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X. M. Fan,* H. M. Chen, J. Tan, C. X. Xu, Y. M. Zhang, Y. X. Huang, and M. S. Kang
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 kernel rows per ear (RE), number of kernels per row (KR), and 1000-kernel weight (TKW), and (ii) to estimate general combining 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.
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 (email@example.com). 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 alternative 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 tolerance 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 distributed 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
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the classification and determination if the new introduced TRMGs belongs to a new alternative heterotic group becomes critical in our maize breeding programs. 2002). This study suggested that the China local maize germplasms were genetically different from U. This result further suggested that TRMGs might be genetically different from TEMGs originating from both China and the U. The two testers classified 23 of the 38 tested inbred lines into two heterotic groups based on SCA effects and testcross’s mean grain yields. Code no. and China local germplasms.. Fan et al. These lines had been classified into different heterotic groups in previous studies (Zhang et al. RE. and TKW using a line × tester mating design from four TRMGs and 25 TEMGs to see if TRMGs can increase the genetic diversity of China local maize germplasm. and grain texture of 25 TEMGs and four elite TRMGs used for this study are listed in Table 1. Germplasm sources and main characters of 29 maize inbred lines.S. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 Inbred line† CML171 CML166 YML145 YML146 81515 Qi319 Yuzi87–1 P138 X178 Shen136 543 Tie7922 Ye107 Luyuan92 Zheng58 DS01 Yun147 Liao3180 K22 Lai3189 Mo17 13247 Zi330 Tie9010 Dan340 Zong3 Xi502 K1218 Huangzao4 Heterotic Source group Pool25QPM Tropical Pop66QPM Tropical Improved yellow Tuxpeno Tropical Suwan1 Population of Thailand Tropical Huangzao4 × TSPT (HuaFeng100 × Ai C103) Pioneer78599 Reid Hybrid 87001 from US Reid Pioneer78599 Reid Pioneer78599 Reid Double cross hybrid from US Reid Pioneer78599 Reid Pioneer3382 Reid Foreign Hybrid XL80 Reid Yuanqi123 × 1137 Reid Mutation of Ye 478 Reid Hybrid CM190 from US Reid Hybrid from US Reid Foreign Hybrid Reid K11 × Ye478 Reid 5003 × U8112 TSPT C103 × 187–2 Lancaster 1324 × Ye478 Lancaster OH43 × Keli 67 Lancaster Kang1 × Dan340 LDRC λ irradiation mutant LDRC Baizhoulu9 × Pod corn synthetic cultivar N/A Dan340 × Huangzao4 TSPT Variation of K12 TSPT Tangsipingtou TSPT Grain texture Flint Dent Dent Flint Flint Flint Middle Flint Flint Flint Flint Dent Semi-dent Semi-dent Middle Dent Flint Flint Flint Flint Semi-dent Middle Dent Middle Middle Dent Dent Flint Flint effects. These can then be effectively used to synthesize populations. EL. In addition. ED. Yuan et al. MATERIALS AND METHODS Experimental Materials The sources. Barata and Carena (2006) attempted to classify some elite North Dakota maize inbred lines into current U. 1988. 2002a. Corn Belt heterotic groups and evaluated the consistency between SSR grouping and testcross data analysis.S. Huang and Li.were grouped into six heterotic groups: of which three heterotic groups were related to Mo17. Golden Glow pedigree and all the 13 North Dakota inbred lines were screened with 49 SSR markers. These 25 TEMGs are the major parental lines for the commercial maize hybrids currently grown in China. contrary to what had previously been suggested. two groups were classified into two China local heterotic groups.. whereas CML166 and YML145 are tropical dent inbred lines. Genetic diversity is a basis for maize breeding program to achieve success (Hallauer and Miranda. (2004) used two testers representing the flint and dent heterotic patterns to test 38 tropical maize inbred lines.S. Yuan et al. they recommended that the marker-based grouping techniques might only serve as the basis to design and carry out combining ability studies in the field to establish clearly defined heterotic groups with a greater GS within groups. Thus. 2002b. Huang and Li. 2002.. For effective utilization. respectively. Menkir