Documente Academic
Documente Profesional
Documente Cultură
ABSTRACT
1
INTRODUCTION
The Guinea savannas of West and Central (WCA) Africa has the highest potential for
increased maize production and productivity due to high solar radiation, low night
temperatures and low incidence of diseases. However, maize production in this ecology is
seriously constrained by low soil fertility, especially low levels of N., recurrent drought,
and Striga hermonthica parasitism. On the average, drought occurs two to three times per
increase in temperature and decrease in precipitation resulting from global warming are
likely to increase the intensity and unpredictability of drought and decrease the length of
the growing season in WCA (DNRP-GAPCC, 2000). The occurrence of drought from a
few days before anthesis to the beginning of grain filling, which are the most sensitive
stages of the maize crop, can reduce grain yield by as much as 90% (NeSmith and
Ritchie, 1992). Unfortunately, drought occurrence is hardly predictable and most severely
affect grain yield at flowering and grain-filling period in maize. Consequently, breeding
for early and extra-early maturing maize cultivars with better tolerance to drought stress
and resistance to Striga that fit into areas with short growing period is essential to
The parasitic weed, Striga hermonthica is a major threat to maize production and
productivity in the savannas of WCA. The levels of infestation are often so high that
maize can suffer 100% yield loss. Increased population pressure on available land has led
soil fertility (Vogt et al., 1991), a condition highly suitable for Striga infestation. Many
2
maize farmers in the savannas of WCA have been forced to abandon their farmlands
because of Striga infestation that seem to have defied cultural control options (Bebawi et
al., 1984; Odhiambo and Ransom, 1994; Shaxson and Riches, 1998). Use of host plant
which is the ability of the host plant to produce yield despite infestation (Kim 1994), are
considered the most economically feasible and sustainable approach for reducing the
effects of Striga (DeVries, 2000; Badu-Apraku et al., 2004). A major strategy of the IITA
Maize Program to stabilize maize yield in the WCA therefore, is to develop maize
Another major strategy of the IITA Maize Program to stabilize maize yield in the
sub region is to breed for drought tolerant maize cultivars. Two approaches have been
adopted. The first is to breed for early and extra-early maturing cultivars that escape
drought before the onset of drought stress. These are referred to as drought escaping
cultivars. The second strategy is to breed for drought tolerant cultivars that possess
drought tolerant genes and can withstand drought which occurs during the flowering and
grain-filling period.
During the last two decades, the International Institute of Tropical Agriculture
has developed a wide range of high yielding drought tolerant/escaping extra-early Striga
resistant populations (white and yellow endosperm), inbred lines, and cultivars in an
effort to combat the threat posed by Striga hermonthica and recurrent drought in the
savannas of WCA. The extra early populations from which the inbred lines and cultivars
were derived were formed from crosses between local landraces, exotic and introduced
3
germplasm identified through extensive multilocation trials in WCA (Badu-Apraku et al.,
1999, 2001, 2007). They were selected on the basis of high grain yield, earliness, and
resistance to the maize streak virus and above all adaptation to the high temperatures and
drought stress characteristic of the Sudan savanna in Burkina Faso, Mali, Mauritania,
Africa not only low-N tolerant inbreds and hybrids but also, drought-
tolerant extra-early inbreds and hybrids that are not only drought
the low-N and drought tolerant extra-early inbreds and hybrids should
The long-term goal of the IITA maize program is to develop stress tolerant broad-
based populations with varying grain colour, texture, and endosperm types which are
heterotic to each other and could be exploited effectively for population improvement,
development of synthetics and hybrids in the lowland tropics. Classifying inbreds into
heterotic groups is essential for not only maximizing their potential usefulness for the
4
development of productive hybrids and synthetics but also for creating new heterotic
groups. Wellhausen (1978) was the first to make a systematic attempt to classify tropical
germplasm into distinct heterotic groups. He identified Tuxpeno in combination with Eto
and other Carribean flints as the best heterotic pairs. Since then, some progress has been
made towards the identification and formation of new heterotic groups for intermediate
and late maturing maize in the tropics (Vasal et al., 1992 a and b; Kim et al., 1999;
Menkir et al., 2003;). Results of studies conducted in IITA showed that the inbred lines
could not clearly be separated into dent vs flint, tropical vs temperate-derived or Tuxpeno
dent vs Carribean flint, (Kim and Ajala, 1996; Kim et al., 1999) probably due to the
mixing of the source populations without due regard to heterotic groups (Kim et al.,
1999; Wellhausen, 1978). TZB (Carribean mixed) x Across 7721 (Tuxpeno) and Suwan 1
x Across 728 (tropical dent) were identified as the best combinations. Despite the
progress made with the intermediate and late maturity groups in identifying new heterotic
groups, the heterotic patterns and the extent of diversity in the Striga resistant extra-early
germplasm in IITA- maize program have not been determined. Furthermore, no extra-
early inbred testers appropriate for developing stress tolerant hybrids are available in
inbreds would be invaluable to the maize breeding program at IITA and the national
maize programs in WCA. Furthermore, studies have shown that the heterotic patterns of
inbred lines and populations are influenced by the environment under which they are
evaluated (Gutierrez-Gaitan et al., 1986; Kim and Ajala, 1996; Vasal et al., 1993).
patterns of inbred lines and their hybrids under contrasting environmental conditions
5
would be invaluable for the development of an efficient breeding strategy for the growing
conditions in WCA.
information on the combining ability and heterotic patterns of the inbreds in the program.
hybrid program and the selection of parents for the development of synthetic varieties or
combining ability and heterotic patterns of the extra-early inbred lines would be
maximum genetic variability for further selection and the introgression of desirable genes
from diverse germplasm sources into the available genetic base (Thompson et al., 1998).
Diallel analysis provides the most complete and reliable information on general
combining ability (GCA) and specific combining ability (SCA) of inbred lines in hybrid
combinations (Sprague et al., 1942). It is very effective for determining the mode of gene
action, identifying superior parents for hybrid development, classifying inbred lines into
heterotic groups and for identifying appropriate testers for use in a breeding program
(Beck et al.,1991; Vasal et al.,1993; Kim, 1994; Bhatnagar et al., 2004; Menkir et al.,
2003; Gethi and Smith, 2004; Yallou et al., 2009; Badu-Apraku et al., 2011a, b).
However, because there were too many inbred lines to classify into heterotic groups in
6
the IITA Maize Program, the initial strategy was to use the line x tester analysis for the
grouping of our inbred lines. Our earlier attempt to classify extra-early white inbreds in
the IITA Program into heterotic groups using two IITA intermediate maturing inbred
testers, 1368 and 9071, under Striga infested and Striga free conditions showed that
none of the extra-early white inbred lines could be classified into heterotic groups under
any of the evaluation environments because the testers were not sufficiently effective
(Badu-Apraku, 2007, unpublished). The first successful attempt at classifying the extra-
early white and yellow inbreds in the program into heterotic groups was made in 2005
The genotype main effect plus genotype x environment interaction (GGE) biplot
proposed by Yan and Hunt (2002) is a powerful statistical tool for obtaining visual
information on the general combining ability (GCA) and specific combining ability
(SCA) of inbred lines, classifying inbred parents with a similar genetic background into
heterotic groups, and identifying superior hybrids. The GGE biplot allows data to be
presented graphically thus facilitating the understanding of the patterns of the data. As a
result, several researchers have used the GGE Biplot in studies involving diallel crosses
(Narro et al., 2003; Andio et al., 2004; Bhatnagar et al., 2004; Bertoia et al., 2006; Badu-
Apraku et al., 2011 a, b). The objectives of this study were to use a combination of the
conventional diallel analysis and the GGE biplot to (i) classify the extra-early white
endosperm inbred lines into contrasting heterotic groups, (ii) examine the genetic control
and modes of gene action for grain yield and other traits, (iii) identify appropriate inbred
testers for developing stress tolerant hybrids, (iv) determine the relationship between line
7
per se and hybrid performance and (v) study the yield stability of the extra-early hybrids
Genetic materials
Seventeen white endosperm lowland tropical extra-early maize inbred lines derived from
broad based populations formed from local x introduced germplasm were used in diallel
study (Table 1). The inbred lines were developed through the pedigree breeding method
with evaluations of the topcross performance at the S3 stage under Striga-infested and
Striga-free conditions. Details of the methodology adopted for the development of the
Four sets of experiments were conducted between the period 2007 and 2010. In the first
experiment, ninety extra-early white and yellow inbreds (including the seventeen inbreds
used in the diallel crosses) were evaluated for their agronomic performance under
induced moisture stress during the 2007/2008 and 2008/2009 dry seasons as well as
altitude, 1200 mm annual rainfall) and under terminal drought stress at Bagauda (drought
prone environment; 1200N, 822E, 580 msl, 800 mm annual rainfall) during the
growing seasons of 2008. In the second experiment, one hundred and thirty-six single
cross hybrids formed from diallel crosses involving seventeen extra early white inbred
lines plus four early maturing, drought tolerant and Striga resistant open-pollinated
8
check varieties were used for the study. Early maturing drought tolerant and Striga
resistant varieties were used as check varieties because at the time of this study there
were no extra-early maturing varieties that combined drought tolerance at flowering and
grain-filling with Striga resistance. The seventeen inbreds were selected based on their
superior performance under drought stress and well-watered conditions. All possible
crosses among the seventeen inbreds were made at Ibadan, Nigeria in 2008 and seeds
from the reciprocal crosses were bulked for the evaluations assuming no reciprocal effect
on drought tolerance and Striga resistance in maize. The diallel crosses and the checks
were evaluated for their agronomic performance under induced moisture stress during the
2008/2009 and 2009/2010 dry seasons at Ikenne and under well-watered conditions at
Ikenne, Zaria (high yield environment; 1111N, 738E, 640 msl, 1200 mm annual
rainfall), and Mokwa (9о18’N, 5о 4’E, 457 m altitude, 1100 mm annual rainfall) during
the growing seasons of 2009 and 2010. In addition, the diallel crosses were evaluated
under terminal drought stress at Bagauda during the planting seasons of 2009 and 2010. A
randomized incomplete block design with two replications was used in each trial. The
experimental units were one-row plots, each 5 m long with a row spacing of 0.75 m.
