHP00328
grain (Table 3). (Table 3).

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Table 2. Mean squares for five grain susceptibility parameters of the 24 provitamin-A maize lines across
environments.
Source Df Mean squares
Adult Mortality N° F1 Insects MDP IS GD
Environments (E) 2 6622.24*** 2281.20 6622.24*** 49.43** 2250.12*
Rep/E 9 1070.1*** 5022.3*** 82.57*** 12*** 1125.6***
Genotypes (G) 23 1450.84** 6503*** 1450.84** 25.92*** 2403.36***
GxE 46 592.00*** 1521* 592 *** 6.32*** 723.20***
Pooled error 157 -181 116.36 783.22 15.57 2.37 251.76
Grand Mean 17.52 82.53 46.53 9.12 39.54
CV (%) 61.57 33.91 8.48 16.88 40.13

Rep/E= Replications nested to Environments, Adult Mortality= percentage of dead insects after 10 days of
oviposition, MDP=Median development Period, IS= Index of susceptibility, GD= Grain damage. ***Significant at
P<0.001, **significant at P<0.01, *significant at P<0.05.

Table 3. Mean response of the 24 lines and testers for five grain susceptibility parameters across environments.
ENTRY Adultb mortality N° F1 Insects MDP GD IS Category
CML304 18.58 121.49 46.86 62.85 10.3 S
CML486 12.82 102.54 44.36 47.46 10.4 S
CML451 6.97 87.03 46.94 48.54 9.5 S
CLHP00306 25.44 74.05 45.91 31.88 9.3 S
CLHP00478 23.83 57.77 48.25 29.78 8.3 S
CLHP00476 22.75 74.52 45.78 34.11 9.3 S
CLHP0310 7.73 70.17 46.15 39.11 9.1 S
CLHP0290 12.81 62.66 45.52 31.25 9.0 S
CLHP00308 10.77 79.16 46.86 41.51 9.3 S
CLHP0302 17.85 62.94 45.36 34.75 9.2 S
CLHP0352 11.11 127.64 43.48 51.25 11.2 HS
CLHP00294 39.80 61.98 47.00 27.99 8.4 S
CLHP00328 9.90 157.73 45.30 58.59 10.8 HS
CLHP0301 9.02 155.62 44.44 57.94 11.2 HS
CLHP0331 9.45 73.45 46.96 34.50 8.9 S
CLHP0289 16.18 72.67 45.36 39.50 9.4 S
CLHP00434 13.74 81.14 49.02 47.99 9.1 S
CLHP0014 53.10 18.22 43.41 4.75 4.5 MR
CLHP0002 7.64 72.55 44.61 43.52 9.7 S
CLHP0006 12.03 104.27 49.23 60.56 9.9 S
CML300 15.46 102.76 45.80 50.15 10.0 S
CLHP0005 25.42 54.45 53.75 20.25 6.5 MR
CLHP0003 27.40 54.35 48.48 22.60 7.7 S
CLHP0020 10.76 51.49 47.79 28.04 7.9 S
Grand Mean 17.52 82.53 46.53 39.54 9.12
LSD 8.69 22.54 3.18 12.79 1.24

HS= highly susceptible, S= Susceptible, MR= Moderately Resistant.

Responses of single cross hybrids provitamin A showed that hybrid CLHP00478/CLHP0005 had the
maize to S. zeamais lowest adult mortality while hybrid
Significant differences among the single crosses of CLHP00434/CLHP0020 exhibited the highest adult
provitamin-A maize genotypes were obtained for all mortality . The number of F1 insects emerged varied
susceptibility parameters evaluated except the from 22 insects for the hybrid CLHP0331/CLHP0020
Median development period (Table 4). Genotype by
to 107 insects for CML486/CML300. The median
environment interaction had also a significant effect
development period was the lowest in the susceptible
on the performance of the genotypes for all
check (Longe 5, MDP = 41) as compared to the other
parameters across environments (Table 4).
genotypes. On the other hand, grain damage was the
Mean values of the provitamin-A maize hybrids for highest for hybrid CML486/CML300 and the lowest
the five grain susceptibility parameters evaluated for CLHP0331/CLHP0020 (14.28%).
(Table 5),

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 Int. J. Agron. Agri. R.