et al. (2002) employed 51 simple sequence repeat (SSR) markers to analyze 134 maize inbred lines from both temperate regions (China and U. Two of the four tropical inbred lines.. respectively. Therefore. The objectives of this study were to (i) evaluate the combining ability of grain yield and five YCTs. CML171 and YML146 are tropical flint inbred lines. KR. CML171 and CML166. to develop inbred lines and hybrids. Minnesota #13. (ii) analyze the patterns of GCA and SCA effects to see if a new heterotic pattern between tropical and temperature maize can be established. Northwester Dent.S. high yield.S. Previous studies have shown that genetic base of TRMGs was different from that of TEMGs (Fan et al... and one group was identified as tropical group. 2000. 2002. The 134 maize inbred lines were grouped into nine clusters according to genetic similarity (GS) values between the inbred lines. 2005). BSSS. 2002a). 2003). namely. The other two tropical inbred lines. 2002b.S. are improved inbred lines originally introduced from CIMMYT by YAAS (Fan et al. and Oh43. Corn Belt. they suggested that extensive field evaluation was needed to classify unrelated maize inbred lines into a heterotic group. Corn Belt were placed into the remaining eight clusters.-originated maize germplasms and the TRMGs were genetically different from both U. 2002). The tropical lines YML145 and YML146 had been introduced and improved for adaptation. Agronomy Journal • Volume 100. Issue 4 • 2008 † CML and YML refer to the inbred lines of CIMMYT Maize Line and Maize Line from Yunnan Academy of Agricultural Sciences. They found that most of the inbred lines from CIMMYT were assigned to one major cluster whereas all inbred lines that originated from both China and U. inbred lines representing Lancaster Sure Crop. Significant GCA and SCA effects for grain yield were detected in tested inbred lines.. Yuan et al. Diversity analysis of all 40 maize inbred lines using amplified fragment length polymorphism (AFLP) and SSR markers classified the tested lines into different heterotic groups obtained by mean grain yield and SCA Table 1. disease resistance and other stress resistances in the last decade by our institution (Fan et al. The study showed that heterotic groups of genetically similar germplasms could not be identified accurately and reliably with molecular markers even when the available germplasm was diverse. the heterotic group. YML145 and YML146. 2002a. Thirteen North Dakota maize inbred lines representing diverse genetic background were crossed in a diallel mating design in 2000. Melani and Carena. 918 . B73. Corn Belt) and CIMMYT. were selected from tropical populations of Tuxpeno and Suwan1.
RE.58** 198 2135.64.74 2. Location name Parameter/cultural practice Kunming Dehong Linchang Elevation.6 16. A randomized complete-block design with three replications was used at all locations.01** 69. The hybrid seeds were hand-planted at the three locations. Mo17.19** 457. SCA × Loc.84** 0. 2. and TKW are 6.56** 273735.88 99 2392. Zheng58.39** 0. As the variances of crosses and crosses-by-locations interaction were significant. Tie9010. Among the TEMGs.7 m and length of 5 m. Climatic Parameters for the Three Locations Three locations with different climatic and geographical conditions were selected to test grain yield and other differences among hybrids developed by crossing local lines with CIMMYT lines. previous crop.82** 0.8 kg Fertilizer applied at ear development (N by urea) per mu †.56.85** 2227.97. μ = population mean. MGCA: general combining ability for male parent. The GCA. Rep(Loc. sunshine. 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).21** 1. GCA × locations.) Crosses MGCA FGCA M × F. the average grain yield from these crosses was significantly higher than the yields from the crosses of CML166 and CML171. 4.21** 12477. CV for Yield. 