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Identifying Mega-Environments and Targeting

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 Published March, 1997
Identifying Mega-Environments and Targeting Genotypes
Hugh G. Gauch, Jr.,*
ABSTRACT
To maximizeyield throughout a crops heterogeneous growing re-
gion, despite differences in cultivar rankings from place to place due
to genotype-environmentinteractions, frequently it is necessary to
subdivide a growingregion into several relatively homogeneous mega-
environments and to breed and target adapted genotypes for each
mega-environment. Theobjectives of this study are to identify relevant
criteria for evaluating mega-environmentanalyses and to apply the
Additive Main Effects and Muitiplicative Interaction (AMMI) model
to mega-environmentanalysis. The proposed analysis is illustrated
using a Louisianacorn (Zea maysL.) trial. Statistical strategies for
identifying mega-environments should meet four criteria: flexibility
in handlingyield trials with various designs, focus on that fraction of
the total variation that is relevant for identifying mega-environments,
duality in giving integrated informationon both genotypesandenviron-
ments, and relevance for the primary objective of showing which
genotypes win where. The AMMI modelmeets these criteria effectively
whenthe usual biplots are supplemented with several new types of
graphs designed to address questions about mega-environments. Pre-
liminary results indicate that a small and workablenumberof mega-
environmentsoften suffices to exploit interactions andincrease yields.
N A REGIONAL YIELD TRIAL with heterogeneous envi-
.ronments and numerous genotypes, different genotypes
are superior in different locations (or years or both).
optimize growers yields, despite genotype-environment
interactions that cause no one genotype to win every-
where and always, the growing region must be subdivided
into relatively homogeneousmega-environments and ap-
propriate genotypes must be targeted for each of these
mega-environments. A mega-environment is defined as
a portion (not necessarily contiguous) of a crop species
growing region with a fairly homogeneousenvironment
that causes similar genotypes to perform best. With no
subdivision, only broad adaptation can be exploited,
but with subdivision, narrow adaptations can also be
exploited. Subdivision of a crops growing region into
several mega-environments implies more work for plant
breeders and seed producers, but subdivision also implies
higher heritabilities and faster progress for plant breed-
ers, potentially stronger competitiveness for seed produc-
ers, and higher yields for growers. For example, Annic-
chiarico (1992) compared alfalfa (Medicago sativa L.)
yields in three mega-environments in Italy with yields for
the overall regional averages. The spread and statistical
significance of yield differences Wasgreater in the indi-
vidual mega-environmentsthan in the overall averages,
and the highest yielders in each mega-environmentwere
different from the overall winner. Also, results from a
given location had good predictive reliability for other
H.G. Gauch, Jr., Soil, Crop and Atmospheric Sciences, Cornell Univ.,
1021 Bradfield Hall, Ithaca, NY14853-1901; and R.W. Zobel, USDA-
ARS-NAA, 1017 Bradfield Hall, Ithaca, NY14853-1901. Received 19
April 1996. *Corresponding author (hggl@comell.edu).
Published in Crop Sci. 37:311-326 (1997).
311
and Richard W. Zobel
locations in the same mega-environment (but of course
not outside the locations own mega-environment).
The term mega-environment appears to have been
coined by researchers at CIMMYT. "Mega-environments
are broad, not necessarily contiguousareas, usually inter-
national and frequently transcontinental, defined by simi-
lar biotic and abiotic stresses, cropping system require-
ments, consumer preferences, and, for convenience, by
a volume of production of the relevant crop sufficient
to justify ... attention," for example, "tropical lowland,
late-maturing, white dent" corn with relevant disease
resistances, which occupies 3.8 million hectares across
18 countries (CIMMYT,1989, p. 58). Note that this
definition encompasses environmental, genotypic, geo-
graphical, and even economic aspects of mega-envi-
ronments. Mega-environments are used to allocate re-
sources in a breeding or research program, to rationalize
germplasm and information exchanges between breeding
programs (allowing even small programs to progress by
focusing on the most promising material), to increase
heritabilities within relatively well-defined and predict-
able environments, to increase the efficiency of testing
and breeding programs, and to target genotypes to appro-
priate production areas (Brown et al., 1983; Peterson
and Pfeiffer, 1989; Abdalla et al., 1996). Manyother
terms, however, have essentially the same meaning, such
as agroclimatic or ecogeographic regions. Because the
need for mega-environments arises from genotype-envi-
ronment interactions and furthermore these interactions
involve both genotypes and environments, there are also
corresponding terms applied to genotypes for identifying
adaptation traits, such as the maturity groups in corn
and soybean [Glycine max (L.) Merr.] that are used
for delineating adaptation zones. Virtually the entire
literature on interaction, clustering environments, yield
stability and dependability, wide and narrow adaptation,
and heritability is relevant for understanding and charac-
terizing mega-environments, whether or not the word
mega-environment appears in a given paper.
Subdivision of a crops growing region into several
mega-environments could be avoided if genotypes could
be found with yield superiority throughout the region,
meaning that cultivars bred in favorable environments
wouldalso perform best in different or unfavorable envi-
ronments. But this expectation assumes that "the same
genetic system controls yield" in diverse environments,
contrary to extensive evidence from genetics, physiol-
ogy, and yield trials (Ceccarelli and Grando, 1993; Cec-
carelli, 1989; Simmonds,1991). Consequently, for many
crops, acceptable yields have necessitated targeting nar-
rowly adapted genotypes to several different, well-
Abbreviations:AMMI, Additive MainEffects and Multiplicative Interac-
tion; AOV,analysis of variance; CIMMYT, Centro Internacional de
Mejoramientode Maizy Trigo; df, degrees of freedom; IPCA,interaction
PCAaxis; PCA,principal componentsanalysis; S/N, signal-to-noise; SS,
sum of squares; G, genotype; E, environment; L, location; Y, year; R,
replication.
312 CROPSCIENCE, VOL. 37, MARCH-APRIL
defined mega-environments.Indeed, one can hardly ex-
pect a single cultivar of a crop to flourish the world
over, under all environmentsand management practices
(Rosielle and Hamblin,1981; Ceccarelli, 1989). A culti-
var planted outside its mega-environmentfrequently
suffers yield reductions. Furthermore,evenif the breed-
ing goal is wide adaptation (rather than mega-envi-
ronmentdirected breeding as in Abdallaet alo, 1996),
the best strategy could be to identify several mega-
environmentsand place a test location in each to select
for wide adaptation.
For several reasons, identifying mega-environments
has attracted greater attention recently. Interest has
grownin providing adapted material for marginal envi-
ronments,particularly becausemillions of people live
in such places and yet they often lack the economic
meansneededto purchaselarge quantities of grain from
moreproductiveagricultural areas. Thesemarginalenvi-
ronmentsare stressed in a variety of waysthat typically
generate large genotype-environmentinteractions, so
genotypesthat win in highly favorable environmentsmay
rank poorly there (Ceccarelli, 1989; Simmonds,1991;
Nachitet al., 1992;Zavala-Garcia et al., 1992;Ceccarelli
and Grando, 1993; Annicchiarico and Perenzin, 1994;
Falconer, 1989, p. 313-335). On the other hand, within
the favorable environmentsof major production areas,
conventional managementprocedures have produced in
the past a rather coherent environmenteasily targeted
by plant breeders. In the future, however,it is likely
that greater concernabout long-term soil conservation
and about loweringpesticide and fertilizer usages will
stimulate a greater diversity of management practices,
hence creating a variety of mega-environmentswithin
major crop regions previously managedin a muchmore
uniform manner(Smith and Zobel, 1991). Finally and
moregenerally, most plant breeders feel that they are
exploiting rather than ignoring the potential for yield
increases that resides in genotype-environment interac-
tions (Cooperet al., 1993). Nevertheless,despite breed-
ers strong interests in interactions andspecific adapta-
tions, one can question whetherthey have routinely had
adequatestatistical tools to exploit interactions aggres-
sively and to grasp clearly the inevitable implications
for identifying mega-environments.
The primary proposal adopted here for interlinking
mega-environmentsand genotypes is to define a mega-
environmentas a portion of a crops overall growing
region in whicha particular genotype wins. A comple-
mentaryproposalis to define mega-environments in terms
of major environmentalfactors, such as irrigated low-
lands and nonirrigated uplands. These genotype-based
and environment-basedproposals are interdependent and
complementary(Peterson and Pfeiffer, 1989; DeLacyet
al., 1994). The genotype-basedconcept is relevant be-
cause planting the winning genotype in each mega-
environmentoptimizes yield. Andthe environment-based
concept is helpful for assigning individual farms to an
appropriate mega-environment (and hence genotype rec-
ommendation), especially if the mainenvironmentalfac-
tor has an obviousand simple geographicaldistribution.
This interlinking of mega-environmentsand genotypes
1997
is especially clear whendevelopmentof more widely
adaptedcultivars revises andsimplifies a previousmega-
environment classification. Indeed,if differencesin envi-
ronmentalfactors fromplace to place (and year to year)
affected only environmentmaineffects but not genotype-
environment interactions, then these environmental
differences wouldbe of interest to crop physiologists
and agricultural economistsbut not plant breeders. Mega-
environmentdifferences exert their importance - for
plant breeders - by changing genotype rankings from
one mega-environmentto another. Accordingly, this
studys title presents "identifying mega-environments"
and "targeting genotypes"as deeply and intrinsically
interrelated tasks.
The mainstatistical strategy for groupinglocations
into mega-environments has been various classification
and ordination methods(or both, whichis termedpattern
analysis). Amongthese numerousordination methods,
the AMMI model has provided biplot graphs useful for
delineating mega-environments(Annicchiarico, 1992;
Annicchiaricoand Perenzin, 1994; Annicchiaricoet al.,
1995). Despite this success, however,the usual biplot
graphsreveal only a portion of the informationavailable
from AMMI that bears on mega-environments.In partic-
ular, these graphs fail to makeevident whichgenotype
has the highest yield in various domainsof the AMMI
graphs. Accordingly, this paper presents several new
extensions of AMMI that enhance its informativeness.
