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桃金娘科(Myrtaceae)桉属(Eucalyptus L'Hérit.)、杯果木属(Angophora Cav.)和伞房属(Corymbia K.D. Hill & L.A.S. Johnson)树种统称桉树,在全球热带、亚热带和温带地区广为种植,面积超过2 100万hm2[1]。桉树也是华南地区重要的短周期人工林树种,种植面积达450万hm2[2],其木材广泛用于制造纸浆、胶合板、纤维板和家具等。相应地,生长和木材性质是重要的经济性状[3-4],一直以来都是桉树育种改良的主要目标性状。
木材密度是重要的材性性状,既影响森林产出木材的生物量,从而影响碳汇的估算[5-6],也与木材强度、结构和化学成份等木材品质相关,是一个材性的综合经济指标[7]。木材密度测定的传统方法包括容积法(或饱和排水法)和X射线法,存在耗时长、费用高的缺点[8],很难用于大量样品的测定。目前,一些快速、无损地测定木材密度的间接方法得到发展,如Pilodyn针击法[9]、近红外光谱法[10]和Resistograph钻刺法[8-11],其中,Resistograph钻刺法基于电钻连接的Resistograph钻针(直径仅1.5 mm)匀速穿刺树径,即时记录扭力曲线作为木材密度的指示[8]。Resistograph钻刺法已有效用于针叶树木材密度的间接测定,如火炬松(Pinus taeda L.)[11]和海岸松(P. pinaster Ait.)[12],但阔叶树中只笔者报道了其对尾叶桉(E. urophylla S. T. Blake)×细叶桉(E. tereticornis Smith)杂种(简称尾细桉)的木材密度测定[13],而对Resistograph钻刺法与标准方法测定的木材密度的相关性尚无研究。
尾细桉在华南沿海广泛栽培,其速生、高产且具有较好的抗风[14]和耐寒潜力[15-16]。目前,尾细桉选育研究主要针对生长和耐寒性[14-16],进一步的选育工作需要纳入材性性状。本研究基于不完全析因交配设计产生的56个尾细桉杂交组合,建立了Resistograph钻刺法与容积法测定木材密度的相关性,分析了7.5年生尾细桉生长和木材密度的遗传变异和性状相关性,并结合两类性状选择了杂交组合和单株,旨在为培育速生性好、木材密度高的桉树良种提供材料基础。
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7.5年生时利用Vertex Ⅲ测高器(瑞典Haglof)和胸径尺观测树高(H)和胸径(D),单株材积(V)基于树高和胸径计算:V=H×D2/30 000。
7.5年生时还利用Resistograph F-400S (德国Instrumenta Mechanik Labor GmbH)间接测定木材密度,即钻刺木材密度(WDi)。对胸径大于8 cm的单株进行树皮-树皮的钻刺,共56个杂种488株,钻刺位置为1.3 m高处,避免节疤,即时记录的钻刺扭力曲线自动保存于仪器。扭力曲线下载到电脑后,利用厂家提供的Resistograph软件查看。由于扭力会随钻刺进度而增大,只取一端树皮内开始的5 cm钻刺曲线进行单株平均扭力(%)的估算[12],单株WDi为平均扭力的平方根[13]。此外,为验证钻刺法间接测定木材密度的可靠性,随机选取53个杂交组合的79株及9个自由授粉家系的14株,利用生长锥(直径5 mm)钻取木芯,通过容积法测定木材基本密度(WD),测定方法参考国家标准GB/T 1933-2009。
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利用R软件进行各性状的平均值和变异系数(CV, %)计算及方差分析和多重比较,杂种与自由授粉家系的差异显著性利用t检验进行检测。方差分析基于个体的混合线性模型:
$ Y_{i j k}=\mu+M_{i}+F_{j}+M F_{i j}+B_{k}+E_{i j k} $
式中:Yijk为重复k中母本i和父本j的杂种单株表型值,μ为总体平均值,Mi、Fj、MFij是随机效应,分别为母本i的遗传效应、父本j的遗传效应、母本i和父本j的互作效应,Bk是固定效应,为重复k的效应,Eijk是误差。利用ASReml程序包[18]估算性状间的表型相关系数(rp)和遗传相关系数(rg)。WDi与WD的相关分析基于容积法测定的53个杂交组合的79株进行。
杂交组合和单株的选择采用独立淘汰法。组合的选择基于材积和钻刺木材密度的多重比较(Duncan法,α=0.01),两性状值均高于对照无性系的杂种入选。单株的选择以对照的最好单株作为标准,材积和木材密度均高于标准的单株入选。
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表 1表明:