Table 1. Mean squares for five grain susceptibility parameters for the 76 genotypes evaluated for resistance to S.
zeamais across environments in Uganda.
Source Df Mean squares
Adult Mortality N° F1 Insects MDP GD IS

Environments (E) 1 2597.12 36941.56* 13620.9*** 26275.56* 838.73**

Rep/E 6 2335.07*** 4900.4*** 141.67*** 2799.6*** 25.30***

Genotypes (G) 75 343.84** 2395.88*** 26.68 980.9*** 6.80**

GxE 75 185.08*** 986.2*** 23.70*** 489.2*** 3.43***

Pooled error 412-444 88.25 387.42 11.53 153.77 11.48

Grand Mean 13.96 54.34 46.78 32.55 8.41

SEM 3.83 8.03 1.39 5.06 0.50

CV (%) 67.28 36.22 7.26 38.10 14.45

Adult Mortality= percentage of dead insects after 10 days of oviposition. MDP=Median development Period, IS=
Index of susceptibility, GD= Grain damage. ***Significant at P<0.001, **significant at P<0.01, *significant at
P<0.05.

Table 2. Mean response of the 15ahybrids provitamin-A maize genotypes for five grain susceptibility parameters
assessed across environments in 2015.
Genotypes Adult Mortality N° F1 Insects MDP GD IS Category

CLHP00434/CLHP0020 34.72 23.71 46.56 15.63 6.28 MR

CLHP00306/CLHP0005 20.84 29.21 49.41 17.35 6.50 MR

CLHP00434/CLHP0003 13.06 25.34 47.37 16.19 6.74 MR

CML486/CLHP0005 18.15 28.34 48.69 19.15 6.76 MR

CLHP0331/CLHP0020 15.59 21.59 43.98 14.28 6.91 MR

CLHP00478/CLHP0005 2.50 36.81 48.90 25.69 7.28 S

CML486/CLHP0020 13.06 38.59 49.93 23.88 7.28 S

CLHP0002/CLHP0020 18.01 36.96 46.93 22.18 7.34 S

CLHP0289/CLHP0005 4.53 36.59 49.40 21.79 7.34 S

CLHP0006/CLHP0005 15.89 44.59 49.19 31.27 7.39 S

CML486/CLHP0003 12.93 83.59 45.08 52.48 9.91 HS

CLHP0352/CML300 18.83 92.34 45.78 46.49 10.08 HS

CLHP0352/CLHP0003 20.81 89.34 44.67 44.89 10.15 HS

CLHP0002/CML300 7.09 81.59 42.83 55.22 10.41 HS

CML486/CML300 8.98 106.84 44.91 61.47 10.44 HS

Susceptible check

Longer 5 6.45 74.84 41.00 43.05 10.52 HS

Grand Mean 13.96 54.34 46.78 32.55 8.41 -

LSD 9.23 19.34 - 12.19 1.19 -

HS= highly susceptible, S= Susceptible, MR= Moderately Resistant. 15a= 5 highly susceptible genotypes, 10
susceptible genotypes and 5 Moderately Resistant genotypes.

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Index of susceptibility ranged from 6.28 in resistance to weevils showed that genotypes had a
CLHP00434/CLHP0020 to 10.52 in the susceptible distribution skewed toward susceptibility (Fig. 1). The
check (Longe 5). Genotypes Longe 10H, provitamin-A hybrids maize
CML312y/CML444y and CML202y/CML395y had an CLHP00434/CLHP0020, CLHP00306/CLHP0005,
index of susceptibility of 10.2, 9.08 and 8.27, CLHP00434/CLHP0003, CML486/CLHP0005 and
respectively. A classification of the 72 provitamin-A CLHP0331/CLHP0020 were moderately resistant to
maize hybrids in different category of the maize weevils attack.

Table 6. Mean squares for kernel colour of the hybrids provitamin-A maize genotypes.
Source Df Mean squares
Kernel colourd
Environments (E) 1 1.26
Rep/E 2 2.85**
Genotypes (G) 74 7.15***
GxE 74 0.52
Pooled error 80 0.81
Grand Mean 6.75
SEM 0.21
CV (%) 13.38

d=Kernel colour scored on a scale of 1(lightest yellow) to 12 (darkest orange).