The merits of this new methodologyare (i) focus
that fractionof the total yield variationthat is relevantfor
identifying mega-environments and targeting genotypes,
(ii) duality in giving integratedinformationon both geno-
types and environments,and (iii) relevancefor the pri-
mary objective of showingwhich genotypes win where.
This analysis is also flexible in handlingyield trials with
various designs. Preliminaryresults indicate that a small
and workable number of mega-environments often
suffices to exploit interactions and increase yields.
FOUR CRITERIA
Rational assessments of mega-environmentanalyses
are possibleonly after specifyingthe criteria for evalua-
tion. Fourkeycriteria are emphasized here: (i) selective
focuson the variation in yield that is relevant for identi-
fying mega-environments, (ii) relevance to agronomists
and breeders questions, especially "Whatwinswhere?",
(iii) dual analysis of both genotypesand environments,
and(iv) flexibility with handlingvariousdata structures.
First, mega-environment analyses should focus on rel-
evantvariationin the data andignoreirrelevant variation.
Considera regional variety trial fromthe perspective of
a breeder makingselections or an agronomist offering
recommendations.Analysis of variance partitions the
treatment variation into three basic sources: genotypes,
environments, and genotype-environmentinteraction.
Here a treatment is defined as a genotypeand environ-
mentcombination.The genotypeand interaction sources
affect genotype rankings within each environmentand
hence are relevant for identifying mega-environments
and targeting genotypes. But the environment main
 GAUCH& ZOBEL: IDENTIFYING MEGA-ENVIRONMENTS
effects are wholly irrelevant for genotype rankings and
hence for present objectives. Because of the central goal
of makinggood selections, "the breeder is particularly
interested in the rank orders of the genotypes in the
different environments" (Hiihn et al., 1993). Fox and
Rosielle (1982), Gauch (1992, p. 220-230), Cooper
DeLacy(1994), and Annicchiarico et al. (1995) strongly
emphasize that breeding decisions and mega-environment
choices dependon genotype and interaction effects exclu-
sively (Yau, 1991). Often these relevant effects account
for just 10 to 40%of the treatment variation (because
environmentmain effects account for the remaining varia-
tion), so a statistical analysis modelingall of the variation
is mostly capturing irrelevant features of the data, which
greatly hinders effective mega-environment identification.
For example, Kang (1993) kindly supplied us the data
from a Louisiana corn yield trial. This experiment had a
randomizedcomplete-block design with four replications
(except that 32 treatments had only three replicates and
1 had only two replicates). Because of the missing data,
for convenience, the analysis is approximated here as a
completely randomized design [but Piepho (1994) ex-
plains the calculations for randomized complete blocks
without missing data]. Table 1 gives the analysis of
variance table for the AMMI-1 model. The SS for treat-
ments is 3842.13. However, the goal of a mega-
environment analysis should not be to account for this
entire treatment SS. Rather, the relevant portion of this
SS, ignoring the environmentmain effect, is only 238.93
+ 738.03 = 976.96, or 25.4%.
Furthermore, errors from uncontrolled variation
within experimental fields are also irrelevant for present
concerns and unfortunately cause inaccuracy or noise in
yield estimates. Because interaction has many more df
than the genotype and environment main effects, most
of this noise appears in the interaction. (The grand mean,
genotype deviations, and environment deviations also
contain some noise, but these problems are neglected
here because this other noise is small comparedwith the
interaction noise and because there is no way to removea
sizable portion of this other noise.) Noisegives yield-trial
results a spurious complexity, thereby multiplying the
number of mega-environments needlessly. Conse-
quently, reducing noise helps when identifying mega-
environments.
For this corn trial, the error mean square is 2.035
and the interaction has 225 df, so the interaction contains
approximately 225 2.035 = 457.88 noise SS and
738.03 - 457.88 = 280.15 real structure SS or 62.0%
noise and 38.0% pattern (Gauch, 1992, p. 147). Dis-
counting for this interaction noise, the relevant variation
is only 238.93 + 280.15 = 519.08 or 13.5% of the
treatment SS. So, an ideal mega-environment analysis
would focus on this 13.5%of relevant variation in the
data, and ignore the other 86.5 %pertaining to irrelevant
environment effects and interaction noise. By contrast,
capturing less than this 13.5 %underfits real and relevant
patterns in the data, and capturing morethan this 13.5 %
overfits noise and captures irrelevant features in the data.
One gathers from extensive results in Talbot (1984),
DeLacyet al. (1990), Braun et al. (1992), and HiJhn
313
Table 1. AMMI-1analysis of variance table for the Lousiana corn
yield trial. The grand mean is "6.69324 Mgha-~.~
Source df SS MS Probability
Total 989 5336.17 5.396
Treatments 255 3842.13 15.067 0.0000000
Genotypes 15 238.93 15.929 0.0000000
Environments 15 2865.17 191.011 0.0000000
G x E 225 738.03 3.280 0.1)000019
IPCA 1 29 229.81 7.925 0.0000000
Residual 196 508.22 2.593 0.0139627
Error 734 1494.04 2.035
AMMI = Additive M__ainEffects and Multiplieative I_nteraction; df =
degrees of freedom; SS = sum of squares; MS= mean square; IPCA=
interaction principal components
analysis axis.
al. (1993) that this exampleis typical. Again,the relevant
variation for identifying mega-environmentsis often only
~ 10 to 40%of a yield trials treatment variation.
Incidentally, because the interaction has most of the
treatment df and hence noise but has only a fraction of
the treatment SS, interaction is muchnoisier than the
data (yield averages across replications). The data
of 3842.13 with 255 df contain 255 2.035 = 518.93
noise and 3323.20 pattern, so its S/N ratio is 3323.20/
518.93 = 6.40. But the interaction S/N ratio is only
280.15/457.88 = 0.61, which is ten times smaller. The
general lesson to be learned here is that the interaction
gets buried by noise more quickly than genotype and
environment main effects. Consequently, the breeding
strategy of sampling more locations with fewer replica-
tions to characterize interactions better can be self-
defeating if there is not enoughaccuracy to makeinterac-
tions rise abovethe noise. Yield trials with elite genotypes
having relatively small interactions that are conducted
as unreplicated tests are especially prone to this problem.
Mega-environment analysis should capture genotype
and (real) interaction effects but should also distinguish
these two effects for reasons that are agricultural rather
than statistical. Genotypeeffects control wide adapta-
tions, whereasinteraction effects control narrow adapta-
tions, and these two kinds of adaptations offer breeders
different problems and opportunities, especially in re-
gards to designing testing systems and making good
selections. Also, genotype and interaction effects often
relate to different environmental and genetic systems.
For example, genotype main effects may involve water
and nutrient uptake, whereas interaction effects may
involve photoperiod response. For these reasons, it is
preferable for a mega-environmentanalysis not to con-
found the genotype and interaction SS but rather to
treat them separately. Althoughgenotype and interaction
effects should be distinguished, they should also be inte-
grated in regards to their impact on yields because yield
rankings are affected by these two effects jointly. So,
genotype and interaction effects should be distinguished,
not confounded, and should be integrated, not divorced.
Second, mega-environment analyses should provide
relevant answers to researchers primary questions. The
obvious question is: What wins where? An answer to
this question provides a basis for grouping locations with
identical (or at least similar) winners into a mega-
environment and for targeting suitable genotypes for
each mega-environment. Mega-environment analyses
314 CROPSCIENCE, VOL. 37, MARCH-APRIL
that answer questions other than "Whatwins where?"
should be carefully checkedto determinewhether their
answersbear effectively on the objective of subdividing
a crops growingregion to increase yields. Otherwise,
laborious calculations mayoptimizea mathematicalprop-
erty of dubiousor tangential relevanceto the agricultural
task at hand.
Interactionsdiffer in their relevancefor plant breeders.
Breedershave emphasizedcrossover interactions because
noncrossover interactions do not change rankings or
selections (Crossa, 1990). But this insight should
supplementedwith a second emphasis, that crossovers
amongwinninggenotypes are moreimportant than cross-
overs amonglosers. For a trial with manygenotypes,
only a tiny fraction of the crossovers involve winners,
so equal emphasison all crossovers diffuses attention
from the mainaction and multiplies mega-environments
needlessly. Furthermore, crossover interactions have
three sources-spurious noise, unpredictable environ-
mental causes, and predictable environmental causes-
of whichonly the last can be exploited for identifying
mega-environments. In summary, mega-environment
analysis should emphasizecrossover interactions among
winninggenotypescaused by reasonably predictable en-
vironmentalfactors.
Third, mega-environment analyses should provide an
integrated understandingof the genotypesand environ-
ments. The underlying concept of genotype-environment
interaction, and the derived concept of mega-envi-
ronmentsare fundamentallydual, involving both geno-
types and environments.Hence,the objectives of identi-
fying mega-environmentsand targeting genotypes are
inherently and deeply interrelated and must be pursued
jointly, as this studystitle suggests.Werethe roster of
genotypeschangedin a regional trial, it is possible,
and even probable, that appropriate mega-environments
wouldalso change. Likewise,were the definition or the
numberof mega-environmentschanged, the number of
recommended genotypes and the particular recommenda-
tion for each mega-environmentwould probably also
change. Accordingly,an ideal analysis or clustering of
mega-environmentsdoes not analyze just genotypes or
just environmentsseparately but rather both genotypes
and environmentsin a single integrated model.It would
be strange for a clustering analysis applied to mega-
environmentsto analyze only environmentsor only geno-
types since genotype-environmentinteraction involves
both.
Fourth and finally, mega-environment analyses needto
be flexible regardingthe structure of availableyield-trial
data. The data mayhave a two-wayGEstructure or a
three-way GLYstructure. The trial maybe replicated,
resulting in a GERor GLYR structure, or it maybe
unreplicated. Experimentsmayhave completeor incom-
plete factorial designs, and there maybe somemissing
data. So, the least generoussetup is a GEdesign with
somemissing data and no replication, whereasthe most
generous setup is a GLYR experiment with a complete
factorial design,replication, andnomissingdata. Statisti-
cal ahalyses for identifying mega-environments should
be flexible, accepting data across this wholerange of
1997
possibilities. Naturally, moreextensive data allow more
extensive results, but smaller experimentsalso merit
effective analysis. For a trial with GLY structure, re-
peated across both locations and years, mega-envi-
ronmentanalysis should discriminate betweenpredict-
able and nonpredictableinteractions, largely associated
with locations andyears, respectively.
