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Journal of Biology, Agriculture and Healthcare
ISSN 2224-3208 (Paper) ISSN 2225-093X (Online)
Vol.3, No.12, 2013

www.iiste.org

Assessment of Genotype by Environment interactions and Grain
Yield Performance of Extra-Early Maize (Zea Mays L.) Hybrids
G.B. Adu1*, R. Akromah2, M.S. Abdulai1, K. Obeng-Antwi3, A.W. Kena2, K.M.L. Tengan3 and H. Alidu1
1.

CSIR-Savanna Agricultural Research Institute, Nyankpala, Tamale, Ghana

2.

Department of crops and soil sciences, Faculty of agriculture, Kwame Nkrumah University of
Science and Technology, Kumasi, Ghana

3.

CSIR-Crops Research Institute, Kumasi, Ghana

* E-mail of the corresponding author: gloriaboakyewaa@yahoo.com
Abstract
Maize (Zea mays L.) is one of the most important cereal crops of Ghana in terms of production and consumption.
Currently, it is produced in all the agro-ecological zones of the country. In Ghana, Genotype by Environment
interactions (GxE) effects on maize grain yield is usually significant due to the diverse environmental conditions
at growing sites. A proper understanding of the effects of GxE on variety evaluation and cultivar
recommendations is vital. The study was conducted in 2011 at three locations in Ghana to (i) determine the
presence of GxE of 100 extra-early maize genotypes and (ii) To use the GGE biplot methodology to determine
grain yield performance and stability of the genotypes evaluated across three environments. The effects of
genotype and environment were significant (P < 0.01) for grain yield. However, GxE was not significant for the
same trait. TZEEI 8 x TZEEI 51, TZEEI 5 x TZEEI 53, TZEEI 21 x TZEEI 39, TZEEI 27 x TZEEI 36 and
TZEEI 4 x TZEEI 6 were identified as high yielding and most stable hybrids. Therefore, these hybrids have the
potential for production across the test locations as well as others within the same agro-ecological zones. On the
contrary, DODZI, TZEEI 23 x TZEEI 6, TZEEI 19 x TZEEI 24, TZEEI 11 x TZEEI 24 and TZEEI 20 x TZEEI
39 were not only low yielding but also among the least stable genotypes.
Keywords: Zea mays, single-cross, stability, multiple environments, GxE
1. Introduction
Maize (Zea mays L.) is produced in all the agro-ecological zones of Ghana and its highest production is in the
transition zone. Maize yields in Ghana are well below their attainable levels; averaging approximately 0.5 metric
tons per hectare. However, yields as high as 5.5 metric tons per hectare have been realized by farmers using
improved seeds, fertilizer, mechanization and irrigation (MiDA 2010). Lower yields have been attributed to the
use of low-yielding varieties, use of self produced seed, poor soil fertility and limited use of fertilizers, low plant
population, and inappropriate weed control methods. Moreover, significant potential improvements in yields
could be achieved through the use of hybrid maize varieties.
Crop breeders have been striving to develop genotypes with superior grain yield, quality and other desirable
characteristics over a wide range of different environmental conditions. Genotype by environment interaction
(GxE) makes it difficult to select the best performing and most stable genotypes. GxE refers to the differential
ranking of genotype among locations or years. It is an important consideration in plant breeding programmes
because it impedes progress from selection in any given environment (Yau 1995). In West and Central Africa,
GxE interaction effects have been reported in maize cultivars (Fakorede & Adeyemo 1986; Badu-Apraku et al.
2007; 2008). Various studies have been conducted to study the effects of genotype by environment interaction in
Sub-Saharan Africa and on Ghanaian maize varieties. However, the changing environmental conditions, the
expansion of maize into new agro-ecologies, coupled with inadequate maize varieties available for the different
environments necessitate a rigorous and continuous study of GxE interaction effects for a dynamic crop
improvement programme.
There are many statistical methods available to analyse GxE: for example, combined ANOVA, stability analysis
and multivariate methods. Combined analysis of variance (ANOVA) is more often used to identify the existence
of GxE interactions in multi-environmental experiments. However, the main limitation of this analysis is the
assumption of homogeneity of variance among environments required to determine genotype differences.
Although this analysis allows the determination of the components of variance arising from different factors
(genotype, environment and the GxE), it does not allow exploring the response of the genotypes in the non-

