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Abstract
The objective of this study was to investigate field trends in agronomic characters of soybean (Glycine max [L.]
Merr.) performance trials. Field experiments were arranged as lattice designs, and for each plot within an experiment,
an environmental covariate representing the spatial trend was calculated using neighbour analysis. Correlations
between environmental covariates of different characters were then used to analyse the degree of similarity in their
spatial trends. Grain yield, seed protein content, and seed size were more influenced by spatial variations than time
to flowering, time to maturity, and oil content. Significant correlations between environmental covariates were found,
which demonstrate the similarity of trend patterns in different characters within particular experiments. In field trials
exhibiting a high degree of spatial heterogeneity, correlations between environmental covariates of grain yield and
protein content were highly positive. This indicates that field conditions, which promote grain yield, would also
enhance protein content. Moreover, the observed trend patterns considerably affected the calculation of phenotypic
coefficients of correlation between grain yield and seed protein content. The results suggest that spatial analysis should
be applied to all characters of interest, when field conditions are not homogeneous. 2000 Elsevier Science B.V. All
rights reserved.
Keywords: Grain yield; Neighbour analysis; Seed protein content; Soybean; Spatial heterogeneity
1161-0301/00/$ - see front matter 2000 Elsevier Science B.V. All rights reserved.
PII: S1 1 6 1 -0 3 0 1 ( 9 9 ) 0 0 04 2 - 8
14 J. Vollmann et al. / European Journal of Agronomy 12 (2000) 1322
Moreover, estimates of heritability and other of correlation between agronomic and seed quality
genetic parameters might be biased by field hetero- characters was investigated.
geneity (Rosielle, 1980; Magnussen, 1993; Helms
et al., 1995). In agronomic trials, treatment effects
could be masked by spatial variations, thus prohib- 2. Materials and methods
iting the identification of the most favourable
agronomic treatment (Scharf and Alley, 1993). 2.1. Experiments
In order to monitor and control spatial field
variation statistically, a number of different con- During the present study, seven different soy-
cepts and procedures, such as incomplete block bean performance trials ( Table 1), which had been
designs, neighbour analysis, trend analysis, or grown at Raasdorf, Lower Austria (10 km east of
correlated error models, have recently been dis- Vienna, Austria, 4812 N, 1632 E), between 1995
cussed and evaluated comparatively using yield and 1997, were analysed for spatial field variations
data (e.g. Brownie et al., 1993; Stroup et al., 1994; in different agronomic characters. The soil type at
Grondona et al., 1996; Gleeson, 1997). Although the experimental site was classified as a chernozem
substantial effects of spatial heterogeneity on grain of alluvial origin. Each performance trial was
yield have been demonstrated, spatial methods of originally planted as a generalized lattice design in
analysis are rarely being considered for characters two replications at a plot size of 5.51.25 m with
other than grain yield. In wheat, field trends in four rows per plot. The trials designated EXP1,
seed size, test weight, seed protein content, element EXP2, and EXP5 in Table 1 comprised genotypes
of soybean maturity groups 0000 ( Fehr and
concentration of seed, and plant height have been
Caviness, 1980), which were F -derived lines in
reported from production fields and from plot 2
F or F generations from crosses between high-
trials (Rosielle, 1980; Mulla et al., 1992; 5 6
yielding and high-protein parents. The trials desig-
Berndtsson and Bahri, 1995). In soybean trials,
nated EXP3, EXP4, EXP6, and EXP7 involved
characters such as seed size, seed protein and oil
F -derived breeding lines of soybean maturity
content, symbiotically fixed nitrogen and fibrous 5
group 00 in F or later generations, which had
root area have been adjusted for spatial hetero- 7
been pre-selected for yield performance in earlier
geneity in order to enhance the precision of experi-
experiments.
mental results (Herridge and Rose, 1994;
In all experiments, soybean seeds were
Pantalone et al., 1996; Rosielle, 1980; Rebetzke inoculated with Nodular-G (Serbios, Badia
et al., 1998). In barley, environmental trends have Polesine, Italy), a commercial preparation of
been observed among scores of powdery mildew Bradyrhizobium japonicum ( Kirchner) Jordan, in
in an experiment on disease control (Hackett order to promote nodulation and symbiotic nitro-
et al., 1995). gen fixation. Mineral nitrogen was applied at a
Despite the numerous reports on spatial vari- rate of 30 kg ha1 prior to sowing. In addition,
ability in field trials, the related effects of spatial phosphorus and potassium were provided prior to
heterogeneity on different agronomic characters seed bed preparation at rates of 70 kg ha1 P O
within the same experiment have never been con- 2 5
and 140 kg ha1 K O, respectively.