Table 7. Kernel colour scores of the first 5 deepest orange and last 5 lightest yellow hybrids provitamin-A maize
genotypes across environments in Uganda in 2015.
Hybrids Kernel colour scores
CLHP00306/CLHP0020 9.25
CLHP0310/CLHP0020 9.00
CLHP0352/CLHP0020 9.00
CLHP00294/CLHP0020 9.00
CLHP0302/CLHP0020 8.75
CLHP0301/CLHP0020 8.65
CML486/CLHP0020 8.50
CLHP0289 /CLHP0005 8.50
CLHP0290/CLHP0020 8.25
CLHP00434 /CLHP0020 8.25
CLHP00328/CLHP0005 5.75
CLHP00328/CLHP0003 5.75
CLHP0002/CLHP0003 5.75
CLHP00476/CLHP0005 5.63
CLHP00308/CLHP0005 5.38
CLHP00328/CML300 5.25
CLHP0331 /CLHP0005 5.25
CML451/CML300 5.08
CML451/CLHP0003 4.57
CML451/CLHP0005 3.75
Grand Mean 6.25
LSD 0.59

Thirty-height genotypes were classified in the four times more F1 weevils and have considerably
susceptible group while the rest of the genotypes were higher percentage of grain damage than moderately
highly susceptible (Fig.1). The high susceptible resistant ones (Table 5).
hybrids harboured three to

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There were significant differences in kernel colour relationship between the number of F1 insects
(P<0.001) between provitamin-A hybrids screened emerged and index of susceptibility (R2= 0.86,
(Table 6). Hybrid CML451/CLHP0005 was the P<0.001). The index of susceptibility was also
lightest with the kernel colour score of 3.75 whilst the
positively correlated with Grain damage (R2= 0.69,
deepest orange kernel (9.25) was obtained from the
P<0.001). Grain damage was strongly correlated with
cross between the lines CLHP00306 and CLHP0020
(Table 7). number of F1 insects emerged (R2= 0.74, P<0.001).
Negative correlation was obtained between the
The frequency distribution of the hybrids into four
Median development period and the index of
kernel colour classes showed that genotypes were
susceptibility (r= -0.6, P<0.001) while no significant
normally distributed (Fig. 2). On average the
linear relationship was found between IS and adult
genotypes were orange in colour with the kernel
colour score mean of 6.75. mortality on the grains of the provitamin-A hybrids
maize.
Relationship between grain susceptibility
parameters There was no consistent relationship between kernel
Correlations analysis between grain susceptibility colour of the genotypes and their index of
parameters (Table 8) showed strong positive susceptibility (r=0.004ns).

Table 8. Relationship between grain susceptibility parameters.

Parameters Coefficient of correlation (r) Coefficient of determination (R2)
Adult Mortality vs. IS -0.21 0.04ns
F1 insects emerged vs. IS 0.93 0.86***
MDP vs. IS -0.6 0.4***
GD vs. IS 0.83 0.69***
GD vs. F1 insects emerged 0.86 0.74***
Kernel colour vs. IS 0.004 1.60E-05 ns

***Significant at P<0.001, **significant at P<0.01, *significant at P<0.05, ns = non-significant.

Discussion ability of a particular genotype to resist weevil

Results from the laboratory screening of the inbred infestation (Abebe et al., 2009, Ajayi and Soyelu,
lines and resultant crosses for resistance to storage 2013). There was a significant genotype by
weevils showed considerable variation among the environment interaction effect, indicating that the
genotypes for the grain susceptibility parameters environment plays a role in the resistance of the
evaluated. These differences indicate the inherent maize varieties to weevils.

Fig. 1. Frequency distribution of the 72 hybrids provitamin-A maize into different susceptibility classes.

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 Int. J. Agron. Agri. R.

This finding is supported by Kim and Kossou (2003). This response developed against the maize weevils in
Whereas the screened lines were in general the moderate resistant genotypes could have been
susceptible to the weevils; two genotypes (CLHP0014 triggered by intrinsic resistance mechanisms in their
and CLHP005) out of the 24 inbred lines had a grains. Studies have identified antibiosis and non-

moderate resistance to weevil’s infestation. This preference as basis of grain resistance to S. zeamais
(Hall, 1975; Derera et al., 2001a, 2001b), and this
response among inbred lines may explain the high
suggests further experiments in order to elucidate the
rate of susceptibility observed in their resulting
type of mechanism present in these moderately
crosses. These hybrids crosses are yellow to orange
resistant genotypes. The genotypes assessed in the
maize genotypes and their index of susceptibility
test for resistance to weevils generally showed a
range from 6.24 to 10.44; which was not correlated to
relatively long median development period. The
the variation in the hybrids kernel colour inbred line CLHP0005 had the longest Median
scores(Table 8). Differential responses of development period and was moderately resistant.
yellow/orange maize genotypes have been reported The inbred lines differed in their median
by previous studies where the index of susceptibility development period but did not induce substantial
varied from 7.8 to 15.2 in the yellow/orange kernel genetic variation in their progenies for the same
coloured maize varieties(Dobie, 1974, Santos et al., parameter. The median development period of the
2006). Although most single crosses based on the hybrids was negatively correlated with the index of
Dobie’s index of suceptibility were classified as susceptibility. Similar result was reported by Abebe et