Distance between two adjacent plants within the row was 0.40 m in all trials. Three seeds
were planted per hill. The maize plants were thinned to two per hill about 2 weeks after
emergence to give a final plant population density of about 66,000 plants ha–1.
The induced drought stress was obtained by withdrawing irrigation water from 21
days after planting (DAP) until maturity so that the maize plants relied on stored water in
the soil for growth and development. During the first three weeks of growth, the plants
were irrigated using a sprinkler irrigation system which provided 12 mm of water per
9
week. The soil in the experiment station at Ikenne is eutric nitrosol (FAO classification)
and the experimental fields at the station are flat and fairly uniform, with high water-
holding capacity. The well-watered (rainfed) experiment at Ikenne was conducted during
the growing season. At Zaria, a high-yield and non-stress environment, the plants relied
to natural terminal drought stress. Except for the well-watered treatments, all
management practices were the same for both well-watered and drought stress studies.
Fertilizer was applied to the well-watered and drought stress plots at the rate of 60 kg ha –1
trials were kept weed-free with the application of Atrazine and Gramoxone as pre- and
by hand weeding.
In the third experiment, the 136 single cross hybrids plus the four check varieties
were evaluated separately under artificial Striga infestation at Mokwa and Abuja (90 16′
N 70 20′ E, 300 m altitude, 1500 mm rainfall) in the southern Guinea savanna of Nigeria
in 2010. The fourth experiment involved evaluation of 25 extra-early white and yellow
inbreds (including seven of the extra-early inbreds involved in the 136 single crosses)
were evaluated under artificial Striga infestation were evaluated at Mokwa and Abuja in
2008 and 2009. In each trial, a randomized incomplete block design with four replicates
was used. Each experimental unit was evaluated in two-row plots, 5 m long, spaced 0.75
m apart with 0.4 m between hills. Three maize seeds were sown per hill and thinned to
two plants per hill two weeks after planting to give a final population density of 66,000
plants ha-1. The Striga infestation was done according to the method of Kim, (1991);
10
Kim and Winslow (1991). Striga hermonthica seeds collected from sorghum fields in the
preceding season and stored for at least six months were used for the infestations.
Ethylene gas was injected into the soil before two weeks before artificial infestation in
order to stimulate suicidal germination of existing Striga seeds in the field. The plots
were arranged in such a way that Striga-infested blocks were placed back-to-back in
strips with non-infested blocks across the field. This arrangement reduced the movement
of Striga seeds into the non-infested plots. Apart from Striga infestation, management
practices were the same for both infested and non-infested plots. Fertilizer application
was delayed until about 21-25 days after planting when 30 kg ha -1 each of N, P, and K
were applied as NPK 15-15-15. Weeds other than Striga were controlled by hand
weeding.
Field Measurements
In the first and second experiments, data were recorded on both drought stress and
well-watered plots for days to 50% silking (DS), and days to anthesis (DA) as the number
of days from planting to when 50% of the plants had emerged silks and had shed pollen.
The anthesis-silking interval (ASI) was calculated as the difference between days to 50%
silking and 50% anthesis. Plant (PLHT) and ear heights (EHT) were measured as the
distance from the base of the plant to the height of the first tassel branch and the node
bearing the upper ear. Root lodging (the percentage of plants leaning more than 30 from
the vertical), and stalk lodging (the percentage of plants broken at or below the highest
ear node) were recorded. Stay green characteristic (LD) was recorded for the drought-
stress experiments at 70 DAP on a scale of 1 to 9, where 1 = almost all leaves green and 9
11
= virtually all leaves dead. Plant aspect (PASP) was recorded on a scale of 1 to 5 based on
overall plant type, where 1 = excellent plant type and 5 = poor plant type. Husk cover
was rated on a scale of 1 to 5, where 1 = husks tightly arranged and extended beyond the
ear tip and 5 = ear tips exposed. Ear aspect (EASP) was based on a scale of 1 to 5, where
1 = clean, uniform, large, and well-filled ears and 5 = ears with undesirable features. EPP
was computed by dividing the total number of ears harvested per plot by the number of
plants in a plot. For trials conducted under drought stress, harvested ears from each plot
were shelled to determine the percentage grain moisture. Grain yield in kg ha -1 was
computed from the shelled grain weight, adjusted to 15% moisture. For the well-watered
experiments a shelling percentage of 80% was assumed for all cultivars and grain yield
(obtained from ear weight and converted to kg ha-1) was adjusted to 15% moisture.
The observations made in the third experiment were the same as those outlined
for the first experiment except that no data were taken on the stay green characteristic. In
addition, host plant damage syndrome rating (Kim, 1991) and emerged Striga counts
were taken at 8 and 10 weeks after planting (WAP) (56 and 70 DAP) in the Striga–
infested plots. Striga damage syndrome ratings were scored per plot on a scale of 1–9
where 1 = no damage, indicating normal plant growth and high resistance, and 9 =
complete collapse or death of the maize plant, i.e., highly susceptible (Kim, 1991). A
shelling percentage of 80% was assumed for all cultivars and grain yield (obtained from
ear weight and converted to kg ha-1) was adjusted to 15% moisture. Even though data
were collected on several traits, only those on the most important traits in the studies are
12
Statistical analysis
Separate analyses of variance (ANOVA) were performed on data collected across years
and locations for each research condition (drought stress, well-watered, and Striga-
infested environments) with PROC GLM in SAS using a RANDOM statement with the
TEST option (SAS, 2001). Subsequently, combined ANOVA were carried out across all
10 test environments for grain yield, DS, ASI, EPP, PLHT, EASP; in eight test
environments for PASP; in four test environments for stay green characteristic and in two
test environments for Striga damage ratings and number of emerged Striga plants at 8
and 10 WAP. In the combined ANOVA, the location-year combinations, replicates, and
blocks were considered as random factors while entries (136 hybrids and 4 checks) were
considered as fixed effects. Means were separated using the LSD. General combining
ability (GCA) effects of the parents and specific combining ability (SCA) effects of the
crosses, as well as their mean squares in each environment and across environments,
were estimated on the 136 diallel crosses, excluding the checks, following
Griffing’s method 4 model 1 (fixed model) (Griffing, 1956) and the DIALLEL-SAS
program developed by Zhang et al. (2005) adapted to SAS software version 9.2 (SAS,
2001). Effects of GCA and SCA for the measured traits were computed from the mean
values adjusted for the block effects for each environment and across environments.
The statistical model used for the combined diallel analysis across environments is as
follows:
13
Yijk = μ + Ee + gi + gj + sij + gEeg + sEes + εijk
where Yijk is the observed measurement for the ijth cross grown in the kth
environment; μ is the grand mean; gi and gj are the GCA effects; sij is the SCA effect;
gEeg is the interaction effect between GCA and the Environment; sE es is the
interaction effect between SCA and the Environment, and εijk is the error term
associated with the ijth cross evaluated in the kth replication and Ee environment
(Hallauer and Miranda, 1988). The following restrictions were imposed on the
combining ability effects: gi=0, and sij=0 for each j (Griffing, 1956). GCA and
SCA effects were tested for significance using t-test. The standard errors of the GCA
and SCA effects were estimated as the square root of the GCA and SCA variances
(Griffing, 1956).