Whathappenswhenthese four criteria are neglected?
(i) Clusteranalysisof the yield data withoutfirst removing
environmentmaineffects causesweakresults that mostly
reflect irrelevant features of the data. Likewise,mistaking
spuriousnoise for real interactions causesirrelevant and
unrepeatable results and multiplies mega-environments
needlessly. (ii) Not groupinglocations by their winning
genotypecausesindividual clusters with a mixof winners
or separate clusters with the samewinner or both prob-
lems, so these clusters are not suitable for identifying
mega-environments or for targeting genotypes.(iii) Not
analyzing both genotypes and environmentsmeansthat
a satisfactory understanding of genotype-environment
interactions and their implications for mega-envi-
ronments is just not possible. (iv) Nothandlinga diversity
of generousor sparce datasets limits the applicablity of
an analysis. Furthermore, combinations of several of
these problemsat once producevery confusing and mis-
leading conclusions. For example,clustering yield data
with large environment maineffects anda sizable interac-
tion that is actually just noise could divide a region
into several clusters or mega-environments, and yet this
subdivisionreflects only irrelevant environment effects.
Unfortunately,were such a poor analysis taken seriously
becauseits problemswere not recognized, it could make
a sensible analysis look bad becauseit reaches different
conclusions.Clearly, it is essential for mega-environment
analyses to be evaluated accordingto relevant criteria.
THE ADDITIVE MAIN EFFECTS
AND MULTIPLICATIVE INTERACTION
MODEL FAMILY
Considera yield trial with a two-wayfactorial design
of G genotypesand E environments,with R replications,
denoting yield observations by Yger- The AMMI model
combines AOVwith additive parameters and PCAwith
multiplicative parametersinto a single analysis. Having
first removedthe grand meanI~ from the data, AOV
partitions the varianceinto three sources: genotypedevia-
tions from the grand meantg, environmentdeviations
from the grand mean I~e, and genotype-environment
interactions 0g~. ThenPCAfurther partitions the interac-
tions 0g~into NaxesEn~.n3,gn~5~ with ~.n the singularvalue
for axis n, 3g~ the gth value in the genotypeeigenvector
for axis n, and 6~n the eth value in the environment
eigenvector,plus residuals pg~if Nis less than the full
model.If the experimentis replicated, there is also an
error term, eg~r, further partitioned in accordancewith
the particular experimental design. The AMMI model
equation is
Yger = l-i, q- (tg + ~e + ,~_~nLn~gnSend- Pge d- I~ger
AMMI
is not a single modelbut rather a modelfamily,
 GAUCH& ZOBEL: IDENTIFYING MEGA-ENVIRONMENTS 315

with that model having N axes denoted by AMMI-N. broad implications for multivariate models in general.
(In the same way, the more familiar polynomial models This potential for gaining accuracy was first exploited
comprise a modelfamily including constant, linear, quad- for AMMI by Gauch (1988). Subsequently, agronomists
ratic, cubic, and higher-order models.) The most simple and breeders have used AMMIextensively for both of
and most complex extremes are AMMI-0with no axes, these purposes: understanding interactions and gaining
equivalent to AOV,and the full model AMMI-Fwith accuracy (Gauch and Zobel, 1996a). Both are relevant
F = min(G - 1, E - 1) axes, equivalent to means across in the present context of identifying mega-environments.
replications. Hence, AMMIanalysis provides models
intermediate to the extremes of the additive model and
ADDITIVE MAIN EFFECTS
the raw data.
AND MULTIPLICATIVE INTERACTION
With replication, data splitting can be used to diagnose
MEGA-ENVIRONMENT ANALYSIS
the most predictively accurate member of the AMMI
family for a given dataset (Gauch, 1988, 1992, p. 134- Mega-environmentanalysis is illustrated here using
153; Gauch and Zobel, 1988). As will be emphasized yield data from a Louisiana corn trial analyzed previously
momentarily, effective use of AMMI requires careful by Kang (1993). Parameters of the AMMI-1model for
model diagnosis. The Expectation-Maximization imple- these data are given in Table 2. This yield trial has 16
mentation of AMMIcan handle and impute missing genotypes tested in 16 environments comprised of four
data (Gauch and Zobel, 1990; Gauch and Furnas, 1991; locations for four years, with four replications (or occa-
Gauch, 1992, p. 157-162). A convenient scaling for the sionally only two or three, but to avoid excessively
multiplicative parameters, for a given IPCAaxis, is difficult computations, an approximate solution is ob-
~,0.5~g and ~,5~5obecause the product of these interaction tained here by treating these data as if they were bal-
scores gives the interactions expected value directly, anced). The genotype codes are transparent abbreviations
without needing further multiplication by the singular of the full names in Kang(1993), the location codes are
value. For balanced data, AOVfollowed by PCA of AXfor Alexandria, BR for Baton Rouge, BC for Bossier
the interaction provides the least-squares solution. The City, and SJ for St. Joseph, and the years are 1985 to
AMMI solution is unique up to simultaneous sign changes 1988. For example, using this table and the above model
of a given genotype and environment eigenvector pair equation, the AMMI-1model estimates the yield for
(because the product ~,3g~e remains unchanged), Genotype 3165 in Environment BR87as 6.693 + 0.472
different software (or different orderings of the genotypes
and environmentsin the dataset) can give reversed signs.
This possibility emphasizes that contrast and trend from Table 2. AMMI-1model for the Louisiana corn yield trial with
16 genotypes and 16 environments. Deviations are in units of
positive to negative scores is meaningful but not the Mgha -1, IPCA scores are in units of (Mg ha-~) -s, and the
happenstancethat a particular score is positive or negative grand meanis 6.693 Mgha- 1. Entries are listed in rank order
in the output from a given software package. A worked of the IPCAscores.~"
example of AMMI calculations is given by Gauch (1992, Entity Deviation IPCA 1
p. 85-88). The AMMIanalysis was done by MATMO-
DEL l, which includes a model validation procedure for 3165 0.472 1.295
3147 0.665 0.783
diagnosing the most predictively accurate memberof the 1860 0.619 0.745
AMMIfamily for a given dataset (Gauch and Furnas, CK21 0.182 0.554
8990 - 1.046 0.371
1991). 4765 - 0.105 0.322
The AOV and PCA portions of the AMMImodel 8951 - 0.519 0.231
3389 - 0.255 0.183
were invented by Fisher (1918) and Pearson (1901). 0.003
8172 0.840
Fisher and Mackenzie (1923) applied both AOVand 3320 - 0.259 - 0.021
PCAseparately to the same potato (Solanum tuberosum 4673 0.280 - 0.093
4733 - 0.202 - 0.475
L.) yield trial. But the two components of AMMI were 1802 - 0.332 - 0.793
not combinedinto a single analysis until 1952 by Williams 4522 - 0.431 - 0.985
(1952) and Pike and Silverberg (1952). Further develop- 8150 - 0.050 - 0.987
1827 0.144 - 1.135
ments are reviewed by Gauch (1992, p. 8-12), but the BC88 - 1.184 1.338
incisive paper by Kempton(1984) first brought AMMI AX87 0.625 1.181
AX88 - 2.422 0.723
to the widespread attention of agronomists and breeders BC86 - 3.436 0.593
for the purpose of understanding complex genotype- BC85 - 1.608 0.297
environment interactions. Coincidentally, at about the SJ87 1.884 0.189
BC87 - 1.142 0.136
same time that AMMI was invented, Stein (1955) first SJ85 0.164 0.122
demonstrated that estimators from a multivariate model SJ86 0.541 0.008
could be more accurate and efficient than the usual aver- AX85 2.177 - 0.186
BR87 2.130 - 0.620
age across replications, and this intriguing result had AX86 2.478 - 0.622
BR88 - 0.242 - 0.639
SJ88 - 1.127 - 0.699
~ Mentionof a trademark or proprietary product does not constitute a BR85 0.327 - 0.873
BR86 0.834 - 0.947
guarantee or warranty of the product by the USDA or Cornell University
and does not imply its approval to the exclusion of other products that AMMI = Additive M._ainEffects and M._ultiplicative I_nteraction; IPCA=
mayalso be suitable. interaction principal componentsanalysis axis.
316 CROPSCIENCE, VOL. 37, MARCH-APRIL
1997

+ 2.130 + [1.295 (-0.620)] = 8.492 Mg -l ,
which approximates the actual yield of 8.554.
The AMMI-1model was diagnosed for this particular
dataset by validation calculations that identified AMMI-1
as the most predictively accurate memberof the AMMI
family. This diagnosis is confirmed by a much simpler
calculation: estimation of the interaction noise. Recall
that the interaction SS is 738.03, which includes 457.88
noise and 280.15 real interaction. The AMMI analysis
first three IPCAcapture 229.81, 133.99, and 94.87.
Hence, the AMMI model that comes closest to the goal
of capturing 280.15 of the interaction is AMMI-1. Recall
that the target for mega-environment analysis of this
Louisiana corn trial is 13.5% of the treatment SS not
100%. The AMMI-1model captures 12.2%, which is
close to the target. This modelfocuses on relevant geno-
type and real interaction effects while ignoring irrelevant
environment effects and muchinteraction noise.
The AMMI parameters in Table 2 can also be presented
in a biplot graph, so namedbecause it has two kinds of
markers, one for genotypes and one for environments
(Gabriel, 1971, 1981; Bradu and Gabriel, 1978). Two
different biplot axis systems are common:the abscissa
and ordinate showing main effects and IPCA1 (Zobel
et al., 1988; Gauch, 1992, p. 92) or else IPCA1 and
IPCA 2 (Kempton, 1984; Gauch, 1992, p. 94). Figure
1 showsthis first kind of biplot.
