A New Maize Heterotic Pattern between Temperate and

Tropical Germplasms
X. M. Fan,* H. M. Chen, J. Tan, C. X. Xu, Y. M. Zhang, Y. X. Huang, and M. S. Kang

ABSTRACT
New heterotic groups are needed for increasing genetic diversity and productivity in maize (Zea mays L.). A study was conducted
with the following objectives: (i) to determine if exotic tropical maize germplasm (TRMG) can increase diversity of Chinese
maize germplasm for grain yield and five yield component traits (YCTs), namely, ear length (EL), ear diameter (ED), number of

Corn
kernel rows per ear (RE), number of kernels per row (KR), and 1000-kernel weight (TKW), and (ii) to estimate general combin-
ing ability (GCA) and specific combining ability (SCA) effects to identify a possible new heterotic pattern. Twenty-five temperate
maize germplasm (TEMG) from both China and the United States were crossed with four TRMG from the International Maize
and Wheat Improvement Center (CIMMYT). The 100 testcrosses were evaluated in field trials for grain yield and five YCTs at
three locations in Yunnan, China. Tropical lines YML146 and YML145 showed, in crosses with most of the TEMG, significant
positive SCA effects for grain yield and most of the YCTs, indicating that these lines had a different genetic base from that of
TEMG and could increase genetic diversity of Chinese maize germplasm. The GCA effects were more important and reliable
than SCA effects for heterotic pattern classification. Exotic line YML146 (derived from Suwan1) was identified as a new heterotic
group, different from Reid, Lancaster, Tangsipingtou (TSPT), and Luda Red Cob (LDRC). A new heterotic pattern of temperate
× Suwan1 was postulated, which could be as important as the widely accepted fl int × dent heterotic pattern.

N arrow genetic base is one of the most important

limiting factors for maize yield improvement and is a
bottleneck in China maize breeding programs (Zeng, 1990;
Wang et al., 1997; Xu, 2003). A study showed that 71% of
the commercial hybrids grown in China came from four
inbred lines or their derivatives, namely, Mo17, Huangzao 4,
Dan 340, and Yie 478 (Zhang et al., 2000). Therefore, it is
necessary to broaden the genetic base for improving maize
productivity to meet continuously increasing food demand in
China. An effective way to solve this problem is to introduce
exotic maize germplasms and introgress some useful genes
into locally adapted germplasms (Albrecht and Dudley, 1987;
Abadassi and Hervé, 2000; Li et al., 2001; Fan et al., 2002a,
2002b; Reif et al., 2004).
Maize heterotic patterns and heterotic groups have been
extensively studied all over the world (Kauffman et al., 1982;
Vasal et al., 1992; Fan et al., 2002a, 2002b; Huang and Li,
2002; Yuan et al., 2002; Menkir et al., 2004; Melani and
Carena, 2005; Barata and Carena, 2006) and new heterotic
patterns have been widely searched and studied in the United
States (Kauffman et al., 1982; Goodman, 1985; Godshalk
X.M. Fan, H.M. Chen, J. Tan, C.X. Xu, and Y.X. Huang, Institute of
Food Crops, Yunnan Academy of Agricultural Sciences, Kunming 650205,
Yunnan Province, China; Y.M. Zhang, Research Analyst, Parts Distribution
Center, John Deere Co., Milan, IL 61264; and M.S. Kang, Vice Chancellor,
Punjab Agricultural Univ., Ludhiana 141 004, India. Received 5 Sept. 2007.
*Corresponding author (fanxingm@public.km.yn.cn).
Published in Agron. J. 100:917–923 (2008).
doi:10.2134/agronj2007.0298
Copyright © 2008 by the American Society of Agronomy,
677 South Segoe Road, Madison, WI 53711. All rights
reserved. No part of this periodical may be reproduced
or transmitted in any form or by any means, electronic
or mechanical, including photocopying, recording, or
any information storage and retrieval system, without
permission in writing from the publisher.
and Kauffman, 1995; Melani and Carena, 2005). Two new
heterotic patterns, namely, Leaming × Reid Yellow Dent
and Leaming × Lancaster, were found by Kauffman et al.
(1982). Melani and Carena (2005) identified BS21(R)C7 ×
CGL(S1-S2)C5 and BS22(R)C5 × LEAMING(S)C5 as alter-
native heterotic patterns to Iowa Stiff Stalk Synthetic (BSSS) ×
Lancaster. The identification of these new heterotic groups has
greatly helped in development of high grain yield maize hybrids
and inbred lines with high levels of abiotic and biotic stress tol-
erance for the U.S. Corn Belt.
Fan et al. (2002a) used a diallel mating design to evaluate
the combining ability of 10 quality protein maize lines. Five
of these lines were introduced from CIMMYT during 1980s
and have undergone domestication and improvement for
adaptation to local environment by the Yunnan Academy of
Agriculture Science (YAAS). The other five lines were domestic
elite quality protein maize lines selected from across China.
Based on the SCA effects and mean grain yields of 45 crosses,
these10 quality protein maize lines were classified into four
heterotic groups with CIMMYT germplasms being classified
into groups different from China local maize lines, indicating
different genetic base of CIMMYT lines from that of China
local lines.
Huang and Li (2002) evaluated 45 maize inbred lines from
China, U.S. Cornbelt, and tropical regions with 44 restriction
fragment length polymorphism (RFLP) markers equally dis-
tributed on all 10 chromosomes of maize. The 45 inbred lines
Abbreviations: AFLP, amplified fragment length polymorphism; BSSS,
Iowa Stiff Stalk Synthetic; CIMMYT, International Maize and Wheat
Improvement Center; YAAS, Yunnan Academy of Agriculture Science; ED,
ear diameter; EL, ear length ; GCA, general combining ability; GS, genetic
similarity; KR, number of kernel per row; LDRC, Luda Red Cob; RFLP,
restriction fragment length polymorphism; RE, number of kernel row per
ear; SCA, specific combining ability; SSR, simple sequence repeat; TEMG,
temperate maize germplasm; TKW, thousand kernel weight; TRMG, tropical
maize germplasm; TSPT, Tangsipingtou; YCT, yield component traits.

