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Research Article
Newly developed varieties can only contribute to increased productivity if high producing
varieties are released in production niches they are adapted to. In order to enhance adoption of
new improved cassava varieties in western Kenya, a study was conducted to evaluate the effects
of genotype by environment interaction (GEI) on agronomic and farmer preferred traits of cassava
and to asses yield stability of 16 cassava genotypes. The study was conducted in randomized
complete block design with three replications across five different environments of western
Kenya. AMMI analysis of variance identified highly significant (P= 0.001) GEI effects for plant
height, height at first branching, and fresh root yield. Generally, GEI effects accounted for 14.98%,
24.64% and 28.3% variability in PH, HB, and FRY respectively. GGE biplot analysis shows that
MM06/0138, MM96/9308, MM97/0293, MM98/3567, MM06/0074, MM96/4271 were high yielding and
stable genotypes. AMMI stability value revealed that genotype MM06/0143 combined high stability
for plant height, height at first branching, number of storage roots and fresh root yield. Genotypes
MM06/0138, MM98/3567, MM96/9308, MM97/0293, and MM06/0074 outperformed the check in
storage roots yield exhibited high yields in farmer preferred traits and were classified as stable
genotypes. Therefore, recommended for release to farmers.
Key Words: Elite cassava, Farmer preferred traits, Genotype X Environment interaction, AMMI analysis, GGE-
biplot analysis
INTRODUCTION
Cassava (Manihot esculenta Crantz) is an important staple (Mwango’mbe, et al., 2013). Cassava mosaic disease
grown for its starchy tuberous roots. Its roots and leaves (CMD) and cassava brown streak disease (CBSD) are the
are suitable for human consumption as well as animal leading yield limiting biotic constraints for cassava
feed. The tuberous roots are an important source of production causing an estimated loss of more than US$ 14
carbohydrates while the leaves are cheap valuable source million per annum in CMD (Alabi, et al., 2015). Cassava
of proteins, minerals and vitamins A, B and C (Montagnac genetic improvement has been difficult due to the biology
et al., 2009). The storage roots are also used as industrial of the crop (Ceballos et al., 2004).
raw materials like starch extractions for various industrial
uses, breweries, pharmaceutical, and biofuel among other
uses (Nweke, 2004; Jackson et al., 2014). Cassava is the
second most important food crop after maize in Western *Corresponding Author: Rotich C. Damaris, Department
and coastal regions of Kenya (Njeru & Munga, 2003). of Biotechnology University of Eldoret P.O Box 1125-
However, production level in Kenya is 11 t/ha, below the 30100, Eldoret Kenya. Email: (rotich.damaris@gmail.com
potential of 90 t/ha, which is attributed to low yield of Tel: +254703276027. Co-Authors Email:
2
popular varieties, poor access to quality planting material, kiplagatoliver@yahoo.com, Tel: +254723967672,
3
lack of well adapted varieties, pests and diseases vwoyengo@gmail.com, Tel: +254729981023
Genotype by Environment Interaction on Cassava Components and Yield Stability Analysis of Elite Cassava Genotypes in Western Kenya
Rotich et al. 362
Use of improved varieties is the current leading tool for years frequently remains too large (Crossa 1990).
solving viral disease challanges in cassava (Alabi, et al., Moreover, Woyengo and Omari, (2014) clearly pointend
2015), hence continuous deployments of elite resistant out that it is not feasiable to breed for specific adaptation
cultivars are necessary as CMDs are known to evolve with current eratic climatic conditions and effect of
producing virulent strains while different strains of CBSD climimate change hence breeding for stable varieties
are being reported in Kenya (Mware, 2009). However, low remains as the only viable option. Many statistical
adoption of the new improved varieties has been reported procedures have been advocated for the basis of analysis
in western Kenya (Odendo et al., 2010; Woyengo & Omari, of GEI and stability of genotypes. Studies show that
2014). stability of performance are expected to become more
relevant issues as greater emphasis is placed on
In the last three decades, cassava breeding has majorly sustainability of agricultural systems (Kang et al., 2012).
concentrated on increasing yields and resistance to pest The objective of this study therefore, was to evaluate GEI
and disease (Ceballos et al., 2004). However, there has effects on agronomic and farmer preferred traits of
been lack of focus on farmer preferred traits by breeding cassava and to assess yield stability of 16 cassava
programmes which has been observed to be the major genotypes across five environments of western Kenya.
