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Jos Marcelo Soriano Viana
13 PUBLICATIONS 55 CITATIONS
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CSIRO PUBLISHING
www.publish.csiro.au/journals/cp
Universidade Federal de Vic osa, Departamento de Biologia Geral, 36570-000, Vic osa, MG, Brazil.
Embrapa Florestas; Universidade Federal de Vic osa, Departamento de Engenharia Florestal, 36570-000,
Vic osa, MG, Brazil.
C
Universidade Federal de Vic osa, Departamento de Estatstica, 36570-000, Vic osa, MG, Brazil.
D
Corresponding author. Email: jmsviana@ufv.br
B
Abstract. The testcross method is considered efcient for identifying inbred families with superior general combining
ability. The objective of the present study was to assess the relative importance of the performance per se and in crossing in the
selection of inbred progenies using bi-trait best linear unbiased prediction. We analysed data for expansion volume (EV) and
grain yield from three tests of popcorn (Zea mays L. ssp. everta) S3 families and seven testcross trials, using the ASRemL
software. Four selection strategies were assessed based on: breeding value (strategy 1), general combining ability effect
(GCA) (strategy 2), additive value and GCA from strategies 1 and 2 (strategy 3), and breeding value and GCA predicted by
bi-trait analyses considering EV and yield of the families and testcrosses as different traits (strategy 4). The bi-trait analyses of
the same characteristic assessed in S3 families and topcrosses were generally more accurate and had greater heritabilites. The
greatest predicted gains in EV were obtained using strategy 4, which was inferior to the other strategies for the yield predicted
gains. Strategies 1 and 2 differed most for the families selected. Selection based on GCA maximised heterosis. All of the
strategies resulted in comparable realised gains, especially the strategies 3 and 4 based on breeding value and GCA. Selection
on S3 based on the additive value and GCA (strategies 3 and 4) resulted in inbred lines superior in number of favourable genes
and in general combining ability.
Additional keywords: inbred family, testcross, bi-trait analysis.
Introduction
The testcross method aims to assess the potential of inbred
families to produce superior hybrids from crosses with at least
one tester. It was suggested by Davis (1927) and became common
in maize hybrid breeding programs. In the 1980s, 78% of maize
breeders assessed general combining ability (GCA) by testcross,
using one of the three rst selng generations (Bauman 1981).
Many theoretical and applied results with maize showed that
testcross might be used efciently in inbred family selection.
Baktash et al. (1981) obtained positive signicant correlations
between the performances of maize testcrosses and singlecross hybrids for grain yield and other traits. In the studies of
Bernardo (1991), Lile and Hallauer (1994), and Bordes et al.
(2007) the genotypic correlation between the performance of the
testcrosses in early and late generations and that of their direct
homozygous descendants ranged from 0.71 for S1 lines, to 0.99
for S6 lines.
The choice of the tester is an important challenge to hybrid
crop breeders (Hallauer and Miranda Filho 1988). Hull (1945)
suggested the use of an inbred line with predominantly recessive
genes or a population with low favourable gene frequencies.
This kind of tester maximises the genotypic variance among
testcross families (Wright 1980; Cowen 1987). Comparisons
CSIRO 2011
1836-0947/11/060515
516
nil. The objective of the present study was to assess the relative
importance of the performance per se and in crossing in the
selection of inbred progenies, using the BLUP methodology.
Materials and methods
Testcross and testers
The genotypic means of the testcrosses are:
GOA1 A1 xt MT qat MT GCAA1 A1
MT at =2aAA1 A1
q p
at MT GCAA1 A2
2
MT at =2aAA1 A2
GOA1 A2 xt MT
J. M. S. Viana et al.
fixed effectf
X
fixed effectf
f
517
518
J. M. S. Viana et al.
rA;A~
s
PEVA~
1
~ 2A
2Fs
and rGCA;GCA
~
s
PEVGCA
~
1
~ 2GCA
s
Table 1. Parameters estimated from the bi-trait BLUP/REML analyses of expansion volume (EV, mL/g) and grain yield (kg/ha) of S3 families and
testcross hybrids from the popcorn populations Vic osa and Beija-Flor
Parameter
Vic osa
EV
Beija-Flor
Yield
Program 1
EV
Additive variance
Cond. interval 95%
Dominance variance
Conf. interval 95%
Block
Error
Accuracy (A)
h2
Genetic correlation
Gain/yearA
Gain/yearB
Gain/yearC
S3 mean
GCA
Cond. interval 95%
GCA Env.
