Field Crops Research 100 (2007) 61–72

www.elsevier.com/locate/fcr

Progress over 20 years of sunflower breeding in central Argentina

Abelardo J. de la Vega a,*, Ian H. DeLacy b, Scott C. Chapman c
a
Advanta Semillas S.A.I.C., Ruta Nac. 33 Km 636, C.C. 559, 2600 Venado Tuerto, Argentina
b
School of Land and Food Sciences and A.C.P.F.G., The University of Queensland, Brisbane, Qld 4072, Australia
c
CSIRO Plant Industry, Queensland Bioscience Precinct, 306 Carmody Rd., St. Lucia, Qld 4067, Australia
Received 11 January 2006; received in revised form 16 May 2006; accepted 17 May 2006

Abstract
This paper applies linear mixed model analysis to 122 on-farm trials of commercial and near-commercial sunflower (Helianthus annuus L.)
hybrids grown over 15 years in 32 locations of central Argentina to quantify increases in oil yield and to determine the contributions of change in
both biotic stress resistance and yielding ability in favourable environments. The best linear unbiased predictors (BLUPs) from this analysis can be
regarded as measures of ‘relative peak performance’ of hybrids in environments for which they were selected, and are a better measure of their
adaptation compared to small trial sets of ‘historical’ hybrids. The BLUPs of 49 commercial hybrids released between 1983 and 2005 showed a
genetic gain for oil yield of 11.9 kg ha1 yr1. Special purpose hybrids that were converted for single traits or that were developed for low-
technology markets lagged by 5–15 years in terms of genetic gain. Genetic gains came about due to both an increase in the number of hybrids with
resistance to the major biotic stress (Verticillium dahliae Klebahn) and a genetic gain in oil yield of 14.4 kg ha1 yr1 within these hybrids. Based
on the data and the estimated time lag between commercial release and peak use, the improvement in oil and grain yield of conventional hybrids in
central Argentina will be sustained until at least 2010, with evidence that the new germplasm pools still have substantial genetic variance to be
exploited.
# 2006 Elsevier B.V. All rights reserved.

Keywords: Genetic gain; Helianthus annuus L.; Meta-analysis; Mixed model analysis; Relative peak performance; Repeatability; Sunflower; Variance components

1. Introduction commercialization in Argentina during the last 10 years

Commercial sunflower (Helianthus annuus L.) production in
Argentina commenced in the early 1930s with open-pollinated
varieties. From a maximum production area of about
3.5 million ha in the mid 1990s, current production has
stabilised to 1.8–2.2 million ha. A major portion of this area
(ca. 1.0 million ha) is planted in central Argentina in the region
between latitudes 33.58 and 36.58 S and longitudes 618 and 658
W. The average grain yield remained around 700 kg ha1 for a
long period from 1930 to the early 1970s, when the first hybrids
were released (López Pereira et al., 1999), and increased at a
rate of 49 kg ha1 yr1 from the early 1970s to the mid 1990s
(Fig. 1) during the period of hybrid adoption by farmers.
Although almost 250 sunflower hybrids were registered for
DOI of original article: 10.1016/j.fcr.2006.05.007.
* Corresponding author. Tel.: +54 3462 435235; fax: +54 3462 435231.
E-mail addresses: avega@waycom.com.ar (A.J. de la Vega),
i.delacy@uq.edu.au (I.H. DeLacy), scott.chapman@csiro.au (S.C. Chapman).
0378-4290/$ – see front matter # 2006 Elsevier B.V. All rights reserved.
doi:10.1016/j.fcr.2006.05.012
(Comisión Nacional de Semillas, unpublished data), average
national grain yield has not increased (Fig. 1). This apparent
slow down of the progress in yield improvement of sunflower,
particularly when compared with maize and soybean (SAG-
PyA, 2005), emphasises the need to examine the role of current
plant breeding and crop management practices in the
development of a sustainable sunflower production system.
Two observations can be made to this respect. Firstly, oil
yield (i.e., the product of grain yield and grain-oil concentra-
tion) is the main selection criterion of most sunflower breeding
programs, and best represents the real return to farmers.
National data (Fig. 1) only accounts for mean grain yield over
time, and effectively assumes that oil content of grain does not
change. However, the relative magnitude of the impact of
increases in grain yield and grain-oil concentration on the
genetic gains achieved for oil yield may differ depending on the
period of time considered. Secondly, the explosive growth of
soybean in Argentina, which increased from 6.0 million ha
planted area in 1994 to 14.5 million ha in 2004 (SAGPyA,
2005), has pushed sunflower production toward more marginal
62 A.J. de la Vega et al. / Field Crops Research 100 (2007) 61–72
Fig. 1. Bi-lineal relationship between mean sunflower grain yield for Argentina
and year. Data obtained from SAGPyA (2005).
western environments and the genetic progress for oil yield may
have been masked in national figures by the decline in
‘agronomic quality’ of the target growing region.
The relative contributions of plant breeding and crop
management to yield improvement over time in a given
cropping region can be separated (e.g., Slafer et al., 1993;
Duvick and Cassman, 1999). Genetic gains can be estimated
by comparing an historic set of cultivars with uniform
management or from the trial data collected by breeding
programs. This gain in relative terms is subtracted from the
total gain in farmers’ fields and the residual is assumed to be
due to changes in management practices (Bell et al., 1995).
Using the first approach, López Pereira et al. (1999) found no
significant increase in yield potential of 11 sunflower (H.
annuus L.) hybrids released in Argentina between 1973 and
1995. These authors hypothesized that the historic require-
ment for disease tolerance and grain quality, together with a
rather narrow genetic base, has imposed restrictions for
improvement in yield potential. In further studies, in which
four sets of hybrids released between 1983 and 1998 were
tested in four experiments, Sadras et al. (2000) found a
positive association between oil yield and year of commercial
release, which was related to both resistance to the fungal
disease Verticillium wilt (Verticillium dahliae Klebahn) and
response to intraspecific competition (measured as the yield
response when competition between rows was reduced by
uprooting plants in buffer rows).
Two main drawbacks could bias studies of genetic gain
based on comparisons of historical sets of cultivars grown under
uniform management (Bell et al., 1995). Firstly, agronomic
practices and incidence of pests and diseases change with time,
such that performance of older hybrids under today’s growing
conditions may be different to their performance when they
were at their peak of use and popularity (Duvick et al., 2004).
Sencondly, environmental conditions during the period of
experimental estimation of relative yields of historic cultivars
may not be representative of the target population of
environments, so that components of genotype-by-environment
(G  E) interactions are not considered.
These and other issues can be addressed through meta-analysis
of the multi-environment trials (METs) that are routinely
conducted as part of the breeding programs (e.g., Frensham
et al., 1998) or long series of productivity data (e.g., Ellis et al.,
2001). The data collected from METs over time can be utilised to
develop a historical database which greatly expands the sample of
environments and genotypes considered in the analysis of
genotypic adaptation (DeLacy et al., 1996a and references
therein). Therefore, many questions that cannot be addressed in
one or a few years of experiments can be investigated using a
larger database that represents the crop’s target population of
environments and in which every commercial cultivar was tested
at the environment for which it was created. Thus, in this type of
analysis, the hybrid estimates become a measure of their
performance at the environment of their peak of use and will be
named as relative peak performance in this study.
Analysis of historical data has been severely complicated in
the past by the imbalance endemic to METs in the way that
commercial checks, experimental cultivars and trial locations
change over time. Different methods described by DeLacy et al.
(1996a) for dealing with imbalance in METs have been devised.
The general mixed model (Henderson, 1963, 1977) using either
general least square (GLS) or residual maximum likelihood
(REML, Patterson and Thompson, 1975) provides a powerful
method for analysing unbalanced (or balanced) MET data as it
enables the analysis of any linear model with or without cov-
ariates. Kempton (1984) described how fixed effect models could
be designed to better accommodate genotype and G  E effects
than do standard ANOVA. Smith et al. (2005) have recently
reviewed the role of these types of models in a linear mixed model
context for application to cultivar breeding and evaluation trials.
In sunflower, mixed model analyses were used to compare
testing strategies (Chapman and de la Vega, 2002) and to assess
the convenience of dividing the growing region of Argentina into
subregions (de la Vega and Chapman, 2006a). This paper reports
on a sunflower MET dataset consisting of 122 advanced trials
grown in 2–15 locations per year over 15 years in the central
sunflower subregion of Argentina. The analyses described here
were used to measure genetic progress of sunflower breeding
programs through the removal of environment effects from the
datasets. The objectives are: (1) to extend the mixed model
analysis described by Chapman and de la Vega (2002) to a much
larger MET dataset and to compare estimates with those from
smaller trials of fixed entries; (2) to use MET data to estimate the
contribution of more than 20 years of sunflower breeding to oil
yield and grain yield progresses in Argentina; (3) to predict what
the industry should expect in the future from current hybrids in
terms of improved performance; (4) to examine how changes in
relative performance (product longevity) of hybrids relate to their
time of release and tolerance to major stresses.
2. Materials and methods
2.1. Study zone
The study zone comprises the central-west and north-west
portions of the Argentinian Pampas. Previous studies (de la
 A.J. de la Vega et al. / Field Crops Research 100 (2007) 61–72 63