Figure 1 is extremely informative, showing both main
and interaction effects for both genotypes and environ-
ments. This biplot captures the genotype SS of 238.93
and the environment SS of 2865.17, and IPCA1 captures
229.81 of the interaction SS of 738.03, so the biplot
reveals 86.8% of the treatment SS. Note that nearly as
much variation in yield is caused by the interaction
pattern captured in IPCA 1 as by the genotype main
effect, so interaction is important for this dataset. In
other words, specific and wide adaptations are equally
important. Genotypesnear the top of Fig. 1 are mostly
long-season varieties, those in the middle medium-season
varieties, and those near the bottom short-season varie-
ties, and correspondingly the environments near the top
have longer growing seasons, so the interaction appears
to be caused by different maturity requirements (M.S.
Kang, 1996, personal communication). Incidentally, al-
though the environment main effects are irrelevant for
delineating mega-environments, they may be of interest
for other reasons (such as looking for correlations or
relationships between environment main effects and envi-
ronment IPCA1 scores). At any rate, they can also be
shown in this biplot without any confounding or extra
effort because one kind of marker is for environments
and one of the axes is for main effects.
In this graph, distances in the direction of the abscissa
showmain effect differences, whereas the ordinate shows
interaction differences. For example, consider the three
environments SJ87, BC87, and BR87. SJ87 and BC87
differ markedly in main effects (1.884 vs. -1.142) but
have similar interaction scores (0.189 and 0.136),
whereasSJ87 and BR87differ in interaction scores (0.189
vs. -0.620) but have similar main effects (1.884 and
2.130), and BC87and BR87differ in both since they
lie in diametrically opposite quadrats. Interactions are
estimated by multiplying a genotype and environment
IPCA 1 score. For example, the AMMI-1model esti-
mates the interaction for Genotype 3165 in Environment
BR87 as 1.295 (-0.620) = -0.803 Mg -t, which
approximates the observed interaction effect of -0.742.
Data in Fig. 2 show the AMMI-1nominal yields for
each of the genotypes as a function of the environment

BC88 0 3165
AX87
1.0
03147
AX88 O 1860
BC86
0.5 O CK21
O 8990 4765
BC85 SJ87
8951O,-, $J85
BC87 338~ s..J8~8172
0
3320 AX85
-0.8 -0.4 0 0.4 0.8 1.2
-0.5 4733 0
BR88~ AX86 05
EnvironmentIPCA1 Score (Mg/ha)
SJ88~18020 BR87
BR85 Fig. 2. Additive__MainEffects and Multiplicativelnteraction (AMMI)-I
-I.0 4522 O0 8150 BR86 nominal yields for a Louisiana corn trial as a function of the
O~ 1827 = ~ environmentinteraction principal componentsanalysis axis (IPCA)
4 5 6 7 8 9 I score. Thescores for the 16 environmentsare indicated by triangles
along the abscism. The four genotypes that win over someregion of
Mean(Mg/ha) this score are shownwith thick lines andare identified individually,
Fig. 1. Additive Main Effects and Multiplicative Interaction (AMMI)-I whereas the remaining 12 genotypes that never win are shownwith
biplnt for a Louisiana corn trial, showingboth mainandinteraction thin lines. Hypothetical Genotype A or any line above it would be
effects for both genotypes and environments. Opencircles denote a universal winner. Hypothetical GenotypeB is the universal winner
genotypes; closed circles denote environments(IPCA= interaction with a genotype IPCAI score of 0 andthe smallest possible genotype
principal componentsanalysis axis). mean.
 GAUCH& ZOBEL: IDENTIFYING MEGA-ENVIRONMENTS 317

IPCA1 score, where the nominal yield is defined as relatively well or relatively poorly." By providing an
the yield from the AMMImodel equation without the effective summaryor overview of yield trial results,
environment deviation IBe. Hence, nominal yields repre- standard biplots can provide helpful insight for delineat-
sent an average environment (with IBe = 0), and they ing mega-environments. However, such graphs fail to
can be adjusted to give actual AMMI yield estimates for make evident what wins where, which limits their value
an individual environment by adding that environments for analyzing mega-environments. Accordingly, several
deviation. However, this adjustment merely shifts all new kinds of AMMIgraphs are developed here that do
yields within a given environment up or downequally, so address mega-environment issues directly. The key is
the genotype rankings and overall picture are unchanged. combiningfeatures of Fig. 1 and 2 and decreasing atten-
Again, environment deviations are irrelevant for present tion on losing genotypes while focusing attention on
concerns because they do not affect genotype rankings winning genotypes.
within each environment. Genotypes affect mega-envi- Figure 2 showed that the winning genotype in each
ronments through their main and interaction effects, but environment, e, is determinable for the AMMI-1model
environments affect mega-environments through their from a single parameter, the environment IPCA1 score
interaction effects only. In Fig. 2, the four genotypes (~,0.56e) shownon the abscissa, and that in this particular
that win in portions of the IPCA1 space are identified case, there are four winners. That information is depicted
individually, whereas the other 12 genotypes that never differently in Fig. 3, where this environment score is
win are represented by thin lines. Scores for the 16 nowon the ordinate instead of the abscissa.
environments are shown by the 16 triangles along the Figure 3 shows the four mega-environmentsidentified
abscissa (which are the same scores listed previously in by AMMI.Note for example that in Fig. 2, the lines
Table 2 and shown in the ordinate of Fig. 1). Note for for winning Genotypes 3147 and 3165 cross at an envi-
example, that Genotype 3165 has the highest AMMI-1 ronment score of 0.376. Accordingly, there is a line
estimated yield for four environments, namely those at across Fig. 3 at this score of 0.376, and Genotype3165
environment IPCA1 scores of 0.593, 0.723, 1.181, and wins in the four environments that happen to have scores
1.338 (which can be identified more specifically in Fig. above this value. Likewise, from 0.376 to 0.224, Geno-
1 as BC86, AX88, AX87, and BC88). type 3147 has 1 win; from 0.244 to -0.612, Genotype
The mathematical relationship between Fig. 1 and 2 8172 has 5 wins; and below -0.612, Genotype 1827
should be understood clearly. Correspondingto the above has 6 wins. On the right side of Fig. 3, nominal yields
AMMI model equation, Table 2 gives the AMMI-1model are indicated, copied from the previous figures ordinate.
parameters for this Louisiana corn trial, with two values For example, from Fig. 2, at an environment IPCA 1
for each genotype (and each environment). For example, score of 1.0, the winning Genotype 3165 has a nominal
Genotype 3165 has a mean of 6.693 + 0.472 = 7.166 yield of 8.461 Mgha-~, as is also indicated in Fig. 3.
Mgha -1 5.
and an IPCA 1 score of 1.295 (Mg ha-I) Nominal yields form a response surface in Fig. 3
Figure 1 uses these two values directly in its abscissa
and ordinate to place its marker for Genotype3165. But
Fig. 2 uses this same information differently, depicting 1.4
[] BC88 8.899
each genotypeby a straight line (instead of a point). For 1.2 3165 Wins 8.8
O AX87
example, Genotype 3165 is depicted by a line with the 8.6
1.o 8.4
equation y = 7.166 + 1.295x, where y is the nominal
yield and x is the environment IPCAscore, so this line O AX88 8.2
uses these same two parameters. The left-most x of 0.8 ] BC86 8.o
-0.947 implies a yield of 5.940 Mgha-l, whereas the
right-most score of 1.338 implies 8.899, and these two 7.653
3147 WinsOBC85 7.534
0.2
coordinate pairs give two points that define the line for 8172 Wins [3 BC87 SJ85 SJ87
Genotype 3165, as shown in Fig. 2. More generally, 13.0 SJ86
each point in Fig. 1 corresponds to a line in Fig. 2. -13.2 AX85 O
Movinga point left or right in Fig. 1 equates to moving
a line downor up in Fig. 2, and moving a point up or -0.4
downin Fig. 1 equates to tilting a lines slope upward 7.531
7.6
or downwardin Fig. 2. -13.8 1827 Wins SJ88 BR88 BR87 (3
BR85 AX8,~ 7.8
Whena biplot of the AMMI-1model, as in Fig. 1, 7.912
-1.13
fails to capture all of the real structure in the interaction, 4 5 6 7 8 9
another kind of biplot using IPCA1 and IPCA2 as its Environment
Mean(Mg/ha)
two axes can showmore of the interaction, as in Kempton
Fig. 3. ~dditive MainEffects and Multiplicative Interaction (AMMI)
(1984) and Gauch (1992, p. 94, 222). But Westcott mega-environmentanalysis for a Louisiana corn trial. There are
(1987) remarks that Kemptonsbiplot shows interactions, four genotypes that win in various regions of the AMMI-1 models
not actual performance meaningyield, in line with Kemp- parameter space, such as Genotype 8172 that wins in the five
tons ownremarkthat his biplot "indicates whichvarieties environments with an interaction principal componentsanalysis
axis (IPCA) score between 0.224 and - 0. 612. The fo ur Louisiana
have similar and which have very different patterns of locations are shownwith different symbolsto facilitate comprehend-
response over the environments"but "it does not identify ing the results for each location across the four years. Onthe right
the particular environments in which a variety performs is the nominalyield, which correspondsto the values in Fig. 2.
318 CROPSCIENCE, VOL. 37, MARCH-APRIL
composedof a convex set of planes, with yields increas-
ing both above and below the lowest yield of 7.531 Mg
ha -] located at an IPCAscore of -0.612. The yield
response surface for Genotype8172 is nearly flat (as is
also evident in Fig. 2), whereas 1827 and 3165 are much
steeper. The lowest yield for a winning genotype always
occurs for that genotype with the highest main effect
and hence an automatic win at an environment IPCA1
score of zero, which is Genotype 8172 in this case.
Higher yields are achieved by positive interactions that
more than compensate for comparatively small reduc-
tions in genotype main effects. Hence, Fig. 3 shows why
specific adaptations (interaction effects) are important
well as broad adaptations (main effects) in the pursuit
of high yields.