Block/Rep./Env.
Error
Accuracy (GCA)
h2
Genetic correlation
Gain/yearB
Gain/yearA
Gain/yearC
Testcross mean
Hybrid mean
A
0.0166
2.5102
0.1359 4.8845
0.0000
14.4213
1E17
97 675
47 315 148 036
0
391 817
0.6181
0.6744
0.4374
0.4979
0.1224 0.3368
1.46
39.75
0.02
9.15
0.81
13.47
27.9
0.52940.0265
01.0800
0.0000
4.8461
1165.0
35 9960.0151
072 355
14 228
283 106
0.5739
0.5896
0.3656
0.3780
0.3811 0.3647
0.22
7.86
0.01
2.97
0.01
4.76
27.0
30.9
3243.2
3218.5
Program 2
Yield
0.0506
4.2106
09.1028
0.8651
28.3837
0.0773
23 876
055 195
6394
201 385
0.6115
0.5712
0.4314
0.3936
0.1882 0.4471
1.99
59.25
0.94
52.55
1.13
79.67
22.7
11.29451E18
6.449616.1394
1.99530.1310
0.2424
8.0713
1055.4
155 9151E18
84 386 227 444
15 9281E06
53 594
287 532
0.9228
0.9067
0.9703
0.8564
0.8452 0.0630
1.32
73.73
0.26
5.63
1.43
72.25
25.2
33.1
2545.5
2581.7
EV
Yield
0.0001
4.0082
014.4127
2.66100.8415
024.0649
0.0000
12.9116
21 8310.0003
636.329643 025
6059
129 363
0.6353
0.5839
0.4274
0.3806
0.2097 0.4541
1.69
53.81
0.43
5.30
0.77
18.31
23.8
1.94281E05
0.63983.2459
2.53591E11
0.2871
6.2324
783.2
13 2610.0110
026 903
30 0782E05
10 581
143 958
0.7579
0.6384
0.2317
0.2453
0.3194 0.2815
0.21
16.61
0.04
2.02
0.05
13.04
24.4
31.5
1769.9
1961.0
519
Table 2. Parameters estimated from the bi-trait BLUP/REML analyses of expansion volume (EV, mL/g) and grain yield (kg/ha), measured in S3
families and testcross hybrids from the popcorn populations Vic osa and Beija-Flor, considering the same trait measured in inbred families and
testcrosses as distinct variables
Parameter
Vic osa
EV
Beija-Flor
Yield
Program 1
EV
Additive variance
Cond. interval 95%
Dominance variance
Cond. interval 95%
Block
Error
Accuracy (A)
h2
Gain/yearA
GCA
Cond. interval 95%
GCA Env.
Block/Rep./Env.