Vega et al., 2001; de la Vega and Chapman, 2006a) reported
that this geographical zone corresponds to a single mega-
environment for sunflower, i.e., it shows a pattern of
genotypic discrimination consistently different to other
sunflower regions. Soil, climate and technology of this
region have been described by Hall et al. (1992), Mercau et al.
(2001) and Chapman and de la Vega (2002). Briefly, the
typical sunflower growing soils are deep sandy mollisols
(Entic Haplustoll, Entic Hapludoll, Typic Hapludoll) formed
over loessic sediments. Moving from the northeast to the
southwest of the region, October–March total rainfall
decreases from over 700 to 500 mm, although large seasonal
and spatial variation in total water availability is common
within a zone (Mercau et al., 2001). Fungal diseases are the
main biotic stress in this sunflower cropping system: in
particular Verticillium wilt and Sclerotinia head rot (Scler-
otinia sclerotiorum (Lib.) de Bary) (Sadras et al., 2000).
Diseases produced by the fungi Albugo tragopogonis Pers.,
Alternaria helianthi (Hansf.) Tubaki & Nishihara, and Phoma
oleracea var. helianthi tuberosi Sacc. also reduce yield in
humid years (Chapman and de la Vega, 2002). Standard
sowing dates are from the end of September through to early
December. Industry production figures show sunflower grain
yield in this region averaged 1.8 t ha1 (1995–2004).
2.2. Trial dataset
The dataset was 122 sunflower hybrid trials conducted at 32
locations of central Argentina during the period 1990/1991–
2004/2005 (16,468 data points for oil yield) (Table 1). The
entries were the market-leading hybrids (from all major
companies) and experimental hybrids that were being evaluated
by Advanta Semillas for commercial release. These trials are
the final stage of approximately 4 years of successively
extensive testing prior to commercial consideration and
therefore do not represent the entire genetic variation available,
i.e., the germplasm has undergone selection for oil and grain
yield, grain-oil concentration, disease tolerance, maturity and
other agronomic traits and represents ‘near-elite’ hybrids.
Trials affected by spontaneous disease infections were retained

Table 1
Locations used in the Advanta Semillas hybrid testing program (32 locations with 16,468 data points for oil yield across 122 three-replicate trials conducted from
1991 to 2005)
Location Latitude (S) Longitude (W) Trial number (total) Years
A. Ledesma 33.6 62.6 1 2002
América 35.5 63.0 2 1999, 2002
Arboledas 36.9 61.5 2 2004, 2005
Bolı́var 36.2 61.1 1 1998
C. Casares 35.6 61.4 1 1994
C. de Areco 34.4 59.8 6 2000–2005
Daireaux 36.6 61.8 12 1991–1993, 1996,
1998–2005
G. Moreno 35.6 63.4 1 2005
G. Pico 35.7 63.7 8 1991, 1994, 1997,
1998, 2002–2005
G. Pinto 34.8 61.9 2 1995, 1996
G. Villegas 35.0 63.0 7 1991, 1993, 1995, 1996,
2000, 2003, 2005
Henderson 36.3 61.7 3 2002–2004
H. Lagos 35.0 64.1 1 2001
H. Renancó 34.9 64.4 2 2003, 2005
I. Alvear 35.2 63.6 2 2000, 2001
Junı́n 34.6 61.0 6 1994–1997, 2000, 2003
M. Lauquen 36.2 63.0 2 2002, 2003
9 de Julio 35.5 60.9 7 1994, 1996–2001
Pehuajó 35.8 61.9 3 1991, 1999–2000
Piedritas 34.8 63.0 4 1996–1999
Q. Quemu 36.1 63.6 3 2001, 2002, 2005
Riestra 35.3 59.8 2 1991, 1997
Sampacho 33.4 64.7 9 1991, 1995–2000,
2002, 2003
S. Basilio 33.5 64.3 7 1996–2002
S. Rosa 36.6 64.3 1 1991
S. Spiritu 34.0 62.3 5 1998–2002
30 de Agosto 36.3 62.5 1 2004
T. Lauquen 36.0 62.7 4 1999, 2000, 2002, 2005
Trili 35.9 63.7 2 2002, 2003
V. Maza 36.8 63.3 2 2004, 2005
V. Tuerto 33.7 62.0 12 1992–1998, 2000–2002,
2004, 2005
V. Valeria 34.3 64.9 1 2002
64 A.J. de la Vega et al. / Field Crops Research 100 (2007) 61–72