7
Journal of Biology, Agriculture and Healthcare
ISSN 2224-3208 (Paper) ISSN 2225-093X (Online)
Vol.3, No.12, 2013

www.iiste.org

additive term: the GxE (Zobel et al. 1988). Among the statistical analyses proposed for the interpretation of the
GxE based on the use of biplots, the AMMI (additive main effect and multiplicative interaction) model stands
out due to the largest group of technical interpretations available (Duarte & Vencovsky 1999). AMMI analysis
interprets the effect of the genotype (G) and sites (E) as additive effects plus the GxE as a multiplicative
component and submits it to principal component analysis. Yan et al. (2000) proposed a modification of the
conventional AMMI analysis called GGE that has been used for GxE analysis. The GGE analysis pools genotype
effect (G) with GE (multiplicative effect) and submits these effects to principal component analysis. This biplot
is identified as a GGE biplot. The GGE biplot has been recognized as an innovative methodology in biplot
graphic analysis to be applied in plant breeding. Fan et al. (2007) showed that the GGE biplot methodology was
a useful tool for identifying locations that optimized hybrid genotypes performance and for making better use of
limited resources available for the maize testing programmes.
The objectives of this study were (i) to determine the presence of GxE of 100 extra-early maize genotypes and
(ii) to use the Genotype main effect plus Genotype by Environment interaction (GGE) biplot methodology to
determine grain yield performance and stability of the genotypes evaluated across three environments.
2. Materials and Methods
2.1 Field evaluations
Ninety-eight single-cross maize hybrids, one local three-way hybrid (Akposoe) and an open pollinated variety
(Dodzi) (Table 1) were evaluated at Fumesua, Ejura and Kpeve from April, 2011 to July, 2011 major season.
The evaluation sites are located in the deciduous forest, transition and coastal Savannah zones of Ghana,
respectively (Table 2). The hybrids were developed from 61 extra-early maturing white and yellow endosperm
inbred lines developed for grain yield, drought and Striga hermonthica tolerance. A standard protocol was
adopted at each site during the period of evaluation. Evaluations were done under rain fed conditions and fields
were planted when the rains at each experimental site had become fully established. The genotypes were planted
in an incomplete block design (10 x 10) with two replications. Each plot consisted 2-rows of 5 m long, an interrow spacing of 75 cm and an intra-row spacing of 40 cm. Three seeds were sown per hill and seedlings later
thinned to two after emergence and seedling establishment. Pre-emergence chemical weed control was practised
and comprised an application of a combination of Pendimethalin [N-(1-ethylpropyl)-3, 4-dimethyl-2, 6dinitrobenzenamine] and Gesaprim [2-chloro-4-(ethylamino)-6-(isopropylamino)-s-triazine] at 1.5 Lha-1 and 1.0
Lha-1 active ingredient, respectively at planting. Hand weeding was also done when necessary to control weeds
during the growing period. NPK 15-15-15 fertilizer was applied at the rate of 30 kg N ha-1 and 60 kg P2O5 ha-1 as
basal fertilizer at 1-2 weeks after planting and top-dressed with additional N at 60 kg N ha-1 at four weeks after
planting. Other management practices were done according to the recommendations of the specific areas.
2.2 Collection of agronomic data
Data recorded include days to anthesis, days to silking, plant and ear heights, root lodging (number of plants
leaning more than 300 from vertical), stalk lodging (stalks broken at or below highest ear node), ear aspect,
number of plants harvested, number of ears harvested and percent moisture in the grains. Days to anthesis and
days to silking were calculated as the number of days from planting to when 50 % of the plants had shed pollen
and had emerged silks, respectively. Anthesis-silking interval was determined as the difference between days to
silking and days to anthesis. Plant and ear heights were measured as the distance from the base of the plant to the
height of the flag leaf and the node bearing the upper ear, respectively. Husk cover was rated on a scale of 1 to 5,
where 1= husk tightly arranged and extended beyond the ear tip and 5 = ear tips exposed. Ear aspect was based
on a scale of 1 to 5, where 1= clean, uniform, large, and well-filled ears and 5 = ears with undesirable features.
Number of ears per plant was obtained by dividing the total number of ears per plot by the number of plants
harvested. The grain yield in kilograms per plot recorded was converted to grain yield in tons per hectare at 15 %
grain moisture based on 80% shelling percentage. Even though data were collected on several traits, only those
on the most important trait in the study are presented in the results.

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Journal of Biology, Agriculture and Healthcare
ISSN 2224-3208 (Paper) ISSN 2225-093X (Online)
Vol.3, No.12, 2013

www.iiste.org

Table 1. Description of extra-early maturing maize genotypes tested across three environments in 2011
Entry

Entry name

Entry

Entry name

Entry

Entry name

1

TZEEI 1 x TZEEI 2

34

TZEEI 9 x TZEEI 58

67

TZEEI 20 x TZEEI 22

2

TZEEI 1 x TZEEI 4

35

TZEEI 9 x TZEEI 59

68

TZEEI 20 x TZEEI 24

3

TZEEI 1 x TZEEI 5

36

TZEEI 9 x TZEEI 60

69

TZEEI 20 x TZEEI 39

4

TZEEI 1 x TZEEI 13

37

TZEEI 10 x TZEEI 22

70

TZEEI 21xTZEEI 20

5

TZEEI 1 x TZEEI 14

38

TZEEI 11 x TZEEI 8

71

TZEEI 21 x TZEEI 24

6

TZEEI 1 x TZEEI 22

39

TZEEI 11 x TZEEI 22

72

TZEEI 21 x TZEEI 39

7

TZEEI 1 x TZEEI 23

40

TZEEI 11 x TZEEI 24

73

TZEEI 22 x TZEEI 6

8

TZEEI 1 x TZEEI 50

41

TZEEI 12 x TZEEI 19

74

TZEEI 22 x TZEEI 24

9

TZEEI 2 x TZEEI 1

42

TZEEI 13 x TZEEI 1

75

TZEEI 22 x TZEEI 36

10

TZEEI 2 x TZEEI 11

43

TZEEI 13 x TZEEI 6

76

TZEEI 22 x TZEEI 51

11

TZEEI 2 x TZEEI 14

44

TZEEI 13 x TZEEI 12

77

TZEEI 23 x TZEEI 4

12

TZEEI 2 x TZEEI 24

45

TZEEI 13 x TZEEI 22

78

TZEEI 23 x TZEEI 5

13

TZEEI 4 x TZEEI 6

46

TZEEI 14 x TZEEI 4

79

TZEEI 23 x TZEEI 6

14

TZEEI 4 x TZEEI 7

47

TZEEI 14 x TZEEI 6

80

TZEEI 23 x TZEEI 36

15

TZEEI 4 x TZEEI 24

48

TZEEI 14 x TZEEI 22

81

TZEEI 23 x TZEEI 39

16

TZEEI 4 x TZEEI 39

49

TZEEI 14 x TZEEI 39

82

TZEEI 23 x TZEEI 40

17

TZEEI 4 x TZEEI 51

50

TZEEI 14 x TZEEI 49

83

TZEEI 23 x TZEEI 50

18

TZEEI 4 x TZEEI 90

51

TZEEI 14 x TZEEI 90

84

TZEEI 23 x TZEEI 90

19

TZEEI 5 x TZEEI 4

52

TZEEI 15 x TZEEI 6

85

TZEEI 24 x TZEEI 51

20

TZEEI 5 x TZEEI 23

53

TZEEI 1 x TZEEI 8

86

TZEEI 26 x TZEEI 24

21

TZEEI 5 x TZEEI 39

54

TZEEI 15 x TZEEI 19

87

TZEEI26 x TZEEI 51

22

TZEEI 5 x TZEEI 40

55

TZEEI 15 x TZEEI 21

88

TZEEI 26 x TZEEI 53

23

TZEEI 5xTZEEI 50

56

TZEEI 15 x TZEEI 22

89

TZEEI 27 x TZEEI 36

24

TZEEI 5 x TZEEI 53

57

TZEEI 15 x TZEEI 24

90

TZEEI 27 x TZEEI 51

25

TZEEI 6 x TZEEI 4

58

TZEEI 15 x TZEEI 39

91

TZEEI 29 x TZEEI 26

26

TZEEI 6 x TZEEI 5

59

TZEEI 19 x TZEEI 6

92

TZEEI 29 x TZEEI 33

27

TZEEI 6 x TZEEI 23

60

TZEEI 19 x TZEEI 8

93

TZEEI 29 x TZEEI 36

28

TZEEI 6 x TZEEI 36

61

TZEEI 19 x TZEEI 21

94

TZEEI 30 x TZEEI 4

29

TZEEI 6 x TZEEI 39

62

TZEEI 19 x TZEEI 22

95

TZEEI 30 x TZEEI 23

30

TZEEI 6 x TZEEI 40

63

TZEEI 19 x TZEEI 24

96

TZEEI 30 x TZEEI 36

31

TZEEI 6 x TZEEI 90

64

TZEEI 20 x TZEEI 8

97

TZEEI 30 x TZEEI 39

32

TZEEI 8 x TZEEI 24

65

TZEEI 20 x TZEEI 19

98

TZEEI 31 x TZEEI 8

33

TZEEI 8 x TZEEI 51

66

TZEEI 20 x TZEEI 21

AKPOSOE (Three-way)