The estimated GCA effects of the inbred parents for each environment were
environments, and inbred x environment interaction. In addition, the GGE biplot was
used on the means adjusted for block effects to obtain information on the
performance of the single-cross hybrids under Striga infestation, drought stress, and
optimum growing environments, examine the combining ability of the inbreds, place
them in heterotic or tester groups, and identify the best testers across test
environments. Furthermore, the GGE biplot was used to obtain information on the
performance and yield stability of the single cross hybrids. The analyses were done
using GGE biplot, a Windows application that fully automates biplot analysis,
Where
Yij is the genetic value of the combination (pure line or hybrid) between Entry i and
Tester j for the trait of interest; j is the mean of all combinations involving Tester j; 1,
and 2 are the singular values for PC1 and PC2; ξi 1 and ξi2 are the PC1 and PC2
eigenvectors, respectively, for Entry i; ηj1 and ηj2 are the PC1 and PC2 eigenvectors,
respectively, for Tester j; and εij is the residual of the model associated with the
combination of Entry i and Tester j. Since in a diallel cross data, each genotype is an
entry and a tester, i and j can refer to the same or different genotypes.
stress conditions, a selection index was computed separately for drought stress and Striga
infestation using standardized data for selected variables. The best 20 and the worst ten
single crosses under Striga infestation were selected using a base index similar to that
used by Menkir and Kling (2007) and Badu-Apraku et al. (2010, 2011a, b), which
integrate grain yield, Striga emergence counts, Striga damage syndrome rating, and EPP
measured under infested conditions. The means of the selected traits were expressed on
standard deviation units and the index scores computed as I = [(2 x YLI) + EPP –
(STRA8 + STRA10) – 0.5 (STC8 + STC10)], where YLI was the yield of Striga-infested
plots, EPP was the number of ears at harvest in the Striga-infested plots, STRA8 and
STRA10 were Striga damage ratings at 8 and 10 WAP, STC8 and STC10 were number of
emerged Striga plants at 8 and 10 WAP. Similarly, the best and worst single crosses under
managed drought stress were identified, based on the index values calculated using the
following equation:
15
I = [(2 x Yield) + EPP - ASI - PASP - EASP -LD].
Since under both Striga infestation and managed drought stress each trait was
standardized, with a mean of zero and standard deviation of 1 to minimize the effects
of different scales, a positive value was an indication of the tolerance of the inbred
RESULTS
General and Specific Combining abilities of grain yield and other traits of Extra-
The analysis of variance (ANOVA) of the diallel crosses showed that the mean squares
for entries (G) and environments (E) were significant for all measured traits except the
Striga damage ratings at 8 WAP (Table 3). The mean squares for G x E interaction (GEI)
was significant for all traits except number of emerged Striga plants at 8 and 10 WAP and
Striga damage ratings at 8 and 10 WAP. Partitioning of the entries into GCA and SCA
components revealed that the mean squares for GCA and SCA were significant for all
traits except the SCA for the number of emerged plants at 8 and 10 WAP, Striga damage
rating at 8 WAP, and stay green characteristics. The GCA mean squares were larger than
those of the SCA for all measured traits across research environments. It is interesting to
note that the mean squares for GCA was about three times that of the SCA for grain
yield, ASI, EASP,EPP,number of emerged Striga plants at 8 and 10 WAP, and the stay
green characteristic. The GCA mean square was about five times that of the SCA for
Striga damage at 8 and 10 WAP. Both GCA x E and SCA x E mean squares were
16
significant for most traits with the exception of the number of emerged Striga plants and
The GCA effects of inbreds for grain yield were smaller than that of the SCA
under drought stress but higher under Striga infestation, optimal growing conditions and
across research environments (Fig.1). It is worth noting that the relative importance of
GCA decreased under drought stress compared to optimal conditions but increased under
Striga infestation.
The general combining ability effects of the extra-early inbreds for grain yield and
other traits under drought stress, Striga infestation, optimum growing conditions and
across research environments are presented in Table 4. Inbred TZEEI 39, TZEEI 13,
TZEEI 29 and TZEEI TZEEI 13 had the highest GCA effects for grain yield under
Drought stress, Striga infestation, and optimum growing conditions and across
environments, respectively. Positive and significant GCA effects were detected for
Striga damage and number of emerged Striga plants at 8 and 10 WAP for the inbreds,
TZEEI 55, TZEEI 90 and TZEEI 32, the only exception being Striga damage at 10
WAP which was not significant for TZEEI 32 . On the other hand, TZEEI 38 had
significant negative GCA effects for the Striga damage at 8 WAP and number of emerged
Striga plants at 8 and 10 WAP. No significant GCA effects were observed for the stay
green characteristic except TZEEI 21 which_ had a negative and significant GCA effects.
Environments.
17
The biplot of the diallel data accounted for 69.3% (50.6% for PC1 and 18.7% for PC2) of
the total variation across ten environments for the GCA effects of the single cross hybrids
(Figs. 2 and 3.). In the polygon view displayed in Fig. 2, TZEEI 13 was the vertex inbred
in the sector where environments MKSTR, IK2WW, ABSTR, IK1WW, IK2DT, IK1DT,
and BG1DT were located. Similarly, the inbred TZEEI 3 was the vertex inbred for the
environment, BG2DT, TZEEI 21 for ZR1WW, and TZEEI 29 for ZR2WW. There were
no environments found in the sector with TZEEI 38, and TZEEI 32 as the vertex inbreds
indicating that the hybrid combinations involving TZEEI 38 and TZEEI 49, TZEEI 4, and
TZEEI 39; TZEEI 32 and TZEEI 90 were not adapted to any of the test environments.
In the GGE biplot analysis, the absolute length of the projection of an inbred on to
the ATC y axis (horizontal broken lines originating from zero on the y axis) is a measure
of its stability. The shorter the projection, the more stable the inbred. Thus, the inbreds
TZEEI 2, TZEEI 46, and TZEEI 54were the most stable across test environments and had
yield performance which was above average (Fig. 3). The inbred TZEEI 13 had the
highest GCA effects across test environments but was one of the least stable inbreds.
According to Yan et al. (2000) the ideal genotype should have a high mean yield and high
stability. Based on this TZEEI 29 was identified as the ideal inbred across test
environments. The vector view of the genotype plus genotype x environment interaction
biplot showing the ranking of the inbreds based on both the discriminating and
18
The biplot dispay in Fig. 4 was used to investigate the relationships among test
environments. This biplot accounted for a total of 69.3% of the total variation in grain
yield (50.6% for PC1 and 18.7% for PC2). In the biplot figure, the rays connecting the
test environments to the biplot origin are referred to as test environment vectors. The
vector length of a test environment measures the magnitude (or discriminating power) of
its ability to asses the performance of cultivars under the test environments.Shorte
environmental vectors denote that the specific environments were not strongly correlated
with environments with longer vectors and that they were probably not strongly
correlated with one another either (Yan et al., 2010). Based on this interpretation, the
short vector test environments, IK1DT, IK2DT, BG1DT, and BG2DT all of which were
stressed environments except IK1WW may be considered as unique and independent test
had long vector lengths and were therefore more powerful in discriminating among the
cultivars. It is interesting to note that the test environments MKSTR, ABSTR, IK1WWW
and IK2WW had angles which were <900 and were therefore positively correlated. The
implication is that even though these environments had different stresses they were
expected to discriminate consistently among the inbreds. The test environments ZR1WW
and ZR2WW had long vector lengths and an angle which was <900 indicating that they
were closely related and will discriminate consistently among the genotypes.
Furthermore, the GGE biplot revealed very weak association between the test
environments ZR1WW, and ZR2WW on one hand and MKSTR, ABSTR, IK1WW and
IK2WW on the other.The fact that in general the drought stress, Striga-infested, and the
optimal growing conditions were each separated by the GGE biplot, justified our decision
19
to compare separately the diallel crosses under each research condition as well as across
General and Specific Combing Abilities of Inbreds under Drought stress, Striga-
Yan and Kang (2003) indicated that the SCA effects refer to crosses and
represented a residual that was not explained by the GCA effects. The small circle in the
average tester ordination view of the GGE biplot in Figs. 5, 7 and 8, represents an
average tester, which is defined by the average PC1 and PC2 values of all testers. The
line passing through the biplot origin and the average tester, with an arrow pointing to the
average tester, is referred to as the average tester axis or average tester coordinate (ATC)
absicissa. The length of the average tester vector (distance from biplot origin and the
average tester) determines the relative importance of the GCA versus entry × tester
interaction. The longer the average tester vector, the more important the GCA and the
more meaningful the ranking of parents based on GCA. In the present study, the length of
the average tester vector under drought stress and optimal growing environments was
short, and located near the biplot origin with testers on both sides of the biplot origin
relative to the ATC abscissa (Figs. 7 and 8) This implied that GCA effects of the inbred
lines were of minor importance uinder the two research environments. Therefore due to
the prepondearance of SCA, there were no inbreds that had significantly higher GCA than
inappropriate to classify the inbreds into distinct heterotic groups based on GCA and
SCA. Thus, the tester-vector views of the biplots have been used to provide invaluable
information on the relationship among the testers and their discriminating power under
20
drought stress and optimal growing environments. In contrast, the length of the average
tester vector in the average tester ordination view under the Striga-infested environments
was long, and located far from the biplot origin with testers on only one side of the biplot
origin relative to the ATC abscissa (Fig.5). Therefore due to the over dominating GCA,
the inbred lines could be placed in appropriate heterotic groups in the following sections.
