Preliminary study on the interaction effect between genotypes and environment of growth traits in Pinus massoniana clones
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摘要:目的 马尾松是我国南方重要速生用材树种之一。研究10个马尾松无性系在不同试验点胸径和树高生长性状的变异特征和遗传稳定性，筛选高遗传增益的优良基因型推广利用，为马尾松适地适基因型高效栽培和人工林提质增效提供试验依据。方法 以在广西宁明县（E1）、柳州市（E2）、博白县（E3）和钦州市（E4）营建的10个3年生马尾松无性系（M1 ~ M10）区域试验林为对象，测定胸径和树高生长，分析不同基因型生长性状差异和变异规律，开展无性系稳定性分析和综合评价，筛选生产力高和稳定性好的优良基因型。结果 马尾松无性系生长变异丰富，同一地点无性系间总变异和同一无性系地点间总变异均表现为胸径大于树高。多点联合方差分析表明：基因型、环境条件、基因型与环境互作效应对马尾松无性系胸径和树高生长都有极显著影响（P < 0.01），试验点内无性系间生长表现不同，无性系在不同试验点间生长表现也存在较大差异。不同地点无性系胸径和树高的重复力处于0.59 ~ 0.89之间，生长性状主要受遗传控制。遗传稳定性分析结果表明：M1、M2、M4、M5和M7等5个无性系胸径和树高生长速度较快且稳定性良好，M8对E1试验点具有特殊适应性。对4个试验点的马尾松无性系胸径和树高生长进行综合评价，筛选出M1、M2、M4、M5、M7 5个优良无性系，胸径和树高遗传增益分别比种子园良种实生苗提高37.90% ~ 45.65%和21.92% ~ 24.38%。结论 马尾松胸径和树高生长性状在无性系间变异丰富，基因型、环境和基因型与环境的互作效应对马尾松无性系的生长性状产生了极显著影响；M1、M2、M4、M5和M7等5个无性系生长迅速且稳定，遗传增益较高，可在马尾松栽培区适当推广。Abstract:Objective Pinus massoniana is one of the important fast-growing timber species in south China. In this study, phenotypic and genetic stability of DBH and tree height traits of 10 Pinus massoniana clones were studied at multiple sites, and excellent genotypes with high genetic gain were screened, so as to provide experimental basis for efficient cultivation of Pinus massoniana with suitable genotypes and quality and efficiency improvement of plantation.Method The DBH and tree height of 10 3-year-old Pinus massoniana clones (M1 − M10) were measured from regional experimental forest in Ningming County (E1), Liuzhou City (E2), Bobai County (E3) and Qinzhou City (E4) of Guangxi, southern China. The growth traits and phenotypic variation of different genotypes were analyzed. Clone stability analysis and comprehensive evaluation were carried out to screen excellent genotypes with high productivity and good stability.Result The growth variation of Pinus massoniana clones was rich, the total varation of DBH between clones at the same site and between clones at the same site was greater than tree height. Multi-point joint variance analysis showed that genotype, environmental conditions and genotype-environment interaction had significant effects on DBH and tree height growth of Pinus massoniana clones (P < 0.01). The growth performance of clones within the test site was different, and the growth performance of clones among different test sites was also different. The repeatability of DBH and tree height of clones at different sites was between 0.59 and 0.89, indicating they were mainly controlled by heredity. The results of genetic stability analysis showed that the DBH and tree height of M1, M2, M4, M5 and M7 clones grew faster and had good stability. Clone M8 had a special adaptability to the test site at Ningming County. Five superior clones (M1, M2, M4, M5, M7) were selected by comprehensive evaluation of DBH and tree height across four experimental sites. The genetic gains of DBH and tree height were increased by 37.90%−45.65% and 21.92%−24.38%, respectively, compared with the performance of seed orchard seedlings.Conclusion The DBH and tree height of Pinus massoniana vary greatly among clones. Genotype, environment and the interaction between genotype and environment have significant effects on the growth traits of Pinus massoniana clones. The five clones, M1, M2, M4, M5 and M7, grow rapidly and stably, and bring high genetic gain, which could be appropriately promoted in Pinus massoniana cultivation areas.
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Keywords:
- Pinus massoniana /
- clone /
- interaction effect /
- genetic variation /
- growth trait
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图 1 试验点在广西区内的分布情况
E1. 宁明县；E2. 柳州市；E3. 博白县；E4. 钦州市。下同。E1, Ningming County; E2, Liuzhou City; E3, Bobai County; E4, Qinzhou City. The same below.