A g r o n o my J o u r n a l • Vo l u m e 10 0 , I s s u e 4 • 2008 917
were grouped into six heterotic groups: of which three heterotic effects. Thus, they recommended that the marker-based grouping
groups were related to Mo17, B73, and Oh43; two groups were techniques might only serve as the basis to design and carry out
classified into two China local heterotic groups; and one group combining ability studies in the field to establish clearly defined
was identified as tropical group. This study suggested that the heterotic groups with a greater GS within groups.
China local maize germplasms were genetically different from Barata and Carena (2006) attempted to classify some elite
U.S.-originated maize germplasms and the TRMGs were geneti- North Dakota maize inbred lines into current U.S. Corn Belt
cally different from both U.S. and China local germplasms. heterotic groups and evaluated the consistency between SSR
Yuan et al. (2002) employed 51 simple sequence repeat (SSR) grouping and testcross data analysis. Thirteen North Dakota
markers to analyze 134 maize inbred lines from both temper-
maize inbred lines representing diverse genetic background
ate regions (China and U.S. Corn Belt) and CIMMYT. The
were crossed in a diallel mating design in 2000. In addi-
134 maize inbred lines were grouped into nine clusters accord-
tion, inbred lines representing Lancaster Sure Crop, BSSS,
ing to genetic similarity (GS) values between the inbred lines.
They found that most of the inbred lines from CIMMYT were Minnesota #13, Northwester Dent, Golden Glow pedigree and
assigned to one major cluster whereas all inbred lines that origi- all the 13 North Dakota inbred lines were screened with 49 SSR
nated from both China and U.S. Corn Belt were placed into markers. The study showed that heterotic groups of genetically
the remaining eight clusters. This result further suggested that similar germplasms could not be identified accurately and reliably
TRMGs might be genetically different from TEMGs originat- with molecular markers even when the available germplasm was
ing from both China and the U.S. Corn Belt. diverse, contrary to what had previously been suggested. Therefore,
Menkir et al. (2004) used two testers representing the flint they suggested that extensive field evaluation was needed to clas-
and dent heterotic patterns to test 38 tropical maize inbred lines. sify unrelated maize inbred lines into a heterotic group.
Significant GCA and SCA effects for grain yield were detected Genetic diversity is a basis for maize breeding program
in tested inbred lines. The two testers classified 23 of the 38 to achieve success (Hallauer and Miranda, 1988; Fan et al.,
tested inbred lines into two heterotic groups based on SCA 2002b; Melani and Carena, 2005). Previous studies have
effects and testcross’s mean grain yields. Diversity analysis of all shown that genetic base of TRMGs was different from that
40 maize inbred lines using amplified fragment length polymor- of TEMGs (Fan et al., 2002a; Huang and Li, 2002; Yuan et
phism (AFLP) and SSR markers classified the tested lines into al., 2002). The tropical lines YML145 and YML146 had been
different heterotic groups obtained by mean grain yield and SCA introduced and improved for adaptation, high yield, disease
resistance and other stress resistances in the last decade by our
Table 1. Germplasm sources and main characters of 29 maize institution (Fan et al., 2002a, 2002b, 2003). For effective uti-
inbred lines.
lization, the classification and determination if the new intro-
Code Inbred Heterotic Grain
no. line† Source group texture duced TRMGs belongs to a new alternative heterotic group
1 CML171 Pool25QPM Tropical Flint becomes critical in our maize breeding programs. These can
2 CML166 Pop66QPM Tropical Dent then be effectively used to synthesize populations, to develop
3 YML145 Improved yellow Tuxpeno Tropical Dent inbred lines and hybrids. The objectives of this study were to
4 YML146 Suwan1 Population of Thailand Tropical Flint
Huangzao4 × (i) evaluate the combining ability of grain yield and five YCTs,
5 81515 TSPT Flint
(HuaFeng100 × Ai C103) namely, EL, ED, RE, KR, and TKW using a line × tester
6 Qi319 Pioneer78599 Reid Flint
mating design from four TRMGs and 25 TEMGs to see if
7 Yuzi87–1 Hybrid 87001 from US Reid Middle
8 P138 Pioneer78599 Reid Flint TRMGs can increase the genetic diversity of China local maize
9 X178 Pioneer78599 Reid Flint germplasm; (ii) analyze the patterns of GCA and SCA effects
10 Shen136 Double cross hybrid from US Reid Flint to see if a new heterotic pattern between tropical and tempera-
11 543 Pioneer78599 Reid Flint
12 Tie7922 Pioneer3382 Reid Dent
ture maize can be established.
13 Ye107 Foreign Hybrid XL80 Reid Semi-dent
14 Luyuan92 Yuanqi123 × 1137 Reid Semi-dent MATERIALS AND METHODS
15 Zheng58 Mutation of Ye 478 Reid Middle Experimental Materials
16 DS01 Hybrid CM190 from US Reid Dent
17 Yun147 Hybrid from US Reid Flint
The sources, the heterotic group, and grain texture of 25
18 Liao3180 Foreign Hybrid Reid Flint TEMGs and four elite TRMGs used for this study are listed in
19 K22 K11 × Ye478 Reid Flint Table 1. These 25 TEMGs are the major parental lines for the
20 Lai3189 5003 × U8112 TSPT Flint commercial maize hybrids currently grown in China. These
21 Mo17 C103 × 187–2 Lancaster Semi-dent
22 13247 1324 × Ye478 Lancaster Middle lines had been classified into different heterotic groups in previ-
23 Zi330 OH43 × Keli 67 Lancaster Dent ous studies (Zhang et al., 2000; Huang and Li, 2002; Yuan et
24 Tie9010 Kang1 × Dan340 LDRC Middle al., 2002). Two of the four tropical inbred lines, CML171 and
λ irradiation mutant
25 Dan340 LDRC Middle CML166, are improved inbred lines originally introduced from
Baizhoulu9 × Pod corn
26 Zong3 synthetic cultivar N/A Dent CIMMYT by YAAS (Fan et al., 2002a). The other two tropi-
27 Xi502 Dan340 × Huangzao4 TSPT Dent cal inbred lines, YML145 and YML146, were selected from
28 K1218 Variation of K12 TSPT Flint
29 Huangzao4 Tangsipingtou TSPT Flint
tropical populations of Tuxpeno and Suwan1, respectively.
† CML and YML refer to the inbred lines of CIMMYT Maize Line and Maize Line CML171 and YML146 are tropical flint inbred lines, whereas
from Yunnan Academy of Agricultural Sciences, respectively. CML166 and YML145 are tropical dent inbred lines.