in the dataset, but those that suffered significant bird, insect
pest, or hail damage were excluded.
Within years, the same hybrids were grown in all locations.
Across years, the data were unbalanced and no single genotype
was grown every year (Table 2). In total, 216 experimental and
88 commercial (Table 3) hybrids were tested. Since not all the
locations were represented in each of the 15 years of this study,
the terms ‘trial’ and ‘environment’ will be used here to define a
particular location in a given year. In fact, sunflower
production, and the evaluation locations, were pushed further
west during the rapid expansion of soybean production over this
time (Fig. 6, discussed later in text).
Throughout this paper, single years in tables or figures (e.g.,
1991) refer to the summer season of that year (i.e., 1990/1991).
As for most datasets of this type, the ‘location’ is a loose spatial
reference, as, in different seasons, the location (identified by a
town name) may actually be different paddocks, farms and/or
soil types subject to slightly different management regimes.
There was only a single trial at a location in any 1 year. Most
trials were located on-farm, although one site per year was
always on the Advanta Argentina research station (V. Tuerto).
Crops were sown within the normal sowing window at each
location (i.e., ca. end of September to early December). In all
trials, hybrids were laid out in alpha- or square-lattice designs,
with three replicates and 30–49 entries per trial. The trials were
over-planted and thinned to 47,600 plants ha1. A plot size of
three or four rows  6 m and inter-row spacing of 0.70 m was
used. Conventional tillage practices were used in all trials until
2002. Afterwards, an increasing proportion of testing sites were
under zero-tillage (now comprising about 80% of trials). All
trials were rain-fed and nutrient deficiencies were generally
avoided through fertilization. Weeds and insect pests were
controlled chemically, but fungal diseases were not controlled.
Plot data of grain (achene) yield were determined by either
hand harvesting of 4.0 m2 (central row, discarding the border
plants) or machine harvesting of 8.4 m2 (four-row plot trials,
two central rows). All yield data are presented at 11% grain
moisture. Grain-oil concentration (% dry matter) was
determined on a plot basis by nuclear magnetic resonance
(Granlund and Zimmerman, 1975) using 10 g oven-dried
samples. Oil yield was calculated as the product of grain yield
and grain-oil concentration. Time to anthesis (days), defined as
the time at which 50% of the plot plant population reached full
anthesis (R-5.5, Schneiter and Miller, 1981), was recorded for
46 out of the 122 trials.
2.3. Statistical analyses
2.3.1. Hybrid performance across trial seasons
To accommodate the imbalance of the dataset, the data for
grain yield (kg ha1, at 11% grain moisture) and oil yield
(kg ha1) were analysed as mixed models with separate
residual terms for the different trials (van Eeuwijk et al., 2001;
Smith et al., 2005). The phenotypic observation yijmn on hybrid i
in incomplete block n of replicate m of environment j was
modelled as:
yi jmn ¼ m þ e j þ ðr=eÞ jm þ ðb=r=eÞ jmn þ gi þ ðgeÞi j þ ei jmn
(1)
where m is the grand mean; ej the fixed effect of the environ-
ment j; (r/e)jm the random effect of the replicate m nested within
the environment j and is NIDð0; s 2r Þ, m = 1, . . ., r; (b/r/e)jmn
the random effect of the incomplete block n nested within the
replicate m of environment j and is NIDð0; s 2b Þ, n = 1, . . ., b; gi
the random effect of hybrid i and is NIDð0; s 2g Þ, i = 1, . . ., g;
(ge)ij the random effect of the interaction between the hybrid i
and environment j and is NIDð0; s 2ge Þ; eijmn is the random
residual effect for hybrid i in the incomplete block n of replicate
m of environment j (experimental error) and is NIDð0; s 2eð jÞ Þ.

Table 2
Number of hybrids common across years in the Advanta Semillas sunflower trial dataset (diagonal entries are numbers for individual years)
Year 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005
1991 36
1992 19 30
1993 13 19 36
1994 11 14 21 42
1995 10 13 19 29 42
1996 8 8 13 21 26 42
1997 8 8 10 13 16 24 49
1998 5 5 5 9 11 17 30 42
1999 2 2 2 2 4 9 16 17 42
2000 5 5 6 6 9 13 19 20 21 49
2001 2 2 2 2 5 8 12 13 16 33 49
2002 1 1 1 1 3 6 8 8 8 16 25 49
2003 0 0 0 0 2 4 6 6 7 13 16 21 49
2004 0 0 0 0 2 3 3 3 3 9 11 15 24 49
2005 0 0 0 0 1 2 2 2 3 8 10 11 13 27 49
Number of locations 7 2 3 5 5 9 8 9 9 12 9 15 10 8 11
Oil yield data points 756 178 323 600 628 1124 1176 1088 1127 1764 1315 2204 1458 1142 1585
Mean oil yield (kg ha1) 1346 1492 1095 1871 1540 1591 1111 1038 1664 1516 1571 1891 1545 1746 1615
Number of locations, data points and arithmetic means for oil yield (kg ha1) per year are given in the table. Across the entire dataset, average oil yield was
1551 kg ha1.
 A.J. de la Vega et al. / Field Crops Research 100 (2007) 61–72 65

Table 3
Commercial sunflower hybrids (alphabetically ordered) from the Advanta Semillas multi-environment trials 1991–2005
Hybrida Release Year Hybrida Release Year Hybrida Release Year
ACA 864 2003 1 DK 3920 2002 2 Paraı́so 27 2003 1
ACA 872 1999 2 DK 4000 CL 2003 1 Paraı́so 30 1999 5
ACA 884 1991 9 DK 4030 1993 7 Paraı́so 33 2003 2
ACA 885 1999 2 DK 4040 1997 6 Paraı́so 35 2003 1
Agrobel 910 1993 1 DK 4050 1999 5 Paraı́so 40 2000 1
Agrobel 920 1995 1 DK 4200 2003 1 Paraı́so 50 2001 2
Aguará 1997 3 DK G100 1984 4 Paraı́so 101CL 2003 1
Asgrow 548 1989 3 DK G103 1987 6 Puelche 1991 3
Atuel 1999 2 MG 2 1997 6 SPS 3130 1986 7
CF 11 1997 3 MG 4 1999 1 Super 407 1988 1
CF 13 1999 3 MG 50 2002 3 Super 515 1991 1
CF 17 1997 10 Morgan 731 1984 4 T 600 1998 2
CF 17 DMR 2004 1 Morgan 733 1988 2 TC 2000 1991 9
CF 19 1997 11 Morgan 734 1989 5 TC 2001 1994 5
CF 21 1997 7 Morgan 742 1996 8 TC 3001 1991 6
CF 23 CL 2005 1 Mycosol 2 1995 1 TC 3002 1991 1
CF 25 1997 7 Olisun 2002 3 TC 3003 1992 4
CF 27 2004 3 Olisun 2 2005 1 VDH 370 2005 2
CF 29 2004 4 P 6440 1988 4 VDH 475 1993 2
CF 31 2005 2 P 64A41 1996 1 VDH 480 1993 5
CF 3 Negro 1997 3 P 6510 1989 5 VDH 481 2003 3
Contiflor 15 1989 11 P 6520 1993 2 VDH 483 1998 4
Contiflor 3 1983 10 Paraı́so 1990 5 VDH 485 2000 4
Contiflor 7 1988 9 Paraı́so 2 1990 5 VDH 487 2004 3
Contiflor 8 1985 7 Paraı́so 3 1993 5 VDH 488 1999 7
Contiflor 9 1988 11 Paraı́so 4 1993 1 VDH 93 1999 7
DK 3880 CL 2003 2 Paraı́so 5 1993 1 VDH 96 1994 4
DK 3881 1993 4 Paraı́so 6 1993 4 Zenith 1993 1
DK 3900 1997 4 Paraı́so 20 1997 6
DK 3915 1997 6 Paraı́so 22 2003 2
Years of testing (yr) in the Advanta dataset are indicated.
a
Seed companies and hybrid codes are: Asociación de Cooperativas Argentinas (ACA), La Tijereta (Agrobel), Nidera (Asgrow, Paraı́so), Syngenta (Atuel,
Puelche), Advanta (Aguará, CF, Contiflor, Olisun, VDH), Monsanto (DK, Super), Dow (MG, Morgan, Mycosol, T), SPS (SPS), Agroatar (TC), and Sursem (Zenith).