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.09 0.33** 22.56** 24 3200.25** 31. ED.52** 2.35** 0. After harvest. EL.98 ** Refers to 0.62** 599. The locations.24** 4006. China. Tie 7922. Zi330. For all other significance tests.27** 31. longitude.Table 2. Experimental ﬁeld location. vij = F1 hybrid effect = gi + g j + sij [where gi = general combining ability (GCA) for the ith parental line. RE: number of kernel row per ear.06** 6 567. Dan340. Rep(Loc. significance of location variance was tested against replication-within-location entity.14** 1.23 m and the population density was approximately 60. Mean square from ANOVA analysis of 100 test-crosses for yield and yield components under three environmental conditions of Yunnan province in 2005. °C 14.81 108. Because the three locations selected for this experiment were not a random sample of all possible locations within Yunnan.72** 144 750. g j = GCA effect of jth tester.17** 2.77** 0. SCA.9 Previous crop wheat corn oil crop Soil texture loam loam loam Planting date 3 May 2 July 12 June Fertilizer applied at planting 20 kg 40 kg 20 kg (NPK-21–6-13) per mu† Fertilizer applied at seedling 9.1 17. Graphs were made using Statistica 7. Liao 3180.91** 9. sij = specific combining ability (SCA) for the ijth F1 hybrid].7 20.01 probability level for all traits studied. 2002). at Kunming. X178. Only replication was considered to be a random factor. whereas Lai3189.34** 4504. TKW: thousand kernel weight.22** 114.00** 11550. and KR were recorded from each harvested ear. Inc. Information on elevation. Experimental Design for Field Trial In 2004.74. Issue 4 • 2008 between ith F1 hybrid and lth location.2 kg 9.57** 17. and DS01 had significantly positive GCA effects. soil properties. KR. eijkl = residual effect. The EL.16** 3 34632.01. which yielded 100 testcrosses. 1 mu = 1/6 acre. 919 . and SCA × locations were also significant for all traits analyzed (Table 3).92** 1. KR: number of kernel per row. The tropical inbred lines YML146 and YML145 had significantly positive GCA effects for grain yield. preparation. crosses.1 (SAS Institute.02 0.2 kg stage (N by urea) per mu 9.75** 0. At maturity.000 plants ha–1. Thus. 2006) and Microsoft Excel 2003.: location. °C 16. we treated locations as a fi xed factor.02 0.46 0. bkl = block or replication effect within each location.28** 44997. αl = location effect. rainfall.92** 2132. 10 ears were harvested from 10 consecutive plants in the middle of each row.45** 9.66** 48 1903. Distance between two adjacent plants was 0. with all TEMGs. the 25 TEMG’s were used as female parents and were crossed with the four TRMG.70** 594 82. 4. latitude.08** 6. † Loc.49 257.60** 39.01 signiﬁcance probability level. FGCA: general combining ability for female parent.1 Mean annual rainfall. A widely grown commercial hybrid. K1218. EL: ear length. which had significantly negative GCA effects.2 kg 13.86** 5.42** 68. mm 1031 1653 1750 Mean rainfall for 2005. MGCA × Loc. 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. In 2005.9 19.67 3. experiment error term was used (Table 3).73** 25.67** 6 37249. and then TKW and grain yield per plant were determined. and 5. 5. the crosses variation was further partitioned into GCA and SCA and the interaction variation was further partitioned into GCA × location and SCA × location interactions.09** 2. temperature.† Sources Loc. ED: ear diameter.60. SCA Cross × Loc.62** 140.53** 259. When these two lines were crossed with all TEMGs. consequently. GCA and SCA effects were regarded as fi xed effects.56** 29. (αv)ijl = interaction effect Agronomy Journal • Volume 100.06** 1. Yun147.57** 45.01** 227869.2 kg 9.38** 4. Yunyou 21. ED. Each experimental unit was a single row plot with a row spacing of 0. RE. the 100 testcrosses were fieldtested for grain yield and five YCTs at three locations in Yunnan. Data collected from the trials were first analyzed via the GLM procedure of SAS 9. Table 3.06** 177. Hybrid or cross effect and.38** 34. Error Yield DF plant–1 EL ED RE KR TKW 2 22920.): replications under each location.2 Mean temperature in 2005. Luyuan92. h 2054 2385 2115. and weather information.01** 0.97** 72 779. respectively.12** 5. m 1960 913 1750 Longitude 102°27´0˝ 98°20´60˝ 100°1´12˝ Latitude 25°1´12˝ 24°15´36˝ 23°25´12˝ Mean annual temperature. the kernels were air-dried until constant moisture of 130 g kg–1 was achieved. and crosses × locations sources of variation were significant at the 0. and fertilizer application for the three locations is presented in Table 2. FGCA × Loc.8 kg 13. The GCA and SCA were analyzed via a specific SAS program developed by our team.91** 5.1 (StatSoft. was employed as a check. mm 976 1405 1588 Annual sunshine. DF: degree of freedom.