918 Agronomy Journal • Volume 100, Issue 4 • 2008

Table 2. Experimental field location, preparation, and Table 3. Mean square from ANOVA analysis of 100 test-crosses for yield
weather information. and yield components under three environmental conditions of Yunnan
province in 2005.†
Parameter/cultural Location name
practice Kunming Dehong Linchang Yield
Sources DF plant–1 EL ED RE KR TKW
Elevation, m 1960 913 1750
Loc. 2 22920.06** 177.08** 6.91** 9.53** 259.56** 273735.06**
Longitude 102°27´0˝ 98°20´60˝ 100°1´12˝
Rep(Loc.) 6 567.21** 1.35** 0.02 0.67 3.81 108.88
Latitude 25°1´12˝ 24°15´36˝ 23°25´12˝
Crosses 99 2392.57** 17.39** 0.73** 25.01** 69.00** 11550.16**
Mean annual temperature, °C 14.9 19.1 17.2
MGCA 3 34632.42** 68.45** 9.19** 457.62** 599.01** 227869.56**
Mean temperature in 2005, °C 16.7 20.6 16.1
FGCA 24 3200.57** 45.92** 1.60** 39.22** 114.21** 12477.97**
Mean annual rainfall, mm 1031 1653 1750
M × F, SCA 72 779.91** 5.75** 0.09** 2.25** 31.85** 2227.58**
Mean rainfall for 2005, mm 976 1405 1588
Cross × Loc. 198 2135.86** 5.84** 0.14** 1.56** 29.24** 4006.67**
Annual sunshine, h 2054 2385 2115.9
MGCA × Loc. 6 37249.27** 31.52** 2.01** 0.62** 140.28** 44997.66**
Previous crop wheat corn oil crop
FGCA × Loc. 48 1903.12** 5.82** 0.17** 2.38** 34.34** 4504.72**
Soil texture loam loam loam
SCA × Loc. 144 750.38** 4.77** 0.06** 1.33** 22.92** 2132.70**
Planting date 3 May 2 July 12 June
Fertilizer applied at planting 20 kg 40 kg 20 kg Error 594 82.09 0.46 0.02 0.74 2.49 257.98
(NPK-21–6-13) per mu† ** Refers to 0.01 significance probability level.
Fertilizer applied at seedling 9.2 kg 9.2 kg 9.2 kg † Loc.: location; Rep(Loc.): replications under each location; MGCA: general combining ability
for male parent; FGCA: general combining ability for female parent; DF: degree of freedom;
stage (N by urea) per mu EL: ear length; ED: ear diameter; RE: number of kernel row per ear; KR: number of kernel per
Fertilizer applied at ear develop- 9.2 kg 13.8 kg 13.8 kg row; TKW: thousand kernel weight. CV for Yield, EL, ED, RE, KR, and TKW are 6.60, 4.01,
ment (N by urea) per mu 2.97, 5.64, 4.56, and 5.74, respectively.
†, 1 mu = 1/6 acre.

Experimental Design for Field Trial
In 2004, at Kunming, China, the 25 TEMG’s were used as
female parents and were crossed with the four TRMG, which
yielded 100 testcrosses. In 2005, the 100 testcrosses were field-
tested for grain yield and five YCTs at three locations in Yunnan.
A widely grown commercial hybrid, Yunyou 21, was employed
as a check. A randomized complete-block design with three
replications was used at all locations. Each experimental unit
was a single row plot with a row spacing of 0.7 m and length of
5 m. Distance between two adjacent plants was 0.23 m and the
population density was approximately 60,000 plants ha–1. At
maturity, 10 ears were harvested from 10 consecutive plants in
the middle of each row. The EL, ED, RE, and KR were recorded
from each harvested ear. After harvest, the kernels were air-dried
until constant moisture of 130 g kg–1 was achieved, and then
TKW and grain yield per plant were determined.
Climatic Parameters for the Three Locations