Another model was also tested with the environment term
partitioned into years (Y), locations (L) and Y  L interaction.
Since 75% of the total G  E interaction observed for oil and
grain yield was accounted for in the G  Y  L interaction
(results not shown) and a high proportion of trial locations
varied over years (Table 1), this model produced very similar
results when compared with model (1), the latter being finally
selected on the basis of parsimony. REML (Patterson and
Thompson, 1975) using the sparse Average Information algo-
rithm (Gilmour et al., 1995) was used to estimate the variance
components and standard errors of random terms, as imple-
mented in GenStat 8.2 (2005).
The use of breeding programs’ METs to estimate the genetic
gain of a cropping system was compared to the typical approach
of designing specific trials by comparing hybrid-mean
repeatability and reliability of the estimated genotypic variance
components for oil yield of both strategies. The components of
variance estimated for the four agronomic traits by the REML
analysis were used to estimate hybrid-mean repeatability (Fehr,
1987, p. 97) as:
s 2g
H¼
s 2g þ s 2ge =e þ s 2eð jÞ =ðerÞ
where s 2g is the G variance component, s 2ge the G  E inter-
action variance component, s 2eð jÞ the residual variance compo-
nent and e and r are a given number of environments and
replicates, respectively. The estimates of residual variance from
each trial were used to compute a pooled estimate of residual
variance, which was used to calculate repeatability. Another
model was also used to calculate repeatability in which the
effect of the environment was partitioned into years and trials
within years. Hybrid-mean repeatability was estimated as:
s 2g
H¼ (3)
s 2g þ s 2gy =y þ s 2ge=y =ðyeÞ þ s 2eð jÞ =ðyerÞ
where s 2gy is the G  Y interaction variance component, s 2ge=y
the within-year G  E interaction variance component pooled
across years, and y, e and r are a given number of years, trials
per year and replicates, respectively.
The oil yield datasets of two specifically designed sunflower
experiments were re-analysed using model (1), to assess the
reliability of the estimated genotypic variance components
when compared to the breeding program’s MET data. The
datasets were: (1) a reference set of 10 differentially adapted
(2) sunflower hybrids tested across 7 years in 22 trials (de la Vega
and Chapman, 2006a) and (2) a sunflower North Carolina
66 A.J. de la Vega et al. / Field Crops Research 100 (2007) 61–72
Design II (4 females  4 males) tested across five locations in
one season (de la Vega and Chapman, 2006b). The two datasets
included trials conducted in other sunflower regions of
Argentina, but during re-analysis here, only those grown in
the central region were retained.
In order to conduct a genetic gain analysis for a particular
trait, it is necessary to obtain a single estimate for each cultivar,
which best predicts the true genotypic effect. By definition, this
implies the use of best linear unbiased predictors (BLUPs)
(Robinson, 1991), so that the genotypic effect should be
regarded as random (Smith et al., 2005; Piepho and Möhring,
2006), as in model (1). In this study, all experimental hybrids
were developed by the sunflower program of Advanta Semillas
but the commercial checks were developed by different seed
companies. Thus, treating the collection of hybrids as a sample
from one reference population is a first approximation to the
study of the genotypic variation of this dataset. Using GenStat
8.2 (2005), the BLUPs of the genotypic effects (i.e., predictors
that were adjusted for the unbalanced nature of the data) were
computed from REML analysis and allowed, under certain
assumptions (Smith et al., 2005), comparison of genotypes that
were never tested together in the same trial via their relative
performance for a particular trait across the target population of
environments. These BLUPs are therefore estimates of the
relative peak performance of genotypes, as evaluated in their
environments of selection and release. As trials, years and trials
nested within years were defined as fixed effects in models (1)
and (4), their estimates computed from REML are best linear
unbiased estimates (BLUEs).
A key point in genetic gain studies is the inference to be
made from the set of genotypes that are included in the
analysis. In crops like maize, for which yield and yield stability
are by far the most important selection criteria used by
breeders, the most applied criterion is to retain the hybrids that
were widely grown and popular with farmers in their time (e.g.,
Duvick and Cassman, 1999). In the case of sunflower, the
central objective of commercial breeding programs is to
increase oil yield and oil yield stability. However, secondary,
special purpose targets may lead to the release of a cultivar that
does not outperform the existing ones (in yield or stability), at
least under uniform management conditions, and is briefly
illustrated in the results. We propose that these types of
cultivars should be excluded when estimating the contribution
of breeding to yield gains in the farmers’ fields. In this study, a
set of 49 commercial hybrids out of the 88 tested in the Advanta
Semillas MET (Table 3) was selected. The hybrids excluded
from the estimation of genetic gain were those that were: (1)
expressly developed for sunflower regions other than central
Argentina (e.g., Aguará and CF 13, developed for the northern
and southern regions, respectively); (2) developed for a low-
price/low-technology seed market (e.g., TC 3001); (3) created
through the backcross conversion of a previously released
hybrid to incorporate disease resistance (e.g., CF 17 DMR),
differential oil quality (e.g., Olisun), or herbicide resistance
(e.g., CF 23 CL); or (4) evaluated for less than 3 years, with the
exception of the last releases in the dataset for which 2 years of
evaluation were accepted. The BLUPs for oil yield were
plotted against the year of hybrid commercial release and
genetic gain estimates were computed as the slopes of fitted
linear regressions.
2.3.2. Past and future grain and oil yields in farmers’ fields
To evaluate if mean increases for oil yield during the period
1995–2005 were mostly achieved by increasing grain-oil
concentration rather than grain yield, we compared the BLUPs
of the hybrid effects for oil yield and grain yield derived from
model (1) against year of peak of use. For these relationships we
only used the commercial hybrids that had a high level of
impact on the market (i.e., >5% of market share). Due to the
lack of official data for hybrid market share, the selection of
these hybrids was made on the basis of the empirical knowledge
of the commercial team of Advanta Semillas, and thus may be
subject to some level of error. Similarly, it was also considered
that a successful hybrid reaches its peak of use and popularity 4
years after its commercial release (Jorge Moutous, Advanta
Semillas, personal communication). The BLUPs of the recently
released hybrids, which have not yet reached their peak of use,
were used as prediction values for their performance 4 years
after commercial release (6 years after their first testing in pre-
commercial trials).
2.3.3. Hybrid performance within trial season
In order to study the dynamics of the performance of a
genotype over time, it is necessary to obtain a single estimate
for each genotype at every trial season in which it was tested.
The BLUPs of the within-year genotypic effects (i.e.,
genotype + genotype-by-year interaction effects) for oil yield
(kg ha1) were computed from REML (Patterson and
Thompson, 1975) analysis, using GenStat 8.2 (2005). The
data were analysed as mixed models with separate residual
terms for the different trials (van Eeuwijk et al., 2001). The
phenotypic observation yikjmn on hybrid i in incomplete block n
of replicate m of trial j in year k was modelled as:
yik jmn ¼ m þ yk þ ðe=yÞk j þ ðr=e=yÞk jm þ ðb=r=e=yÞk jmn
þ ðgyÞik þ ðge=yÞki j þ eik jmn (4)
where m is the grand mean; yk the fixed effect of year k; (e/y)kj
the fixed effect of the trial j nested within year k; (r/e/y)kjm the
random effect of the replicate m nested within the trial j and
year k and is NIDð0; s 2r Þ, m = 1, . . ., r; (b/r/e/y)kjmn the
random effect of the incomplete block n nested within the
replicate m of trial j and year k and is NIDð0; s 2b Þ, n = 1, . . ., b;
(gy)ik the random effect of the interaction between the hybrid i
and year k and is NIDð0; s 2gy Þ; (ge/y)kij the random effect of
the interaction between the hybrid i and trial j nested within
year k and is NIDð0; s 2ge=y Þ and eijmn is the random residual
effect for hybrid i in the incomplete block n of replicate m of
environment j and year k (experimental error) and is
NIDð0; s 2eðk jÞ Þ. REML using the sparse Average Information
algorithm (Gilmour et al., 1995) was used to estimate the
variance components and standard errors of random terms,
as implemented in GenStat 8.2 (2005).
 A.J. de la Vega et al. / Field Crops Research 100 (2007) 61–72 67