17 Yun147 134.66 3.60 –16.34 8.48 109.81** 1.78 13 Ye107 139.41 131.64 9.01 0.51** –1.14** –2.51 139.41* –11.8 130.Table 4.12* 3.65 111.66** 0.28** –7.65** 4.66** 2.54** 1.09** –0. RE.62 137.13** 1.5 0.29** –0. Because these two lines had significantly positive GCA effects for most of the YCTs as well as for grain yield.21** –8.49** 0.01 signiﬁcance probability level.58** 0.14 155.03 –0.69** –4. and TKW.88 132.01 signiﬁcance probability level.40 4.54 131. RE.47 –8.87 6.02** 104.28 12.66 164. Whereas those with significantly negative GCA effect for grain yield generally had more YCTs showing significantly negative GCA effects (Table 4).94 10.41 149.46 149.27 9. Twenty-seven testcrosses showed significantly positive SCA effects and the * Refers to 0. YML145.72** 2.12 –2.41 7.24** 1.62 –19.95 9.89 157.36** –7.19 9.71** 1.68 –7. for the four tropical lines of YML146. TKW: thousand kernel weight. This result suggested that though 9 X178 117. respectively.05** –0.82** 144. 7. numbers of crosses with significantly positive SCA effects ** Refers to 0.71 –1.44** 0.02** Shen136.50** –1.25** 162. RE: number of kernel row per ear. that is.08 122. lines plant–1 9.16** –30.93 –3.56 128.54** –0.40 146.74 –0. A line which had significantly positive GCA effect for grain yield.77** –4.38** 1.65* 18 19 20 21 22 23 24 25 26 27 28 29 Liao3180 K22 Lai3189 Mo17 13247 Zi330 Tie9010 Dan340 Zong3 Xi502 K1218 Huangzao4 143.05 signiﬁcance probability level.03 7. and TKW.33** 0.77 3.55 9.18* effects.52* 1. 1).93* 170.97** 2.48 8.11** –0.17 14.67 137.41** RE –1.40 –6.08** 8.61 –0.30** 3.84 –2.02 1.90 16.06* 0. and TKW.22** 1.64* –6.91** 142. respectively. had usually more YCTs showing significantly positive GCA effects.00** 0.35 117.82** –0. K22.52* –2.19 140.92 127.11** 31.99 147.13** 0.25* 0.84** –1.67 4.38 –3.45** 0.44** –1.37 5.58** –3.23** 0. and 6.52** 137.22** –0.09** 0.06** 1.36 9.98 115.32** 0.67** –2.83 –1.20** 132.24** 0.07** 0.39 8.14* 115.74 151.59* the GCA effects for the four lines were 10 Shen136 125.79** 147.65** 147. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 Inbred lines CML171 CML166 YML145 YML146 81515 Qi319 Yuzi87–1 P138 X178 Shen136 543 Tie7922 Ye107 Luyuan92 Zheng58 DS01 Yun147 Liao3180 K22 Lai3189 Mo17 13247 Zi330 Tie9010 Dan340 Zong3 Xi502 K1218 Huangzao4 Yield plant–1 –4.98 g plant–1) were crosses between the two tropical exotic • Volume 100.75 * Refers to 0.18** –44. respectively.39** 10.54** 28.11 146.58 –8.42** 4.25 0.87** –1.89 124.04** 142.73** –0.46** 136.69** 0.64 115.82 29. indicating that the direction and 14 Luyuan92 126.31 3.29** –8.28** 0.45** 0. 8.03** –13.30 160.05 signiﬁcance probability level.36** 3.82** –1.83** 2.00 –11.34 different (Table 4). SCA effects and the numbers of crosses Temperate Tropical inbred lines with significantly negative SCA effects inbred lines for the four tropical lines of YML146.08 140.56** –0.93** –0.04 159.91 –5.12 9.02** 24.97** 9.87** –0.06 –2.01 –0.30 –9.13 125.54 0.34 –6.36** 18.25** –0.09 –0.42 –4.87 3.77 160.50 –2.97* –8.05** 5.61 2.36 –2.82 EL† –0.20** 0.36 118. and CML171 were Yield Yield Yield Line Inbred Yield SCA plant–1 SCA plant–1 SCA plant–1 SCA no.33** –2. ** Refers to 0. Thus.44** line seemed not to be related to the SCA 12 Tie7922 135.25** –0. The GCA effects for the five YCTs are also listed in Table 4.19 132.40 