Although Fig. 3 has the advantage, unlike standard
biplots, of revealing genotypes winning domains, it has
the disadvantage of providing information on main and
interaction effects in different scales and units, namely
yield and the square root of yield. Consequently, the
relative importance of main and interaction effects is
not apparent. Accordingly, Fig. 4 presents the AMMI
mega-environmentresults in still another format.
The abscissa of Fig. 4 shows the environment main
effect plus the genotype main effect for the particular
genotype that wins in each environment (as identified in
the right of Fig. 4, as well as earlier in Fig. 3). The
ordinate shows the interaction effect, again for each
environment using its winning genotype. For example,
Environment BC88 has winning Genotype 3165 and
hence an AMMI-1estimated interaction that equals the
product of this environment and genotypes IPCA 1
scores, namely 1.338 1.295 = 1.733 Mgha -]. Main
effects and interaction effects are shownon the abscissa
and ordinate to the same scale, to make evident their
relative importance. The AMMI-1yield estimate for
each environment is simply the sum of its two values
shown here. For example, for its winning Genotype
3165, Environment BC88has main effects of 6.693 +
0.472 - 1.184 = 5.981 and an interaction effect of
1.295 1.338 = 1.733, resulting in a yield of 5.981
+ 1.733 = 7.714 Mgha-]. This ordinate preserves the
convex topology of the ordinate of Fig. 3 (which has
lowest nominal yield at an environment IPCA1 score
\
\ \ rl BC88 \ T
~ ~BC85[] BC87. J87 .--
/ and empirical reasons for giving priority to the AMMI-1
/ SJ88i/ I BR85 BR8710
4 5 6 7 8 9 10
Geno~peand EnvironmentMeans(Mg~a)
Fig. 4. Maineffectsand interaction effects for a Louisiana corn trial,
showing yields for each environmentwhenplanted to its genotype
winner. Isoyield lines forma herringbonepattern. Mainand interac-
tion effects are shownto the same scale to make their relative
importance evident.
1997
of -0.612 and increasing yields in either direction away.
from this value). Consequently, the thin isoyield lines
indicating equal yields form a herringbone pattern. For
example, environments BC88, SJ86, and BR85 all have
nearly equal yields of ~ 8 Mgha- i, but they achieve this
yield in different ways: SJ86 has almost no interaction,
whereas BC88and BR85have sizable and opposite inter-
actions, which is why these three environments have
three different genotype winners and hence fall into three
different mega-environments.
A sequence of mega-environment assignments with
one to many mega-environments can be determined on
the basis of the above AMMI results, while calculating
the implications for yield. Wereeach of the 16 environ-
ments planted to its own winner according to the raw
data, there would need to be eight different genotypes
and hence eight mega-environments, and this highest
possible average yield would be 8.416 Mg ha-I. But
were each environment planted instead to its AMMI-1
winner indicated in the above figures, then only four
genotypes or mega-environments are required, and the
average yield based on raw means would be 7.585 Mg
ha- or a 9.9% yield drop. Were the entire region
declared a single mega-environment and planted to the
overall winner 8172, the average yield based on raw
means would be 7.533 Mgha-l, or a 10.5% yield drop.
So, according to the raw yield data, one, four, and
eight mega-environments allow average yields of 7.533,
7.585, and 8.416 Mg ha -~. In particular, based on
AMMI-F, switching from the eight AMMI-Fwinners
to the four AMMI- 1 winners causes an estimated average
yield loss from 8.416 to 7.585 Mg ha-l or 12.4% of
the grand mean.
However, were AMMI- 1 yield estimates trusted across
AMMI-Fraw data because the AMMI-1model is more
predictively accurate than its data, then switching to
AMMI-1winners causes an estimated yield gain from
7.527 to 7.806 Mg ha -] or 4.2% of the grand mean.
AMMI-Fand AMMI-1pick the same winner in only 2
of the 16 environments. Also, according to AMMI-1
estimates, were the one Genotype 8172 planted every-
where, the average yield would drop substantially to
7.533 Mg ha -]. AMMI-0 and AMMI-1 pick the same
winner in only five environments. So, according to
AMMI-1yield estimates, one, four, and eight mega-
environments allow average yields of 7.533, 7.806, and
7.527 Mgha-]. AMMI-0is worse than AMMI-1because
it underfits real interaction patterns in the data, whereas
AMMI-Fis worse than AMMI-1because it overfits
spurious noise in the data.
Because AMMI-1estimates are more predictively ac-
curate than AMMI-F estimates, there are strong rational
assessment. But even without accepting that argument,
the AMMI-1 analysis with four mega-environments could
be regarded as a useful simplification if practical consid-
erations impose a limit of about four mega-environments
(rather than allowing the rather numerous eight mega-
environments of the AMMI-Fresults).
Figure 5 reviews the AMMI-1yield estimates and
AMMI-Fraw data for average yields achievable with
 GAUCH& ZOBEL: IDENTIFYING MEGA-ENVIRONMENTS
one mega-environmentusing the AMMI-0 winner 8172,
four mega-environments using the AMMI-1 winners,
and eight mega-environmentsusing the AMMI-F win-
ners. TheAMMI- 1 curve reflects a predictive perspective
in which the data are deemeda sample from a larger
populationof inference, with optimalyield at rather few
mega-environments. Theshape of this curve is a typical
Ockhamshill, with optimal performancefor a fairly
parsimoniousmodeland penalty to the left for underfit-
ting real patterns and penaltyto the right for overfitting
spurious noise (Jefferys and Berger, 1992;Gauch,1993).
But the AMMI-F curve reflects a postdictive perspective
in whichthe data are deemedthe entire population of
inference, so increasingly complexmodelsautomatically
seem better. The AMMI-1 and AMMI-Fyields are identi-
cal for one mega-environmentplanted to the overall
winner8172because discarding part of the interaction
by the AMMI-1model has no effect on an average
computedacross all environments.Note that predictive
assessmentbased on AMMI-1 is qualitatively different
from postdictive assessmentbased on AMMI-F raw data.
For the postdictive perspective, more mega-environ-
mentscontinueto increase yields until there are enough
genotypesfor every environmentto be assigned its win-
ner, which number eight in this case. But from the
predictive perspective, both too few and too manymega-
environments result in a yield loss, and four mega-
environmentsare ideal in this case.
Figure 5 emphasizesthat different membersof the
AMMI family differ in their predictive accuracy and in
their value for delineating mega-environments. For this
particular yield trial, AMMI-1 happensto be best, but
for another dataset, the optimal modelmightbe AMMI-2
or some other member.If the optimal AMMI model for
a given dataset has not been diagnosed, then selection
of somearbitrary memberof the AMMI family is likely
to give a suboptimal analysis, which weakens AMMI
results and confuses mega-environment choices. For ex-
ample,if the interaction is essentially all noise, thenthe
PCAportion of AMMI will find a large number of
small eigenvalues,exactly as describedby Mandel(1969,
1995). Thenit will look like AMMI fails to capture the
interaction effectively, since numerousaxes are required
to get most of the interaction SS, such as the AMMI-5
modelto capturejust 60 %of the interaction. This verdict
leads to a complex model with numerous mega-
environments.But the truth of the matter, whichwould
be revealed by not neglecting modeldiagnosis, is the
exact opposite. Theinteraction is buried in the noise,
so only maineffects are statistically significant. Conse-
quently, only one winner is detected, the main-effect
winner, whichleads to no mega-environment subdivision
whatsoever.If no distinction is madebetweenreal inter-
actions and spurious noise, mega-environment analysis
becomesa disaster.
It mayseemsurprising that these two models, AMMI-1
and AMMI-F, differ by such a large amount, 0.889 Mg
ha- ~, at eight mega-environments. Butthis discrepancyis
easily explained, showingthat AMMI-Fis unrealistically
high, rather than AMMI-1unduly low. The AMMI-F
yield estimates are simplythe averages across the four
319
8.2
AM
8.0
78 "~
1 2 3 4 5 6 7 8
Number
of Mega-environments
Fig. 5. Averageyield for a Louisian corn trial using one, four, or
eight mega-environmentsand estimating yields by the Additive
_MainEffects and Multiplicative I_nteraction (AMMI)-Imodel(solid
line) or the AMMI-F averages across replications (dashed line).
The AMMI-Imodel merits more serious consideration because it
is morepredictively accurate than AMMI-F.Unfortunately, the
AMMI-Fraw data are doubly confusing because they both overesti-
mate the yield with eight mega-environmentsand underestimate
the yield with four mega-environments. Noise encourages a choice
of too many mega-environments, which both increases costs and
decreases yields.
replications, whichhave, for this Louisianacorn trial,
a standard error of 0.713 Mgha-l. Theseaverages are,
fromstatistical theory, unbiasedestimators of the true
means.However,after ranking the 16 genotypesyields
within each environment,the highest yields are no longer
unbiasedbecausethey tend to havepositive errors, and
likewise the lowest yields tend to have negative errors
(Gauch and Zobel, 1989; Gauch, 1992, p. 171-177).
The AMMI-F value is larger than AMMI-1 by 0.889/
0.713 = 1.247 standard errors. Nowthe first order
statistic for 16 drawsfromthe standardnormaldistribu-
tion is 1.766, so wereall 16 genotypesessentially equal,
the upwardbias in the winners wouldapproximate1.766
x 0.713 = 1.259 Mgha-~. However,the 16 genotypes
are not all nearly equal, and if the top group of real
contenderswere morelike about six entries, that first
order statistic of 1.267 wouldvery nearly explain the
observed discrepancy. So, the AMMI-Fexcessively large
yield at eight mega-environments is easily attributed to
spurious noise, to positive samplingerrors accumulated
after rankingthe data. Butif the effects of experimental
noise are not understoodproperly, researchers will be
tempted to identify too manymega-environments,which
will actually reduce averageyields, while also imposing
unnecessary burdens on plant breeders and seed pro-
ducers.