Table 4
Estimated variance components (standard errors) for oil yield derived from the trials (e), hybrids (g), replicates (r), incomplete blocks (b) and experimental error (e)
model applied to the Advanta Semillas sunflower hybrid trial dataset and two sunflower trials designed for specific studies
Source of variation Oil yield (kg ha1) Oil yielda (kg ha1) Oil yieldb (kg ha1)

s 2r=e 2496  372 2636  1057 2945  2505

s 2b=r=e 4442  352 0 2478  2829
s 2g 9897  1066 17026  8541 12102  6205
s 2ge 14540  592 14682  2489 12546  4559
s 2e c 42596  6410 31204  9756 31187  8432
G  E to G ratio 1.47 0.86 1.04
a
Reference set of 10 sunflower hybrids tested across 7 years in 22 trials in the central region of Argentina (de la Vega and Chapman, 2006a).
b
Sunflower North Carolina Design II (4 females  4 males) tested in 5 trials in the central region of Argentina (de la Vega and Chapman, 2006b).
c
Variance components and standard errors were averaged across environments.

Commercial hybrids tested across 5 years or more (30 out of
88) were used to study the dynamics of hybrid performance for
oil yield over time, i.e., ‘hybrid longevity’. The BLUPs of the
within-year genotypic effects for oil yield were plotted against
trial season and the rates of performance change were computed
as the slopes of the linear regression functions fitted to those
relationships.
calculated repeatabilities for two trials are 0.25 and 0.26,
provided they are conducted across 1 or 2 years, respectively
(Fig. 2b). For three trials, repeatabilities are 0.32 and 0.35 for
trials conducted across 1 and 3 years, respectively. This
observation confirms previous studies (de la Vega and
Chapman, 2006a) and indicates that in the central sunflower