113.27 –8.79 148.51* 7.05 111.59** 5.86 3.64** –21.60 –6.06 115. CML166.23* –0. Dent × Flint Heterotic Pattern in the Crosses between Tropical Maize Germplasms and Temperate Maize Germplasms Grain yields of all crosses are represented in a scatter plot (Fig.78** –1.87** –2.83 20.44** –5.50** –0. The GCA effects for grain yield did not reach significant level for remainder of the TEMGs.27** –0.83 0. CML166.61** 0.79** 140.55 130.18** 134.10** –0.45** 0. it is best to improve all of its YCTs.88 6.74** 141.38** 0. KR.74** 0.36 144.39** –0.07 0.77** –20.65** KR –0.62 –6.47** –1.82 1.37 1.50 0.38 1.51** 152.92** and negative SCA effects did not show any 6 Qi319 140. The number of g g g g crosses with both significantly positive 5 81515 142.49 7.49** 0.99** 10.56** –0.16** 0.62 131.66 –7.13** 32.57 136. ED: ear diameter.38** –0. the tropical lines CML166 and CML171 could be used for line or population improvement for ED.33** 157.09** –0. General combining ability (GCA) effects of grain yield and yield component traits of 25 temperate inbred lines and four tropical testers.92* 134.93** 2.65** –1. and 5.29 148.46** 0.31** 21.55** –23.32** 114.70 –12.41** –0.57 –18.02 –0.29** –10.59 153.42** –1.61** 115.90** 5.08** 0.59** 8 P138 132.89** 1.85* 12.79 statistical difference for the four tropical 7 Yuzi87–1 135.24** 14.32 116.04** –2. had significantly positive GCA effects for ED. 6.52** –0.34 –11.15 1.95 1.71** 3.32** 1. the GCA effect of a 11 543 127.01 –0. Grain yield and speciﬁc combining ability (SCA) effects of all crosses.60 2.89 159.66** not be predicted from the GCA effects of 16 DS01 141. Code no.48 –3.24 0.72 –0.27 0.26 139. It shows that most of the crosses with grain yield higher than the check (average yield for the check was 141.38** 1.53 162.29** –1.57 12.62** –0.01 10.19** 0.56** 146.21** 1.23** –10.24** ED –0.34** –4.55 2.80** 147. Issue 4 • 2008 920 Agronomy Journal .40** 1.81 –5. CML171 CML166 YML145 YML146 YML145.59 1.23** –0.7 0.19 125.48** –0.45 –14. † EL: ear length.59 152. EL.32** 0.86** 190.79 168. The two other tropical lines.82** –0.31 –8.17** –14.02 –0.40** –0.40** –1.61** –19.06** TKW 10.55 –5.81 –16.96** 0.48 –21.13** –0. These results suggested that grain yield was closely related to the YCTs and for the development of an inbred line having higher positive GCA effect for grain yield.65** 140.28** –1.37** magnitude of SCA effect of a cross could 15 Zheng58 144.23** –4.75** –0.75 110.51** 1.94** 0.03 134.49 11.39** 2. CML166 and CML171.27 –2.87 –10.23** –0.32 –12. The YML146 had significantly positive GCA effects for all YCTs except KR and YML145 showed significantly positive GCA effects for three of the five YCTS.23 –13.81 120. Twenty-nine testcrosses showed significantly negative Table 5.99** –1.24** 19.59 154.63** 2.07 0.34** 1.14** 0.46** 0.38 123.69** 3.91 –0.54 122.73 –1. respectively. 8.43** the parental lines.11* 148.76** –14.89 6.10** 0.83** lines. and P138 had significantly negative GCA effects. Speciﬁc Combining Ability Analysis The SCA effects of the crosses of 25 TEMGs with four TRMGs for grain yield are given in Table 5. and CML171 were 7.21 –3.00 125.81 –3.90 143.66 121.01 0.17** 0.84** –0.91 –16.25** –7. KR: number of kernel per row.05 –0. these two tropical lines might be used in maize hybrid development programs directly.13 –1.34** 162.26 104.