Three locations with different climatic and geographical
conditions were selected to test grain yield and other differ-
ences among hybrids developed by crossing local lines with
CIMMYT lines. Information on elevation, longitude, latitude,
temperature, rainfall, sunshine, soil properties, previous crop,
and fertilizer application for the three locations is presented
in Table 2. The hybrid seeds were hand-planted at the three
locations.
Statistical Model and Analyses
The following statistical model was used for the data
analysis:
Yijkl = μ + αl + bkl + vij + (αυ)ijl + eijkl
vij = gi + g j + sij
where Yijkl = observed value from each experimental unit; μ =
population mean; αl = location effect; bkl = block or replication
effect within each location; vij = F1 hybrid effect = gi + g j + sij
[where gi = general combining ability (GCA) for the ith paren-
tal line; g j = GCA effect of jth tester; sij = specific combining
ability (SCA) for the ijth F1 hybrid]; (αv)ijl = interaction effect
between ith F1 hybrid and lth location; eijkl = residual effect.
Because the three locations selected for this experiment were
not a random sample of all possible locations within Yunnan,
we treated locations as a fi xed factor. Hybrid or cross effect and,
consequently, GCA and SCA effects were regarded as fi xed
effects. Only replication was considered to be a random factor.
Thus, significance of location variance was tested against rep-
lication-within-location entity. For all other significance tests,
experiment error term was used (Table 3).
Data collected from the trials were first analyzed via the
GLM procedure of SAS 9.1 (SAS Institute, 2002). The GCA
and SCA were analyzed via a specific SAS program developed
by our team. Graphs were made using Statistica 7.1 (StatSoft,
Inc. 2006) and Microsoft Excel 2003.
RESULTS AND DISCUSSION
Analysis of Variance of the 100 Testcrosses for
Grain Yield and Yield Component Traits
The data on grain yield and the five YCTs from the three
locations were subjected to an ANOVA (Table 3). The loca-
tions, crosses, and crosses × locations sources of variation were
significant at the 0.01 probability level for all traits studied.
As the variances of crosses and crosses-by-locations interaction
were significant, the crosses variation was further partitioned
into GCA and SCA and the interaction variation was further
partitioned into GCA × location and SCA × location interac-
tions. The GCA, SCA, GCA × locations, and SCA × locations
were also significant for all traits analyzed (Table 3).
General Combining Ability Analysis
The GCA effects for grain yield and the five YCTs for the 25
TEMGs and four TRMGs are listed in Table 4. The tropical inbred
lines YML146 and YML145 had significantly positive GCA
effects for grain yield. When these two lines were crossed with all
TEMGs, the average grain yield from these crosses was significantly
higher than the yields from the crosses of CML166 and CML171,
which had significantly negative GCA effects, with all TEMGs.
Among the TEMGs, Zheng58, Tie 7922, Liao 3180, K1218,
Zi330, Dan340, Yun147, and DS01 had significantly positive
GCA effects, whereas Lai3189, Luyuan92, X178, Mo17, Tie9010,