The biplot of the inbreds across the Striga-infested environments explained 52.4% of the
total variation (35% for PC1 and 17.4% for PC2) (Fig. 5). Th e axis of the AEC abscissa
or “average environment axis” is the single-arrowed line that passes through the biplot
origin and the “average environment,” which is at the center of the small circle. In the
average tester ordination view of the GGE biplot (Fig. 5), the ATC abscissa (single-
arrowed horizontal line) approximates the average GCA effects of all entries. Points
located above this line represent positive GCA effects while those below the line are
negative GCA effects. The ATC ordinate (doubled-arrowed vertical line) and the ATC
abscissa divide the biplot into four right-angled triangles. The top right triangle contains
points with high positive GCA while the top left contains low positive GCA. Similarly,
the bottom right triangle contains high negative GCA effects and the bottom left
triangle contains low negative GCA effects. Values for the entries increase as the
concentric cycles become larger. Based on these interpretations, the entries TZEEI 13 and
TZEEI 14 had the highest positive GCA effects while entry TZEEI 3 and TZEEI 32 had
the lowest. The GCA effects of the entries were ranked as follows: TZEEI 13 ≈ TZEEI 14
21
TZEEI 3.The polygon view in Figure 6 revealed that Entry TZEEI 13 had the highest
GCA effect because it combined well with testers TZEEI 21, TZEEI 55, TZEEI 4, TZEEI
39, TZEEI 32, TZEEI 46, TZEEI 14 and TZEEI 90. On the other hand, Entry 14 had the
highest GCA because it combined well with Testers TZEEI 29, TZEEI 49, TZEEI 37,
TZEEI 6, TZEEI 57 and TZEEI 3. It is striking that Entries 13 and 14 had the highest
GCA in a wider range of environments than other inbreds (Fig. 2). The projections of the
entries onto the ATC ordinate (doubled-arrowed vertical line) approximated their SCA
effects (the deviation from the average effects), which indicated the tendency of the
entries to produce superior hybrids with specific testers. Entry TZEEI 13 had the highest
SCA effects (largest projection onto the ATC ordinate). In contrast, entries TZEEI 37 and
TZEEI 38 had the smallest SCA effects (smallest projection onto the ATC ordinate) (Fig.
5). Two heterotic groups could be identified, based on the biplot in Fig. 5. Inbreds
TZEEI 13, TZEEI 29, TZEEI 57, TZEEI 54 and TZEEI 3 constituted the first; TZEEI 14,
TZEEI 46, TZEEI 21, TZEEI 4, TZEEI 6, TZEEI 55, TZEEI 32 and TZEEI 39
constituted the second. Thus, forty crosses [TZEEI 13, TZEEI 29, TZEEI 57, TZEEI 54,
and TZEEI 3] × [TZEEI 14, TZEEI 46, TZEEI 21, TZEEI 4, TZEEI 6, TZEEI 55, TZEEI
32, TZEEI 39] are expected to exhibit hybrid vigor. Entries TZEEI 37, TZEEI 38, TZEEI
49, and TZEEI 90 were located near the ATC abscissa and did not belong to either of the
heterotic groups. These results imply that the inbreds in each heterotic group could be
selection. Subsequently, inbred lines could be extracted from each population for the
development of hybrid and synthetic cultivars. An ideal tester should have the largest
vector of all testers (i.e., be most discriminating) and have zero projection onto the ATC
22
ordinate (i.e., be most representative of testers). Thus the closer a tester’s marker was to
the ideal tester, the better it was. On this basis, TZEEI 14 was identified to be the closest
to the ideal tester (Figure not shown). The best tester identified in this study, TZEEI 14,
may be used in line × tester analysis to classify the numerous extra-early maturing white
The biplot of the extra-early inbreds under drought stress contributed 43.1% (26.2% of
the PC1 and 16.9% of the PC2) of the total variation (Fig. 7). In the average tester
ordination view of the GGE biplot shown in Fig. 7, the cosine of the angle between any
two tester vectors provides an approximation of the correlation coefficient between the
testers. The smaller the angle between any two testers, the more closely related they are.
Thus, TZEEI 21, TZEEI 46, TZEEI 14, TZEEI 13 and TZEEI 49, had angles less than
90° indicating positive correlation. On the other hand, TZEEI 38, TZEEI 6, TZEEI 37,
TZEEI 3, and TZEEI 39, and TZEEI 21, TZEEI 46, TZEEI 14, TZEEI 13 and TZEEI 49
had angles closer to 180° indicating that they were negatively correlated (Yan and Kang,
2003). In the biplot display, the rays connecting the tester label to the biplot origin
referred to as tester vectors. The vector length of a tester indicates the magnitude
(discriminating power) of its ability to assess the GCA of an inbred line. Testers with
shorter vectors were not strongly correlated with those with longer vectors and they were
probably not strongly correlated with one another either (Yan et al. (2010). Therefore, the
short-vector testers TZEEI 32, TZEEI 4, and TZEEI 29 may be regarded probably as
independent and unique testers. In contrast, the long-vector testers, TZEEI 55, TZEEI 90,
TZEEI 21, TZEEI 13, TZEEI 3, TZEEI 37, and TZEEI 6 were more powerful in
23
discriminating among the inbred lines under drought stress. The inbreds within each of
the three tester groups TZEEI 21, TZEEI 46, TZEEI 14, TZEEI 13 and TZEEI 49;
TZEEI 38, TZEEI 6, TZEEI 37, TZEEI 3, and TZEEI 39; TZEEI 90, TZEEI 57, TZEEI
55 and TZEEI 54 were identified as similar under drought stress (Fig. 7). Therefore, no
heterosis would be expected in hybrids formed by inbreds within any group. On the other
hand, hybrids among the three contrasting groups would be expected to show heterosis.
Testers TZEEI 32, TZEEI 4, and TZEEI 29 either had short vectors or were located near
the ATC abscissa and did not belong to any of the identified tester groups.
16.7% of the PC2) of the total variation (Fig. 8). The inbreds TZEEI 4, TZEEI 32, TZEEI
55, and TZEEI 90 had angles less than 90° indicating positive correlation. In contrast,
TZEEI 4, TZEEI 55 and TZEEI 32 had angles closer to 180° with TZEEI 14, TZEEI 49
and TZEEI 13 indicating that they were negatively correlated. The short-vector testers
TZEEI 38, TZEEI 39, TZEEI 54, TZEEI 6, TZEEI 29, and TZEEI 21 may be regarded as
independent and unique testers. On the other hand, the long-vector testers, TZEEI 3,
TZEEI 32, and TZEEI 46 were more powerful in discriminating among the inbred lines
under optimal growing conditions. The inbreds within each of the three tester groups
TZEEI 4, TZEEI 32, TZEEI 55, and TZEEI 90; TZEEI 46, TZEEI 49, TZEEI 13, TZEEI
and TZEEI 14; TZEEI 57, TZEEI 3 and TZEEI 37 were identified as similar under
hybrids formed by inbreds within any group. On the other hand, hybrids among the three
contrasting groups would be expected to show heterosis. Testers TZEEI 38, TZEEI 39,
24
TZEEI 54, TZEEI 6, TZEEI 29, and TZEEI 21 either had short vectors or were located
near the ATC abscissa and did not belong to any of the identified tester groups.
The combined ANOVA of the genotypes evaluated under drought stressed- and
significant for all traits except GEI for EPP under optimum growing environments (Table
4). Under artificial Striga infestation, G, and E mean squares were significant for all
measured traits except E for ASI and Striga damage at 8 WAP (Table 5). In contrast,
Under drought stress, the mean grain yield of the hybrids ranged from 1114 kg
ha-1 for TZEEI 14 x TZEEI 13 to 2734 kg ha -1 for TZEEEI 29 x TZEEI 21. The top-
ranking hybrid, TZEEI 29 x TZEEI 21 out-yielded the best Striga resistant and drought
differences in the grain yield of TZEEI 29 x TZEEI 57 and the check, TZE-W DT STR
C4. Overall, grain yield reduction of the genotypes under drought stress represented 56%
25
of what the average yield would have been in the same environment under well-watered
conditions.
Under artificial Striga infestation, TZEEI 29 x TZEEI 14 was the top ranking
hybrid and out-yielded the best Striga and drought tolerant early open-pollinated check
cultivar, TZE-W DT STR QPM by 24 % (Table 5). Grain yield reduction of the
genotypes under artificial Striga infestation was 62 % of what the average yield would
have been in the same environment under optimum management (Tables 4 and 5).
Stability analysis of the top 20 and worst five single cross hybrids and four open-
pollinated early check varieties, revealed TZEEI 29 x TZEEI 14 as the second highest
yielding and most stable single crosses across research environments while the highest-
yielding single cross hybrid, TZEEI6 x TZEEI 14 was the least stable (Fig.5). The
vector view of the genotype plus genotype x environment biplot which ranked the
hybrids based on the discriminating ability and representativeness revealed the hybrid
TZEEI 29 x TZEEI 14 as the closest to the ideal (Figure not shown). The best open-
pollinated early cultivar, TZE Comp 3 C3 produced grain yield which was above the
mean grain yield of the genotypes and was among the most stable across test
environments.
Discussion
The identification of parental inbred lines that form superior hybrids is the most costly
inbred lines does not predict the performance of maize hybrid for grain yield (Hallauer of
26
and Miranda, 1988). Predictors of single-cross hybrid value or heterosis between parental
inbred lines could therefore increase the efficiency of hybrid breeding programs.