100

DODZI (OPV)

Check
99

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Journal of Biology, Agriculture and Healthcare
ISSN 2224-3208 (Paper) ISSN 2225-093X (Online)
Vol.3, No.12, 2013

www.iiste.org

Table 2. Description of the test environments used in the study
Location

Latitude

Longitude
1 o 37’E

Altitude (m
ASL)
228.7

Mean
Seasonal
Rainfall *(mm)
599.7

Agroecological zone
Transition

Ejura

7o 38’N

Fumesua

6o 43’N

1o 36’W

228

626.86

3o 20’N

0 o 17’E

69

519.11

Deciduous
forest
Coastal
savannah

Kpeve

Soil Type
Forest/savannah
ochrosols
Ferric acrisols
Savannah achrosols

*: mean rainfall during April to July, 2011
2.3 Statistical analysis
The data were analyzed separately for each location, and then combined and analyzed across locations for grain
yield with PROC GLM in SAS using a RANDOM statement with the TEST option (SAS 2001) to determine if
GxE interaction effects were significant. In the combined analysis of variance, genotypes were considered as
fixed effects, while environments, replications, genotype by environment interaction and all other sources of
variation were considered as random effects. Means were separated using the LSD at P < 0.05. Subsequently, the
data on the grain yield were subjected to GGE biplot analysis to determine grain yield stability and the pattern of
response of genotypes evaluated across the three environments. The GGE biplots were constructed using the first
two principal components (PC1 and PC2) that were derived from subjecting environment centered trait means
for each environment to singular value decomposition. The data were not transformed (Transform = 0),
standardized (Scale = 1), and were environment-centered (Centering = 2). This provided information on the
cultivars that were suitable for the different environments and investigation of stability of genotypes in the
various environments. The analyses were done and biplots generated using the GGEbiplot software (Version
6.5). The GGE biplot Model 3 equation used is as follows:
Where:
Yj is the average yield across all genotypes in environment j; λ1 and λ2 are the singular values for PC1 and PC2,
respectively; ξi1 and ξi2 are the PC1 and PC2 scores, respectively, for genotype i; ηj1 and ηj2 are the PC1 and PC2
scores, respectively, for environment j; and εij is the residual of the model associated with the genotype i in
environment j.
3. Results and Discussion
3.1 Analysis of variance
The combined ANOVA showed differences among environments (E) and genotypes (G) to be significant
indicating that they were diverse. However, genotype by environment interactions for grain yield was not
significant. The proportions of the total variance in grain yield attributable to the environments were the highest
(87.27 %) while genotypes and GxE contributed 5.45 % and 0.84 %, respectively (Tables 3). This result is
similar to the findings of Badu-Apraku et al. (1995; 2003) and Mohammadi et al. (2009), who reported that the
largest proportion of total variation in multi-environment trials is attributed to locations, whereas G and G×E
sources of variation are relatively smaller. The lack of significant GxE mean square for grain yield indicated that
the expression of this trait would be consistent across the test environments. Mean grain yield of all the entries
evaluated at the three locations was 5.21 t/ha (Table 3). The grain yields recorded for 54 of the genotypes were
above the average yield (Figure 1).

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Journal of Biology, Agriculture and Healthcare
ISSN 2224-3208 (Paper) ISSN 2225-093X (Online)
Vol.3, No.12, 2013

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Table 3. Mean square values from the combined analyses of variance of grain yield (t/ha) of 100 extra-early
maturing maize genotypes evaluated across three environments in Ghana

Source of variation
Environment
Reps(Environment)
Blocks (Environment*Reps)
Genotype
Environment*Genotype
Error
Mean
CV%
*

DF
2
3
54
99
198
243

Mean square
87.27**
12.47**
1.96**
5.45**
0.84ns
0.75
5.2
16.65

= P ≤ 0.05, ** = P ≤ 0.01, ns = not significant, respectively

Figure 1. The frequency distribution of grain yield (t/ha) of the 100 extra-early maturing maize genotypes
evaluated across three environments in Ghana
3.2 GGE biplot analysis of grain yield response and stability of the 100 extra-early maturing maize genotypes
The biplots in Figure 2 and 4 were based on genotype-focused singular value partitioning (SVP = 2) and is
therefore appropriate for visualizing the relationships among environments. Also, the biplot in Figure 3 was
based on environment focused-singular value partitioning (SVP = 1) and is therefore appropriate for visualizing
the relationships among genotypes. The principal component (PC) axis 1 explained 74.4 % of total variation;
while PC2 explained 17 %. Thus, these two axes accounted for 91.4 % of the G+G×E variation for grain yield
(Figure 2, 3 and 4). The entry names of entries used in this section are shown in Table 1. The results are
presented as three sections. Section one presents the results of “which won-where” to identify the best genotypes
for each environment. Section two; the results of hybrids’ performance and their stability; section three gives the
discriminating power and representativeness of the test environments.
i.
The “which-won-where” patterns
The GGE biplot is an invaluable statistical tool for examining the performance of genotypes tested in different
environments. The polygon view of the GGE biplot in Figure 2 indicated the best genotype in each environment.
The presence of two or more environments within a sector indicates that a single genotype has the highest yield
in those environments. If environments fall into different sectors, it means that different genotypes won in
different environments (Yan et al. 2005; 2010). Based on the above information, entry 25 was the vertex
genotype where Ejura and Fumesua fell while entry 78 was the highest yielding genotype at Kpeve. No

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Journal of Biology, Agriculture and Healthcare
ISSN 2224-3208 (Paper) ISSN 2225-093X (Online)
Vol.3, No.12, 2013

www.iiste.org

environment fell into the sector where entries 100, 44, 86, 79 and 70 were the vertex genotypes, indicating that
these genotypes were the lowest-yielding genotypes at all or some locations. Genotypes within the polygon,
particularly those located near entries 87, 77, 33, 90, 95, 80, 70, 90, 96 and 48 were less responsive than the
vertex genotypes.