course of low adoption of improved varieties despite the
high yield and resistance to common pests and diseases
(Woyengo, 2011). Moreover, some of the improved MATERIALS AND METHODS
varieties fail to perform well in target production niches due
to lack of detailed stability studies of these traits. It’s The study was conducted across five environments
therefore a prerequisite that cassava varieties should not (Kakamega, Sang’alo, Alupe, Kibos and Migori), which
only be released on the basis of average yield and reaction represent major cassava growing zones of western Kenya,
to diseases pests but also on the presence of farmer between 2014 and 2015. The experimental material
preferred traits and stability. The work of Achepong et al. consisted of 15 elite cassava clones (G1=MH95/0183
(2013), identified longevity and disease resistance as two G3=MM06/0013, G4=MM06/0046, G5=MM06/0074,
major attributes of cassava that influence adoption of G6=MM06/0082, G7=MM06/0083, G8=MM06/0131
improved varieties in Ghana. Njukwe et al. (2013) G9=MM06/0138, G10=MM06/0139, G11=MM0H6/0143,
observed regional differences for farmer preference in G12=MM96/2480, G13=MM96/4271, G14=MM96/9308,
cassava attributes and cassava genotypes in Cameroon, G15= MM97/0293, G16=MM98/3567) in advanced stage
for instance farmers in Ebolowa and Bertoua preferred of yield trials performance and one local check
leafy, sweet roots and early branching varieties while (G2=migyera). Improved clones’ seeds were developed
those in Bamenda and Ngaoundere preferred tall, drought and introduced from International Institute of Tropical
tolerant and in some cases flowering varieties. In Kenya Agriculture (IITA). The clones were derived from half-sib
the work of Were (2011), identified farmer preferred traits progenies of elite varieties. They have been tested by
that encourage adoption of improved cassava genotypes Kenya agricultural livestock and research organization
by order of preference as high root yield, tall plants and (KALRO) Kakamega for resistance to CMD and CBSD.
lower height of first branching. However, no study has The experiment was laid in randomised complete block
been reported on influence of environment on farmer design with three replications and established under rain
preferred traits and stability of new improved cassava fed conditions. No fertilizer nor pesticide were applied.
genotypes. Each experimental plot had six rows and 30 plants spaced
of 1m by 1m between plants and rows.
Genotype stability and adaptability are ultimate resources
for achieving food security which is an allusive goal for Data Collection
Kenya and Sub-Saharan Africa at large (Muzari et al.,
2012). Lobell (2009) stated that agricultural adaptability Data was collected on agronomic traits (Dry matter content
should be a priority in meeting food security presently and and starch content) and traits preferred by farmers in
in future in the face of sever climate change. This is western Kenya as identified by Woyengo, (2011) which
achieved by development of stable varieties of crops. include plant height, height at first branching, number of
Stability of performance of quantitative traits is influenced storage roots per plant and fresh root yield. Data was
by genotype, environment and genotype by environment collected at twelve months after the date of planting and at
interaction (GEI) effects. GEI is important in plant breeding harvesting. Data were recorded on plant and plot basis.
because it complicates demonstration of a superiority of a Five plants per plot were sampled from the inner rows.
variety. An effective method which has been used to Data were recorded on plant height (PH), height at first
reduce GEI is stratification of environment such that the branching (HB), number of storage roots per plant
sub-region in which the breeder is developing improved (NSR)and fresh root yield converted to tonnes per hectare
varieties are somehow similar (specific adaptation). (FRY). Data on dry matter content (DMC) and starch
However, this is not mineable to breeding since even with content was collected according to the methodology
the refinement of this technique the interaction of described by Fukuda et al., (2010) converted to DM% and
genotypes within a location in a sub-region and with Starch % as follows:
environments encountered at the same location in different
Genotype by Environment Interaction on Cassava Components and Yield Stability Analysis of Elite Cassava Genotypes in Western Kenya
Int. J. Plant Breed. Crop Sci. 363
Root sample weighing 3-5kg was prepared, the weight of Another important point was further explained by Yan et al.