The inability of the GGE biplot to separate the inbreds TZEI 22, TZEI 1, and
TZEI 26 into well-defied heterotic groups could be due to several reasons. Th e number
of inbred lines used in this study (nine) was probably too few and the genetic diversity
among the lines might have been too small. According to Beck et al. (1991) and
Bhatnagar et al. (2004), the genetic interpretation of a diallel with a few parental inbreds
erefore, combining abilities of the inbred lines used in this study could have been biased
by the correlation of gene frequencies. Another plausible explanation is that the GGE
biplot accounted for only 67.5% of the total variation in grain yield among the inbreds,
probably due to the complexity of the genetics among the inbreds in their resistance to
the stresses. Th ere is a need to employ molecular markers to characterize the inbred lines
References
Beck, D. L., S.K. Vasal, and J. Crossa. 1991. Heterosis and combining ability among
subtropical and temperate intermediate maturity maize germplasm. Crop Sci. 31:
68-73.
27
Bertoia , L., C. Lopez, and R. Burak. 2006. Biplot analysis of forage combining ability in
Griffing, B., 1956. Concept of general and specific combining ability in relation to diallel
Kim, S.K., 1994. Genetics of maize tolerance of Striga hermonthica. Crop Sci. 34:900-
907.
Betran, F.J., J.M. Ribaut, D. Beck, and D. Gonzalez De Leon. 2003. Genetic diversity,
Specific combining ability, and heterosis in tropical maize under stress and
Bhatnagar, S., F.J. Betran, and L.W. Rooney. 2004. Combining abilities of quality protein
Edmeades, G.O., M. Banzinger, S.C. Chapman, J.M. Ribaut, and J. Bolaños.1995. Recent
Gethi, J.G., and M.E. Smith. 2004. Genetic responses of single crosses of maize to Striga
hermonthica (Del.) Benth. and Striga asiatica (L.) kuntze. Crop Sci. 44, 2068-2077.
Kim, S.K., 1991. Breeding maize for Striga tolerance and the development of a field
infestation technique, p. 96–108. In: S.K Kim (ed.), Combating Striga in Africa.
28
Proceedings of the International Workshop organized by IITA, ICRISAT, and
Kim, S.K. 1994. Genetics of maize tolerance of Striga hermonthica. Crop Sci. 34: 900–
907.
Kim, S.K., and M.D. Winslow. 1991. Progress in breeding maize for Striga-tolerance/
resistance at IITA, p. 494-499. In: J.K. Ransom (ed.) Proc. 5th Int. Symposium on
Griffing, B. 1956. Concept of general and specific combining ability in relation to diallel
Yallou, C.G., A. Menkir, V.O. Adetimirin, and J.G.Kling. 2009. Combining ability of
maize inbred lines containing genes from Zea diploperennis for resistance to Striga
Sparry. 2010. Identifying essential test locations for oat breeding in eastern Canada. Crop
Yan, W., and M.S. Kang. 2003. GGE biplot analysis: A graphical
Yan, W., and L.A. Hunt. 2001. Interpretation of genotype by environment interaction for
29
Yan, W., and L.A. Hunt. 2002. Biplot analysis of diallel data. Crop Sci. 42:21-30.
Yan, W., L.A. Hunt, Q. Sheng, and Z. Szlavnics. 2000. Cultivar evaluation and mega-
Narro, L., S. Pande, J. Crossa, C. De Léon, and F. Salazar. 2003. Using line × tester
interaction for the formation of yellow maize synthetics tolerant to acid soils.
Euphytica 135:355-360.
30
Table 1. Description of the seventeen white-grained extra-early maize inbred lines used in
this study.
Reaction
Inbred Drought stress Striga
designation Pedigree hermonthica
TZEEI 3 TZEE-W SR BC5 x 1368 STR S7 Inb. 76 Tolerant Resistant
TZEEI 4 TZEE-W SR BC5 x 1368 STR S7 Inb. 85 Tolerant Resistant
TZEEI 6 TZEE-W SR BC5 x 1368 STR S7 Inb. 100 Tolerant Resistant
TZEEI 13 TZEE-W Pop x LD S6 (Set A) Inb. 27 Susceptible Resistant
TZEEI 14 TZEE-W Pop x LD S6 (Set A) Inb. 41 Tolerant Resistant
TZEEI 21 TZEE-W Pop x LD S6 (Set B) Inb. 44 Tolerant Resistant
TZEEI 29 TZEE-W SR BC5 x 1368 STR S7 Inb. 27 Tolerant Resistant
TZEEI 32 TZEE-W SR BC5 x 1368 STR S7 Inb 72. Tolerant Resistant
TZEEI 37 TZEE-W SR BC5 x 1368 STR S7 Inb. 82 Susceptible Resistant
TZEEI 38 TZEE-W SR BC5 x 1368 STR S7 Inb. 92 Tolerant Susceptible
TZEEI 39 TZEE-W SR BC5 x 1368 STR S7 Inb. 98 Tolerant Resistant
TZEEI 46 TZEE-W Pop x LD S6 (Set A) Inb. 17 Tolerant Unknown
TZEEI 49 TZEE-W Pop x LD S6 (Set A) Inb. 43A Tolerant Unknown
TZEEI 54 TZEE-W Pop x LD S6 (Set B) Inb. 5-2-2 Tolerant Unknown
TZEEI 55 TZEE-W Pop x LD S6 (Set B) Inb. 23-2-4 Susceptible Resistant
TZEEI 57 TZEEW- Pop x LDS6(Set B) Inb 23 Tolerant Unknown
TZEEI 90 TZEE-W Pop Co S6 Inb. 22-2-4 Susceptible Susceptible
31
Table 2. Characteristics of the experimental sites for the evaluation of hybrids formed
from a diallel among seventeen white-grained extra-early maize inbred lines.
32
Table 3. Mean squares from the combined analysis of variance of grain yield, days to silk, anthesis-silking interval, ear aspect and
ears per plant (across 10 environments), plant aspect (across eight environments), Striga rating at 8 and 10 WAP and Striga count at 8
and 10 WAP (across two environments) and leaf death scores (across four environments) of extra-early maturing white-grained inbred
lines evaluated in Nigeria, 2007-2009.
Source DF Grain yield, Days to Days to ASI Plant Ear Ears DF Plant DF Striga Striga Striga Striga D
kg ha-1 silk anthesis height, aspect per aspect emerge emerge damage damage
cm plant nce count nce count rating at rating at
at 8WAP at 10 8 WAP 10 WAP
WAP
Environment 9 777037629** 3180.5** 3754.8** 372.9** 138330** 566.0** 5.80** 7 33.4** 1 22.6** 44.8** 0.9ns 5.1* 3
(ENV)
Rep (ENV) 10 7459192** 56.5** 48.9** 6.9** 3489** 8.1** 0.03ns 8 2.6** 2 4.3** 3.7** 18.0** 11.0** 4
Entry 135 2508767** 33.6** 24.2** 4.6** 815** 1.4** 0.05** 135 0.5** 135 0.9** 0.6** 1.7** 2.4** 13
GCA 16 5791649.98** 130.6** 130.6** 13.5** 2923** 3.1** 0.12** 16 0.6** 16 2.1** 1.5** 6.4** 8.6** 16
SCA 119 2067371.47** 9.9** 9.9** 3.5** 532** 1.1** 0.04** 119 0.5** 119 0.7ns 0.5ns 1.0ns 1.5* 11
ENV*Entry 1215 1039808** 7.9** 5.5** 3.2** 264** 0.7** 0.03** 945 0.3** 135 0.5ns 0.3ns 1.0ns 1.4ns 40
GCA*ENV 144 1481459.9** 7.6** 7.6** 5.0** 349** 1.0** 0.05** 112 0.4** 16 0.6ns 0.4ns 0.6ns 1.2ns 48
SCA*ENV 1071 980426.3** 5.2** 5.2** 3.0** 253** 0.6** 0.02** 833 0.3** 119 0.5ns 0.3ns 1.1ns 1.4ns 35
Error 1350 693257 5.4 3.3 2.4 186 0.2 0.02 1080 0.2 270 0.6 0.4 1.0 1.2 53
*,** Significant F test at 0.05 and 0.01 levels of probability, respectively .