susceptble/higly susceptible to weevils in this study, al(2009) who pointed out that the weevils on varieties
having a high index of susceptibility displayed
the cross combinations CLHP00434/CLHP0020,
reduced periods of completion and that a
CLHP00306/CLHP0005, CLHP00434/CLHP0003,
prolongation of development periods will also result
CML486/CLHP0005 and CLHP0331/CLHP0020 had
in reduction of number of generations of the weevils
moderate resistance to weevils may be improved for
in the season.
reducing the threat due to the maize weevils.

Fig. 2. Frequency distribution of the 72 hybrids provitamin-A maize into four colour classes.

In the screening of the inbred lines, CLHP0014 had parameters (F1 insects emerged, GD, IS) in the
the lowest number of F1 emergency and low response of hybrids to the maize weevils attack. Thus,
percentage of grain damage (GD) with the lowest highly susceptible hybrids had more F1 weevils and
index of susceptibility. A strong positive correlation have considerably higher percentages of grain
was also found between these three damage than the moderately resistant ones.

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 Int. J. Agron. Agri. R.

In maize grains, weevils grow from larvae to adult Alleoni B, Ferreira W. 2006. Control of Sitophilus
eating the maize grains from inside to out, creating zeamais Mots., 1958 and Sitophilus oryzae (L ., 1763 )
holes on the grain, thus leading to the susceptibility weevils (Coleoptera , Curculionidae) in stored rice
status of the genotypes. Many studies found that the grain (Oryza sativa l.) with insecticide pirimiphos
number of F1 insects emerged is high in susceptible methyl (Actellic 500 CE). 9th International Working
genotypes and is positively correlated to grain
Conference on Stored Product Protection, 1234–1241.
damage (Abebe et al., 2009; Siwale et al., 2009; Keba
and Sori, 2013). In the present study adult mortality
Bouis HE,Welch RM. 2010. Biofortification--A
was in average low among the hybrids maize and was
Sustainable Agricultural Strategy for Reducing
not significantly correlated with susceptibility. Thus,
Micronutrient Malnutrition in the Global South. Crop
this parameter may not be an adequate factor in
discriminating the genotypes in their response to Science 50(1), 20–32.

weevils’ attack (Dobie, 1974), indicating the

usefulness of considering multiple traits in assessing Caneppele MAB, Caneppele C, Lázzari FA,
resistance to S. zeamais in maize germplasm. Lázzari SMN. 2003. Correlation between the
infestation level of Sitophilus zeamais Motschulsky ,
Conclusion 1855 (Coleoptera , Curculionidae ) and the quality
Overall, this study showed inbred lines and hybrids factors of stored corn , Zea mays L . (Poaceae).
that are moderately resistant to the maize weevils and
Revista Brasileira de Enthomologia 47(4), 625–630.
the need for their improvement for use in the
developing areas would reduce the high incidence of
Chandler K, Lipka AE, Owens BF, Li H,
weevil’s damage. This would help in preserving the
Buckler ES, Rocheford T, Gore MA. 2013.
nutritional quality of the stored maize grains up to the
Genetic Analysis of Visually Scored Orange Kernel
consumers and contribute to fight-off micronutrients
Color in Maize. Crop Science 53(1), 189–200.
deficiency. Most of the prtovitamin A genotypes
screened exhibited a susceptible response toward the
maize weevils’ attack, suggesting that breeding Derera J, Giga DP, Pixley KV. 2001a. Resistance
activity for developing provitamin-A maize varieties of Maize to the Weevill: II. Non-Preference. African
should be conducted concurrently with effort of Crop Science journal 9(2), 444–450.
introgressing weevils’ resistance genes into the maize
varieties. Derera J, Pixley KV, Giga DP, Makanda I. 2014.
Resistance of maize to the maize weevil : III . Grain
Acknowledgement weight loss assessment and implications for breeding.
This work was supported by RUFORUM through the Journal of Stored Products Research 59, 24–35.
Share Intra- ACP Scheme Project and the National
Crop Resources Research Institute (NaCRRI).
Derera J, Pixley KV, Giga DP. 2001b. Resistance
of Maize to the Maize Weevil :I. Antibiosis. African
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