Error
2E06
7E23
5.5167
2.63858.3950
0.0000
10.0892
108370
56491 160 249
0
348 639.0
0.7925
0.7440
1.41
0.7377
0.5475
12.18
0.55250.0090
01.1339
0.0000
4.7322
6E21
1E17
0.9228
0.9041
2.44
44 0170.0062
316284 872
10 926
291 426
10.50187E21
6.008614.9950
1.97301E05
0.3084
6.3165
0.6289
0.3928
0.09
0.6884
0.4913
6.57
0.9018
0.9680
1.04
Correl. A, GCA
0.6160 0.2333
0.8069 0.2281
0.8182 0.0711
0.9651
0.7156
0.9022
0.8106
0.8826
0.9272
Correl. A, A
Correl. GCA, GCAB
EV
24.9701
315 975
15.025234.9150 169 016 462 934
0.0000
0
12.264
226 476
Accuracy (GCA)
h2
Gain/yearA
Program 2
Yield
4.6712
0.69588.6467
9.57237E06
3.837815.3068
0.0000
5.3639
0.8926
0.9542
79.06
106 9734E15
50 865163 081
10 3770.3537
58 138
369 680
Yield
0.0006
16 6180.0024
035 643
0
160 973
0.6977
0.5086
0.90
1.78608E08
0.55293.0190
2.52263E12
0.2923
6.9638
0.8235
0.7907
83.60
0.5067
0.1863
8.65
14 5090.0070
35528 664
27 3200.0002
9944
127 660
0.7517
0.1642
0.12
0.6419
0.2839
10.04
0.5136 0.1438
0.5500 0.2434
0.1967 0.4718
0.6090
0.9161
0.9648
0.9754
0.9649
0.9790
Table 3. Average heterosis, heterosis of selected S3 families, and realised gains for expansion volume (EV, mL/g) and grain yield (kg/ha), and
coincidence (%) of selected families, relative to four selection strategies
Selection strategies: 1, based on breeding value (A); 2, based on GCA effect; 3, based on A and GCA effect; 4, based on A and GCA effect from modied bi-trait
BLUP analysis
Pop.
Vic osa
B-Flor
Prog. 1
B-Flor
Prog. 2
Average heterosis
EV
Yield
0.4
(1.3%)
0.8
(3.2%)
0.1
(0.6%)
639.5
(24.6%)
299.7
(13.3%)
72.6
(4.3%)
Selection strategy:
1
EV
Yield
EV
Yield
EV
Yield
EV
Yield
0.3
1.2
2.7
623.7
24.0
630.0
0.8
3.0
2.3
597.9
23.0
653.5
18.8
0.2
0.6
3.0
664.7
25.5
653.8
91.3
21.7
0.1
0.3
2.8
666.2
25.6
615.3
82.6
26.1
88.4
0.3
1.1
6.2
312.9
13.9
156.6
1.9
7.2
6.7
127.5
5.7
259.3
45.3
2.1
8.2
7.7
131.0
5.8
307.0
56.6
69.8
1.3
5.1
6.9
88.3
3.9
267.2
69.8
62.3
58.5
0.2
0.8
1.5
55.7
3.3
114.7
1.6
6.7
2.9
208.0
12.3
158.8
1.4
5.6
2.5
133.0
7.8
163.9
1.3
5.4
2.4
143.3
8.4
163.1
41.2
64.7
72.5
66.7
74.5
96.1
520
J. M. S. Viana et al.
References
Baktash FY, Younis MA, Al-Younis AH (1981) Correlation coefcients
between topcrosses and diallel crosses in corn (Zea mays L.).
Mesopotamia Journal of Agriculture 15, 5158.
Bauman LF (1981) Review of methods used by breeders to develop superior
corn inbreds. In Proceedings of the 36th Annual Corn and Sorghum
Industry Research Conference. pp. 199208. (American Seed Trade
Association: Washington, DC)
Bekavac G, Purar B, Jockovic D (2008) Relationships between line per se and
testcross performance for agronomic traits in two broad-based populations
of maize. Euphytica 162, 363369. doi:10.1007/s10681-007-9527-y
Bernardo R (1991) Correlation between testcross performance of lines at early
and late selng generations. Theoretical and Applied Genetics 82, 1721.
doi:10.1007/BF00231272
Bernardo R (1994) Prediction of maize single-cross performance using
RFLPs and information from related hybrids. Crop Science 34, 2025.
doi:10.2135/cropsci1994.0011183X003400010003x
Bohn M, Magg T, Klein D, Melchinger AE (2003) Breeding early maturing
European dent maize (Zea mays L.) for improved agronomic performance
and resistance against the European corn borer (Ostrinia nubilalis HB.).