Agronomy Journal • Volume 100, Issue 4 • 2008 919

Table 4. General combining ability (GCA) effects of grain yield Shen136, K22, and P138 had significantly negative GCA effects.
and yield component traits of 25 temperate inbred lines and four
tropical testers. The GCA effects for grain yield did not reach significant level for
Code Inbred Yield
remainder of the TEMGs.
no. lines plant–1 EL† ED RE KR TKW The GCA effects for the five YCTs are also listed in Table 4.
1 CML171 –4.76** –0.50** –0.23** –1.24** –0.02 10.18** The YML146 had significantly positive GCA effects for all
2 CML166 –14.59** –0.45** 0.05** 1.84** –0.83** –44.30** YCTs except KR and YML145 showed significantly positive
3 YML145 5.24** 0.46** –0.07** –1.00** 2.29** 3.11**
4 YML146 14.10** 0.49** 0.25** 0.40** –1.44** 31.02** GCA effects for three of the five YCTS, that is, EL, KR, and
5 81515 0.72 0.51** –0.22** –1.40** 0.89** 24.13** TKW. Because these two lines had significantly positive GCA
6 Qi319 –0.61 1.54** –0.06* –0.58** 1.28** 32.54** effects for most of the YCTs as well as for grain yield, these
7 Yuzi87–1 2.81 –0.67** 0.24** 0.73** –1.33** 28.77** two tropical lines might be used in maize hybrid development
8 P138 –3.41* –2.56** 0.11** –0.45** –2.25** –4.48
9 X178 –11.34** –0.62** –0.13** 0.82** –0.52* –21.64** programs directly. The two other tropical lines, CML166 and
10 Shen136 –4.87** –0.66** –0.01 –0.44** 1.13** –21.87** CML171, had significantly positive GCA effects for ED, RE,
11 543 –2.57 0.32** –0.01 –1.28** 1.71** –1.60 and TKW, respectively. Thus, the tropical lines CML166 and
12 Tie7922 12.14** 0.25* 0.03 0.50** 3.34** –6.99**
CML171 could be used for line or population improvement
13 Ye107 –2.57 0.32** –0.01 –1.29** 1.71** –1.60
14 Luyuan92 –18.36** 0.32** –0.16** –0.45** 1.66** –16.31** for ED, RE, and TKW, respectively.
15 Zheng58 18.69** 1.08** 0.09** 0.02 2.54** 21.93** A line which had significantly positive GCA effect for grain
16 DS01 3.12* 0.97** –0.09** –0.65** 1.93** –0.48 yield, had usually more YCTs showing significantly positive
17 Yun147 3.99** 2.06** –0.33** –1.49** 2.52** 8.24**
18 Liao3180 10.69** 1.39** 0.09** 0.15 –0.48 19.17**
GCA effects. Whereas those with significantly negative GCA
19 K22 –4.55** –0.74** 0.23** 1.78** –3.27** –14.77** effect for grain yield generally had more YCTs showing signifi-
20 Lai3189 –23.29** 0.23* –0.41** –1.42** –0.09 –20.23** cantly negative GCA effects (Table 4). These results suggested
21 Mo17 –8.63** –0.38** –0.38** –1.22** –0.13 –10.25** that grain yield was closely related to the YCTs and for the
22 13247 2.36 –0.96** 0.23** 1.46** –1.56** –7.03**
23 Zi330 9.23** 0.84** –0.13** 0.38** –0.75** –13.16**
development of an inbred line having higher positive GCA
24 Tie9010 –4.97** –0.87** 0.14** 1.72** –0.82** –30.39** effect for grain yield, it is best to improve all of its YCTs.
25 Dan340 9.17** –0.38** 0.48** 2.10** –0.94** 2.50
26 Zong3 0.27 1.51** –0.01 –0.25 0.52* 0.60 Specific Combining Ability Analysis
27 Xi502 0.19 –1.61** 0.20** 0.07 –2.82** 2.82
28 K1218 9.36** 0.06 0.19** 0.81** –1.58** 29.61**The SCA effects of the crosses of 25 TEMGs with four
29 Huangzao4 3.82 –2.24** 0.41** 1.65** –3.06** TRMGs for grain yield are given in Table 5. Twenty-seven
–19.02**
* Refers to 0.05 significance probability level. testcrosses showed significantly positive SCA effects and the