33
Table 4. GCA effects of inbred parents for grain yield and other agronomic traits evaluated under and
across drought stress, artificial Striga infestation, and optimal conditions between 2007 and 2009
Leaf death Striga damage Striga damage Striga emergence Striga emergence
Inbred Grain yield scores rating at 8 WAP rating at 10 WAP count at 8 WAP count at 10 WAP
DT STR OPT Across DT STR STR STR STR
TZEEI 6 86 -106 29 25ns -0.01ns -0.03ns 0.14ns -0.01ns -0.03ns
TZEEI 4 68 171 -85 28ns -0.14ns -0.11ns -0.36** -0.13ns -0.11ns
TZEEI 32 -101 -319 -188 -179ns 0.11ns 0.21* 0.21ns 0.23* 0.21*
TZEEI 29 66 176 320* 189ns -0.12ns 0.06ns -0.17ns 0.02ns 0.06ns
TZEEI 3 -77 -466* -17 -131ns 0.11ns 0.17ns 0.53** 0.24* 0.17ns
TZEEI 55 -12 -295 -26 -74ns -0.06ns 0.20* 0.43** 0.24* 0.20*
TZEEI 21 111 181 172 149ns -0.33** -0.05ns -0.35* -0.07ns -0.05ns
TZEEI 46 34 155 223 134ns -0.11ns -0.04ns -0.06ns -0.06ns -0.04ns
TZEEI 49 50 -56 -255 -93ns -0.05ns -0.16ns 0.32* -0.15ns -0.16ns
TZEEI 39 169 -208 -230 -66ns -0.01ns 0.08ns 0.34* 0.11ns 0.08ns
TZEEI 90 -83 -265 -234 -180ns 0.14ns 0.19* 0.41** 0.22* 0.19*
TZEEI 14 -50 581** 276* 207* 0.04ns -0.13ns -0.64** -0.18ns -0.13ns
TZEEI 13 6 676** 245* 236* 0.18ns -0.06ns -0.77** -0.14ns -0.06ns
TZEEI 57 -40 -13 35 -5ns 0.19ns -0.15ns 0.03ns -0.17ns -0.15ns
TZEEI 38 -142 -29 -257 -166ns -0.02ns -0.36** -0.26ns -0.40** -0.36**
TZEEI 37 -23 -227 -49 -74ns -0.09ns 0.14ns 0.19ns 0.17ns 0.14ns
TZEEI 54 -61 44 41 1ns 0.17ns 0.05ns 0.01ns 0.07ns 0.05ns
Standard Error 130 165 132 106 0.103 0.09 0.15 0.102 0.09
Relative
importance of
GCA over SCA 0.008 0.140 0.035 0.06 0.32 0.77 0.46 0.18 0.20
*,** Significant F test at 0.05 and 0.01 levels of probability, respectively
34
Table 5. Grain yield and other traits of extra-early maturing hybrids (the best 20 and the worst 10 based on the base index) and checks
evaluated under drought stress environments in Nigeria between 2007 and 2009.
HYBRID Category‡ Grain yield Days to Days to ASI Plant height, Ears per Plant Leaf
(kg ha-1 ) anthesis silk cm plant aspect death BA
Ear aspect scores IN
DT WW DT WW DT WW DT WW DT WW DT WW DT WW DT WW DT
TZEEI 29 x TZEEI 21 TxT 2734 5243 52 53 53 53 1 0.8 147 174 0.86 0.98 2.7 2.2 2.6 2.5 3.6 12
TZEEI 55 x TZEEI 49 SxT 2674 4368 48 51 50 51 1 0.6 142 173 0.88 0.96 3.0 2.5 3.0 2.9 3.8 7.8
TZEEI 39 x TZEEI 90 TxS 2667 4078 49 53 51 53 2 0.7 138 169 0.94 0.93 2.7 2.5 2.4 2.5 4.1 11
TZEEI 29 x TZEEI 49 TxT 2576 4960 50 51 51 52 2 1.1 145 171 0.89 0.93 2.7 2.7 2.7 2.7 3.5 10
TZEEI 3 x TZEEI 46 TxT 2576 5385 53 53 55 53 2 0.6 146 171 0.87 0.96 2.6 2.4 2.7 2.3 3.9 9.2
TZEEI 6 x TZEEI 21 TxT 2520 5080 51 52 53 53 2 0.9 152 184 0.78 0.93 2.4 2.1 2.9 2.8 3.7 7.1
TZEEI 39 x TZEEI 14 TxT 2498 4974 51 53 53 53 2 0.6 144 169 0.84 0.94 2.8 2.3 2.6 2.3 4.3 7.5
TZEEI 32 x TZEEI 13 TxT 2485 4606 51 52 54 53 3 0.7 152 175 0.88 1.01 2.7 2.4 2.7 2.6 4.1 7.4
TZEEI 29 x TZEEI 57 TxT 2436 5620 52 53 54 53 3 0.8 131 171 0.85 1.02 2.7 2.4 2.7 2.4 4.4 5.8
TZEEI 29 x TZEEI 3 TxT 2418 5081 52 52 54 52 2 1.1 130 176 0.84 0.98 2.7 2.4 2.8 2.4 3.8 6.7
TZEEI 55 x TZEEI 46 TxT 2410 4694 51 53 54 52 3 0.3 143 179 0.76 0.94 2.5 2.4 2.8 2.6 3.5 5.7
TZE-W DT STR C4 2391 5635 54 53 57 54 3 0.9 145 176 0.83 0.96 2.6 2.1 2.6 2.6 3.9 7.2
TZEEI 21 x TZEEI 39 TxT 2361 4522 51 52 52 53 1 0.8 146 167 0.81 1.02 2.5 2.5 2.5 2.6 3.9 9.7
TZEEI 4 x TZEEI 21 TxT 2360 5246 50 51 52 51 2 0.7 150 189 0.90 0.97 2.3 2.2 2.7 2.6 3.2 11
TZEEI 6 x TZEEI 46 TxT 2340 5264 51 52 53 52 2 0.9 140 177 0.77 0.98 2.6 2.3 2.7 2.4 3.5 7.0
TZEEI 46 x TZEEI 90 TxS 2336 3974 50 52 52 53 2 1.0 139 174 0.90 0.95 2.7 2.3 2.7 2.7 4.6 5.9
TZEEI 32 x TZEEI 54 TxT 2278 4973 52 53 55 54 3 1.2 144 180 0.86 1.04 2.8 2.3 2.8 2.5 3.9 5.6
TZEEI 29 x TZEEI 90 TxS 2275 5191 51 52 55 53 4 1.0 144 172 0.85 1.07 2.4 2.4 2.7 2.5 4.0 5.5
TZEEI 14 x TZEEI 37 TxS 2270 5052 53 52 55 52 3 1.0 145 173 0.87 1.02 2.8 2.5 2.8 2.6 3.9 5.3
TZEEI 46 x TZEEI 37 TxS 2232 4886 51 52 53 52 1 0.7 147 179 0.84 0.96 2.6 2.5 2.8 2.5 3.9 7.4
TZEEI 3 x TZEEI 49 TxT 2064 5051 49 50 51 51 2 1.0 135 162 0.89 1.00 3.0 2.6 2.8 2.7 3.7 5.1
TZE-W DT STR QPM C0 2012 4670 54 54 58 55 4 1.1 137 180 0.68 0.91 3.2 2.4 3.1 2.8 4.4 -5
AC 90 Pool 16 DT STR 1794 5367 53 53 56 55 3 1.4 149 177 0.79 0.91 2.6 2.3 3.1 2.6 3.9 -0
TZEEI 3 x TZEEI 39 TxT 1746 4209 50 51 54 52 4 0.8 130 159 0.63 0.91 3.0 2.6 3.1 2.8 4.3 -7
TZE Comp 3 C3 1731 6158 54 54 57 55 3 0.7 141 179 0.69 0.97 3.0 2.1 2.9 2.2 3.6 -1
35
TZEEI 6 x TZEEI 39 TxT 1637 3234 52 52 56 53 3 1.1 119 156 0.82 0.82 3.5 2.7 3.2 3.0 4.8 -8
TZEEI 38 x TZEEI 37 TxS 1508 3566 52 52 55 52 3 1.1 137 175 0.61 0.88 3.3 2.6 3.2 2.8 4.4 -9
TZEEI 4 x TZEEI 54 TxT 1380 4938 53 54 57 54 3 0.7 131 183 0.62 0.95 3.2 2.4 3.1 2.7 4.4 -9
TZEEI 3 x TZEEI 38 TxT 1337 3849 51 51 54 52 3 0.9 132 173 0.77 0.99 3.2 2.8 3.1 2.9 4.4 -7
TZEEI 55 x TZEEI 90 SxS 1306 3731 52 53 56 54 4 0.9 124 173 0.73 0.93 2.9 2.7 3.0 3.2 4.5 -7
TZEEI 21 x TZEEI 46 TxS 1282 3742 52 53 55 53 3 0.7 127 170 0.71 0.83 3.4 2.8 3.2 3.2 4.2 -9
TZEEI 3 x TZEEI 37 TxS 1245 3244 53 53 58 55 5 2.0 129 159 0.72 0.87 3.3 2.6 3.1 2.8 4.0 -1
TZEEI 57 x TZEEI 54 TxT 1131 3527 53 55 56 55 3 0.7 125 165 0.60 0.93 3.5 2.7 3.2 3.1 4.8 -1
TZEEI 14 x TZEEI 13 TxS 1114 2749 55 54 58 55 3 0.7 124 145 0.69 0.95 3.2 3.0 3.1 3.4 4.4 -8
Mean 2015 4583 52 52 54 53 3 0.9 138 173 0.77 0.95 2.9 2.5 2.9 2.7 4.2
LSD 643 693 1.6 1.2 2.3 1.3 1.4 0.6 11 8.9 0.13 0.11 0.5 0.33 0.4 0.37 0.7
P stat for Genotype (G) ** ** ** ** ** ** ** * ** ** ** ** ** ** ** ** **
P stat for Environment ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** **
(E)
P stat for GxE ** ** ** ** ** ** ** * ** ** ** ns ** ** * ** **
*, Significant different at 0.05 level of probability.
**, Significant different at 0.01 level of probability.
‡ T = Tolerant to drought stress, S = Susceptible to drought stress
36
Table 6. Grain yield and other traits of extra-early maturing hybrids (the best 20 and the worst
10 based on the base index) and checks evaluated under Striga infestation in Nigeria between
2008 and 2009.