Maydica 48, 239247.
Bordes J, Charmet G, de Vaulx RD, Lapierre A, Pollacsek M, Beckert M,
Gallais A (2007) Doubled-haploid versus single-seed descent and S-1family variation. Euphytica 154, 4151. doi:10.1007/s10681-006-9266-5
Cowen NM (1987) Testcross selection theory. Theoretical and Applied
Genetics 73, 371378. doi:10.1007/BF00262503
Cullis BR, Smith AB, Coombes NE (2006) On the design of early generation
variety trials with correlated data. Journal of Agricultural, Biological &
Environmental Statistics 11, 381393. doi:10.1198/108571106X154443
Davis RL (1927) Report of the plant breeder. In Annual reports. pp. 1415.
(Agricultural Experiment Station: Puerto Rico)
de Boer IJM, van Arendonk JAM (1992) Prediction of additive and dominance
effects in selected or unselected populations with inbreeding. Theoretical
and Applied Genetics 84, 451459.
Elias HT, Carvalho SP, Andr CGM (2000) Comparac o de testadores
na avaliac o de famlias S2 de milho. Pesquisa Agropecuaria
Brasileira 35, 11351142. doi:10.1590/S0100-204X2000000600009
Gallais A (1997) Combined testcross and S-1 selection for the improvement
of testcross and inbred performances. Crop Science 37, 11261133.
doi:10.2135/cropsci1997.0011183X003700040016x
521
Gilmour AR, Gogel BJ, Cullis BR, Thompson R (2009) ASReml User Guide
Release 3.0. (VSN International Ltd: Hemel Hempstead, UK)
Gustafson TJ, Coors JC, de Leon N (2010) Selection for forage yield and
composition on the Wisconsin Quality Synthetic maize population. Crop
Science 50, 17951804. doi:10.2135/cropsci2009.12.0725
Hallauer AR, Lopez-Perez E (1979) Comparisons among testers for
evaluating lines of corn. In Proceedings of the 34th Annual Hybrid
Corn Industry Research Conference. pp. 5775. (Pioneer Hi-Bred:
Washington, DC)
Hallauer AR, Miranda Filho JB (1988) Quantitative genetics in maize
breeding. 2nd edn (Iowa State University Press: Ames, IA)
Henderson CR (1974) General exibility of linear model techniques for sire
evaluation. Journal of Dairy Science 57, 963972. doi:10.3168/jds.
S0022-0302(74)84993-3
Hull FH (1945) Recurrent selection for specic combining ability in corn.
Journal - American Society of Agronomy 37, 134145.
Lile SM, Hallauer AR (1994) Relation between S2 and later generation
testcrosses of two corn populations. The Journal of the Iowa Academy
of Science 101, 1923.
Littell RC, Milliken GA, Stroup WW, Wolnger RD, Schabenberger O (2006)
SAS for mixed models. 2nd edn (SAS Institute Inc.: Cary, NC)
Menkir A, Kling JG (1999) Effect of reciprocal recurrent selection on grain
yield and other traits in two early-maturing maize populations. Maydica
44, 159165.
Mihaljevic R, Schon CC, Utz HF, Melchinger AE (2005) Correlations and
QTL correspondence between line per se and testcross performance for
agronomic traits in four populations of European maize. Crop Science
45, 114122.
Mrode RA (2005) Linear models for the prediction of animal breeding
values. 2nd edn (CABI Publishing: Oxfordshire, UK)
Mulamba NN, Mock JJ (1978) Improvement of yield potential of Eto Blanco
maize (Zea mays L.) population by breeding for plant traits. Egyptian
Journal of Genetics and Cytology 1, 4057.