sidered explicitly, and trends occurring in several 2
plant traits have not been compared, although 2.2. Data collection
they might be caused by variations in the same
soil parameters. Therefore, the objective of this Field data of time to flowering, time to maturity,
study was to analyse the influence of spatial trends duration of reproductive phase and plant height
on agronomic and seed compositional traits within were collected as one single reading per experimen-
soybean performance trials by using covariates tal plot. Time to flowering and time to maturity
from a neighbour analysis procedure. Further- were expressed as the number of days required to
more, the influence of field trends on the estimates reach the R1 (beginning of bloom) and R8 (full
J. Vollmann et al. / European Journal of Agronomy 12 (2000) 1322 15
Table 1
Experimental designs of seven different soybean performance trials grown in three years, and the efficiency (%) of lattice analysis as
an indication of the presence or absence of effects of spatial variations on agronomic and seed compositional characters
Design/character Experiment
maturity) developmental stages, respectively, (RCB) designs in order to compare the two
according to Fehr and Caviness (1980). designs. Moreover, the efficiency (%) of lattice
For determination of protein and oil content, a analysis relative to randomised complete blocks
25 g sample of dry seeds from each plot was finely (Cochran and Cox, 1957) was calculated to com-
ground and scanned by near-infrared reflectance pare the degree of spatial variation in the different
spectroscopy (NIRS) using an InfraAlyzer model characters within each experiment, as detectable
450 spectrophotometer and IDAS calibration soft- by lattice analysis.
ware (Bran and Luebbe, Norderstedt, Germany). For the application of neighbour analysis, indi-
Seed protein content, oil content, and the sum of vidual plot residuals were calculated for each
protein and oil content were expressed in g kg1 experimental plot and for each character as:
on a dry matter basis. Seed size [weight of 1000
e =x x: ,
seeds (g)] and grain yield (kg ha1) were expressed ij ij i
on an 8% seed moisture basis. where e denotes the plot residual, x the observed
ij ij
character expression of genotype i in replication j,
2.3. Statistical analysis and x: the arithmetic mean of genotype i. Plot
i
residuals of particular neighbour plots (Fig. 1)
Experimental data were subjected to a lattice were then used to calculate environmental covari-
model of ANOVA, and the F-test was applied to ates for each experimental plot. Following the
examine the statistical significance of genotype designation introduced by Stroup et al. (1994) for
differences for the characters investigated. The long and narrow plots, longitudinal neighbour
generalized lattices were also regarded as random- covariates were referred to as eastwest ( EW )
ised complete block designs with additional restric- covariates. Neighbour plot covariate EW1 was
tions within replications (Cochran and Cox, 1957). calculated as the arithmetic mean of the eastern
For this reason, the lattice experiments were fur- and the western nearest-neighbour plots, which is
ther analysed as randomised complete block identical to the earliest nearest neighbour analysis
16 J. Vollmann et al. / European Journal of Agronomy 12 (2000) 1322
Table 2
Comparison of various ANOVA models for the control of spatial variation in four characters of the EXP2 experiment
ANOVA model Grain yield Protein content Oil content Seed size
RCBc 175 126 0.001 17.5 421 0.179 7.0 145 0.001 5.3 74.0 <0.001 5.5
66 latticed 83 023 <0.001 12.0 234 0.038 5.2 124 <0.001 4.9 36.7 <0.001 3.9
EW1e 71 751 <0.001 11.2 334 0.181 6.2 131 <0.001 5.0 46.2 <0.001 4.4
EW2e 77 168 <0.001 11.6 272 0.040 5.6 137 <0.001 5.2 36.3 <0.001 3.9
EW3e 86 096 <0.001 12.3 263 0.061 5.5 126 <0.001 4.9 33.5 <0.001 3.7
Table 3
Coefficients of correlation (r) between EW2 residuals from a neighbour analysis of different characters in three performance trials
grown in different years
Character Time to maturity Grain yield Protein content Oil content Protein plus oila
EXP1
Time to maturity
Grain yield +0.47**
Protein content +0.33** +0.69**
Oil content 0.18 0.45** 0.34**
Protein plus oila +0.26** +0.51** +0.89** +0.13
Seed size +0.29** +0.41** +0.48** 0.05 +0.48**
EXP2
Time to maturity
Grain yield +0.09
Protein content 0.28* +0.83**
Oil content +0.21 0.57** 0.48**
Protein plus oila 0.18 +0.61** +0.85** +0.07
Seed size +0.51** +0.80** +0.58** 0.36** +0.44**
EXP5
Time to maturity
Grain yield +0.43**
Protein content +0.04 +0.29**
Oil content +0.06 0.31** 0.84**
Protein plus oila +0.17 0.02 +0.37** +0.24
Seed size +0.65** +0.53** +0.14 0.02 +0.21*
tions in grain yield only. However, variations in side of a test plot was often more efficient than
soil parameters have been reported from pro- using only one nearest neighbour for covering a
duction fields, which affected different plant field trend ( Table 2), which is in agreement with
parameters such as element concentrations in grain earlier findings ( Vollmann et al., 1996a,b).