the sample was measured in air (Wa) using a digital (2007) that genotype and genotype-by-environment
weighing balance. The same sample was also measured effects must be considered simultaneously to make a
in water (Ww). Specific gravity(x) was computed at: meaningful decision in selection. Significant genotype by
environment interaction was also analyzed by GGE biplot
Ww/ (Wa-Ww) which was also useful in ranking genotypes based on their
average performance and stability for farmer preferred
%DMC was computed using the formula DMC = (158.3x- traits in cassava. The model for the GGE biplot based on
142)100 singular value decomposition (SVD) of first two principal
components is:
%starch was computed using the formula starch = (112.1x-
106.4)100 Yij j 1i1 j1 2i2 j2 ij
Genotype by Environment Interaction on Cassava Components and Yield Stability Analysis of Elite Cassava Genotypes in Western Kenya
Rotich et al. 364
Table 2: AMMI analysis of variance for16 cassava clones evaluated across five agroecological zones of Western Kenya
PH (cm) HB(cm) NSR FRY (t/ha) DMC (%) STARCH
Source Df MS %SS MS %SS MS %SS MS %SS MS %SS MS %SS
Total 239 2923 557 10.37 100.3 11.24 73.8
Treatments 79 8567*** 1584*** 28.72*** 288.9*** 20.84*** 162.5***
genotype 15 9119*** 20.20 4247*** 50.9 47.29*** 64.83 370.6*** 24.35 75.37*** 68.67 661.9*** 77.33
Environ 4 109651*** 64.81 7651*** 24.46 252.19*** 23.07 2701.7*** 47.35 19.76* 4.84 422.6*** 13.16
Block 10 474ns 43ns 1.05ns 7.1ns 7.08ns 19.7ns
GEI 60 1690*** 14.98 514*** 24.64 9.18ns 12.09 107.7*** 28.30 7.28ns 26.52 20.3ns 9.5
IPCA 1 18 3172*** 56.31 1132*** 66.13 15.18ns 49.59 178.7*** 49.79 15.25ns 30.00 37.5ns 55.46
IPCA 2 16 2104*** 33.2 524*** 27.20 9.49ns 27.55 104*** 25.72 5.22ns 26.67 18.4ns 24.2
IPCA 3 14 486*** 6.72 125** 5.69 6.64ns 16.86 84.8*** 18.38 4.68ns 23.33 11.7ns 13.45
IPCA 4 12 319** 3.77 25ns 0.98 2.74ns 6.00 32.7*** 6.11 1.10ns 20.00 7.1ns 6.97
Residuals 0 0 0
Error 150 114 50 1.33 7.1 6.46 30.6
*** Significant at (P≤0,001), ** (P≤ 0.01), *(P≤ 0.05) and ns = not significant respectively; PH = plant height, HB = plant height at first
branching, NSR = number of storage roots per plant, FRY =Fresh root yield, DMC = dry matter content, SS=% sum of squares,
IPCA=interaction principle component.
Table 3: Ranking of 16 cassava genotypes according to their AMMI stability value evaluated for PH, HB, SR and FRY
Genotype PH R HB R NSR R FRY R
MH95/0183 3.57 9 5.29 9 0.39 3 4.53 15
MIGHERA 9.32 14 7.59 13 3.1 16 4.01 14
MM06/0013 3.24 7 2.1 4 0.88 8 2.96 10
MM06/0046 1.07 1 1.5 2 1.04 9 2.84 9
MM06/0074 4.63 12 9.5 15 0.65 7 0.25 1
MM06/0082 9.83 15 3.35 6 1.6 11 3.75 13
MM06/0083 2.11 3 0.51 1 2.96 15 1.51 3
MM06/0131 13.4 16 6.86 12 0.5 5 5.5 16
MM06/0138 5.02 13 4.98 8 1.91 13 1.94 6
MM06/0139 2.6 5 10.5 16 1.65 12 3.61 12
MM06/0143 1.98 2 1.65 3 0.35 1 2.05 7
MM96/2480 4.1 11 3.73 7 1.36 10 2.49 8
MM96/4271 3.64 10 2.44 5 0.44 4 3.4 11
MM96/9308 3.3 8 6.8 11 0.62 6 1.55 4
MM97/0293 2.66 6 5.96 10 0.35 2 1.08 2
MM98/3567 2.41 4 7.76 14 2.32 14 1.83 5
Plant height (PH), height at first branching (HB), number of storage roots per plant (NSR) and fresh root yield (FRY)
branching and fresh root respectively, meaning more than locations in which they were evaluated and that they can
24% variability observed in height at first branching and consistently be evaluated under any of the locations used
fresh root yield is due to GEI effects. In spite of this, the for this study in impending performance trials. This view is
magnitude of the genotype sum of squares for plant height in conformity with the view of Peprah et al. (2013) who
and height at first was branching was larger than that of found non-significant GEI effects for dry matter content
GEI (20.2% and 50.9%) respectively which indicates and reported that fewer environments may be needed to
presence of moderate control of genotype effects over distinguish clones with high and stable performance for
genotype by environment interaction effects for these this trait. In other words, evaluating genotypes for these
traits. traits concurrently in the various locations used for these
studies in consequent evaluation trials might not be
On the other hand, there was non-significant GEI effects important. Thereby, offering an opportunity to manage
for number of storage roots, dry matter content and starch. inadequate means available for testing programme (Tonk
The phenomenon was also the same as reported by et al., 2011).