respectively. which is flint. brown. 2. Speciﬁc Combining Ability Effects for To see the difference of GCA and SCA effect when the two Heterotic Group Classiﬁcation tropical lines. If a line generally has high positive GCA flint also exits in the crosses between TRMGs and TEMGs. YML146 and YML145. YML146 and YML145. Agronomy Journal • Volume 100.lines. The SCA effects of YML146. and CML171. green. Yun147. and TEMGs. Fig. Fig. suggesting that the exotic lines YML146 and YML145 had genetic base different from that of the 25 TEMGs. 4. Melani and Carena. 2005. 2004.. 2006). On the basis of Griffing’s (1956) model. Menkir et al. with six out of the eight flint Reid lines were significantly negative. and K22) are flint. tive except with Yuzi87–1. respectively.. Thus. average GCA and SCA effects of all crosses were plotted and are shown in Fig.e. 3 for YML146 and YML145.e. 3. X178.temperate lines with YML146. the crosses between YML145. CML166. it could suggest that this line might belong to a new heterotic General Combining Ability and group. P138. On the other effects can also be used in classifying maize heterotic groups. a dent line. The average GCA and SCA effects of the crosses of yield consisted of both SCA and GCA effects. YM L145. mean grain groups. and blue circles represent yields of all crosses between 25 crosses of YML146 with other Reid lines were posi. Combining abilities of YML145 with all 25 tested lines. Liao3180. effects with a set of lines from a current maize heterotic group. Grain yield for all crosses between 25 temperate and four tropical lines. Combing abilities of YML146 with lines belong to different heterotic groups. with flint Reid The GCA effect of a line would be different when evaluated lines all had positive SCA effects except with line Qi319. They most likely belong to a maize heterotic group different from the heterotic groups that the 25 TEMGs belong to.. were crossed with the The SCA effects and mean grain yield have been widely lines from different sets of heterotic groups. Fig. Issue 4 • 2008 921 . These with a different set of lines (i. Shen136. different heterotic groups) with results indicated that the heterotic pattern between dent and different genetic base. 5. Red. 543. GCA YML146 and of YML145 with the four different sets of inbred Fig. The temperate Reid group includes 14 inbred lines and eight of them (i. we had conducted used for classifying maize heterotic group (Kauffman et al. combining ability analyses between the two tropical lines and 1982. hand. 1.. Combining abilities of YML146 with all 25 tested lines. 2 and Fig. Fan et the lines belonging to each of the four sets of known heterotic al. 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. different from the current heterotic groups.. which had a grain texture in-between that of flint and Dent. Combing abilities of YML145 with lines belonging to different heterotic groups. All Fig. Qi319.