(Berndtsson and Bahri, 1995) or grain yield and Negative correlations between soybean yield
protein content of wheat (Mulla et al., 1992) in a and protein content within different populations
correlated manner. Therefore, the available meth- have been well established by numerous breeding
ods of spatial analysis could be applied to any studies (e.g. Leffel and Rhodes, 1993; Wilcox and
character of interest, when statistically detectable Cavins, 1995). For this reason, the unexpected
soil trends are present in one trait. Although finding of a highly positive relationship between
spatial trends were very similar for different traits the field trends ( EW2 residuals) of grain yield and
of the same experiment in the present study ( Figs. 2 protein content ( Table 3 and Fig. 3) deserves
and 3), a different statistical model might be most additional consideration. As genotype effects are
adequate for each of the traits in order to reduce omitted through the calculation of neighbour cova-
the residual error mean square ( Table 2). In gene- riates, the correlation between EW2 residuals of
ral, however, lattice analysis and the methods of grain yield and protein content has to be regarded
neighbour analysis were far more efficient than the as a purely environmental correlation (Bos and
randomised complete block analysis in modelling Caligari, 1995). Therefore, a variation in environ-
spatial variations. Moreover, in regular field plots mental parameters such as nitrogen availability
of long and narrow shape, the use of two or three from mineralization, symbiotic N fixation and
2
neighbour plots residuals ( EW2 or EW3) at each many other factors, which can promote both yield
20 J. Vollmann et al. / European Journal of Agronomy 12 (2000) 1322
Table 4
Phenotypic (above diagonal ) and genetica (below diagonal ) coefficients of correlation (r) between different characters of the EXP2
experiment at three ways of analysis of field data
Character Time to maturity Grain yield Protein content Oil content Protein plus oilb Seed size
RCB (unadjusted)c
Time to maturity +0.80** 0.20 +0.38* +0.12 +0.50**
Grain yield +0.96++ 0.18 +0.34* +0.11 +0.61**
Protein content 0.41+ 1.10+ 0.51** +0.64** 0.12
Oil content +0.51++ +0.79++ 0.79++ +0.33 +0.35*
Protein plus oil +0.24 0.29 +0.14 +0.49+ +0.18
Seed size +0.51++ +0.61++ 0.50+ +0.54++ +0.17
Lattice adjustmentd
Time to maturity +0.78** 0.41* +0.41* 0.02 +0.48**
Grain yield +0.83++ 0.49** +0.42* 0.10 +0.56**
Protein content 0.53++ 0.73++ 0.40* +0.61** 0.34*
Oil content +0.48++ +0.62++ 0.44++ +0.48** +0.35*
Protein plus oil 0.04 0.13 +0.54++ +0.51++ 0.03
Seed size +0.49++ +0.57++ 0.46++ +0.43++ 0.06
EW2 adjustmente
Time to maturity +0.83** 0.33 +0.31 0.04 +0.51**
Grain yield +0.92++ 0.42* +0.32 0.12 +0.54**
Protein content 0.53++ 0.96++ 0.46** +0.56** 0.33*
Oil content +0.38++ +0.56++ 0.72++ +0.48** +0.35*
Protein plus oil 0.11 0.38 +0.20 +0.53+ 0.00
Seed size +0.51++ +0.57++ 0.61++ +0.45++ 0.12
a Significance of a genetic coefficient of correlation is expressed as being greater than once (+) or twice (++) its standard error.
b Sum of protein and oil content.
c Correlations based on unadjusted entry means from the randomized complete block analysis.
d Correlations based on entry means adjusted by lattice analysis.
e Correlations based on entry means adjusted by neighbour analysis using the EW2 neighbour covariate.
and protein content of a soybean crop (Soldati, correlation between grain yield and protein content
1995), could explain the positive correlation is evident after appropriate adjustment of the field
between the trends in grain yield and protein data for spatial trends [Fig. 4(b)].
content in all experiments investigated. The empirical results obtained from the current
The effects of spatial variations on the results investigation of different soybean trials demon-
of performance trials have been characterized as strate that spatial variations can affect various
influencing the ranking of genotypes (Stroup et al., agronomic characters, revealing similar patterns
1994), thus reducing selection efficiency, and of trend in each of the traits, and affecting the
inflating residual error variance, which reduces the estimates of phenotypic correlation between traits.
power of statistical tests and biases the estimates Different statistical methods such as lattice analysis
of heritability as well as other parameters (Ball and various neighbour or trend analysis techniques
et al., 1993). In the case of spatial variations in are available for an efficient monitoring of field
several characters, coefficients of phenotypic corre- variations and for adjusting treatment means.
lation ( Table 4) and estimates of correlated Apart from covering fertility trends in yield trials,
response to selection can also be biased. Using the adjustment for spatial trends in all agronomic
unadjusted field data, the relationship between characters of interest could be useful in different
yield and protein content [Fig. 4(a)] would suggest fields of agronomic and plant breeding research,
that a selection for one character would not influ- e.g. in selecting for seed quality characters such as
ence the other character. However, a negative protein and oil content.
J. Vollmann et al. / European Journal of Agronomy 12 (2000) 1322 21
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