Figure 2. ‘which-won-where’ or ‘which is best for what’ based on a genotype x environment yield data of the
100 extra-early maturing maize genotypes evaluated in three environments in Ghana during the 2011 growing
season
ii.
Performance of genotypes and their stability across environments
In the entry/tester view of the GGE biplot of grain yield of the 100 extra-early maturing maize genotypes
evaluated in three environments in Ghana (Figure 3). The genotypes were ranked along the average-tester axis
(ATC abscissa), with an arrow pointing to a greater value based on their mean performance across all
environments. The double-arrowed line separates entries with below-average means from those with aboveaverage means. The average yield of the genotypes is approximated by the projections of their markers on the
average-tester axis. In the GGE biplot analysis, the average-tester coordinate (ATC) approximates the
genotypes’ contributions to G×E, which is a measure of their instability. The stability of the genotypes is
measured by their projections onto the double-arrow line (average-tester coordinate [ATC] y axis). The greater
the absolute length of the projection of a genotype, the less stable it is (Yan et al. 2000; 2010). Based on this,
entries 32, 24, 72, 89 and 13 were the most stable with an above average performance. Since they were located
away from the ATC abscissa and had a near zero projection onto the ATC coordinate. This implies that their
rankings were highly consistent across locations. For selection for broad adaptation in maize production, an ideal
genotype should have both high mean performance and high stability (Badu-Apraku et al. 2011). Therefore,
entries 32, 24, 72, 89 and 13 were closest to the ideal genotype and may be considered as best genotypes. These
five hybrids are suitable for production in Ejura, Fumesua and Kpeve. On the other hand, ideal genotypes for
specific environments are those that have high mean yield and respond best to the particular environments. Thus,
entries 25 and 78 are especially suitable for production at Ejura and Fumesua, and Kpeve, respectively. Entries
70, 78, 23, 83, 57, 49 and 39 were the least stable highest yielding hybrids. On the contrary, entries 86, 17, 66,
28, 19 and 91 were lowest yielding but very stable hybrids. However, 100, 79, 63, 40, 69, 47, 27, 16 and 44 were
not only low yielding but also among the least stable genotypes. Thus, they may not be good candidates for
commercial production across these environments.

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ISSN 2224-3208 (Paper) ISSN 2225-093X (Online)
Vol.3, No.12, 2013

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Figure 3. The ‘mean vs. stability’ view of the GGE biplot based on a genotype x environment yield data of the
100 extra-early maturing maize genotypes evaluated in three environments in Ghana during the 2011 growing
season
iii.
Discriminating power and representativeness of the test environments
The three test environments used in this study were Ejura, Fumesua and Kpeve representing the transition,
deciduous forest and coastal savannah zones of Ghana. The purpose of test-environment evaluation is to identify
environments that effectively identify superior genotypes in a group of environments. The representativeness and
discriminating power view of GGE biplot analysis are presented in Figure 4. Ejura had the longest vectors
followed by Kpeve while Fumesua had the shortest vector. Fumesua was at the smallest angle to the averageenvironment axis (AEA) followed by Kpeve while Ejura was at the largest angles to it (Figure 4). Since the AEC
abscissa is the average--environment axis, test environments at smaller angles to the AEA are more
representative of the group of environments than those at larger angles to it. Therefore, the cosine of the angle
between any environment vector and the average-environment axis approximates the correlation coefficient
between the genotype values in that environment and the genotype means across the environments (Yan et al.
2007). The small circle is the average-environment and the arrow pointing to it is used to indicate the direction
of the AEA (Yan & Tinker 2005). The absolute length of the projection from the marker of an environment onto
the AEA is a measure of its representativeness: the shorter the projection, the more representative the
environment. In contrast, the absolute length of the projection from the marker of an environment onto the AEA
is a measure of its discriminative ability: the longer the projection, the more discriminative the environment.
Based on these requirements, Ejura was highly discriminating but least representative of the test environments.
Kpeve was most representative and discriminating of the test environments. On the other hand, Fumesua was the
least discriminating but most representatives of the test environments. An ideal test environment should
effectively discriminate genotypes and represent the environments (Yan & Rajcan 2002). This indicated that
amongst the three locations, Kpeve located in the coastal savannah zone, represented the ideal testing
environment for these set of genotypes. This location would therefore be the most appropriate for selecting
superior genotypes. Similar result was reported by Abdulai et al. (2007).

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Figure 4. The ‘discriminating power and representativeness’ view of GGE biplot based on a genotype x
environment yield data of the 100 extra-early maturing maize genotypes evaluated in three environments in
Ghana during the 2011 growing season
4. Conclusion
The non significant GxE interaction effects for grain yield suggests that a promising genotype selected in one of
these locations will also be suitable for production in the other locations in the same agro-ecological zone.
Environments were found to contribute greatly to the variations in performance of genotypes. This indicates that,
unpredictable environmental conditions are one of the major constraints to selecting superior and widely adapted
maize varieties. The use of GGE biplot analyses provided clear bases for determining stability and performance
of the 100 extra-early maize genotypes. Based on the analyses, entries 32, 24, 72, 89 and 13 were the highest
yielding and most stable hybrids. They were the closest to the ideal genotype and may be considered as the best
hybrids. These five hybrids have the potential for production in Ejura, Fumesua and Kpeve and other locations
within the same agro-ecological zones. Entries 86, 17, 66, 28, 19 and 91 were lowest yielding but stable. Thus,
the performance of these genotypes would be predictable in less favourable environments. Entry 25 was
identified as the most promising for production in Ejura and Fumesua, and entry 78 in Kpeve. Kpeve located in
the coastal savannah zone, was identified as the ideal testing environment for this set of genotypes.
Acknowledgements
G. B. Adu is grateful to the Alliance for a Green Revolution in Africa (AGRA), for sponsoring her M.Sc.
training at the Kwame Nkrumah University of Science and Technology, Kumasi, Ghana. Also, we thank Dr.
Baffour Badu-Apraku for his comments on an earlier draft of this paper.
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15

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Assessment of genotype by environment interactions and grain