Figure 1. Distribution of experimental points in Guangxi region
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图 2 参试马尾松无性系生长性状散点分布图
Figure 2. Scatter distribution of growth traits in tested clones of P. massoniana
表 1 各试验点参试无性系胸径和树高多重比较
Table 1 Multiple comparison of DBH and tree height of tested clones in different sites
地点
Site无性系
Clone胸径
DBH/cm树高
Tree height/m地点 Site 无性系
Clone胸径
DBH/cm树高
Tree height/mE1 M1 7.4 ± 1.1 aAB 4.75 ± 0.52 abAB E3 M1 4.5 ± 1.4 aABC 3.53 ± 0.58 aAB M2 6.7 ± 1.3 cdBCD 4.58 ± 0.53 bcABCD M2 4.4 ± 1.2 aABC 3.41 ± 0.69 abAB M3 6.3 ± 1.3 defCDE 4.37 ± 0.69 cdCDE M3 3.1 ± 0.9 cD 2.67 ± 0.52 cC M4 6.8 ± 1.5 bcdABC 4.57 ± 0.59 bcABCD M4 4.6 ± 1.6 aAB 3.40 ± 0.84 abAB M5 7.5 ± 1.2 aA 4.78 ± 0.54 abAB M5 4.3 ± 1.4 aABC 3.35 ± 0.63 abAB M6 5.8 ± 1.3 fgEF 4.35 ± 0.49 cdCDE M6 4.0 ± 1.4 abABCD 3.23 ± 0.59 abAB M7 7.0 ± 1.3 abcABC 4.65 ± 0.54 bABC M7 4.7 ± 1.5 aA 3.58 ± 0.78 aA M8 7.3 ± 1.2 abAB 4.90 ± 0.58 aA M8 3.9 ± 1.1 abcABCD 2.98 ± 0.55 bcBC M9 6.4 ± 1.2 deCDE 4.54 ± 0.50 bcBCD M9 4.0 ± 1.4 abABCD 3.21 ± 0.70 abABC M10 6.0 ± 1.2 efgDEF 4.19 ± 0.59 deE M10 4.4 ± 1.3 aABC 3.27 ± 0.69 abAB CK 5.5 ± 1.2 gF 4.07 ± 0.64 eE CK 3.4 ± 1.3 bcCD 2.67 ± 0.48 cC E2 M1 4.9 ± 1.2 abA 3.68 ± 0.67 abAB E4 M1 6.1 ± 1.6 aA 3.71 ± 0.39 bcdAB M2 4.7 ± 0.9 abAB 3.63 ± 0.57 abcAB M2 6.0 ± 0.9 aA 3.96 ± 0.34 abA M3 4.8 ± 1.2 abA 3.65 ± 0.67 abAB M3 5.4 ± 1.8 abcABC 3.66 ± 0.48 bcdAB M4 4.8 ± 1.1 abA 3.72 ± 0.66 aA M4 6.1 ± 1.6 aA 3.93 ± 0.56 abcA M5 5.1 ± 1.5 aA 3.74 ± 0.72 aA M5 6.2 ± 1.3 aA 4.05 ± 0.49 aA M6 3.6 ± 1.2 cdCD 3.12 ± 0.58 defBC M6 4.4 ± 0.9 deCD 3.43 ± 0.33 deBC M7 4.7 ± 1.5 abAB 3.52 ± 0.82 abcdAB M7 5.8 ± 1.1 abAB 3.86 ± 0.37 abcAB M8 4.2 ± 1.2 bcABC 3.21 ± 0.64 cdefABC M8 4.8 ± 0.7 bcdeABCD 3.47 ± 0.36 deBC M9 3.7 ± 1.5 cdBCD 3.28 ± 0.79 bcdeABC M9 4.9 ± 1.3 bcdeABCD 3.65 ± 0.31 bcdAB M10 4.6 ± 1.5 abAB 3.52 ± 0.7 6abcdAB M10 5.3 ± 1.4 abcdABCD 3.61 ± 0.49 cdAB CK 3.2 ± 1.0 dCD 2.89 ± 0.59 efC CK 4.0 ± 0.6 eD 3.15 ± 0.47 eC 注：不同小写字母和大写字母分别表示同一地点不同无性系之间每个性状在0.05和0.01水平上差异显著。Notes: different lowercase and uppercase letters indicate significant differences in each trait between different clones at the same site at the level of 0.05 and 0.01, respectively. 
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表 2 各试验点马尾松无性系生长性状表型变异系数
Table 2 Phenotypic variation coefficients of growth traits of P. massoniana clones at different sites
% 无性系
Clone胸径 DBH 总变异
Total variation树高 Tree height 总变异
Total variationE1 E2 E3 E4 E1 E2 E3 E4 M1 15.27 25.10 31.33 26.07 30.06 10.95 18.21 16.43 10.51 19.30 M2 19.10 20.64 27.95 16.00 27.04 11.57 15.70 20.23 8.59 18.41 M3 20.79 25.63 31.61 33.52 33.08 15.79 18.36 19.48 13.11 22.78 M4 21.62 22.71 35.65 26.07 30.73 12.91 17.74 24.71 14.25 20.52 M5 16.27 28.63 33.49 20.32 30.81 11.30 19.25 18.81 12.10 20.29 M6 22.24 32.78 34.00 21.82 33.76 11.26 18.59 18.27 9.62 20.66 M7 18.29 32.13 31.70 18.10 29.86 11.61 23.30 21.79 9.59 20.65 M8 16.99 27.62 28.72 15.00 34.78 11.84 19.94 18.46 10.37 26.50 M9 19.06 40.27 35.50 26.94 35.66 11.01 24.09 21.81 8.49 22.57 M10 19.83 31.74 29.55 25.85 28.39 14.08 21.59 21.10 13.57 19.91 总变异
Total variation20.45 30.22 33.57 25.64 12.91 20.51 22.39 12.30
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表 3 各试验点无性系生长性状方差分析
Table 3 Variance analyses of growth traits of clones in each test site
性状
Trait地点
Site平均值
Mean标准差
Standard deviationF 无性系重复力
Clonal repeatability胸径 DBH/cm E1 6.7 1.37 9.468** 0.89 E2 4.5 1.36 4.417** 0.77 E3 4.2 1.41 2.466* 0.59 E4 5.5 1.41 3.809** 0.74 树高 Tree height/m E1 4.57 0.59 6.851** 0.85 E2 3.51 0.72 2.994** 0.67 E3 3.93 0.88 3.494** 0.71 E4 3.74 0.46 5.273** 0.81 注：*表示在0.05水平上差异显著，**表示在0.01水平上差异显著，下同。Notes: * means significant difference at P < 0.05 level, ** means significant difference at P < 0.01 level. The same below.