Peprah et al. (2013) who observed non-significant GEI for
dry matter content. This finding also agrees with those of Complementary to previous results, ASV was computed
Aina et al. (2007) who reported non-significant GEI effects for the traits so as to quantify and rank genotypes
for number of storage roots and dry matter content. according to their stability Table 3 shows the ranking of the
Similarly, Benesi et al. (2004) reported non-significant GEI 16 genotypes according to AMMI stability value.
for starch. Genotypes varied in ranking for stability across the studied
traits. However, some genotypes combined satisfactory
An obvious deduction from non-significant GEI effects on results for stability in various traits. For instance, genotype
number of storage roots, dry matter content and starch is MM06/0143 is very stable as it ranked 2 nd, 3rd, 1st, and 7th
that, genotypes might have similar responses across the for plant height, height at first branching, number of
Genotype by Environment Interaction on Cassava Components and Yield Stability Analysis of Elite Cassava Genotypes in Western Kenya
Int. J. Plant Breed. Crop Sci. 365
storage roots and fresh root yield. MM98/3567 combines concentrated around zero for most of the clones
stability for plant height and fresh root yield while suggesting that the candidate clones were stable for this
MM97/0293 combines stability for plant height, number of trait (number of fresh root yield). Though clones G2
storage roots and fresh root yield. Genotypes MM06/0046, (migyera), G11 (MM0h6/0143) and G4 (MM06/0046)
MM06/0083, MM06/0143, and MM06/0074 were stable revealed some displacement on the Y-axis from the origin,
genotypes for plant height, height at first branching, it’s clear that the PC scores for these clones in this Y’-axis
number of storage roots per plant and fresh root yield is less than one (near zero) for the three clones hence it
respectively. These suggest the possibility of identifying was adjudged that all the candidate clones were stable for
clones exhibiting stable performance in both agronomic this trait.
and farmer preferred traits.
Figure 2: Mean performance and stability of 16 cassava Figure 3: “Which won where pattern” of GGE biplot for
genotypes (G1-G16) at five environments (K:Kakamega, sixteen clones (G1-G16) at five environments (K:Kakamega,
S:Sangalo, A:Alupe, B:Kibos, M:Migori) for fresh root yield. S:Sangalo, A:Alupe, B:Kibos, M:Migori) on number of
storage roots.
The polygon view of the GGE biplot explicitly displays the
’’which won where pattern’‘ and hence is a concise In Figure 4, the ‘‘which won where” pattern of the GGE
summary of the GEI pattern (Figures 3 and 4) .The polygon biplot on fresh root yield explained 77.15% of the total
is formed by connecting the markers of the genotypes that variation due to GEI effects, PC1 accounted for 51.7%
are further away from the biplot origin such that all the while PC2 accounted for 25.45%. The biplot revealed the
genotypes are contained in the polygon (Yan et al., 2007; best genotypes across environments and identified the
Akinwale, 2011).Convex-hull are drawn from the biplot best clones with respect to site. The seven rays that divide
origin which divides the biplot into sectors that demarcate the biplot into seven sectors to which five environments fall
mega-environment the vertex genotypes in a sector of into two of them showed that (Alupe, Sanga’lo and Kibos)
environment are considered the most stable for that environments fall into sector one and the vertex genotypes
environment. In Figure 3, the “which won where pattern” of for this sector was G16 (MM98/3567) Similarly, two
the GGE biplot on number of storage roots grouped all the environments (Kakamega and Migori) fell into sector two
environments in one sector. Moreover, the genotypes and the vertex genotypes for this sector was G9
clustered around the origin of the biplot revealing that the (MM06/0131). No environment fell into sectors with G10
genotypes had the same response across the (MM06/0131), G4 (MM06/0046), G2 (migyera) and G8
environments. Generally, the GGE biplot on number of (MM06/0131) as vertices indicating that these cultivars
storage roots accounted for 95.74% of the total GEI were unstable in all the environments. Genotypes G5
variation due to GEI effects on number of storage roots (MM06/0074) and G13 (MM96/4271) were located at the
with PC1 and PC2 accounting for 87.52% and 8.22%, origin of the biplot revealing that they were highly stable
respectively. clones across the sites.
Genotype by Environment Interaction on Cassava Components and Yield Stability Analysis of Elite Cassava Genotypes in Western Kenya
Int. J. Plant Breed. Crop Sci. 367
Genotype by Environment Interaction on Cassava Components and Yield Stability Analysis of Elite Cassava Genotypes in Western Kenya
Int. J. Plant Breed. Crop Sci. 369
Genotype by Environment Interaction on Cassava Components and Yield Stability Analysis of Elite Cassava Genotypes in Western Kenya