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DOI 10.1007/s10681-012-0784-z
Received: 26 March 2012 / Accepted: 20 August 2012 / Published online: 1 September 2012
Springer Science+Business Media B.V. 2012
Introduction
Hybrid breeding was suggested as promising approach
to increase grain yield performance in triticale (X
Triticosecale Wittmack) (Oettler et al. 2003, 2005;
Fischer et al. 2010). Requirements to conduct hybrid
breeding in triticale such as the availability of a
cytoplasmic male sterility (CMS) system are fulfilled
and first commercial hybrids have been released in
France and Germany (www.hybrid-triticale.de). Nevertheless, hybrid breeding in triticale is still hampered
by the time- and resource-demanding production and
phenotypic evaluation of the large number of singlecross combinations. Consequently, there is a strong
demand for methods to predict performance of triticale
hybrids with high accuracy.
For qualitative traits, prediction of single-cross
performance based on mid-parent values is very
accurate (Oettler et al. 2005). For complex traits such
as grain yield, however, accuracy is too low to be of
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224
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Molecular analyses
Genomic DNA of all 23 parental lines was extracted
by using the modified CTAB method (Doyle and
Doyle 1990) and screened with 52 polymorphic SSR
markers distributed throughout the genome following
the protocol described by Tams et al. (2004). The
molecular data was used to estimate the coefficient of
coancestry hij between inbreds i and j as outlined by
Bernardo (2002) with the exception that we assumed a
base-line identical by descent probability equal to the
minimum simple matching coefficient among pairs of
lines observed in our data set. In addition, we applied
principal coordinate analysis (Gower 1966) based on
the modified Rogers distances among pairs of inbred
lines (Wright 1978). Molecular marker analyses were
carried out by using the software Plabsoft (Maurer
et al. 2008) which is linked to the R environment
(R development core team 2010).
Field data analysis and predicting hybrid
performance
225
gj was the GCA effect of the jth male line, sij was the
SCA effect of crosses between lines i and j.
Mid-parent heterosis was calculated as the difference between the performance of a hybrid and the
mean performance of its two parental inbred lines.
Rogers genetic distances (Rogers 1972) were estimated among all pairs of parental inbred lines. Pearson
product moment correlations (r) between Rogers
distances and heterosis were calculated. Significance
tests for correlations were performed using the Mantel
test (Mantel 1967).
Prediction of hybrid performance
Performance of untested hybrids (yU) was predicted
based on the GCA effects of their parents i and j as:
yU l g^i g^j
The performance of hybrids were also predicted by
using the information from the inverse of the variancecovariance of the tested hybrids (C-1
TT ) and the
covariance among untested and tested hybrids (CUT)
as: (Bernardo 2002)
1
yU CUT CTT
y^T
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Results
We observed on average 3.1 alleles per SSR locus with a
minimum of two and a maximum of six alleles. The first
two principal coordinates together explained 22.1 % of
the total genotypic variation. Male lines tended to
cluster together but were not separated from the female
lines (Fig. 1). This was further supported by coancestry
coefficients among male (mean h = 0.49) and female
lines (mean h = 0.55), which were of comparable
magnitude than coancestry coefficients among male
times female lines (mean h = 0.52) (Fig. 2).
The genotypic variances of the parents were
significantly larger than zero (P \ 0.01) for all three
evaluated traits (Table 1). The magnitude of genotypic
variance was higher for the hybrids than for the parents
for all three traits. The estimates of r2GCA-F were
significantly different from zero (P \ 0.05) for all
three traits, whereas estimates of r2GCA-M were only
significantly different from zero for plant height and
heading time. Estimates of r2SCA were significantly
larger than zero for grain yield and plant height. The
r2GCA was predominant over r2SCA for plant height
and heading time. In contrast, for grain yield, the
magnitude of r2SCA was slightly higher than r2GCA.
Heritability was almost similar for hybrids and
parental lines with values above 0.89.
The contrast of hybrids versus parental lines was
significantly different from zero (P \ 0.01) for
grain yield but not for plant height and heading time.
Mid-parent heterosis for grain yield ranged from
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0.1
0.0
0.3 0.2 0.1
PC2 (10.5%)
0.2
Female lines
Male lines
0.3
0.2
0.1
0.0
0.1
0.2
0.3
PC1 (11.6%)
Fig. 1 Principal coordinate analyses of 23 triticale inbred lines
based on modified Rogers distances estimated using 52 SSR
markers. Values in parentheses refer to the proportion of
variance explained by the principal coordinates
8
Female lines
Male lines
Female x male lines
7
6
5
Density
0.3
226
4
3
2
1
0
0.0
0.2
0.4
0.6
0.8
1.0
Coefficient of coancestry
Fig. 2 Histogram of coancestry coefficients estimated based on
marker data among the 18 female lines, the 5 male lines, and
among the male and female lines
227
of general combining ability effects of females; r2SCA = variance of specific combining ability effects; r2GCA-M9L, r2GCA2
F9L, and r SCA9L refer to variance of interactions with
environments), and heritability estimates (H2) evaluated for
grain yield, plant height, and heading time
Mean
7.19
110
58.8
Min
Max
5.82
8.86
91
126
57.5
59.7
r2G
0.59**
167.12**
4.49**
Source
Parents
0.21**
6.80**
1.82**
r2ae
0.24
10.03
0.68
H2
0.90
0.98
0.91
Mean
7.84
113
58.9
Min
5.91
87
56.1
Max
G9L
F1 hybrids
8.94
133
61.0
r2GCA-F
0.27*
111.82***
0.96***
r2GCA-M
0.13
149.32***
4. 83***
r2SCA
0.47***
13.94***
0.09*
r2GCA-F9L
0.07***
2.62***
0.18***
r2GCA-M9L
0.10***
9.05***
1.56***
r2SCA9L
H2
0.07***
0.92
1.01
0.97
0.09**
0.91
0.54
0.05
0.01
* ** *** Significantly different from zero at the 0.05, 0.01 and 0.001 level of probability, respectively
a
we assumed that parents and F1 hybrids have the same error variance
Discussion
Intensive research on hybrid breeding in triticale was
started two decades ago and the first hybrid Hybridel
based on chemical hybridizing agent was released in
1999 and the first CMS based commercial hybrid
HYT Prime was officially released in 2010 (Longin
et al. 2012). Hybrid breeding in triticale is very
resource demanding, as the GCA test for the females
requires introgression of the lines into CMS background. Consequently, methods to pre-select promising single-cross hybrids are central to enhance the
efficiency of hybrid breeding. This stimulated us to use
a data set generated in the hybrid triticale breeding
program of the State Plant Breeding Institute of the
University of Hohenheim to compare the accuracies of
different hybrid prediction approaches.