3. Results and discussion

3.1. Reliability of breeding programs’ METs and

specifically designed trials

Significant G and G  E interaction components of variance

were found for oil yield (Table 4). Although the ratio of G  E
interaction to G variance components was large, this value was
smaller than other ratios reported for large MET datasets in
diverse sunflower genotypes grown over two regions (de la
Vega and Chapman, 2006a) and other crops (e.g., wheat,
Cooper et al., 1996; maize, Chapman et al., 1997; sorghum,
Alagarswamy and Chandra, 1998 and Chapman et al., 2000;
rice, Cooper et al., 1999; quinoa, Bertero et al., 2004). The
relatively low magnitude of the G  E interaction variance
component relative to G observed in this study can be regarded
as the consequence of the restriction of the analysis within a
single mega-environment of Argentina (i.e., the central
subregion) (de la Vega et al., 2001; de la Vega and Chapman,
2006a) and due to the use of advanced elite hybrids. In these
conditions, the predictable G  E interactions (genoty-
pe  subregion) begin to become ‘part of the G effect’
(DeLacy et al., 1996b). If commercial hybrids developed for
other regions, secondary targets and first round of conversions
are removed from the analysis, a small reduction of the
genotypic variance component is observed (i.e., G  E
interaction to G variance components ratio: 1.66). This
suggests that the inclusion of these hybrids in small proportions
in METs do not contribute to artificial inflation of the G effect.
From the MET analysis, for a hybrid evaluation based on 10
trials, the estimates of hybrid-mean repeatability for oil yield
were 0.63 and 0.77, for one and three replicates per trial, Fig. 2. Predicted hybrid-mean repeatability for oil yield in the central subregion
of Argentina as the number of environments of testing in multi-environment
respectively (Fig. 2a). For 25 trials, the calculated repeat- trials are changed, based on the components of variance given in Table 4 (a) and
abilities were 0.82 and 0.90, for the same number of replicates. on those estimated using a mixed model in which the environment effect was
If the trial effect is partitioned into years and trials within years, partitioned into years and trials within years (b).
68 A.J. de la Vega et al. / Field Crops Research 100 (2007) 61–72
region of Argentina there is scope to redefine testing strategies
by replacing years with locations at no costs in ability to predict
genotype performance. Predicted hybrid-mean repeatabilities
for oil yield (Fig. 2) indicate that a small number of trials (e.g.,
1–6) would not provide the minimum environment sampling
needed to adequately accommodate the effect of the
unpredictable G  E interactions on the estimation of cultivar
means. This suggests that caution should be exercised when
using the ‘traditional approach’ of a small number of specific
trials to compare historical sets of cultivars and highlights the
convenience of using existing MET databases to estimate
genetic gain.
The ratios of standard error to genotypic variance
component were around 0.5 for the two specifically designed
trials and 0.1 for the breeding program’s MET dataset (Table 4).
Thus, the mixed model analysis of a much larger sampling of
genotypes and environments in breeding programs’ METs
produced more reliable estimates than trials designed and
Fig. 3. Best linear unbiased predictors (BLUPs) of relative peak performance
for oil yield for commercial sunflower hybrids in the central region of Argentina
plotted against year of commercial registration. Different symbols indicate
commercial hybrids released for primary (conventional) and secondary target
markets (a) or having contrasting behaviour for Verticillium wilt (b). The
regression functions describe the associations between the variables only for the
conventional hybrids (solid line in (a)), only for the highest yielding conven-
tional hybrids released each year (dotted line in (a)), and only for the hybrids
resistant to Verticillium wilt (b).
conducted with specific purposes. The lower ratio of G  E
interaction to G observed in the specifically designed trial
datasets is related to the inclusion of hybrids specifically
adapted to the northern region, which failed in performing
better than the central-adapted hybrids in most central-region
trials, expanding the genotypic effect. However, this should
have a relatively small effect on the relevant point here, which is
the poorer estimate of parameter error in smaller trial sets.
3.2. The effect of different breeding targets on the
estimation of genetic gain
In considering only the 49 conventional commercial hybrids,
i.e., released for the core market, the analysis showed that oil
yield has increased linearly by 11.9 kg ha1 yr1 over the
period from 1983 to 2005 (Fig. 3a). If only the highest yielding
releases of each year are considered, the estimated genetic
progress is 13.0 kg ha1 yr1 (Fig. 3a).
As noted earlier, one aspect of these analyses is that
sunflower breeding programs do not only release hybrids that
outperform the current market cultivars for oil yield. Different
hybrid types are often needed for different markets and end
uses, and comparisons between them are not only based on
yield. For example, a strategy to participate in the low-
technology (i.e., low seed price) market segment has been to
release an easy-to-produce three-way version of an existing
single-cross hybrid and to deliver it to the market through a
license agreement. This new release does not contribute to the
genetic progress, but allows a company to supply different
market segments. Backcross conversions may also be designed
to deal with particular market issues. Disease-resistant
converted forms of an existing successful hybrid will quickly
replace the original version. Converted hybrids carrying the
Pervenets mutation produce high-oleic acid (i.e. >80%) oil
(Fernández-Martı́nez et al., 1989), which is highly suitable for
the industry and reaches higher prices in the market than
conventional high-linoleic sunflower oil. Recently developed
hybrids having resistance to imidazolinone herbicides (Al-
Kathib and Miller, 2000) allow better weed control and,
although they are currently lower yielding than the conven-
tional hybrids, are preferred by farmers in weed-infested fields.
When the BLUPs of the genotypic effects for oil yield were
plotted against year of commercial release (Fig. 3a), it was clear
that the hybrids developed for special purpose secondary targets
and backcross conversions of existing hybrids do not sit on the
same trend of genetic progress over time shown by the
conventional hybrids, leading to an underestimation of genetic
gain. In the case of the conversions, this effect is likely to be a
temporary phenomenon because, once these genes are spread
throughout elite breeding materials, they will become ‘‘back-
ground genes’’ (Duvick and Cassman, 1999). As might be
expected, hybrids that were specifically developed for other
regions rather than that under study, if included in the analysis,
would also produce a similar negative effect on the estimation
of genetic gain (Fig. 3a). Consequently, it is relevant for a
genetic gain study to concentrate only on those cultivars that
were developed with the aim of increasing yield and stability
 A.J. de la Vega et al. / Field Crops Research 100 (2007) 61–72
within a target region. This criterion, together with the number
of years in evaluation, lead us to focus on the BLUPs of the
genotypic effects of 49 out of the 88 commercially released
hybrids.
3.3. Genetic gain for conventional sunflower in central
Argentina
The BLUPs of the genotypic effects for oil yield (Fig. 3a)
indicate a clear and continuous improvement due to plant
breeding during the last 20 years. López Pereira et al. (1999)
and Sadras et al. (2000) proposed that variation in actual yield
among sunflower hybrids released in different years was limited
in environments with low incidence of diseases, in comparison
to the larger variation found in environments dominated by
major diseases, particularly Verticillium wilt. Although in this
study the proportion of hybrids having genetic resistance to
Verticillium wilt and tolerance to other major diseases did
increase with the year of commercial release, it can be shown
that both biotic stress resistance and yielding ability in
favourable growing conditions have been genetically improved
(Fig. 3b). If the 122 trials are grouped on the basis of their
Fig. 4. Best linear unbiased predictors (BLUPs) of relative peak performance
for oil yield (a) and grain yield (b) for commercial sunflower hybrids in the
central region of Argentina as a function of year of peak of use (i.e. year of
commercial release + 4). The hybrids plotted are those that had a high level of
impact on the market. Lineal (a) and bi-lineal (b) regressions describe the
associations between the variables until 2005.
69
BLUEs for oil yield, REML analyses conducted within each
dataset show that the increase in relative peak performance for
oil yield was positive in the high (P < 0.0001), intermediate
(P < 0.0001) and low (P = 0.0051) oil-yield production
environments (data not presented). When the 49 commercial
hybrids are divided on the basis of their resistance to
Verticillium wilt and only the oil yield BLUPs of the resistant
hybrids are regressed on year of commercial release (Fig. 3b),
the trend indicates an overall gain due to an increase in relative
peak performance within the Verticillium-resistant group (25
hybrids) of 14.4 kg ha1 yr1. These resistant hybrids now
dominate the market.
3.4. Past and future grain and oil yields
When only the hybrids with high impact on the market are
considered, the trend for oil yield indicates a continuous
increase in relative peak performance for sunflower in central
Argentina of 11.9 kg ha1 yr1 from 1986 to 2005 (Fig. 4a).
However, the trend for grain yield can be better described by a
bi-linear function (Fig. 4b) that shows a discontinuity in the
increase in relative peak performance for this trait from 1995 to
2005. This trend is quite consistent with the grain yield trend
shown over the same years in the national data (Fig. 1). It is
likely that at least part of the slow down observed in the grain
yield gains in the farmers’ fields during the last 10 years is the
result of a change in the breeding process which, for that period,
increased oil yield mostly through an increase in grain-oil
concentration. In another paper (de la Vega et al., 2007), we
discuss how these changes arose from the merging of two
distinct germplasm pools over the last 20 years. The BLUPs of
the effects of the recently released hybrids for oil yield (Fig. 4a)
predict that improvements in production oil yields are likely in
the period from 2005 to 2010. For grain yield (Fig. 4b), an
improvement in average production yield can be expected from
the removal of lower grain yield hybrids, but the highest
predicted hybrid grain yield appears to be no greater than that
shown for hybrids in previous years.
Fig. 5. Best linear unbiased estimates (BLUEs) of the year effects for oil yield
as a function of trial season.
70 A.J. de la Vega et al. / Field Crops Research 100 (2007) 61–72

3.5. Dynamics of hybrid performance across years
The variance components and standard errors derived from
model (4) were 9494  732, 13212  580 and 42510  6391,
for s 2gy ; s 2ge=y and s 2e , respectively. A strong inter-annual
variation for oil yield was observed in the MET dataset
(Fig. 5). Both the maximum range of the year-to-year variation
and the low relative oil yields observed in the years of the warm
phase of El Niño Southern Oscillation (1992/1993, 1997/1998
and 2002/2003) are consistent with previous reports on the
sunflower central region of Argentina (Magrin et al., 1998;
Chapman and de la Vega, 2002; de la Vega and Chapman,
2006a). In such years, the higher rainfall in late spring and early
summer in these areas of Argentina can reduce sunflower due to
direct water-logging stress; a net decrease in the seasonal
radiation received by the crop and used for photosynthesis
(Cantagallo et al., 1997; Magrin et al., 1998; Cantagallo and
Hall, 2000; Aguirrezábal et al., 2003); and a reduction in the
partitioning of biomass to grain (de la Vega and Hall, 2002).
Despite the seasonal variation observed for oil yield, the
MET data is consistent with the national data for grain yield in
showing no significant trend (fitted regressions not shown) over
the period from 1991 to 2005 (Figs. 1 and 5). The yield plateau
observed in the MET data could also be partially regarded as the
consequence of the dynamics of trial location substitution over
time in the Advanta Semillas testing net (Fig. 6). With the
exception of the trials conducted in V. Tuerto (research station)
and C. de Areco (eastern location used for disease evaluation),
all trials analysed in this study were conducted on commercial
sunflower production paddocks. From 1991 to 2005, but
particularly since 2000, the explosive growth of soybean in
central Argentina pushed the sunflower industry production
toward more marginal, lower rainfall western environments.
The location selection in the Advanta Semillas testing network
reflects the geographical dynamics of the sunflower industry
during the period covered by the MET dataset.
When the noise due to seasonal oil yield variation is
removed from the analysis, it is observed that relative
performance for oil yield of individual hybrids declined over
time (Fig. 7a). This is expected, as new hybrids are tested each
season. However, since no significant trend was observed for oil
yield across years (Fig. 5), the dynamics shown by the BLUPs
of the within-year genotypic effects for oil yield also reflects the
decline in actual oil yield. This phenomenon could be regarded
as a consequence of the evolution of the pathogen populations,
which become more infective on the current cultivars, and/or
changes in the abiotic challenges, due to the movement of the
crop to more marginal environments.
The comparison of the dynamics of oil yield BLUPs over
time for sequentially released hybrids by the same company
(Fig. 7a) highlights the role of plant breeding in making a
production system sustainable. Oil yields across years have
been maintained through the continuous release of hybrids that
yield better than their predecessors, allowing counteraction of
the decline in ‘agronomic quality’ of the sunflower growing
environments.
Sadras et al. (2000) postulated that a major part of the
improved performance of more recent hybrids was related to
increased resistance to Verticillium wilt. In our study, resistant
hybrids were characterised as those that showed no symptoms
or few symptoms of wilt infection in the bottom third of the
plant for their entire production life. The conclusions of Sadras
et al. (2000) are supported in our results by the observation that,
for hybrids released in the same year (i.e., from the same stage
of germplasm pool development), those that have Verticillium
‘resistance’ have a longer product life, i.e., a slower decline in