Striga
Grain Striga damage Striga Striga
yield Plant Ears damage rating emergence emergence
(kg Days to Days to height, Ear per rating at at 10 count at 8 count at BASE
HYBRID ha-1 ) silk anthesis ASI cm aspect plant 8 WAP WAP WAP 10 WAP INDEX
TZEEI 4 x TZEEI 14 3127 55 55 1.5 145 4.0 0.9 3.1 3.5 17 22 11.6
TZEEI 4 x TZEEI 13 3246 56 54 0.4 150 4.7 0.8 3.2 3.9 18 30 11.4
TZEEI 21 x TZEEI 13 2862 54 52 1.6 152 4.4 1.1 3.2 4.3 11 12 10.8
TZEEI 29 x TZEEI 14 3385 56 55 0.4 142 4.6 1.0 3.7 4.4 20 26 10.6
TZEEI 39 x TZEEI 13 3317 53 52 1.2 158 3.8 0.8 3.6 3.9 21 25 9.8
TZEEI 6 x TZEEI 13 2640 54 53 2.4 158 4.2 0.9 3.5 3.9 20 24 8.7
TZEEI 29 x TZEEI 21 2628 57 55 3.9 139 4.9 0.8 3.2 3.9 18 29 8.7
TZEEI 6 x TZEEI 14 2865 53 52 2.0 132 4.2 0.8 3.8 4.3 12 14 8.3
TZEEI 14 x TZEEI 54 2673 56 54 2.0 121 5.0 0.9 3.8 4.2 19 24 8.2
TZEEI 13 x TZEEI 38 2666 55 54 2.5 138 4.7 0.8 4.1 4.2 16 25 8.2
TZEEI 32 x TZEEI 13 2718 56 55 2.9 136 4.8 1.0 3.9 4.1 22 40 8.0
TZEEI 21 x TZEEI 14 2865 55 54 0.9 125 4.6 0.8 4.0 4.5 16 21 7.8
TZEEI 29 x TZEEI 46 2720 54 54 1.8 139 4.6 0.9 3.8 4.9 14 21 7.7
TZEEI 14 x TZEEI 57 2993 55 55 -0.1 140 4.3 0.8 4.2 4.7 15 20 7.6
TZEEI 29 x TZEEI 13 2269 56 54 1.9 153 4.5 0.9 3.7 4.4 19 24 7.3
TZEEI 4 x TZEEI 54 2603 56 56 1.2 136 4.9 0.8 4.3 4.9 16 18 6.6
TZEEI 90 x TZEEI 13 2650 53 52 1.9 137 4.5 0.8 4.1 4.8 27 35 6.6
TZEEI 49 x TZEEI 38 2137 55 52 4.2 135 4.8 0.8 3.9 4.7 7 9 6.6
TZEEI 14 x TZEEI 13 2337 55 54 2.0 110 4.9 0.8 4.2 4.4 18 25 6.2
TZEEI 29 x TZEEI 37 2158 56 54 3.4 144 4.9 0.9 3.5 4.3 23 24 6.2
TZE-W DT STR QPM C0 2586 58 58 6.5 143 5.1 0.8 4.7 5.0 16 22 5.7
AC 90 Pool 16 DT STR 1598 57 58 5.0 151 5.3 0.6 4.8 5.7 19 24 -0.1
TZE-W DT STR C4 1859 56 55 3.0 141 4.8 0.7 5.5 5.9 28 29 -1.9
TZE Comp 3 C3 1396 58 59 6.1 137 5.1 0.5 5.0 5.7 31 32 -3.9
TZEEI 32 x TZEEI 3 838 57 57 5.6 125 5.9 0.5 5.3 6.0 35 45 -7.2
TZEEI 32 x TZEEI 55 1107 53 56 4.3 128 5.1 0.4 5.6 6.2 39 42 -7.4
TZEEI 32 x TZEEI 39 955 56 55 3.3 120 5.5 0.4 5.6 6.7 25 30 -7.7
TZEEI 37 x TZEEI 54 1104 54 58 5.7 124 5.8 0.5 5.5 6.3 49 52 -8.0
TZEEI 90 x TZEEI 37 1104 56 54 1.6 121 5.6 0.4 5.8 6.7 50 54 -8.6
TZEEI 46 x TZEEI 39 1078 52 54 4.1 128 5.9 0.4 5.8 7.2 43 48 -9.3
TZEEI 6 x TZEEI 90 1199 51 54 5.2 133 5.8 0.5 5.5 7.8 58 59 -9.4
TZEEI 3 x TZEEI 37 756 55 57 3.7 108 6.2 0.4 6.1 6.4 36 34 -9.5
TZEEI 13 x TZEEI 54 752 53 55 7.1 117 6.0 0.4 6.3 6.8 47 51 -10.2
TZEEI 3 x TZEEI 49 424 51 50 4.1 110 6.3 0.3 7.2 8.4 32 36 -15.7
Mean 1758 54 54 3 130 5.1 0.67 4.7 5.50 25.0 30.0
LSD 1119 2.9 3.4 3.2 22.5 1 0.2 1.3 1.4 18 20
P stat for Genotype (G) ** ** ** ** ** * ** ** ** * *
P stat for Environment (E) * ** ** ns ** ** ** ns * ** **
P stat for G x E ns ns ns ** ns ns ns ns ns ns ns
*,** Significant F test at 0.05 and 0.01 levels of probability, respectively
37
Table 7. Correlation between mid-parent value of the inbreds and the yield
performance of the single-cross hybrid evaluated under stress and nonstress environments.
Hybrid yield Hybrid yield Hybrid yield
under drought under Striga under optimal
stress infestation conditions
Mid-parent value under drought 0.028ns -0.1 -0.13
Mid-parent value under optimal -0.062 0.062ns -0.08
Mid-parent value under Striga
infestation -0.092ns 0.47* -0.098ns
Hybrid yield under drought . 0.12ns 0.48
Hybrid yield under optimal
conditions 0.48** 0.30** .
38
Figure 1. Proportion of additive (lower bar) and non additive (upper bar) genetic variance for grain yield at four stress and nonstress
environments and across environments in a diallel among seventeen white-grained maize inbred lines.
39
Environment Code
Ikenne, 2008 Drought stress IK1DT
Ikenne, 2009 Drought stress IK2DT
Ikenne, 2009 Well-watered IK1WW
Ikenne, 2010 Well-watered IK2WW
Bagauda, 2009 Terminal drought
stress BG1DT
Bagauda, 2010 Terminal drought
stress BG2DT
Zaria, 2009 Rainfed ZR1WW
Zaria, 2010 Rainfed ZR2WW
Abuja, 2010 Striga infested ABSTR
Mokwa, 2010 Striga infested MKSTR
Figure 2. A “ which wins where or which is best for what” genotype plus genotype x environment interaction
biplot for grain yield of hybrids (GCA effects) among 17 extra-early maturing white-grained maize inbred lines
evaluated at 10 stress and nonstress environments in Nigeria, 2007-2009. The data were not transformed
(‘Transform=0’), not standardized (‘Scale=0), and were environment-centered (‘Centering=2’). The biplot was
based on genotype-focused singular value partitioning (‘SVP = 2) and is therefore appropriate for visualizing the
relationships among genotypes. Principal components (PC1) and (PC2) for model 3 explained 69.3% of yield
variation.
40
Environment Code
Ikenne, 2008 Drought stress IK1DT
Ikenne, 2009 Drought stress IK2DT
Ikenne, 2009 Well-watered IK1WW
Ikenne, 2010 Well-watered IK2WW
Bagauda, 2009 Terminal drought
stress BG1DT
Bagauda, 2010 Terminal drought
stress BG2DT
Zaria, 2009 Rainfed ZR1WW
Zaria, 2010 Rainfed ZR2WW
Abuja, 2010 Striga infested ABSTR
Mokwa, 2010 Striga infested MKSTR
Figure 4. Average tester coordination view of the genotype plus genotype-by-environment (GGE) biplot showing
relationships among test environments based on grain yield of the hybrids in a diallel among 17 extra-early
inbreds.
41
42
Figure 45. The biplot view showing the ranking of the 17 extra-early maturing white-grained maize inbred lines
based on both the discriminating ability and representativeness for grain yield of hybrids (GCA effects) evaluated
under drought stress, Striga infested and optimal growing conditions in Nigeria, 2007-2009. The data were not
transformed (‘Transform=0’), not standardized (‘Scale=0), and were environment-centered (‘Centering=2’). The
biplot was based on genotype-focused singular value partitioning (‘SVP = 2) and is therefore appropriate for
visualizing the relationships among genotypes. Principal components (PC1) and (PC2) for model 3 explained
95.2% of yield variation.
43
Environment Code
Ikenne, 2008 Drought stress IK1DT
Ikenne, 2009 Drought stress IK2DT
Ikenne, 2009 Well-watered IK1WW
Ikenne, 2010 Well-watered IK2WW
Bagauda, 2009 Terminal
drought stress BG1DT
Bagauda, 2010 Terminal
drought stress BG2DT
Zaria, 2009 Rainfed ZR1WW
Zaria, 2010 Rainfed ZR1WW
Abuja, 2010 Striga infested ABSTR
Mokwa, 2010 Striga
infested MKSTR
Figure 3. An entry/tester genotype plus genotype x environment interaction biplot for grain yield in hybrids (GCA
effects) among 17 extra-early maturing white-grained maize inbred lines evaluated at 10 stress and nonstress
environments in Nigeria, 2007-2009. The data were not transformed (‘Transform=0’), not standardized
(‘Scale=0’), and were environment-centered (‘Centering=2’). The biplot was based on environment-focused
singular value partitioning (‘SVP = 1’) and is therefore appropriate for visualizing the relationships among
genotypes. Principal components (PC1) and (PC2) for model 3 explained 69.3% of yield variation.