  • 1. Journal of Biology, Agriculture and Healthcare ISSN 2224-3208 (Paper) ISSN 2225-093X (Online) Vol.3, No.12, 2013 www.iiste.org Assessment of Genotype by Environment interactions and Grain Yield Performance of Extra-Early Maize (Zea Mays L.) Hybrids G.B. Adu1*, R. Akromah2, M.S. Abdulai1, K. Obeng-Antwi3, A.W. Kena2, K.M.L. Tengan3 and H. Alidu1 1. CSIR-Savanna Agricultural Research Institute, Nyankpala, Tamale, Ghana 2. Department of crops and soil sciences, Faculty of agriculture, Kwame Nkrumah University of Science and Technology, Kumasi, Ghana 3. CSIR-Crops Research Institute, Kumasi, Ghana * E-mail of the corresponding author: gloriaboakyewaa@yahoo.com Abstract Maize (Zea mays L.) is one of the most important cereal crops of Ghana in terms of production and consumption. Currently, it is produced in all the agro-ecological zones of the country. In Ghana, Genotype by Environment interactions (GxE) effects on maize grain yield is usually significant due to the diverse environmental conditions at growing sites. A proper understanding of the effects of GxE on variety evaluation and cultivar recommendations is vital. The study was conducted in 2011 at three locations in Ghana to (i) determine the presence of GxE of 100 extra-early maize genotypes and (ii) To use the GGE biplot methodology to determine grain yield performance and stability of the genotypes evaluated across three environments. The effects of genotype and environment were significant (P < 0.01) for grain yield. However, GxE was not significant for the same trait. TZEEI 8 x TZEEI 51, TZEEI 5 x TZEEI 53, TZEEI 21 x TZEEI 39, TZEEI 27 x TZEEI 36 and TZEEI 4 x TZEEI 6 were identified as high yielding and most stable hybrids. Therefore, these hybrids have the potential for production across the test locations as well as others within the same agro-ecological zones. On the contrary, DODZI, TZEEI 23 x TZEEI 6, TZEEI 19 x TZEEI 24, TZEEI 11 x TZEEI 24 and TZEEI 20 x TZEEI 39 were not only low yielding but also among the least stable genotypes. Keywords: Zea mays, single-cross, stability, multiple environments, GxE 1. Introduction Maize (Zea mays L.) is produced in all the agro-ecological zones of Ghana and its highest production is in the transition zone. Maize yields in Ghana are well below their attainable levels; averaging approximately 0.5 metric tons per hectare. However, yields as high as 5.5 metric tons per hectare have been realized by farmers using improved seeds, fertilizer, mechanization and irrigation (MiDA 2010). Lower yields have been attributed to the use of low-yielding varieties, use of self produced seed, poor soil fertility and limited use of fertilizers, low plant population, and inappropriate weed control methods. Moreover, significant potential improvements in yields could be achieved through the use of hybrid maize varieties. Crop breeders have been striving to develop genotypes with superior grain yield, quality and other desirable characteristics over a wide range of different environmental conditions. Genotype by environment interaction (GxE) makes it difficult to select the best performing and most stable genotypes. GxE refers to the differential ranking of genotype among locations or years. It is an important consideration in plant breeding programmes because it impedes progress from selection in any given environment (Yau 1995). In West and Central Africa, GxE interaction effects have been reported in maize cultivars (Fakorede & Adeyemo 1986; Badu-Apraku et al. 2007; 2008). Various studies have been conducted to study the effects of genotype by environment interaction in Sub-Saharan Africa and on Ghanaian maize varieties. However, the changing environmental conditions, the expansion of maize into new agro-ecologies, coupled with inadequate maize varieties available for the different environments necessitate a rigorous and continuous study of GxE interaction effects for a dynamic crop improvement programme. There are many statistical methods available to analyse GxE: for example, combined ANOVA, stability analysis and multivariate methods. Combined analysis of variance (ANOVA) is more often used to identify the existence of GxE interactions in multi-environmental experiments. However, the main limitation of this analysis is the assumption of homogeneity of variance among environments required to determine genotype differences. Although this analysis allows the determination of the components of variance arising from different factors (genotype, environment and the GxE), it does not allow exploring the response of the genotypes in the non- 7
  • 2. Journal of Biology, Agriculture and Healthcare ISSN 2224-3208 (Paper) ISSN 2225-093X (Online) Vol.3, No.12, 2013 www.iiste.org additive term: the GxE (Zobel et al. 1988). Among the statistical analyses proposed for the interpretation of the GxE based on the use of biplots, the AMMI (additive main effect and multiplicative interaction) model stands out due to the largest group of technical interpretations available (Duarte & Vencovsky 1999). AMMI analysis interprets the effect of the genotype (G) and sites (E) as additive effects plus the GxE as a multiplicative component and submits it to principal component analysis. Yan et al. (2000) proposed a modification of the conventional AMMI analysis called GGE that has been used for GxE analysis. The GGE analysis pools genotype effect (G) with GE (multiplicative effect) and submits these effects to principal component analysis. This biplot is identified as a GGE biplot. The GGE biplot has been recognized as an innovative methodology in biplot graphic analysis to be applied in plant breeding. Fan et al. (2007) showed that the GGE biplot methodology was a useful tool for identifying locations that optimized hybrid genotypes performance and for making better use of limited resources available for the maize testing programmes. The objectives of this study were (i) to determine the presence of GxE of 100 extra-early maize genotypes and (ii) to use the Genotype main effect plus Genotype by Environment interaction (GGE) biplot methodology to determine grain yield performance and stability of the genotypes evaluated across three environments. 