** Refers to 0.01 significance probability level. numbers of crosses with significantly positive SCA effects
† EL: ear length; ED: ear diameter; RE: number of kernel row per ear; KR: number of for the four tropical lines of YML146, YML145, CML166,
kernel per row; TKW: thousand kernel weight. and CML171 were 7, 8, 6, and 6, respectively. Twenty-nine
Table 5. Grain yield and specific combining ability (SCA) effects of all crosses. testcrosses showed significantly negative
Temperate
SCA effects and the numbers of crosses
inbred lines Tropical inbred lines with significantly negative SCA effects
CML171 CML166 YML145 YML146 for the four tropical lines of YML146,
Line Inbred Yield Yield Yield Yield YML145, CML166, and CML171 were
no. lines plant–1 SCA plant–1 SCA plant–1 SCA plant–1 SCA 9, 7, 8, and 5, respectively. The number of
g g g g crosses with both significantly positive
5 81515 142.95 9.61** 115.31 –8.20** 132.00 –11.34** 162.12 9.92**
6 Qi319 140.34 8.32** 114.66 –7.52** 137.42 –4.59 154.66 3.79
and negative SCA effects did not show any
7 Yuzi87–1 135.05 –0.38 123.83 –1.77 160.17 14.74** 141.70 –12.59** statistical difference for the four tropical
8 P138 132.86 3.66 121.37 1.99 147.39 8.18** 134.23 –13.83** lines. This result suggested that though
9 X178 117.93 –3.35 117.37 5.92* 134.31 3.03 134.55 –5.59* the GCA effects for the four lines were
10 Shen136 125.27 –2.48 109.27 –8.65** 147.55 9.79** 147.95 1.34
11 543 127.50 –2.54 131.01 10.79** 140.24 0.19 140.47 –8.44** different (Table 4), the GCA effect of a
12 Tie7922 135.30 –9.46** 136.82 1.89 157.55 2.79 168.40 4.78 line seemed not to be related to the SCA
13 Ye107 139.38 1.56 128.5 0.51 139.58 –8.25** 162.87 6.18* effects, indicating that the direction and
14 Luyuan92 126.28 12.02** 104.7 0.26 104.62 –19.65** 140.49 7.37** magnitude of SCA effect of a cross could
15 Zheng58 144.40 –6.91** 142.59 1.11 146.45 –14.86** 190.83 20.66**
16 DS01 141.88 6.14* 115.87 –10.04** 142.21 –3.53 162.03 7.43** not be predicted from the GCA effects of
17 Yun147 134.73 –1.88 132.89 6.11* 148.02 1.41 149.81 –5.65* the parental lines.
18 Liao3180 143.92 0.61 132.64 –0.84 160.36 7.05** 155.34 –6.82**
19 K22 127.40 –0.67 115.75 –2.49 144.05 5.97* 144.12 –2.81
Dent × Flint Heterotic Pattern
20 Lai3189 113.65 4.32 110.89 11.39** 111.13 –8.21** 120.68 –7.51**
21 Mo17 111.90 –12.08** 124.8 10.65** 125.62 –8.36** 152.64 9.80** in the Crosses between Tropical
22 13247 143.40 8.42** 130.06 4.91 137.59 –7.40** 147.91 –5.93* Maize Germplasms and
23 Zi330 146.62 4.77 115.98 –16.04** 153.57 1.71 170.27 9.56** Temperate Maize Germplasms
24 Tie9010 131.59 3.94 115.19 –2.62 136.29 –1.36 146.54 0.04
25 Dan340 152.08 10.29** 125.32 –6.64* 148.89 –2.90 159.91 –0.74
Grain yields of all crosses are repre-
26 Zong3 122.41 –10.47** 116.55 –6.51* 159.79 16.90** 151.83 0.08 sented in a scatter plot (Fig. 1). It shows
27 Xi502 131.00 –1.81 130.26 7.28** 148.66 5.85* 140.34 –11.33** that most of the crosses with grain yield
28 K1218 125.54 –16.44** 139.36 –7.21** 164.67 12.69** 157.38 –3.46 higher than the check (average yield for
29 Huangzao4 122.07 –5.19 118.74 1.30 137.41 0.14 149.87 3.75
* Refers to 0.05 significance probability level.
the check was 141.98 g plant–1) were
** Refers to 0.01 significance probability level. crosses between the two tropical exotic