But the most interesting feature of Fig. 5 is that
AMMI-1beats AMMI-Fat the optimal four mega-
environments, whichcan be explained as follows. When
AMMI-F yield data are used to calculate the average
yield but each environmentis assigned its AMMI-1 win-
ner rather than its ownAMMI-F winner, only 2 of the
16 environments receive that genotypewhichis also their
AMMI-F winner. Forced away from its ownwinners 14
out of 16 times, the AMMI-F average yield with AMMI-
320 CROP
SCIENCE,
VOL.37, MARCH-APRIL
ls four mega-environments rises only a slight amount
above AMMI-/~s average with only one mega-envi-
ronment. But AMMI-1yield estimates used with its
own winners exploit sizable interactions to achieve a
considerable yield increase when changing from one to
four mega-environments. Indeed, the first IPCAaxis of
AMMIcaptures a SS of 229.81, which is comparable
in magnitude to the genotype main effect of 238.93, so
interactions are quite important for this dataset. Equally
importantly, the AMMI- 1 model leaves behind a residual
SS of 508.22, which is the SS of the differences between
AMMI-1and AMMI-Fyield estimates, and this dis-
carded residual contains mostly noise. So, the differences
between these two models have somewhat more than
twice as muchvariation as do the genotype main effects.
That is, the interaction noise is about twice as large
as the genotype main effects, and this noise is largely
disregarded by AMMI-1,but unfortunately, it is taken
seriously by AMMI-F.Because these two models are
so different, AMMI-ls average is low using AMMI-Fs
eight winners, and AMMI-Fsaverage is low using
AMMI-lsfour winners. Anyway,the net effect of these
factors is that at four mega-environments,AMMI-1 beats
-I.
AMMI-Fby a substantial 0.221 Mg ha
The lesson from Fig. 5 is that from the perspective
of identifying mega-environments,experimental noise is
doubly confusing. Noise causes an overestimate of the
yield increase for an excessive number of mega-
environments and an underestimate of the benefit for a
small and optimal number of mega-environments. Conse-
quently, effective mega-environmentidentification re-
quires aggressive reduction of experimental errors, and a
multivariate model such as AMMI can help considerably.
Furthermore, although this lesson is illustrated here
with a particular Louisiana corn trial, it seems relevant
for manyregional yield trials. This corn trials standard
error is 0.713 and its grand mean is 6.693 Mgha-~, so
this experiment has a middling coefficient of variation
of 10.7%. Hence, the substantial difference in mega-
environment conclusions from choosing the AMMI-1or
AMMI-Fperspective is not due to some enormous or
atypical noise level. Furthermore, manyregional trials
have more than 16 genotypes, making order statistics
problems larger than those encountered here. So, the
general pattern exhibited in Fig. 5 can usually be expected
whenevera trial has substantial interactions and typical
noise. It should not seem too surprising that when at-
tempting to identify rational mega-environments, it
makes a difference whether or not a statistical analysis
is used to filter out someof the noise in the data.
Figure 6 shows the difference between AMMI-F data
and AMMI-1estimates for a typical environment, SJ86.
In this example, Genotype 4673 that ranks first in the
raw data ranks fifth in the AMMI-1model, whereas
Genotype 8172 that ranks first in the AMMI-1model
ranks third in the raw data. Especially because Environ-
ment SJ86 has almost no interaction (Table 2 and Fig.
1), there is a strong case for declaring its winner to be
the main effect winner, Genotype 8172. Furthermore,
that 8172ranks third in the raw data, not first, is easily
explained by noise, given that the experiments coefficient
1997
of variation is 10.7 %. Accordingto the morepredictively
accurate AMMI-1models yield estimates, planting the
AMMI-1winner 8172 instead of the AMMI-Fwinner
4673 results in a yield increasefrom 7.513 to 8.075 Mg
ha -1. Crossa et al. (1991) describe AMMI adjustments
for a bread wheat (Triticutn aestivum L.) trial.
This experiments 16 environments have a factorial
design with four locations and four years, so it is interest-
ing to compile the results for each location across the
years, using the AMMI-1 yield estimates. From Fig. 3,
location BRis always in the winning domainof Genotype
1827. Its average yield in BRis 8.473 Mgha-I. Location
BRis the most predictable, so variety recommendation
is easy. Location SJ gives three wins to 8172 and one
to 1827, with an average winners yield of 7.924 Mg
ha-1. Here the best recommendation, especially in a
predictive context, is to always plant Genotype 8172,
reducing the average yield only imperceptibly to 7.899
Mgha-1. Location BC gives two wins to 3165 and one
each to 3147 and 8172, with an average of 6.147 Mg
ha-~. At this location, these three genotypes average
6.089, 5.979, and 5.693 Mgha-~, respectively. Because
3165 has the most wins and the highest average yield
at BC,the best strategy, especially in a predictive contex.t,
wouldbe to always plant 3165. Finally, the least predict-
able and most problematic site is AX, with winners 3165
(twice), 8172, and 1827 spanning the entire range
interaction patterns, and an average winners yield of
8.682 Mgha -~. Location AXhas large interactions in
hindsight but unpredictable ones in foresight, so the best
predictive strategy is not to try to exploit interactions
but rather to plant the AMMI-0 main effect winner 8172,
resulting in 8.248 Mgha -I. The general outcome is
that predictable interactions, typically associated with
locations, make mega-environments more numerous and
useful, whereas unpredictable interactions, typically as-
sociated with years, make mega-environments less nu-
merous and useful.
In summary, the recommendations are 1827 for BR,
AMMI-F AMMI-1
8172
8.0 ~ 3147
Yield 1860
(ig/ha) 3165
7.5 4673
CK21
1827
(Mean)
81504765
7.0 ~ 4733
33203389
1802
//\ )< 4522
8951
6.5
8990
6.0
Fig. 6. AdditiveMainEffects andMultiplicativel_nteraction
(AMMI)-Fyield dataandAMMI-1 estimatesfor environmentSJ86
of a Louisianacorntrial. Genotype
4673ranksfirst in the raw
databut ranksfifth in the morepredictivelyaccurateAMMI-I
model.AMMI-1 identifies Genotype
8172as the best genotypein
this environment.
 GAUCH& ZOBEL: IDENTIFYING MEGA-ENVIRONMENTS
8172 for SJ and AX, and 3165 for BC, giving an overall
average of 7.677 Mgha-~. In hindsight, planting the
AMMI-1 winner every time results in a somewhat higher
average yield of 7.806 Mgha- 1. But hindsight is clearer
than foresight. The present recommendations capture
98.3% of the potential yield, which seems to be about
the limit of predictive capability, with muchof the penalty
incurred at AXand none at BR. Incidentally, were only
two mega-environments feasible, the recommendation
would be to plant 1827 in BR and 8172 elsewhere,
capturing 98.1%of the potential yield.
Figure 7 uses AMMIparameters to show which geno-
types win over others, as well as to locate the parameter
space that identifies potential new winners. Each of the
16 actual genotypes are located in this graph by their
two AMMI-1parameters, the genotype mean (kt + ~tg)
and the genotype IPCA1 score (~,53,g), as listed
Table 2 and shown in Fig. 1. Also shown are the two
hypothetical genotypes in Fig. 2, with GenotypeA having
a mean of 8.321 and score of 0.432 and Genotype B a
mean of 8.899 and score of 0.
An X is drawn at the markers for the four winning
genotypes but, to avoid clutter, is not indicated for the
other 12 genotypes that never win. The purpose of these
X is to show which genotypes win over which others,
as explained momentarily. Every X has exactly the same
shape, that is, all upwardlines (/) are at the sameslope
and all downwardlines (\) are at the same slope. Casual
inspection might suggest that these upward and down-
ward slopes are of equal magnitude (apart from opposite
signs), but actually the upwardslope is steeper than the
downwardslope, and in general, these two slopes are
not equal.
These two .slopes are calculated as follows. The upward
. C, K2~ / ~ /~. / Universal
809908~1/0034~ ~ ~Winners
~ / 4673/" / ~ ~ ~
oe- -1 ~ential ~~
~)
i ~ ~ i ~ i i i ~ and understood by reference to Fig. 2, which Showsthat
6 7 8 9
Geno~pe Mean (Mg~a)
Fi~. ~. ~dditi~e~alnEff~tsand~ultiplicati~e~nteraction (A~[-])
modelfor the 16 ~enoty~sof a Louisianacorn trial. Four of these
~eeoty~sare the ~nnersthat shadethe other 12 ~enot~s to the
left. Parameter
spaceto the right of the ja~edthick line identi~es
~teetial newwinne~that wouldshadeat least oneof the current
winners,andparameterspacete the H~ht of the s~ondthick line
identifies ~tential ~iver~! winners that would out~rform all
current ~enoty~s. ~noty~s A and B are hy~tbetical universal
winners(IPCA = interaction principal com~nents analysis axis).
321
slope equals the negative reciprocal of the largest negative
environment IPCA1 score. From Table 2, this is -0.947
for Environment BR86, so the upward slope is -1/
-0.947 = 1.056. Similarly, the downwardslope equals
the negative reciprocal of the largest positive environ-
ment IPCA1 score. This is for Environment BC88, so
the downward slope is -1/l.338 = -0.747. The line
with an upward slope (of 1.056) represents the family
of all lines going through the point at the left extreme
of a genotypes line in Fig. 2, whereas the line with a
downwardslope (of -0.747) represents the family
all lines going through the point at the right extreme.