Fan et al. not only the SCA effects of its crosses but also its GCA effects varied. 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. We did not collect data on disease resistance and other important agronomic traits. Further evidence is needed to ascertain to which heterotic group YML145 belongs. 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. China. and the crosses between these tropical lines and TEMGs had higher grain yield than the crosses among TEMGs. but was significantly negative with another Reid dent line luyan92. that is. Furthermore. In contrast. Issue 4 • 2008 . Some studies on heterosis between its derived lines with TEMGs have previously been done (Zhang et al. and TSPT. 4 and Fig. but was significantly positive with another Reid flint line Shen136. was distinguished from other tropical lines and might belong to a new heterotic group. The results from this study suggested that TRMG had a genetic base different from that of TEMG. (2004) using four testers evaluated maize grain yield of 27 maize populations. 5. For example. but also the relative sizes of SCA and GCA effects had changed. Agronomy Journal • Volume 100. Corn Belt. Yield variation was also related to emergence duration or early growth. A new heterotic pattern can be defined as temperate × Suwan1. Suwan1 was identified as a new heterotic group different from the current heterotic groups of Reid. disease resistance and planting-to-silking duration. Comparing the GCA and SCA effects between Fig. and LDRC. Thus. 4 and between Fig. 3 and Fig. we could not relate the difference of the GCA effects to any maize agronomic traits. The authors would like to give great thanks to Prof. 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. Huang Bihua (Dehong Institute of Agricultural Science of Yunnan province) and Prof. 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. 2004. in crosses with lines from the four known heterotic groups. These results strongly suggested that GCA effects should be considered in maize heterotic group classification. 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. Reid dent. 2002b). Their results showed that the GCA effects of the CIMMYT populations were generally higher than those of domestic populations. such as high disease resistance. Reid and Lancaster. YML145 might belong to either Reid dent or Lancaster group based on GCA effect. 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 new heterotic pattern may be as important as the widely accepted heterotic pattern of flint × dent. Lancaster. Another tropical line. 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. respectively. high resistance to drought and other stresses. Li Xuezhi (Lincang Institute of Agricultural Science of Yunnan province) for their help in data collection for this study. Lancaster. (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. YML145. etc. 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.lines are shown in Fig. Zhang et al. Thus. and Lancaster lines for controlling key agronomic traits. Further studies are needed to find which maize agronomic traits are responsible for high GCA effects between YML146/YML145 and TEMGs. no significantly positive GCA effects were found between YML145 and the lines from the Lancaster group..S. but not significant with the Reid Dent lines and Lancaster lines. 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). And the SCA effect of YML146 with Reid flint line P138 was significantly negative. the GCA effects of YML145 with the Reid flint lines were significantly positive. 4). These results suggested that Suwan1 population. TSPT and LDRC. LDRC. Based on the average GCA values of YML146. A new heterotic pattern of temperate × Suwan1 was demonstrated by the crosses of temperature inbred line × YML146 in this study. 2 and Fig.. Fan et al. Because YML146 was derived from Suwan1 population. 5. These two figures showed that when a line was crossed with different sets of maize germplasms grouped by the four heterotic groups. This phenomenon may be due to some different genetic bases between YML146 and YML145. this tropical line seemed to belong to a heterotic group different from the Reid. the GCA effects of YML146 with Lancaster and both Reid flint and Reid dent lines were significantly positive (Fig. strong root system. 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. Because YML146 had a positive average GCA with all lines from the two major temperate heterotic groups in the U. 5). Further systematic studies on temperate × Suwan1 heterotic pattern are warranted to be able to exploit this new heterotic pattern. Giauffret et al. thus. TSPT. we found that not only the absolute values. A New Heterotic Group: Suwan1 YML146 was selected from tropical maize population Suwan1 (Table 1). the parent population of YML146. and from the two major China local heterotic groups. 14 of them were from CIMMYT and 13 of them from China. the SCA effect of YML145 with the Reid dent line Yuzi87–1 was significantly positive. For instance. That difference could cause different levels of gene expression in Reid flint. Suwan1 might be classified as a new heterotic group different from the four heterotic groups. (2002b) employed four testers to evaluate 25 inbred lines from five tropical and subtropical populations. If the average GCA effect is significantly positive.
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