Journal of Plant breeding

Genetic Analysis of Grain yield and other Traits of Extra-Early Inbreds

and their Hybrids under Contrasting Environments

ABSTRACT

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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

decade in sub-Saharan West Africa (DNRP-GAPCC, 2000). Furthermore, the projected

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

improve productivity and ensure stable production of the crop in WCA.

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

to an intensification of crop production, reduced fallow periods, and consequently, to low

soil fertility (Vogt et al., 1991), a condition highly suitable for Striga infestation. Many

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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

resistance exemplified by reduced level of Striga attachment on the host, or tolerance

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

cultivars with resistance to Striga.

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

(IITA)-West and Central Africa Collaborative Maize Research Network (WECAMAN)

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

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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,

Ghana, Nigeria and Niger. Results of evaluation of 90 extra-early inbreds

under low N and drought stress in multiple location trials in Nigeria

over a period of two years to determine their potential for hybrid

production have confirmed that there are now available in sub-Saharan

Africa not only low-N tolerant inbreds and hybrids but also, drought-

tolerant extra-early inbreds and hybrids that are not only drought

escaping (a characteristics of extra earliness), but carry genes that are

tolerant to drought at flowering and grain-filling periods (Badu-Apraku,

2010, unpublished). Furthermore, inbreds and hybrids combining

tolerance to both stresses are also now available. The availability of

the low-N and drought tolerant extra-early inbreds and hybrids should

go a long way in reducing the instability of maize yields in sub-Saharan

Africa due to drought stress and low-N deficiency, especially in the

savannas as well as during the second season in the forest ecologies.

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

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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

WCA. Information on heterotic groups and identification of testers in the extra-early

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).

Therefore, information on the performance, combining ability and stability of heterotic

patterns of inbred lines and their hybrids under contrasting environmental conditions

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would be invaluable for the development of an efficient breeding strategy for the growing

conditions in WCA.

Hybrid development and promotion in maize is a promising strategy to

appreciably increase maize production and productivity and revolutionalize agriculture in

WCA. Presently, there are no commercial extra-early maturing hybrids. An important

requirement for a commercially successful hybrid program is the availability of

information on the combining ability and heterotic patterns of the inbreds in the program.

Accurate assessment of inbred lines in hybrid combinations is crucial to the success of a

hybrid program and the selection of parents for the development of synthetic varieties or

introgression into an elite source population. Knowledge and understanding of the

combining ability and heterotic patterns of the extra-early inbred lines would be

invaluable in planning crosses, assigning inbred lines to specific heterotic groups,

identification of diverse parental combinations to create segregating progenies with

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

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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

using multivariate analysis (Badu-Apraku et al., 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

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per se and hybrid performance and (v) study the yield stability of the extra-early hybrids

across stress and nonstress environments.

MATERIALS AND METHODS

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

inbred lines have been described by Badu-Apraku et al. (2006).

Field Evaluations and Stress Managements

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

under well-watered conditions in 2008 and 2009 at Ikenne (6º53’N, 30º42’E, 60 m

altitude, 1200 mm annual rainfall) and under terminal drought stress at Bagauda (drought

prone environment; 1200N, 822E, 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

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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; 1111N, 738E, 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

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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

on natural rainfall. At Bagauda, a drought-prone environment, the plants were subjected

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

each of N, P and K at planting. An additional 60 kg ha –1 N was top-dressed at 2 WAP. The

trials were kept weed-free with the application of Atrazine and Gramoxone as pre- and

post-emergence herbicides at 5 L/ha each of Primextra and Paraquat, and subsequently,

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

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= 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

presented in the results.

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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:

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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

subjected to GGE biplot analysis to assess the relationships among inbreds,

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,

version 5.4 (Yan 2001). The program is available at www.ggebiplot.com (verified 1

December 2011). The GGE biplot model equation is:

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 Yij - j = λ1ξi1ηj1 + λ2ξi2ηj2 + εij

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.

To identify productive single-cross hybrids for commercial production under

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:

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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

lines to the particular stress, while a negative value indicated susceptibility.

RESULTS

General and Specific Combining abilities of grain yield and other traits of Extra-

early White Inbreds across Growing Environments

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

Striga damage at 8 and 10 WAP.

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.

GGE Biplot Analysis of Stability of Extra-early Inbreds across Contrasting

Environments.

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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

representativeness identified TZEEI 14 as the closest to the ideal genotype across

research environments (Figure not shown).

Relationship among Test environments

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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

environments. In contrast, MKSTR, IK2WW, IK1WW, ABSTR, ZR1WW, and ZR2WW

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

stress and nonstress environments.

General and Specific Combing Abilities of Inbreds under Drought stress, Striga-

infested and Optimal Growing Environments

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

others. Consequently, no inbreds could be identified as ideal testers. It was therefore

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.

Heterotic Groups of Extra-early Inbreds under Striga-infested Environments

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

> TZEEI 29 >TZEEI 46 ≈ TZEEI 21≈ TZEEI 57 ≈ TZEEI 54 ≈ TZEEI 4 ≈ TZEEI 6 ≈

TZEEI 37 ≈ TZEEI 38 ≈ TZEEI 49 ≈ TZEEI 55 ≈ TZEEI 39 ≈ TZEEI 90 > TZEEI 32 ≈

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

recombined to form heterotic populations which could be improved through recurrent

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

inbreds in the IITA Maize Program.

Tester Groupings of Extra-early Inbreds under Drought Environments

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.

Tester Groupings of Extra-early Inbreds under Optimal Growing Environments

Under optimal growing conditions, the biplot contributed 40.9% (24.2% of the PC1 and

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

optimal growing conditions (Fig. 8). 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 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.

Tester Groupings of Extra-early Inbreds Across Environments

Performance of Single Cross Hybrids and Checks under Drought Stressed,

Optimum growing and Striga- infested Environments

The combined ANOVA of the genotypes evaluated under drought stressed- and

optimum-growing environments from 2008-2010 showed that G, E, and GEI were

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,

significant G EI was detected for only ASI.

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

tolerant early maturing open-pollinated variety, TZE-W DT STR C4 by 13 %. Under

well-watered conditions, the top-yielding hybrid was TZEEI 29 x TZEEI 57 (5620 kg ha -

1
) while the lowest was TZEEI 14 x TZEEI 13 (2749 kg ha -1). There were no significant

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

and time-consuming phase in maize hybrid development. Per se performance of maize

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

can be biased by lack ofindependent distribution of genes in the parental lines. Th

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

that could not be placed into distinct heterotic groups.

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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.

Environment Location Year Season Type of environment Code

1 Ikenne 2008 Dry season Severe drought stress IK1DT
2 Ikenne 2009 Dry season Severe drought stress IK2DT
3 Ikenne 2009 Dry season Well watered IK1WW
4 Ikenne 2010 Dry season Well watered IK2WW
5 Bagauda 2008 Rainy season Terminal drought stress BG1DT
6 Bagauda 2009 Rainy season Terminal drought stress BG2DT
7 Zaria 2009 Rainy season Well watered ZR1WW
8 Zaria 2010 Rainy season Well watered ZR2WW
9 Abuja 2010 Rainy season Striga infested ABSTR
10 Mokwa 2010 Rainy season Striga infested MKSTR

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

model 3 explained 93.7% of yield variation. WW Well-watered condition

STR Striga infested condition

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