44
Figure 67. Biplot based on diallel data of 17 extra-early-maturing white-grained maize inbred
lines showing relationship among testers in drought-stressed environments; average tester
coordination view. Exact positions of the entries and testers are at the beginning of the labels.
The circle indicates the average tester. ‘TZ’ followed by a number represents an inbred parent.
45
Figure 78. Average tester ordination view of the Genotype main effect plus genotype-by-
environment interaction (GGE) biplot based on diallel data of 17 white extra-early inbred lines
evaluated under artificial Striga infestation. ‘TZ’ followed by a number represents an inbred
parent.
46
Figure 89. Polygon view of the genotype main effect plus genotype-by-environment interaction
(GGE) biplot based on diallel data of 17 white extra-early inbred lines evaluated under
artificial Striga infestation. ‘TZ’ followed by a number represents an inbred parent.
47
Fiure 910. Biplot based on diallel data of 17 extra-early-maturing white maize inbred lines
showing relationship among testers under optimal environments; average tester coordination
view. Exact positions of the entries and testers are at the beginning of the labels. The circle
indicates the average tester. ‘TZ’ followed by a number represents an inbred parent.
48
Figure 10. A vector view of genotype plus genotype x environment interaction biplot showing the ranking of the
17 extra-early maturing white-grained maize inbred lines based on both the discriminating ability and
representativeness for grain yield of hybrids (GCA effects) evaluated under drought stress, Striga infested and
optimal growing conditions in Nigeria, 2008-2010. The data were not transformed (‘Transform=0’), not
standardized (‘Scale=0), and were environment-centered (‘Centering=2’). The biplot was based on genotype-
focused singular value partitioning (‘SVP = 2) and is therefore appropriate for visualizing the relationships among
genotypes. Principal components (PC1) and (PC2) for model 3 explained 69.3% of yield variation.
49
Code Hybrid
1 TZEEI 29 x TZEEI 14
2 TZEEI 4 x TZEEI 3
3 TZEEI 3 x TZEEI 37
4 TZE-W DT STR QPM C0
5 TZEEI 46 x TZEEI 57
6 TZEEI 14 x TZEEI 13
7 TZEEI 55 x TZEEI 14
8 TZEEI 3 x TZEEI 46
9 TZEEI 6 x TZEEI 57
10 TZEEI 3 x TZEEI 21
11 AC 90 Pool 16 DT STR
12 TZE Comp 3 C3
13 TZEEI 21 x TZEEI 54
14 TZEEI 32 x TZEEI 90
15 TZEEI 6 x TZEEI 13
16 TZEEI 3 x TZEEI 13
17 TZEEI 32 x TZEEI 13
18 TZEEI 4 x TZEEI 13
19 TZEEI 21 x TZEEI 13
20 TZEEI 14 x TZEEI 37
21 TZEEI 29 x TZEEI 54
22 TZEEI 55 x TZEEI 90
23 TZEEI 6 x TZEEI 14
24 TZEEI 39 x TZEEI 13
25 TZEEI 55 x TZEEI 13
26 TZEEI 21 x TZEEI 14
27 TZEEI 29 x TZEEI 57
28 TZEEI 29 x TZEEI 21
29 TZE-W DT STR C4
Figure 11. An entry/tester genotype plus genotype x environment interaction biplot of grain yield for 25
extra-early maturing maize hybrids and four early maturing varieties evaluated under drought stress,
Striga infested, and well-watered conditions in Nigeria, 2008-2010. The data were not transformed Code Environment
DS Drought stress
(‘Transform=0’), not standardized (‘Scale=0’), and were environment-centered (‘Centering=2’). The
biplot was based on environment-focused singular value partitioning (‘SVP = 1’) and is therefore WW Well-watered condition
appropriate for visualizing the relationships among genotypes. Principal component (PC)1 and PC2 for STR Striga infested condition
model 3 explained 93.7% of yield variation.
50
Figure 12. A vector view of genotype plus genotype x environment interaction biplot showing the
ranking of of grain yield for 25 extra-early maturing maize hybrids and four early maturing varieties
evaluated under drought stress, Striga infested, and well-watered conditions in Nigeria, 2008-2010. The
data were not transformed (‘Transform=0’), not standardized (‘Scale=0’), and were environment-
centered (‘Centering=2’). The biplot was based on environment-focused singular value partitioning
(‘SVP = 1’) and is therefore appropriate for visualizing the relationships among genotypes. Principal
component (PC)1 and PC2 for model 3 explained 93.7% of yield variation.
51
Code Hybrid
1 TZEEI 29 x TZEEI 14
2 TZEEI 4 x TZEEI 3
3 TZEEI 3 x TZEEI 37
4 TZE-W DT STR QPM C0
5 TZEEI 46 x TZEEI 57
6 TZEEI 14 x TZEEI 13
7 TZEEI 55 x TZEEI 14
8 TZEEI 3 x TZEEI 46
9 TZEEI 6 x TZEEI 57
10 TZEEI 3 x TZEEI 21
11 AC 90 Pool 16 DT STR
12 TZE Comp 3 C3
13 TZEEI 21 x TZEEI 54
14 TZEEI 32 x TZEEI 90
15 TZEEI 6 x TZEEI 13
16 TZEEI 3 x TZEEI 13
17 TZEEI 32 x TZEEI 13
18 TZEEI 4 x TZEEI 13
19 TZEEI 21 x TZEEI 13
20 TZEEI 14 x TZEEI 37
21 TZEEI 29 x TZEEI 54
22 TZEEI 55 x TZEEI 90
23 TZEEI 6 x TZEEI 14
24 TZEEI 39 x TZEEI 13
25 TZEEI 55 x TZEEI 13
26 TZEEI 21 x TZEEI 14
27 TZEEI 29 x TZEEI 57
Figure 13. Polygon view of genotype plus genotype x environment interaction biplot of grain yield for
25 extra-early maturing maize hybrids and four early maturing varieties evaluated under drought stress, 28 TZEEI 29 x TZEEI 21
Striga infested, and well-watered conditions in Nigeria, 2008-2010. The data were not transformed 29 TZE-W DT STR C4
(‘Transform=0’), not standardized (‘Scale=0’), and were environment-centered (‘Centering=2’). The
biplot was based on environment-focused singular value partitioning (‘SVP = 1’) and is therefore Code Environment
appropriate for visualizing the relationships among genotypes. Principal component (PC)1 and PC2 for DS Drought stress
52
Figure 14. The biplot view showing the ranking of the 17 extra-early maturing white-grained maize inbred lines based on
both the discriminating ability and representativeness for grain yield of hybrids evaluated under Striga infested conditions.
The data were not transformed (‘Transform=0’), not standardized (‘Scale=0), and were environment-centered
(‘Centering=2’). The biplot was based on genotype-focused singular value partitioning (‘SVP = 2) and is therefore
appropriate for visualizing the relationships among genotypes. Principal components (PC1) and (PC2) for model 3 explained
95.2% of yield variation.
53
Figure 16. The biplot view showing the ranking of the 17 extra-early maturing white-grained maize testers based on both the
discriminating ability and representativeness for grain yield of hybrids evaluated under drought stress conditions. The data
were not transformed (‘Transform=0’), not standardized (‘Scale=0), and were environment-centered (‘Centering=2’). The
biplot was based on genotype-focused singular value partitioning (‘SVP = 2) and is therefore appropriate for visualizing the
relationships among genotypes. Principal components (PC1) and (PC2) for model 3 explained 95.2% of yield variation.
54
Figure 15. The biplot view showing the ranking of the 17 extra-early maturing white-grained maize testers based on both the
discriminating ability and representativeness for grain yield of hybrids evaluated under optimal growing conditions. The data
were not transformed (‘Transform=0’), not standardized (‘Scale=0), and were environment-centered (‘Centering=2’). The
biplot was based on genotype-focused singular value partitioning (‘SVP = 2) and is therefore appropriate for visualizing the
relationships among genotypes. Principal components (PC1) and (PC2) for model 3 explained 95.2% of yield variation.
55
Fiure 16. Biplot based on diallel data of 17 extra-early-maturing white maize inbred lines showing
relationship among testers across environments; average tester coordination view. Exact positions of the
entries and testers are at the beginning of the labels. The circle indicates the average tester. ‘TZ’
followed by a number represents an inbred parent.
56
Figure 17. The biplot view showing the ranking of the 17 extra-early maturing white-grained maize testers based on both the
discriminating ability and representativeness for grain yield of hybrids evaluated under across environment. The data were
not transformed (‘Transform=0’), not standardized (‘Scale=0), and were environment-centered (‘Centering=2’). The biplot
was based on genotype-focused singular value partitioning (‘SVP = 2) and is therefore appropriate for visualizing the
relationships among genotypes. Principal components (PC1) and (PC2) for model 3 explained 95.2% of yield variation.
57