2. Materials and Methods 2.1 Field evaluations Ninety-eight single-cross maize hybrids, one local three-way hybrid (Akposoe) and an open pollinated variety (Dodzi) (Table 1) were evaluated at Fumesua, Ejura and Kpeve from April, 2011 to July, 2011 major season. The evaluation sites are located in the deciduous forest, transition and coastal Savannah zones of Ghana, respectively (Table 2). The hybrids were developed from 61 extra-early maturing white and yellow endosperm inbred lines developed for grain yield, drought and Striga hermonthica tolerance. A standard protocol was adopted at each site during the period of evaluation. Evaluations were done under rain fed conditions and fields were planted when the rains at each experimental site had become fully established. The genotypes were planted in an incomplete block design (10 x 10) with two replications. Each plot consisted 2-rows of 5 m long, an interrow spacing of 75 cm and an intra-row spacing of 40 cm. Three seeds were sown per hill and seedlings later thinned to two after emergence and seedling establishment. Pre-emergence chemical weed control was practised and comprised an application of a combination of Pendimethalin [N-(1-ethylpropyl)-3, 4-dimethyl-2, 6dinitrobenzenamine] and Gesaprim [2-chloro-4-(ethylamino)-6-(isopropylamino)-s-triazine] at 1.5 Lha-1 and 1.0 Lha-1 active ingredient, respectively at planting. Hand weeding was also done when necessary to control weeds during the growing period. NPK 15-15-15 fertilizer was applied at the rate of 30 kg N ha-1 and 60 kg P2O5 ha-1 as basal fertilizer at 1-2 weeks after planting and top-dressed with additional N at 60 kg N ha-1 at four weeks after planting. Other management practices were done according to the recommendations of the specific areas. 2.2 Collection of agronomic data Data recorded include days to anthesis, days to silking, plant and ear heights, root lodging (number of plants leaning more than 300 from vertical), stalk lodging (stalks broken at or below highest ear node), ear aspect, number of plants harvested, number of ears harvested and percent moisture in the grains. Days to anthesis and days to silking were calculated as the number of days from planting to when 50 % of the plants had shed pollen and had emerged silks, respectively. Anthesis-silking interval was determined as the difference between days to silking and days to anthesis. Plant and ear heights were measured as the distance from the base of the plant to the height of the flag leaf and the node bearing the upper ear, respectively. Husk cover was rated on a scale of 1 to 5, where 1= husk tightly arranged and extended beyond the ear tip and 5 = ear tips exposed. Ear aspect was based on a scale of 1 to 5, where 1= clean, uniform, large, and well-filled ears and 5 = ears with undesirable features. Number of ears per plant was obtained by dividing the total number of ears per plot by the number of plants harvested. The grain yield in kilograms per plot recorded was converted to grain yield in tons per hectare at 15 % grain moisture based on 80% shelling percentage. Even though data were collected on several traits, only those on the most important trait in the study are presented in the results. 8
  • 3. Journal of Biology, Agriculture and Healthcare ISSN 2224-3208 (Paper) ISSN 2225-093X (Online) Vol.3, No.12, 2013 www.iiste.org Table 1. Description of extra-early maturing maize genotypes tested across three environments in 2011 Entry Entry name Entry Entry name Entry Entry name 1 TZEEI 1 x TZEEI 2 34 TZEEI 9 x TZEEI 58 67 TZEEI 20 x TZEEI 22 2 TZEEI 1 x TZEEI 4 35 TZEEI 9 x TZEEI 59 68 TZEEI 20 x TZEEI 24 3 TZEEI 1 x TZEEI 5 36 TZEEI 9 x TZEEI 60 69 TZEEI 20 x TZEEI 39 4 TZEEI 1 x TZEEI 13 37 TZEEI 10 x TZEEI 22 70 TZEEI 21xTZEEI 20 5 TZEEI 1 x TZEEI 14 38 TZEEI 11 x TZEEI 8 71 TZEEI 21 x TZEEI 24 6 TZEEI 1 x TZEEI 22 39 TZEEI 11 x TZEEI 22 72 TZEEI 21 x TZEEI 39 7 TZEEI 1 x TZEEI 23 40 TZEEI 11 x TZEEI 24 73 TZEEI 22 x TZEEI 6 8 TZEEI 1 x TZEEI 50 41 TZEEI 12 x TZEEI 19 74 TZEEI 22 x TZEEI 24 9 TZEEI 2 x TZEEI 1 42 TZEEI 13 x TZEEI 1 75 TZEEI 22 x TZEEI 36 10 TZEEI 2 x TZEEI 11 43 TZEEI 13 x TZEEI 6 76 TZEEI 22 x TZEEI 51 11 TZEEI 2 x TZEEI 14 44 TZEEI 13 x TZEEI 12 77 TZEEI 23 x TZEEI 4 12 TZEEI 2 x TZEEI 24 45 TZEEI 13 x TZEEI 22 78 TZEEI 23 x TZEEI 5 13 TZEEI 4 x TZEEI 6 46 TZEEI 14 x TZEEI 4 79 TZEEI 23 x TZEEI 6 14 TZEEI 4 x TZEEI 7 47 TZEEI 14 x TZEEI 6 80 TZEEI 23 x TZEEI 36 15 TZEEI 4 x TZEEI 24 48 TZEEI 14 x TZEEI 22 81 TZEEI 23 x TZEEI 39 16 TZEEI 4 x TZEEI 39 49 TZEEI 14 x TZEEI 39 82 TZEEI 23 x TZEEI 40 17 TZEEI 4 x TZEEI 51 50 TZEEI 14 x TZEEI 49 83 TZEEI 23 x TZEEI 50 18 TZEEI 4 x TZEEI 90 51 TZEEI 14 x TZEEI 90 84 TZEEI 23 x TZEEI 90 19 TZEEI 5 x TZEEI 4 52 TZEEI 15 x TZEEI 6 85 TZEEI 24 x TZEEI 51 20 TZEEI 5 x TZEEI 23 53 TZEEI 1 x TZEEI 8 86 TZEEI 26 x TZEEI 24 21 TZEEI 5 x TZEEI 39 54 TZEEI 15 x TZEEI 19 87 TZEEI26 x TZEEI 51 22 TZEEI 5 x TZEEI 40 55 TZEEI 15 x TZEEI 21 88 TZEEI 26 x TZEEI 53 23 TZEEI 5xTZEEI 50 56 TZEEI 15 x TZEEI 22 89 TZEEI 27 x TZEEI 36 24 TZEEI 5 x TZEEI 53 57 TZEEI 15 x TZEEI 24 90 TZEEI 27 x TZEEI 51 25 TZEEI 6 x TZEEI 4 58 TZEEI 15 x TZEEI 39 91 TZEEI 29 x TZEEI 26 26 TZEEI 6 x TZEEI 5 59 TZEEI 19 x TZEEI 6 92 TZEEI 29 x TZEEI 33 27 TZEEI 6 x TZEEI 23 60 TZEEI 19 x TZEEI 8 93 TZEEI 29 x TZEEI 36 28 TZEEI 6 x TZEEI 36 61 TZEEI 19 x TZEEI 21 94 TZEEI 30 x TZEEI 4 29 TZEEI 6 x TZEEI 39 62 TZEEI 19 x TZEEI 22 95 TZEEI 30 x TZEEI 23 30 TZEEI 6 x TZEEI 40 63 TZEEI 19 x TZEEI 24 96 TZEEI 30 x TZEEI 36 31 TZEEI 6 x TZEEI 90 64 TZEEI 20 x TZEEI 8 97 TZEEI 30 x TZEEI 39 32 TZEEI 8 x TZEEI 24 65 TZEEI 20 x TZEEI 19 98 TZEEI 31 x TZEEI 8 33 TZEEI 8 x TZEEI 51 66 TZEEI 20 x TZEEI 21 AKPOSOE (Three-way) 100 DODZI (OPV) Check 99 9
  • 4. Journal of Biology, Agriculture and Healthcare ISSN 2224-3208 (Paper) ISSN 2225-093X (Online) Vol.3, No.12, 2013 www.iiste.org Table 2. Description of the test environments used in the study Location Latitude Longitude 1 o 37’E Altitude (m ASL) 228.7 Mean Seasonal Rainfall *(mm) 599.7 Agroecological zone Transition Ejura 7o 38’N Fumesua 6o 43’N 1o 36’W 228 626.86 3o 20’N 0 o 17’E 69 519.11 Deciduous forest Coastal savannah Kpeve Soil Type Forest/savannah ochrosols Ferric acrisols Savannah achrosols *: mean rainfall during April to July, 2011 2.3 Statistical analysis The data were analyzed separately for each location, and then combined and analyzed across locations for grain yield with PROC GLM in SAS using a RANDOM statement with the TEST option (SAS 2001) to determine if GxE interaction effects were significant. In the combined analysis of variance, genotypes were considered as fixed effects, while environments, replications, genotype by environment interaction and all other sources of variation were considered as random effects. Means were separated using the LSD at P < 0.05. Subsequently, the data on the grain yield were subjected to GGE biplot analysis to determine grain yield stability and the pattern of response of genotypes evaluated across the three environments. The GGE biplots were constructed using the first two principal components (PC1 and PC2) that were derived from