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228
between general combining ability effects and per se performance for grain yield, plant height and heading time for 76
hybrids evaluated in multi-location field trials
Plant height
Heading time
0.14
0.90**
0.77**
r(MP; HP)
0.28
0.88**
0.81**
r(RD; MPH)
-0.29
-0.17
-0.18
Table 3 Accuracy (rG) of predicted hybrid performance (HP) for three traits determined by five-fold cross validation and leave-oneparent-out cross validation
Grain yield
Plant height
Heading time
0.57
0.95
1.00*
Five-fold CV
r(GCA; HP)
r(BLUP-GCA; HP)
0.69
0.96
1.00*
r(BLUP-GCA/SCA; HP)
0.68
0.96
1.00*
Leave-one-male-out CV
r(BLUP-GCA; HP)
0.61
0.65
0.36
r(BLUP-GCA/SCA; HP)
0.55
0.64
0.36
Leave-one-female-out CV
r(BLUP-GCA; HP)
0.56
0.77
0.99
r(BLUP-GCA/SCA; HP)
0.54
0.77
0.99
1.0
0.9
0.8
Accuracy
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0.0
0
15
25
35
45
55
65
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229
References
Barker TC, Varughese G (1992) Combining ability and heterosis among eight complete spring hexaploid triticale lines.
Crop Sci 32:340344
Bernardo R (1994) Prediction of maize single-cross performance using RFLPs and information from related hybrids.
Crop Sci 34:2025
Bernardo R (1996) Best linear unbiased prediction of maize
single-cross performance. Crop Sci 36:5056
123
230
Bernardo R (2002) Breeding for quantitative traits in plants.
Stemma Press, Woodbury
Charcosset A, Bonnisseau B, Touchebeuf O, Burstin J, Dubreuil
P, Barriere Y, Gallais A, Denis JB (1998) Prediction of
maize hybrid silage performance using marker data, comparison of several models for specific combining ability.
Crop Sci 38:3844
Dekkers JCM (2007) Prediction of response to marker-assisted
and genomic selection using selection index theory. J Anim
Breed Genet 124:331341
Doyle JJ, Doyle JL (1990) Isolation of plant DNA from fresh
tissue. Focus 12:1315
Fischer S, Mohring J, Maurer HP, Piepho HP, Thiemt EM,
Schon CC, Melchinger AE, Reif JC (2009) Impact of
genetic divergence on the ratio of variance due to specific
versus general combining ability in winter triticale. Crop
Sci 49:21192122
Fischer S, Maurer HP, Wurschum T, Mohring J, Piepho HP,
Schon CC, Thiemt EM, Dhillon BS, Melchinger AE, Reif
JC (2010) Development of heterotic groups in triticale.
Crop Sci 50:584590
Garcia AA, Wang S, Melchinger AE, Zeng ZB (2008) Quantitative trait loci mapping and the genetic basis of heterosis
in maize and rice. Genetics 180:17071724
Gilmour AR, Gogel BG, Cullis BR, Thompson R (2009) ASReml User Guide Release 3.0. VSN International Ltd,
Hertfordshire. http://www.vsni.co.uk
Gower JC (1966) Some distance properties of latent root and
vector methods used in multivariate analysis. Biometrika
53:325328
Hack H, Bleiholder H, Buhr L, Meier U, Schnock-Fricke U,
Weber E, Witzenberger A (1992) Einheitliche Codierung
der phanologischen Entwicklungsstadien mono- und dikotyler Pflanzen-Erweiterte BBCH-Skala, Allgemein Nachrichtenbl. Deutsch Pflanzenschutz 44:265270
Hallauer AR, Miranda JBF (1988) Quantitative genetics in
maize breeding. Iowa State University Press, Ames
Hallauer AR, Russell WA, Lamkey KR (1988) Corn breeding.
In: Sprague GF, Dudley JW (eds) Corn and corn
improvement, 3rd edn. Agron Monogr 18 ASA, CSSA,
SSSA, Madison, p 463564
Henderson CR (1985) Best linear unbiased prediction of nonadditive genetic merits in non-inbred populations. J Anim
Sci 60:111117
Longin CFH, Muhleisen J, Maurer HP, Zhang H, Gowda M,
Reif JC (2012) Hybrid breeding in autogamous cereals.
Theor Appl Genet (In review)
Mantel N (1967) The detection of disease clustering and a
generalized regression approach. Cancer Res 27:209220
Maurer HP, Melchinger AE, Frisch M (2008) Population genetic
simulation and data analysis with Plabsoft. Euphytica
161:133139
123