Fig. 6. Geographical location of the 32 sites of the Advanta Semillas sunflower multi-environment trial (MET) used in this study. Crosses represent locations
representative of all period covered by the MET. Black circles and open triangles represent locations that have been discontinued and incorporated, respectively,
during the period 2000–2005. October–March total rainfall isohyets (Mercau et al., 2001) are also presented.
 A.J. de la Vega et al. / Field Crops Research 100 (2007) 61–72
Fig. 7. Within-year best linear unbiased predictors (BLUPs) of relative peak
performance for oil yield as a function of trial season. In (a) commercial hybrids
sequentially released by Advanta Semillas are represented. In (b) hybrids
differing in year of release and behaviour against Verticillium wilt are repre-
sented. Inset in (b): box plot of rates of decline in oil yield across years for
hybrids susceptible and resistant to Verticillium wilt.
oil yield (Fig. 7b). When averaged over all of the releases
considered, this rate of decline was less than half of that
observed for Verticilium ‘susceptible’ lines (inset in Fig. 7b).
4. Conclusions
Linear mixed model analysis allows estimates of the genetic
progress in a cropping system using the largely unbalanced
long-term MET datasets that are obtained as part of the routine
activities of breeding programs. Two critical issues should be
considered: (1) for a MET to be useful for this type of analysis,
all commercial hybrids with high impact on the market should
be included in the trials, and most importantly, during their
period of peak of performance; (2) secondary breeding targets
lead to underestimation of the effective genetic gain achieved
by the system (Fig. 3a). Compared to alternative ‘progress
evaluation trials’, linear mixed models allow: (1) expansion of
the sample of genotypes and environments; (2) estimation of
the performance of genotypes at the environment for which
they were bred, i.e., ‘relative peak performance’; (3)
accommodation of G  E interactions (Fig. 2 and Table 4).
Mixed model analysis can also provide useful information for
breeding programs and seed companies, such as the genetic
71
progress of the experimental germplasm or the trends followed
by proprietary and competitor commercial releases.
The analyses reported in this paper show that oil yield gains
in commercial releases of sunflower for central Argentina due
to plant breeding have been continuous over the last 20 years.
Changes in both biotic stress resistance and yielding ability in
favourable environments have contributed to yield mainte-
nance, allowing counteraction of the decline in ‘agronomic
quality’ of the sunflower growing region and ‘hybrid run down’.
It is likely that at least part of the slow down observed in
grain yield gains in the national data during 1995–2005 is the
result of a breeding process which, for that period, increased oil
yield mostly through an increase in grain-oil concentration. In a
companion paper (de la Vega et al., 2007) we explore the role of
changes in grain yield, grain-oil concentration and time to
flowering in improving sunflower oil yield as the initial
germplasm pools were merged and further exploited.
Acknowledgments
This research was supported by Advanta Semillas SAIC.
The authors would like to thank Aldo Martı́nez, Alejandro Dell
Else, Sergio Solián, Carlos Ghanem, Daniel Kennedy, Ney
Flores, César Sánchez and Hugo Baravalle for collaborating in
the field experiments and data management; previous Advanta
sunflower program managers (Ricardo Siciliano and Jorge
Moutous); Advanta sunflower research coordinator (Alan
Scott); Drs. Vı́ctor Sadras, Fernanda Dreccer and Craig
Hardner for review comments.
References
Aguirrezábal, L.A.N., Lavaud, Y., Dosio, G.A.A., Izquierdo, N.G., Andrade,
F.H., González, L.M., 2003. Intercepted solar radiation effect during grain
filling determines sunflower weight per seed and oil concentration. Crop
Sci. 43, 152–161.
Alagarswamy, G., Chandra, S., 1998. Pattern analysis of international sorghum
multi-environment trials for grain-yield adaptation. Theor. Appl. Genet. 96,
397–405.
Al-Kathib, K., Miller, J.F., 2000. Registration of four genetic stocks of sun-
flower resistant to imidazolinone herbicides. Crop Sci. 40, 869–870.
Bell, M.A., Fischer, R.A., Byerlee, D., Sayre, K., 1995. Genetic and agro-
nomic contributions to yield gains: a case study for wheat. Field Crops
Res. 44, 55–65.
Bertero, H.D., de la Vega, A.J., Correa, G., Jacobsen, S.E., Mujica, A., 2004.
Genotype and genotype-by-environment interaction effects for grain yield
and grain size of quinoa (Chenopodium quinoa Willd.) as revealed by
pattern analysis of international multi-environment trials. Field Crops Res.
89, 299–318.
Cantagallo, J.E., Hall, A.J., 2000. Reduction in the number of filled seed in
sunflower (Helianthus annuus L.) by light stress. In: Proceedings of the 15th
International Sunflower Conference, Tolouse, France, pp. D35–D40.
Cantagallo, J.E., Chimenti, C.A., Hall, A.J., 1997. Number of seeds per unit area
in sunflower correlates well with a photothermal quotient. Crop Sci. 37,
1780–1786.
Chapman, S.C., de la Vega, A.J., 2002. Spatial and seasonal effects confounding
interpretation of sunflower yields in Argentina. Field Crops Res. 73, 107–
120.
Chapman, S.C., Crossa, J., Edmeades, G.O., 1997. Genotype by environment
effects and selection for drought tolerance in tropical maize. I. Two mode
pattern analysis of yield. Euphytica 95, 1–9.
72 A.J. de la Vega et al. / Field Crops Research 100 (2007) 61–72
Chapman, S.C., Cooper, M., Butler, D., Henzell, R., 2000. Genotype
by environment interactions affecting grain sorghum. I. Characteristics
that confound interpretation of hybrid yield. Aust. J. Agric. Res. 51,
197–207.
Cooper, M., Brennan, P.S., Sheppard, J.A., 1996. A strategy for yield improve-
ment of wheat which accommodates large genotype by environment inter-
actions. In: Cooper, M., Hammer, G.L. (Eds.), Plant Adaptation and Crop
Improvement. CAB International, ICRISAT and IRRI, Wallingford, UK,
pp. 487–512.
Cooper, M., Rajatasereekul, S., Immark, S., Fukai, S., Basnayake, J., 1999.