subjecting environment centered trait means for each environment to singular value decomposition. The data were not transformed (Transform = 0), standardized (Scale = 1), and were environment-centered (Centering = 2). This provided information on the cultivars that were suitable for the different environments and investigation of stability of genotypes in the various environments. The analyses were done and biplots generated using the GGEbiplot software (Version 6.5). The GGE biplot Model 3 equation used is as follows: Where: Yj is the average yield across all genotypes in environment j; λ1 and λ2 are the singular values for PC1 and PC2, respectively; ξi1 and ξi2 are the PC1 and PC2 scores, respectively, for genotype i; ηj1 and ηj2 are the PC1 and PC2 scores, respectively, for environment j; and εij is the residual of the model associated with the genotype i in environment j. 3. Results and Discussion 3.1 Analysis of variance The combined ANOVA showed differences among environments (E) and genotypes (G) to be significant indicating that they were diverse. However, genotype by environment interactions for grain yield was not significant. The proportions of the total variance in grain yield attributable to the environments were the highest (87.27 %) while genotypes and GxE contributed 5.45 % and 0.84 %, respectively (Tables 3). This result is similar to the findings of Badu-Apraku et al. (1995; 2003) and Mohammadi et al. (2009), who reported that the largest proportion of total variation in multi-environment trials is attributed to locations, whereas G and G×E sources of variation are relatively smaller. The lack of significant GxE mean square for grain yield indicated that the expression of this trait would be consistent across the test environments. Mean grain yield of all the entries evaluated at the three locations was 5.21 t/ha (Table 3). The grain yields recorded for 54 of the genotypes were above the average yield (Figure 1). 10
  • 5. Journal of Biology, Agriculture and Healthcare ISSN 2224-3208 (Paper) ISSN 2225-093X (Online) Vol.3, No.12, 2013 www.iiste.org Table 3. Mean square values from the combined analyses of variance of grain yield (t/ha) of 100 extra-early maturing maize genotypes evaluated across three environments in Ghana Source of variation Environment Reps(Environment) Blocks (Environment*Reps) Genotype Environment*Genotype Error Mean CV% * DF 2 3 54 99 198 243 Mean square 87.27** 12.47** 1.96** 5.45** 0.84ns 0.75 5.2 16.65 = P ≤ 0.05, ** = P ≤ 0.01, ns = not significant, respectively Figure 1. The frequency distribution of grain yield (t/ha) of the 100 extra-early maturing maize genotypes evaluated across three environments in Ghana 3.2 GGE biplot analysis of grain yield response and stability of the 100 extra-early maturing maize genotypes The biplots in Figure 2 and 4 were based on genotype-focused singular value partitioning (SVP = 2) and is therefore appropriate for visualizing the relationships among environments. Also, the biplot in Figure 3 was based on environment focused-singular value partitioning (SVP = 1) and is therefore appropriate for visualizing the relationships among genotypes. The principal component (PC) axis 1 explained 74.4 % of total variation; while PC2 explained 17 %. Thus, these two axes accounted for 91.4 % of the G+G×E variation for grain yield (Figure 2, 3 and 4). The entry names of entries used in this section are shown in Table 1. The results are presented as three sections. Section one presents the results of “which won-where” to identify the best genotypes for each environment. Section two; the results of hybrids’ performance and their stability; section three gives the discriminating power and representativeness of the test environments. i. The “which-won-where” patterns The GGE biplot is an invaluable statistical tool for examining the performance of genotypes tested in different environments. The polygon view of the GGE biplot in Figure 2 indicated the best genotype in each environment. The presence of two or more environments within a sector indicates that a single genotype has the highest yield in those environments. If environments fall into different sectors, it means that different genotypes won in different environments (Yan et al. 2005; 2010). Based on the above information, entry 25 was the vertex genotype where Ejura and Fumesua fell while entry 78 was the highest yielding genotype at Kpeve. No 11
  • 6. Journal of Biology, Agriculture and Healthcare ISSN 2224-3208 (Paper) ISSN 2225-093X (Online) Vol.3, No.12, 2013 www.iiste.org environment fell into the sector where entries 100, 44, 86, 79 and 70 were the vertex genotypes, indicating that these genotypes were the lowest-yielding genotypes at all or some locations. Genotypes within the polygon, particularly those located near entries 87, 77, 33, 90, 95, 80, 70, 90, 96 and 48 were less responsive than the vertex genotypes. Figure 2. ‘which-won-where’ or ‘which is best for what’ based on a genotype x environment yield data of the 100 extra-early maturing maize genotypes evaluated in three environments in Ghana during the 2011 growing season ii. Performance of genotypes and their stability across environments In the entry/tester view of the GGE biplot of grain yield of the 100 extra-early maturing maize genotypes evaluated in three environments in Ghana (Figure 3). The genotypes were ranked along the average-tester axis (ATC abscissa), with an arrow pointing to a greater value based on their mean performance across all environments. The double-arrowed line separates entries with below-average means from those with aboveaverage means. The average yield of the genotypes is approximated by the projections of their markers on the average-tester axis. In the GGE biplot analysis, the average-tester coordinate (ATC) approximates the genotypes’ contributions to G×E, which is a measure of their instability. The stability of the genotypes is measured by their projections onto the double-arrow line (average-tester coordinate [ATC] y axis). The greater the absolute length of the projection of a genotype, the less stable it is (Yan et al. 2000; 2010). Based on this, entries 32, 24, 72, 89 and 13 were the most stable with an above average performance. Since they were located away from the ATC abscissa and had a near zero projection onto the ATC coordinate. This implies that their rankings were highly consistent across locations. For selection for broad adaptation in maize production, an ideal genotype should have both high mean performance and high stability (Badu-Apraku et al. 2011). Therefore, entries 32, 24, 72, 89 and 13 were closest to the ideal genotype and may be considered as best genotypes. These five hybrids are suitable for production in Ejura, Fumesua and Kpeve. On the other hand, ideal genotypes for specific environments are those that have high mean yield and respond best to the particular environments. Thus, entries 25 and 78 are especially suitable for production at Ejura and Fumesua, and Kpeve, respectively. Entries 70, 78, 23, 83, 57, 49 and 39 were the least stable highest yielding hybrids. On the contrary, entries 86, 17, 66, 28, 19 and 91 were lowest yielding but very stable hybrids. However, 100, 79, 63, 40, 69, 47, 27, 16 and 44 were not only low yielding but also among the least stable genotypes. Thus, they may not be good candidates for commercial production across these environments. 12
