Principles of Plant Genetics and Breeding
Principles of Plant Genetics and Breeding
Principles of Plant Genetics and Breeding
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424 CHAPTER 23<br />
example, a significant G × year interaction cannot be<br />
exploited because the climatic conditions that generate<br />
year-to-year environmental variation are not<br />
known in advance. Consequently, the analysis <strong>of</strong><br />
multiple environment yield trials should focus primarily<br />
on G × location interactions, the other interaction<br />
being considered in terms <strong>of</strong> yield stability.<br />
Stability analysis<br />
An ANOVA identifies statistically significant <strong>and</strong> specific<br />
interactions without telling the breeder anything about<br />
them. If a G × E interaction is significant <strong>and</strong> the environmental<br />
variations are unpredictable, the breeder needs<br />
to know which genotypes in the trials are stable. A<br />
stability analysis is used to answer this question. Field<br />
stability may be defined as the ability <strong>of</strong> a genotype to<br />
perform consistently, whether at a high or low level,<br />
across a wide range <strong>of</strong> environments. Stability may be<br />
static or dynamic. Static stability is analogous to the<br />
biological concept <strong>of</strong> homeostasis (i.e., a stable genotype<br />
tends to maintain a constant yield across environments).<br />
Dynamic stability is when a stable genotype<br />
has a yield response in each environment such that it is<br />
always parallel to the mean response <strong>of</strong> the genotypes<br />
evaluated in the trials (i.e., G × E = 0). Static stability is<br />
believed to be more useful than dynamic stability in a<br />
wide range <strong>of</strong> situations, <strong>and</strong> especially in developing<br />
countries. The genotype produces a better response in<br />
unfavorable environments or years. Whenever the G × E<br />
interaction variation is wide, breeding for high-yield<br />
stability is justifiable.<br />
As previously stated, ANOVA only detects the existence<br />
<strong>of</strong> G × E effects. Breeders will benefit from additional<br />
information that indicates the stability <strong>of</strong> genotype<br />
performance under different environmental conditions.<br />
The stability <strong>of</strong> cultivar performance across environments<br />
depends on the genotype <strong>of</strong> individual plants<br />
<strong>and</strong> the genetic relationship among them. Generally,<br />
heterozygous individuals (e.g., F 1 hybrids) are more<br />
stable in their performance than their homozygous<br />
inbred parents.<br />
A variety <strong>of</strong> methods have been proposed for genotype<br />
stability analysis. Examples are regression analysis<br />
<strong>and</strong> the method <strong>of</strong> means.<br />
Regression analysis<br />
This method <strong>of</strong> simple linear regression (also called<br />
joint regression analysis) stability analysis was developed<br />
by K. W. Finlay <strong>and</strong> G. N. Wilkinson (1963)<br />
<strong>and</strong> later by S. A. Eberhart <strong>and</strong> W. A. Russell (1966).<br />
It is preceded by an ANOVA to assess the significance<br />
<strong>of</strong> G × E. The breeder proceeds to the next step <strong>of</strong><br />
regression analysis only if the G × E interaction is<br />
significant.<br />
Statistically, the observed performance (Y ij ) <strong>of</strong> the<br />
ith genotype (i = 1,..., 5) in the jth environment<br />
(j = 1,..., e), may be expressed as:<br />
Y ij =µ+g i + e j + ge ij +ε ij<br />
where µ =gr<strong>and</strong> mean over all genotypes <strong>and</strong> environments,<br />
g i = additive contribution <strong>of</strong> the ith genotype<br />
(calculated as the deviation from µ <strong>of</strong> the mean <strong>of</strong> the ith<br />
genotype averaged over all environments), e j = additive<br />
environmental contribution <strong>of</strong> the jth environment<br />
(calculated as the deviation µ <strong>of</strong> the mean <strong>of</strong> the jth<br />
environment averaged over all genotypes), ge ij = G × E<br />
interaction <strong>of</strong> the ith genotype in the jth environment,<br />
<strong>and</strong> ε ij = error term attached to the ith genotype<br />
in the jth environment. The regression coefficient for a<br />
specific genotype is obtained by regressing its observed<br />
Y ij value against the corresponding mean <strong>of</strong> the jth<br />
environment.<br />
If the yield was conducted over a wide range <strong>of</strong> environments,<br />
<strong>and</strong> hence a wide range <strong>of</strong> yields obtained, it<br />
is reasonable to assume that the individual trial means<br />
sufficiently summarize the effects <strong>of</strong> the environments.<br />
The mean performance <strong>of</strong> each genotype over all the<br />
test environments constitutes the environmental index<br />
(Table 23.2). In effect, this method <strong>of</strong> analysis produces<br />
a scale <strong>of</strong> environmental quality. The results for each<br />
genotype are plotted, trial by trial, against trial means, to<br />
obtain a regression line. According to Eberhart <strong>and</strong><br />
Russell, an average performing genotype will have a<br />
regression coefficient <strong>of</strong> 1.0 <strong>and</strong> deviations from regression<br />
<strong>of</strong> 0.0, since it will tend to agree with the means.<br />
However the genotypes that were responsible for the<br />
G × E interactions detected in the ANOVA, will have<br />
slopes that are unequal to unity. Furthermore, a genotype<br />
that is unresponsive to environments (i.e., a stable<br />
performer) will have a low slope (b < 1), while a genotype<br />
that is responsive to environments (i.e., an unstable<br />
performer) will have a steep slope (b > 1).<br />
The regression analysis technique has certain limitations,<br />
both physiological <strong>and</strong> statistical, that can result<br />
in inaccurate interpretation <strong>and</strong> recommendations for<br />
cultivar release. For example, the use <strong>of</strong> mean yield in<br />
each environment as an environmental index flouts<br />
the statistical requirement <strong>of</strong> an independent variable.<br />
Also, when regression lines do not adequately represent<br />
data, the point <strong>of</strong> intersection <strong>of</strong> these lines<br />
has little biological meaning, unless considered with