Rainfed lowland rice breeding strategies for Northeast Thailand. I. Geno-
typic variation and genotype  environment interactions for grain yield.
Field Crops Res. 64, 131–151.
DeLacy, I.H., Basford, K.E., Cooper, M., Fox, P.N., 1996a. Retrospective
analysis of historical data sets from multi-environment trials—theoretical
development. In: Cooper, M., Hammer, G.L. (Eds.), Plant Adaptation and
Crop Improvement. CAB International, ICRISAT and IRRI, Wallingford,
UK, pp. 243–267.
DeLacy, I.H., Basford, K.E., Cooper, M.C., Bull, J.K., McLaren, C.G., 1996b.
Analysis of multi-environment trials—an historical perspective. In: Cooper,
M., Hammer, G.L. (Eds.), Plant Adaptation and Crop Improvement. CAB
International, ICRISAT and IRRI, Wallingford, UK, pp. 39–124.
de la Vega, A.J., Chapman, S.C., 2006a. Defining sunflower selection strategies
for a highly heterogeneous target population of environments. Crop Sci. 46,
136–144.
de la Vega, A.J., Chapman, S.C., 2006b. Multivariate analyses to display
interactions between environment and general or specific combining ability
in hybrid crops. Crop Sci. 46, 957–967.
de la Vega, A.J., Hall, A.J., 2002. Effects of planting date, genotype and their
interactions on sunflower yield. I. Determinants of oil-corrected grain yield.
Crop Sci. 42, 1191–1201.
de la Vega, A.J., Chapman, S.C., Hall, A.J., 2001. Genotype by environment
interaction and indirect selection for yield in sunflower. I. Two-mode pattern
analysis of oil and biomass yield across environments in Argentina. Field
Crops Res. 72, 17–38.
de la Vega, A.J., DeLacy, I.H., Chapman, S.C., 2007. Changes in agronomic
traits of sunflower hybrids over 20 years of breeding in central Argentina.
Field Crops Res. 100, 73–81.
Duvick, D.N., Cassman, K.G., 1999. Post-green revolution trends in yield
potential of temperate maize in the north-central United States. Crop Sci.
39, 1622–1630.
Duvick, D.N., Smith, J.S.C., Cooper, M., 2004. Long-term selection in a
commercial hybrid maize breeding program. Plant Breeding Rev. 24 (Part
2), 109–151.
Ellis, R.N., Basford, K.E., Cooper, M., Leslie, J.K., Byth, D.E., 2001. A
methodology for analysis of sugarcane productivity trends. I. Analysis
across districts. Aust. J. Agric. Res. 52, 1001–1009.
Fehr, W.R., 1987. Principles of Cultivar Development, vol. 1. Theory and
Technique. MacMillan, New York.
Fernández-Martı́nez, J., Jiménez, A., Domı́nguez, J., Garcı́a, J.M., Garcés, R.,
Mancha, M., 1989. Genetic analysis of the high oleic acid content in
cultivated sunflower (Helianthus annuus L.). Euphytica 41, 39–51.
Frensham, A.B., Barr, A.R., Cullis, B.R., Pelham, S.D., 1998. A mixed model
analysis of 10 years of oat evaluation data: use of agronomic information to
explain genotype by environment interaction. Euphytica 99, 43–56.
GenStat 8.2, 2005. GenStat1 for WindowsTM 8th edition Introduction. VSN
International, Oxford, UK, p. 343.
Gilmour, A.R., Thompson, R., Cullis, B.R., 1995. AI, an efficient algorithm for
REML estimation in linear mixed models. Biometrics 51, 1440–1450.
Granlund, M., Zimmerman, D.C., 1975. Effect of drying conditions on oil
content of sunflower (H. annuus L.) seeds as determined by wide-line
nuclear magnetic resonance (NMR). North Dakota Acad. Sci. Proc. 27 (Pt.
2), 128–132.
Hall, A.J., Rebella, C.M., Ghersa, C.M., Culot, J.P., 1992. Field-crop systems of
the Pampas. In: Pearson, C.J. (Ed.), Field Crop Ecosystems. Elsevier,
Amsterdam, pp. 413–450.
Henderson, C.R., 1963. Selection index and expected genetic advance, in:
Hanson, W.V., Robinson, H.F. (Eds.), Statistical Genetics and Plant Breed-
ing. NAS-NRC, Publication 982, Washington, DC, pp. 141–163.
Henderson, C.R., 1977. Prediction of future records. In: Pollack, E.,
Kempthorne, O., Bailey, T.B. (Eds.), Proceedings of the International
Conference on Quantitative Genetics. Iowa State University Press, Ames,
Iowa, pp. 615–638.
Kempton, R.A., 1984. The use of biplots in intepreting variety by environment
interactions. J. Agr. Sci. Cam. 103, 123–135.
López Pereira, M.L., Sadras, V.O., Trápani, N., 1999. Genetic improvement of
sunflower in Argentina between 1930 and 1995. I. Yield and its components.
Field Crops Res. 62, 157–166.
Magrin, G.O., Grondona, M.O., Travasso, M.I., Boullón, D.R., Rodrı́guez,
G.R., Messina, C.D., 1998. Impacto del fenómeno ‘‘El Niño’’ sobre la
producción de cultivos en la región pampeana. I.N.T.A., Instituto de Clima y
Agua, Castelar, Buenos Aires, Argentina.
Mercau, J.L., Sadras, V.O., Satorre, E.H., Messina, C., Balbi, C., Uribelarrea,
M., Hall, A.J., 2001. On-farm assessment of regional and seasonal variation
in sunflower yield in Argentina. Agr. Syst. 67, 83–103.
Patterson, H.D., Thompson, R., 1975. Maximum likelihood estimation of
components of variance. In: Proceedings of the Eighth International
Biometrics Conference. pp. 197–207.
Piepho, H.P., Möhring, J., 2006. Selection in cultivar trials—Is it ignorable?
Crop Sci. 46, 192–201.
Robinson, G.K., 1991. That BLUP is a good thing: the estimation of random
effects. Stat. Sci. 6, 15–51.
SAGPyA, 2005. Secretarı́a de Agricultura, Ganaderı́a, Pesca y Alimentos.
República Argentina. http://www.sagpya.mecon.gov.ar/new/0-0/agricul-
tura/index.php.
Sadras, V.O., Trápani, N., Pereyra, V.R., López Pereira, M., Quiroz, F.,
Mortarini, M., 2000. Intraspecific competition and fungal diseases as
sources of variation in sunflower yield. Field Crops Res. 67, 51–58.
Schneiter, A.A., Miller, J.F., 1981. Description of sunflower growth stages. Crop
Sci. 21, 901–903.
Slafer, G.A., Satorre, E.H., Andrade, F.H., 1993. Increases in grain yield in
wheat from breeding and associated physiological changes. In: Slafer, G.A.
(Ed.), Genetic Improvement of Field Crops. Marcel Dekker, New York, pp.
1–68.
Smith, A.B., Cullis, B.R., Thompson, R., 2005. The analysis of crop cultivar
breeding and evaluation trials: an overview of current mixed model
approaches. J. Agr. Sci. 143, 1–14.
van Eeuwijk, F.A., Cooper, M., DeLacy, I.H., Ceccarelli, S., Grando, S., 2001.
Some vocabulary and grammar for the analysis of multi-environment trials,
as applied to the analysis of FPB and PPB trials. Euphytica 122, 477–490.