920 Agronomy Journal • Volume 100, Issue 4 • 2008

lines, YML146 and YML145, and TEMGs, suggest-
ing that the exotic lines YML146 and YML145 had
genetic base different from that of the 25 TEMGs.
They most likely belong to a maize heterotic group
different from the heterotic groups that the 25
TEMGs belong to.
To analyze the interaction patterns of the two
tropical lines (YML 146 and YML 145) with
the four current maize heterotic groups of the
25 TEMGs, average GCA and SCA effects of all
crosses were plotted and are shown in Fig. 2 and Fig.
3 for YML146 and YML145, respectively. The tem-
perate Reid group includes 14 inbred lines and eight
of them (i.e., Qi319, P138, X178, Shen136, 543,
Yun147, Liao3180, and K22) are flint. The SCA
effects of YML146, which is flint, with six out of the
eight flint Reid lines were significantly negative. All Fig. 1. Grain yield for all crosses between 25 temperate and four tropical lines.
Red, green, brown, and blue circles represent yields of all crosses between 25
crosses of YML146 with other Reid lines were posi- temperate lines with YML146, YM L145, CML166, and CML171, respectively.
tive except with Yuzi87–1, which had a grain texture
in-between that of flint and Dent. On the other effects can also be used in classifying maize heterotic groups.
hand, the crosses between YML145, a dent line, with flint Reid The GCA effect of a line would be different when evaluated
lines all had positive SCA effects except with line Qi319. These with a different set of lines (i.e., different heterotic groups) with
results indicated that the heterotic pattern between dent and different genetic base. If a line generally has high positive GCA
flint also exits in the crosses between TRMGs and TEMGs. effects with a set of lines from a current maize heterotic group,
it could suggest that this line might belong to a new heterotic
General Combining Ability and group, different from the current heterotic groups.
Specific Combining Ability Effects for To see the difference of GCA and SCA effect when the two
Heterotic Group Classification tropical lines, YML146 and YML145, were crossed with the
The SCA effects and mean grain yield have been widely lines from different sets of heterotic groups, we had conducted
used for classifying maize heterotic group (Kauffman et al., combining ability analyses between the two tropical lines and
1982; Menkir et al., 2004; Melani and Carena, 2005; Fan et the lines belonging to each of the four sets of known heterotic
al., 2006). On the basis of Griffing’s (1956) model, mean grain groups. The average GCA and SCA effects of the crosses of
yield consisted of both SCA and GCA effects. Thus, GCA YML146 and of YML145 with the four different sets of inbred

Fig. 2. Combining abilities of YML146 with all 25 tested lines. Fig. 3. Combining abilities of YML145 with all 25 tested lines.

Fig. 4. Combing abilities of YML146 with lines belong to differ- Fig. 5. Combing abilities of YML145 with lines belonging to dif-
ent heterotic groups. ferent heterotic groups.