Obaidi M, Johnson BE, Van Vleck LD, Kachman SD, Smith OS (1998)
Family per se response to selling and selection in maize based on testcross
performance: a simulation study. Crop Science 38, 367371. doi:10.2135/
cropsci1998.0011183X003800020016x
Patterson HD, Thompson R (1971) Recovery of inter-block information when
block sizes are unequal. Biometrika 58, 545554. doi:10.1093/biomet/
58.3.545
Piepho HP, Mhring J, Melchinger AE, Bchse A (2008) BLUP for
phenotypic selection in plant breeding and variety testing. Euphytica
161, 209228. doi:10.1007/s10681-007-9449-8
Schaeffer LR (1984) Sire and cow evaluation under multiple trait models.
Journal of Dairy Science 67, 15671580. doi:10.3168/jds.S0022-0302
(84)81479-4
Viana JMS, Almeida IF, de Resende MDV, Faria VR, Silva FF (2010a) BLUP
for genetic evaluation of plants in non-inbred families of annual crops.
Euphytica 174, 3139. doi:10.1007/s10681-009-0094-2
Viana JMS, Sobreira FM, Resende MDV, Faria VR (2010b) Multi-trait BLUP
in half-sib selection of annual crops. Plant Breeding 129, 599604.
doi:10.1111/j.1439-0523.2009.01745.x
Viana JMS, Faria VR, Silva FF, Resende MDV (2011) Best linear unbiased
prediction and family selection in crop species. Crop Science, (In press).
Wright AJ (1980) The expected efciencies of half-sib, testcross and S1
progeny testing methods in single population improvement. Heredity 45,
361376. doi:10.1038/hdy.1980.78
522
J. M. S. Viana et al.
Appendix 1.
ASRemL code for bi-trait analysis considering the same trait in S3 family and in testcross
BLUP analysis, S3 families and testcrosses, Beija-Flor population, program 2 (title line)
a 306 !p # additive value/gca effect
avd 306 # average dominance value
env 3 # environment
pop 7 # population
blof 21 # block relative to family test
evf !m0 # ev of S3 family; !m0 recodes 0 as missing values
yf !m0 # yield of S3 family
s # nal stand
m # grain moisture
rep 3 # replication
blot 36 # block relative to testcross test
evt !m0 # ev of testcross
yt !m0 # yield of testcross
ped.txt # pedigree le
A.grm # additive relationship matrix
D.grm # dominance relationship matrix
data.asd !asuv # data le; !asuv allows an error variance other than IS
!ddf # correction of d.f. by the Kenward-Roger method
!continue # more than 10 iterations (default), if necessary
!maxit 100 # maximum number of iterations
evf evt ~Trait Tr.pop at(Trait,2).env at(Trait,2).env.rep !r at(Trait,1).blof at(Trait,2).env.rep.blot Tr.a at(Trait,2).a.env at(Trait,1).avd !f mv # mv estimate
missing values
#yf yt ~Trait at(Trait,2).pop at(Trait,2).env at(Trait,2).env.rep Tr.s Tr.m !r at(Trait,2).env.rep.blot Tr.a at(Trait,2).a.env !f mv
predict a !vpv # !vpv save the prediction error variance matrix
!pin !dene
R gc 7 : 9 # to compute the standard error of the genetic correlation
2 1 5 # one R structure (diagonal) and ve G structures
0 0 id !S2 = 5 # initial value for evf error variance
0 0 id !S2 = 7 # initial value for evt error variance
at(Trait,1).blof 1
at(Trait,1).blof 0 id 1 # 1 is the initial value of var.(blof) for evf
at(Trait,2).env.rep.blot 1
at(Trait,2).env.rep.blot 0 id 1
Tr.a 2
Trait 0 us 5 1 1 # var.(evf) cov(evf, evt) var.(evt); initial values
a 0 ainv # ainv or giv1
at(Trait,2).a.env 1
at(Trait,2).a.env 0 id 2
at(Trait,1).avd 1
at(Trait,1).avd 0 giv2 10
The qualier !m0 is optional, since missing values can be specied as dot (.). In this case the qualier !mvinclude must be included
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