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表 4 各试验点参试无性系生长性状联合方差分析
Table 4 Joint analysis of variance for growth traits of tested clones in different sites
生长性状
Growth trait变异来源
Variation source自由度
Degree of freedom平方和
Sum of square均方
Mean squareF 胸径 DBH 地点 Site 3 1 281.843 427.281 254.208** 无性系 Clone 9 187.378 20.820 12.387** 地点 × 无性系 Site × clone 27 96.461 3.573 2.126** 误差 Error 1 099 1 847.234 1.681 树高 Tree height 地点 Site 3 346.457 115.486 322.876** 无性系 Clone 9 25.568 2.841 7.942** 地点 × 无性系 Site × clone 27 24.176 0.895 2.503** 误差 Error 1 099 393.089 0.358
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表 5 马尾松无性系生长性状与环境互作效应值和适生范围
Table 5 Interaction effect value and suitable range of growth traits and environment of P. massoniana clones
无性系
Clone无性系 × 环境 Clone × environment 适宜范围
Suitable areaE1 E2 E3 E4 M1 0.105 3 0.019 9 0.004 9 −0.130 1 E1、E2、E3 M2 −0.230 2 0.035 3 0.025 3 0.169 5 E2、E3、E4 M3 −0.128 3 0.733 9 −0.637 8 0.218 0 E2、E4 M4 −0.341 7 −0.049 2 0.375 8 0.035 0 E3、E4 M5 0.265 7 −0.017 5 −0.336 6 0.155 9 E1、E4 M6 0.015 0 −0.226 1 0.347 2 −0.193 6 E1、E3 M7 0.085 5 0.017 0 0.329 1 0.153 7 E1、E2、E3、E4 M8 0.484 4 −0.165 9 −0.070 1 −0.313 4 E1 M9 0.317 9 −0.223 5 −0.127 6 0.072 4 E1、E4 M10 −0.359 0 0.052 3 0.265 6 0.023 1 E2、E3、E4
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表 6 各试验点马尾松无性系生长性状综合评价
Table 6 Comprehensive evaluation of growth traits of P. massoniana clones in different sites
E1 E2 E3 E4 平均值 Mean 排名
Ranking无性系
CloneQi 无性系
CloneQi 无性系
CloneQi 无性系
CloneQi 无性系
CloneQi M8 1.25 M5 1.17 M7 1.15 M5 1.26 M5 1.20 1 M5 1.25 M1 1.16 M1 1.14 M4 1.25 M1 1.19 2 M1 1.24 M4 1.15 M4 1.13 M2 1.25 M4 1.18 3 M7 1.22 M3 1.15 M2 1.12 M1 1.23 M7 1.18 4 M4 1.21 M2 1.14 M5 1.11 M7 1.23 M2 1.18 5 M2 1.20 M7 1.13 M10 1.11 M3 1.19 M10 1.14 6 M9 1.19 M10 1.12 M6 1.08 M10 1.18 M8 1.13 7 M3 1.17 M8 1.07 M9 1.08 M9 1.16 M9 1.12 8 M6 1.15 M9 1.05 M8 1.05 M8 1.14 M3 1.12 9 M10 1.14 M6 1.03 M3 0.97 M6 1.12 M6 1.09 10 平均值 Mean 1.20 1.12 1.10 1.20 1.15
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表 7 马尾松优良无性系遗传增益估算
Table 7 Genetic gain estimation of superior clones of P. massoniana
无性系
Clone平均胸径
Average DBH/cm与CK相比
Compared with CK平均树高
Average tree height/m与CK相比
Compared with CK现实增益
Practical gain/%遗传增益
Genetic gain/%现实增益
Practical gain/%遗传增益
Genetic gain/%M1 5.9 47.82 43.99 4.05 26.58 23.12 M2 5.6 41.20 37.90 4.02 25.53 22.21 M4 5.7 42.55 39.14 4.01 25.19 21.92 M5 6.0 49.62 45.65 4.10 28.03 24.38 M7 5.7 43.31 39.85 4.02 25.61 22.28 平均值 Mean 5.8 44.75 41.17 4.04 26.25 22.84
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