Hypothetical GenotypeA is constructed as a line going
through two points, the highest yields at the left and
right extremes of Fig. 2. The point on the left has an
abscissa of -0.947 and ordinate of 7.912, and the point
on the right has coordinates of 1.338 and 8.899. Solving
the equation of a straight line given two points, calling
the environment score x and the nominal yield y, the
equation for GenotypeA is y = 8.321 + 0.432x. Accord-
ingly, this genotype appears in Fig. 7 at exactly these
coordinates, having a nominal yield of 8.321 and score
of 0.432. Incidentally, a line from the winning genotype
with the largest negative IPCA1 score, namely Genotype
1827, connecting to Genotype A defines the upward
slope. Similarly, a line from Genotype 3165 with the
largest positive IPCA1 score connecting to Genotype
A defines the downwardslope. Because Fig. 7 has the
same axes as Fig. 1, every point in Fig. 7 corresponds
to a line in Fig. 2, as explained earlier for Fig. 1 and
2. The points on the line through 3165, A, and B in
Fig. 7 correspondto the family of all lines that go through
the right-most top yield in Fig. 2 (at coordinates x
1.338 and y = 8.899). Likewise, points on the line
through 1827 and A in Fig. 7 correspond to the family
of all lines that go through the left-most top yield in
Fig. 2 (at coordinates x = - 0.947 and y = 7.912). These
two lines in Fig. 7 intersect at one point, hypothetical
Genotype A, which correspondingly is the unique line
in Fig. 2 that goes through both the left-most and right-
most top yields. GenotypeA is also uniquely the universal
winner with the lowest genotype main effect.
To begin the interpretation of this graph, focus on
just one genotype, namely 8172 near the middle. The
X drawn at this genotypes marker divides the parameter
space into four sectors. There are three interpretive
principles. First, the sector to the left is the shadow
cast by 8172 over the other genotypes that it always
outperforms, which include nine genotypes in this case.
These nine complete wins by 8172 can be confirmed
the line for 8172 is always above nine other lines for
the losing genotypes, such as the unlabeled lowest line
with an upwardslope that is actually for Genotype8990.
Second, in Fig. 7, the sectors above and below 8172s
marker identify mixed wins where 8172 sometimes wins
and sometimes loses, which involve six genotypes. For
example, 3147 is in the sector above 8172, and 1827 is
in the sector below 8172. Accordingly, Fig. 2 shows
that the lines for 3147and 1827 cross the line for 8172,
so 8172 sometimes wins and sometimes loses against
322 CROP SCIENCE, VOL. 37,
these competitors. Third and finally, the sector to the
right of 8172s marker identifies genotypes that always
win over 8172. In this case, there are no such actual
genotypes, but there are two hypothetical genotypes, A
and B. Again, it is evident from Fig. 2 that Genotypes
A and B always win over 8172.
This X marking four sectors for Genotype 8172 is
also shownin Fig. 7 for the three other winners, namely
3165, 3147, and 1827. Note that the other 12 gentoypes,
which never win, are always in the shadow of one or
more of the four winners.
Havingexplained the technicalities of Fig. 7, its most
interesting features can nowbe explored. The thick jag-
ged line down the middle of this graph separates the
parameter space to the left that is dominatedby the cur-
rent four winners from the parameter space to the right
that identifies potential newwinners. Note that the other
12 genotypes, which never win, are located to the left
of this thick line, meaning that they are always in the
shadowof one or more of the four winners. On the other
hand, any new genotype with parameters to the right of
this boundary will shadow and outperform at least one
of the current winners. Furthermore, any new genotype
to the right of the second thick line is a universal winner
that will outperform all current genotypes. Note that the
parameter space for universal winners is defined by
extending the line from 3165 to A and the line from
1827 to A because these two current winners have the
largest positive and negative genotype IPCA1 scores
and A is that potential universal winner with the smallest
mean yield.
This figure stimulates insight on possible breeding
strategies. It shows the various combinations of main
and interaction effects that could produce new winners.
It also shows just what sort of new genotype could
outperform two or more current winners, thereby reduc-
ing the number of mega-environments needed to optimize
yield everywhere. For example, consider the point where
two thin lines cross between the labels for 3165 and
3147. A new genotype located near this point would
combine a main effect like 8172 with an interaction
pattern intermediate to 3165 and 3147, and by outper-
forming both 3165 and 3147 (as well as the additional
seven genotypes already shadowedby these two current
winners), it would merge two old mega-environments
into one new mega-environment. Andif another genotype
could shadow 8172 and 1827, then the original entire
yield trial with four mega-environmentswould need only
two mega-environments.
The tradeoff between breeding for main effects or for
interaction effects can be illustrated with hypothetical
Genotypes A and B. Genotype B is most stable, as
indicated by its fiat line in Fig. 2 and correspondingly
its score of 0 in Fig. 7. The current main-effect or
AMMI-0winner, Genotype 8172, is also very stable,
with a genotype IPCA1 score of just 0.003. The differ-
ence in mean yields between B and 8172 is 8.899 -
7.533 = 1.366 Mg ha-~. So, if a new genotype could
be developed with the same stability as 8172 but a main
effect greater by 1.366, it wouldwin everywhere, thereby
eliminating the need to subdivide the region into several
MARCH-APRIL 1997
mega-environments. Assume for sake of argument that
breeders expect to increase main effects by = 1% per
year, which has been typical for many major crops
(Evans, 1980; Russell, 1991). Then 1% of 7.533
0.07533, so the needed increase will take = 1.366/
0.07533 or 18 yr.
But now comparethe idiotype represented by hypothet-
ical GenotypeA. This genotype is not stable but rather
has a substantial interaction. FromFig. 1, this genotypes
interaction is positive for environments like BC88and
AX87 and negative for environments like BR86 and
BR85, which was interpreted earlier as meaning a posi-
tive interaction for yield in relatively long-season envi-
ronments. Although Genotype A is less stable than B,
Genotype A has the merit of requiring considerably less
mean yield to become a universal winner, only 8.321
instead of 8.899 Mgha- t. Assumethat the average main
effect for parental genotypes whose progeny could have
Genotype As target interaction score of ~-0.432 equals
the average for nearby Genotypes 8172 and 3147, namely
7.446. Then gaining 1%of 7.446 surpasses the required
8.321 in 12 yr. So, it is plausible that the target idiotype
A could be reached several years faster than B. The
general lesson here is that breeders should think carefully
about what they really want: stability or superiority.
In essence, to optimize yield despite genotype-envi-
ronment interactions, breeders have two options: to pur-
sue wide adaptation and thereby minimize mega-en-
vironment subdivisions or to exploit narrow adaptations
in several different mega-environments (Brennan and
Byth, 1979). A graph such as Fig. 7 could help breeders
comparethe feasibilities of these two options for a given
breeding program.
By knowing the landscape of the AMMIparameter
space, with the locations of current winners and the
regions for potential and even universal new winners,
desirable idiotypes can be defined. And by knowing
roughly how fast breeders can change main effects or
interaction effects, one can predict roughly howlong it
would take to reach various places in the parameter
space representing potential new winner~. In addition to
providing insight about possible idiotypes, this graph
also gives clear implications for mega-environments.For
example, in comparing the value of various possible
idiotypes, special priority can be given to those that
would shadow several current winners instead of just
one, thereby reducing the number of mega-environments.
Fewer mega-environments simplify and economize a
breeders testing system, concentrate breeding efforts
on fewer needed cultivar types, and streamline seed
production and distribution. So, a graph like Fig. 7
is helpful both for understanding present relationships
amongactual genotypes and future implications of poten-
tial idiotypes. It can help breeders understand where
they have come and where they might go.
Another application of a graph like Fig. 7 could be to
compare the market strengths of various seed companies.
Envision a collection of genotypes not all from a single
breeding program but rather including cultivars sold by
various companies. By identifying the commercial source
of each genotype in this graph, it would be clear which
 GAUCH& ZOBEL: IDENTIFYING MEGA-ENVIRONMENTS
companieshave special strengths or weaknessesin vari-
ous portions of the AMMI parameterspace, in regards to
both current actual winnersand future potential winners.
Another advantage of Fig. 7, which is evident from
the logic of the calculation usedto place the Xover each
winners marker, is that such a graph can be adjusted
or tailored for a different range of target environments.
For example, the present graph showsresults for this
entire Louisianacorn trial, with the interaction extremes
represented by EnvironmentsBR86and BC88,leading
to slopes of 1.056 and -0.747. If instead of the entire
trial, interest focuses on yields across years for just
LocationSJ, then the newinteraction extremesare less
diverse, SJ88 at -0.699 and SJ87 at 0.189, leading to
slopes of 1.431 and -5.291. Consequently, genotypes
cast larger shadowsby their broadersector to the left,
whichalso leads to fewer winnersor mega-environments.
This makessense..From Fig. 2, it is clear that this
smaller range in environmentscores implies only two
winners, 1827and 8172. Moregenerally, rather homoge-
neous environments makegenotype main effects rela-
tively moreimportant than genotype-environment inter-
action effects. Onthe other hand, if interest focuses on
even morediverse environmentsbecausea yield trial or
growingregion is expanded,wider interaction extremes
lead to the opposite response of increasing the sectors
above and belowthe genotypemarkers. This also makes
sense. FromFig. 2, it is clear that were this graph
extendedto the left and right, moreand moreof these
nonparallel lines wouldeventually cross each other, so
there is an increase of mixedwins. So, graphs like Fig.
7 can be adjusted to suggestresults for various subsets
or extensions of environmentsof special interest in the
past or the future.
Theabovebrief remarkssketch only a few of the sorts
of questions that can be framedand addressedby graphs
like Fig. 7 and the earlier graphs. These are highly
informative graphs that can promote insight on past
results and future options. There are other useful kinds
of AMMI graphs not included here. Gauch(1992, p. 222)
extends the AMMI-1winning domains to the AMMI-2
model,in whichdomainsform irregular polygonsinstead
of horizontal bands. Gauch(1992, p. 218) depicts loca-
tions across years as regions, rather than points, in an
AMMI biplot by delineating each locations points across
years. Atypical outcomeis that somelocations differ
from year to year mostlyin maineffects, others mostly
in interactioneffects, others in both, andstill othersare
rather stable. Fig. 2, 3, 4, and7 couldreverse the roles
of genotypes and environmentsto create new kinds of
graphs addressing someother questions related to mega-
environments.