Agronomy Journal • Volume 100, Issue 4 • 2008 921

lines are shown in Fig. 4 and Fig. 5, respectively. These two
figures showed that when a line was crossed with different sets
of maize germplasms grouped by the four heterotic groups, not
only the SCA effects of its crosses but also its GCA effects var-
ied. Comparing the GCA and SCA effects between Fig. 2 and
Fig. 4 and between Fig. 3 and Fig. 5, we found that not only the
absolute values, but also the relative sizes of SCA and GCA effects
had changed. These results strongly suggested that GCA effects
should be considered in maize heterotic group classification.
Figures 4 and 5 revealed that the average GCA effects of
YML146 and YML145 were clearly different when they
crossed with lines from the four known heterotic groups. For
example, the GCA effects of YML146 with Lancaster and both
Reid flint and Reid dent lines were significantly positive (Fig.
4). In contrast, the GCA effects of YML145 with the Reid
flint lines were significantly positive, but not significant with
the Reid Dent lines and Lancaster lines. This phenomenon
may be due to some different genetic bases between YML146
and YML145. That difference could cause different levels of
gene expression in Reid flint, Reid dent, and Lancaster lines
for controlling key agronomic traits, such as high disease
resistance, strong root system, high resistance to drought and
other stresses, etc. Giauffret et al. (2000) studied a diallel with
five temperate and five tropical maize lines and found that
the best crosses with high grain yield were between temperate
and tropical lines and the grain yield variation was related to
two major adaptation factors, that is, disease resistance and
planting-to-silking duration. Yield variation was also related
to emergence duration or early growth. We did not collect data
on disease resistance and other important agronomic traits;
thus, we could not relate the difference of the GCA effects to
any maize agronomic traits. Further studies are needed to find
which maize agronomic traits are responsible for high GCA
effects between YML146/YML145 and TEMGs.
Figures 4 and 5 also revealed that the SCA effects of the
crosses between YML146/YML145 and the lines from the
same heterotic group could be quite different. For instance,
the SCA effect of YML145 with the Reid dent line Yuzi87–1
was significantly positive, but was significantly negative
with another Reid dent line luyan92. And the SCA effect of
YML146 with Reid flint line P138 was significantly nega-
tive, but was significantly positive with another Reid flint line
Shen136. Thus, GCA effect is a reliable criterion and better
than SCA effect for classifying new germplasms into current
heterotic groups because it gave more consistent results.
A New Heterotic Group: Suwan1
YML146 was selected from tropical maize population
Suwan1 (Table 1). Some studies on heterosis between its
derived lines with TEMGs have previously been done (Zhang
et al., 2004; Fan et al., 2002b). Zhang et al. (2004) using four
testers evaluated maize grain yield of 27 maize populations,
14 of them were from CIMMYT and 13 of them from China.
Their results showed that the GCA effects of the CIMMYT
populations were generally higher than those of domestic
populations, and the crosses between these tropical lines
and TEMGs had higher grain yield than the crosses among
TEMGs. Fan et al. (2002b) employed four testers to evaluate
25 inbred lines from five tropical and subtropical populations.
922
They found that the GCA effects of the inbred lines from
population Suwan1 and Pob28 were much higher than those of
other inbred lines. These results suggested that Suwan1 popula-
tion, the parent population of YML146, was distinguished from
other tropical lines and might belong to a new heterotic group.
The way to use GCA effects to classify a new germplasm
into a current heterotic group is to see if a line’s average GCA
effect is significant when it crosses with several lines from
the same heterotic group. If the average GCA effect is signifi-
cantly positive, it indicates that the new line’s additive genes
are complementary to the lines from the heterotic group and
the new line might belong to an alternative heterotic group.
Because YML146 had a positive average GCA with all lines
from the two major temperate heterotic groups in the U.S.
Corn Belt, Reid and Lancaster, and from the two major China
local heterotic groups, TSPT and LDRC, this tropical line
seemed to belong to a heterotic group different from the Reid,
Lancaster, TSPT, and LDRC. Because YML146 was derived
from Suwan1 population, Suwan1 might be classified as a new
heterotic group different from the four heterotic groups. A new
heterotic pattern can be defined as temperate × Suwan1.
Another tropical line, YML145, had higher GCA effects
with flint lines but no significant GCA effect was detected
when it was crossed with dent lines from Reid dent lines
(Fig. 5). It seemed that the high combining abilities between
YML145 and the lines from Reid groups were caused by the
dent and flint heterotic pattern only. Furthermore, no signifi-
cantly positive GCA effects were found between YML145
and the lines from the Lancaster group. Thus, YML145 might
belong to either Reid dent or Lancaster group based on GCA
effect. Further evidence is needed to ascertain to which het-
erotic group YML145 belongs.
SUMMARY AND CONCLUSIONS
This study had shown that both GCA and SCA played a role
in maize heterotic classification with GCA effects being more
important and reliable than SCA effects. The results from this
study suggested that TRMG had a genetic base different from
that of TEMG. Based on the average GCA values of YML146,
in crosses with lines from the four known heterotic groups,
Suwan1 was identified as a new heterotic group different
from the current heterotic groups of Reid, Lancaster, LDRC,
and TSPT. A new heterotic pattern of temperate × Suwan1
was demonstrated by the crosses of temperature inbred line
× YML146 in this study. The new heterotic pattern may be
as important as the widely accepted heterotic pattern of flint
× dent. Further systematic studies on temperate × Suwan1
heterotic pattern are warranted to be able to exploit this new
heterotic pattern.
ACKNOWLEDGMENTS
The authors express their sincere appreciation for the support from
Yunnan Provincial Key Foundation (2004C0011Z) and Yunnan Key
Science and Technology Development Project Foundation for 11th
5-Year Plan (2006NG06), China. The authors would like to give
great thanks to Prof. Huang Bihua (Dehong Institute of Agricultural
Science of Yunnan province) and Prof. Li Xuezhi (Lincang Institute of
Agricultural Science of Yunnan province) for their help in data collec-
tion for this study.
Agronomy Journal • Volume 100, Issue 4 • 2008
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