  • 7. Journal of Biology, Agriculture and Healthcare ISSN 2224-3208 (Paper) ISSN 2225-093X (Online) Vol.3, No.12, 2013 www.iiste.org Figure 3. The ‘mean vs. stability’ view of the GGE biplot based on a genotype x environment yield data of the 100 extra-early maturing maize genotypes evaluated in three environments in Ghana during the 2011 growing season iii. Discriminating power and representativeness of the test environments The three test environments used in this study were Ejura, Fumesua and Kpeve representing the transition, deciduous forest and coastal savannah zones of Ghana. The purpose of test-environment evaluation is to identify environments that effectively identify superior genotypes in a group of environments. The representativeness and discriminating power view of GGE biplot analysis are presented in Figure 4. Ejura had the longest vectors followed by Kpeve while Fumesua had the shortest vector. Fumesua was at the smallest angle to the averageenvironment axis (AEA) followed by Kpeve while Ejura was at the largest angles to it (Figure 4). Since the AEC abscissa is the average--environment axis, test environments at smaller angles to the AEA are more representative of the group of environments than those at larger angles to it. Therefore, the cosine of the angle between any environment vector and the average-environment axis approximates the correlation coefficient between the genotype values in that environment and the genotype means across the environments (Yan et al. 2007). The small circle is the average-environment and the arrow pointing to it is used to indicate the direction of the AEA (Yan & Tinker 2005). The absolute length of the projection from the marker of an environment onto the AEA is a measure of its representativeness: the shorter the projection, the more representative the environment. In contrast, the absolute length of the projection from the marker of an environment onto the AEA is a measure of its discriminative ability: the longer the projection, the more discriminative the environment. Based on these requirements, Ejura was highly discriminating but least representative of the test environments. Kpeve was most representative and discriminating of the test environments. On the other hand, Fumesua was the least discriminating but most representatives of the test environments. An ideal test environment should effectively discriminate genotypes and represent the environments (Yan & Rajcan 2002). This indicated that amongst the three locations, Kpeve located in the coastal savannah zone, represented the ideal testing environment for these set of genotypes. This location would therefore be the most appropriate for selecting superior genotypes. Similar result was reported by Abdulai et al. (2007). 13
  • 8. Journal of Biology, Agriculture and Healthcare ISSN 2224-3208 (Paper) ISSN 2225-093X (Online) Vol.3, No.12, 2013 www.iiste.org Figure 4. The ‘discriminating power and representativeness’ view of GGE biplot based on a genotype x environment yield data of the 100 extra-early maturing maize genotypes evaluated in three environments in Ghana during the 2011 growing season 4. Conclusion The non significant GxE interaction effects for grain yield suggests that a promising genotype selected in one of these locations will also be suitable for production in the other locations in the same agro-ecological zone. Environments were found to contribute greatly to the variations in performance of genotypes. This indicates that, unpredictable environmental conditions are one of the major constraints to selecting superior and widely adapted maize varieties. The use of GGE biplot analyses provided clear bases for determining stability and performance of the 100 extra-early maize genotypes. Based on the analyses, entries 32, 24, 72, 89 and 13 were the highest yielding and most stable hybrids. They were the closest to the ideal genotype and may be considered as the best hybrids. These five hybrids have the potential for production in Ejura, Fumesua and Kpeve and other locations within the same agro-ecological zones. Entries 86, 17, 66, 28, 19 and 91 were lowest yielding but stable. Thus, the performance of these genotypes would be predictable in less favourable environments. Entry 25 was identified as the most promising for production in Ejura and Fumesua, and entry 78 in Kpeve. Kpeve located in the coastal savannah zone, was identified as the ideal testing environment for this set of genotypes. Acknowledgements G. B. Adu is grateful to the Alliance for a Green Revolution in Africa (AGRA), for sponsoring her M.Sc. training at the Kwame Nkrumah University of Science and Technology, Kumasi, Ghana. Also, we thank Dr. Baffour Badu-Apraku for his comments on an earlier draft of this paper. References Abdulai, M.S., Sallah, P.Y.K. & Safo-Kantanka, O. (2007). ‘Maize Grain Yield Stability Analysis in Full Season Lowland Maize in Ghana’. International Journal of Agriculture & Biology 9(1), 41–45. Badu-Apraku, B., Abamu, F.J., Menkir, A., Fakorede, M.A.B., Obeng-Antwi, K. & The’, C. (2003). ‘Genotype by Environment Interactions in the Regional Early Variety Trials in West and Central Africa’. Maydica 48(2), 93–104. Badu-Apraku, B., Fajemisin, J.M. & Diallo, A.O. (1995). The Performance of Early and Extra-Early Varieties across Environments in West and Central Africa. In: Badu-Apraku, B., Akoroda, M.O., Ouedraogo, M., Quin, F.M. (eds) Contributing to food self-sufficiency: Maize research and development in West and Central Africa, Proceedings of a regional maize workshop ’95, Cotonou, Benin republic, 149–159. Badu-Apraku, B., Fakorede, M.A.B. & Lum, A.F. (2007). ‘Evaluation of experimental varieties from recurrent selection for Striga resistance in two extra-early maize populations in the savannas of West and Central Africa’. Experimental Agriculture 43(2):183-200. 14
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