It must be stressed, however, that each and every
analysis of interaction, including AMMI, pertains di-
rectly to that particular collection of genotypesandenvi-
ronmentsin the dataset. Introduction of substantially
different genotypes or environmentsin the future, or
deletion of distinctive genotypesor environments already
in the dataset, can changeresults substantially. It is a
statistical maxim that extrapolationis risky, althoughit
is also an empirical result that both main effects and
323
interaction parameters exhibit some measure of ro-
bustness across different locations, years, cultivars,
crosses, and even crops. So, we wouldcaution against
overinterpreting AMMI graphs whenmakingdecisions
about cultivar recommendations or breeding strategies.
For example,broad features deserve moreconsideration
than fine details. Also, current results can be taken
more seriously whenthe genotypes are experiencing
incremental improvementsrather than drastic changes
from newidiotypes, management procedures, or disease
pressures. Nevertheless,wewouldalso insist that various
kinds of AMMI graphs, as illustrated above, can often
supplementthe understanding and insight that agrono-
mists and breeders have already developed over the
years throughextensivefield experimentation,statistical
analysis, and experience.
DISCUSSION
Subdividing a growing region into several mega-
environmentsbecomesa daunting prospect if the recom-
mendednumberof mega-environmentsis large. By con-
trast, subdivision of a crops growingregion is most
inviting whenmerely a few mcga-cnvironmentssuffice
to capturesubstantialyield increases,as for this Louisiana
corn trial. Fortunately,results for several diverseyield
trials indicate that often remarkablyfew mcga-cnvi-
ronmentsare sufficient (Annicchiarico,1992; Annicchi-
arico and Perenzin, 1994;DcLacyct al., 1994;Annicchi-
arico et al., 1995;Abdallaet al., 1996).
Twofactors largely explain this nice outcome.First,
substantial genotype-year (and/or genotype-location-
year) interactions relative to genotype-locationinterac-
tions reducethe portion of the interaction that has pre-
dictive value for future years. Whena locations points
are spread widely by interaction differences (like AX
but not BRin Fig. 3), less advantage can be taken
of specific adaptations, whichresults in fewer usable
mega-environments. Second,the roster of winnersoften
includes somegenotypes with very small domains, so
the adjacent major winneror winnerscan be substituted
with negligible yield loss to retain just a few big mega-
environments.
However, experience with AMMI mega-environment
analysis is limited, so it is possiblethat there will also
be somesurprising cases requiring morenumerousmega-
environmentsthan expected.To introduce a newstatisti-
cal methodology for exploiting interactions, this study
has used a small, convenient,previouslypublisheddata-
set. But future studies are planned analyzing several
large yield trials.
As agronomistsand breeders consider various propos-
als for exploiting interactions, includingthe statistical
methodologyintroduced here, the maincomparisonmust
be with prevailing current practices, described without
illusions. Simmonds (1991)gives a fair appraisal, saying
that there is "general awareness"of the challengesposed
by genotype-environment interactions but that "little is
donein practice beyondtesting potential newcultivars
in diverse environmentsin the hope of revealing wide
adaptation (i.e., stability) of performanceas well
324 CROP SCIENCE, VOL. 37,
good mean performance." More specifically, breeders
regularly select in high-yielding environments and hope
for good performance in low-yielding environments also,
"even when they knowthat the procedure is suboptimal."
Likewise, breeders often make selection decisions on
yield averages across wide area tests, but this can con-
found mega-environments together, thus obscuring and
diminishing yield differences. Furthermore, as Nielsen
(1992) remarkedin his presidential address to the Ameri-
can Society of Agronomy,"Westill rely on statistical
methods developed more than one-half century ago,"
such as "analysis of variance and regression techniques."
Yet the regression technique developed by Yates and
Cochran (1938) and popularized by Finlay and Wilkinson
(1963) and Eberhart and Russell (1966) remains the
commonmethod for analyzing interactions, even though
an empirical comparison by Yau (1995) shows that
AMMI routinely captures much more of the interaction
(and never less). To meet future needs for crops that
are highly productive in a diversity of environments,
muchof current practices wouldprove inadequate. Inter-
actions must be exploited vigorously, especially when
genotype-location interactions are sizable relative to ge-
notype main effects and genotype-year interactions.
Mega-environmentresults are useful only if they are
reasonably stable-only if somewhatdifferent genotypes
or locations or years would cause little change. More
research is needed to quantify the robustness of the
present methodology, particularly the quantity of data
required to achieve stable results. One simple test would
be to compare results for numerous subsets of several
large datasets. However,there are substantial indications
in the literature of long-term association of locations
(DeLacy et al., 1994), stable mega-environmentsacross
years and even across crops (Annicchiarico and Perenzin,
1994; DeLacy et al., 1994; Abdalla et al., 1996) and
across different subsets of genotypes (Annicchiarico et
al., 1995), and heritability of interaction traits (Pham
and Kang, 1988; Berke et al., 1992; Jalaluddin and
Harrison, 1993; Romagosaet al., 1993). Romagosaand
Fox (1993) claim that multivariate analysis (including
AMMI)of new genotypes with known genotypes based
on just 1 yr of wide testing provides reliable predictions
for future years. Such rapid assessment of a new geno-
types mega-environment is important because commer-
cial cultivars for manycrops have an average lifetime
of just several years. Especially if years are deemeda
randomeffect and a dataset is sizable, a different set of
years is likely to give similar results. However,if the
genotypes change drastically, as by introduction of a
new idiotype, a new assessment of mega-environments
is needed. For example, Braun et al. (1992) report that
the introduction of semi-dwarf wheat increased GEinter-
action. In any case, occasional review and refinement
of mega-environmentsis desirable (DeLacyet al., 1994).
Somedatasets may require a higher-order model than
AMMI-1.Extension to AMMI-2 is relatively straightfor-
ward. The winning domains, which are horizontal bands
for AMMI-1as in Fig. 3, become irregular polygons
for AMMI-2(Gauch, 1992, p. 222). For AMMI-3and
higher models, however, analogous graphs are impracti-
MARCH-APRIL 1997
cal or impossible, so model interpretation must turn to
other options. Fortunately, regardless how many IPCA
axes an AMMI model requires for a given dataset, it is
always possible and simple to list the mega-envi-
ronments, listing the environments won for each winning
genotype (according to yield estimates from the AMMI
model chosen to be most predictively accurate). At-
tempting to relate these mega-environments to evident
geographical or agroecological differences offers the best
hope for a straightforward interpretation. Frequently,
mega-environments delineated by AMMIdo have an
evident geographical or biological interpretation (Annic-
chiarico, 1992; Annicchiarico and Perenzin, 1994; An-
nicchiarico et al., 1995). WhenAMMI-3or higher mod-
els are required, a preliminary clustering may be
considered, hopefully resulting in just a few clusters
being adequate to allow AMMI-1or AMMI-2to suffice
for each data subset. It maybe suspected that high-order
modelswill occur mostly for international scientific yield
trials containing numerous entries that are obviously
nonstarters for local farmers. Finally, when three or
more IPCA axes are recommended, the particular test
procedure or criterion used to reach this conclusion
should be checked to assure its appropriateness relative
to research objectives. For example, F-tests tend to over-
estimate statistical significance relative to predictive tests
(Berger and Berry, 1988; Gauch, 1992, p. 129-153), and
hence F-tests tend to recommendrelatively high-order
models. Likewise, an automatic goal of capturing, say,
70 to 80%of the interaction can be seriously misguided
in the absence of a quantitative estimate of the amount
of signal and noise in a given datasets interaction. Fortu-
nately, "experience has shownthat only very infrequently
there are sufficient grounds for including more than two
axes" (van Eeuwijk, 1995).
Breeding in marginal or stressed mega-environments
calls for exceptional experimental technique to control
errors. Low-productivity environments are prone to large
errors, less differentiation between genotypes, and less
repeatability across years (Braun et al., 1992). Hence,
it is important to select an optimal numberof replications
(Gauch and Zobel, 1996b). Also, AMMIanalysis
regional data can help to control errors and gain accuracy
(Gauch, 1988; Gauch and Zobel, 1996a).
Because historically muchgreater breeding effort has
been expended for major production areas than for mar-
ginal environments, to a considerable extent, any verdict
on the merit of exploiting narrow adaptations is likely
to underestimate the potential. However,Ceccarelli and
Grando(1993) report a rare case in which similar breed-
ing efforts were given to local landraces and elite lines.
They found that selection of locally adapted barley
(Hordeumvulgare L.) landraces in a low-yielding mega-
environment produced superior entries beating the check
variety 30 times as frequently as did selection of elite
barley lines in a high-yielding mega-environment.
Clearly, in other cases, if the genotype roster contains
only genotypes bred in and for highly productive environ-
ments, the results may not be indicative of either the
number of needed mega-environments or the yield boost
from exploiting narrow adaptations that would result
 GAUCH & ZOBEL: IDENTIFYING MEGA-ENVIRONMENTS 325

were the roster expanded to also include genotypes vigor-

ously bred for marginal environments. As a quick diag-
nostic, whenever the genotype-location interaction SS
(discounted for noise) equals or exceeds the genotype
SS and the genotype-year SS, which is a common case,
subdivision of the growing region into several mega-
environments usually cannot be ignored without a sub-
stantial yield loss. If a breeding program in the past
focused on a single highly productive and well-defined
environment but in the future will serve a wider diversity
of mega-environments, then to maintain research effi-
ciency and productivity, stronger statistical tools will be
needed to comprehend interactions, reduce noise, iden-
tify mega-environments, and target genotypes.

ACKNOWLEDGMENTS
We appreciate helpful suggestions on earlier drafts of this
paper from P. Annicchiarico, P.L. Cornelius, J. Crossa, B.E.
Eisenberg, R.A. Fischer, M.S. Kang, H. Pham, H.TP. Piepho,
M. Sorrells, A.F. Troyer, W. Van, K. Yourstone, D. Wallace,
and three anonymous reviewers. The senior author especially
appreciates some penetrating questions from R.A. Fischer
several years ago at CIMMYT, which precipitated the develop-
ment of graphs like Fig. 3. This research was supported by
the Rhizobotany Project of the USDA-ARS.
 326 CROP SCIENCE, VOL. 37, MARCH-APRIL 1997

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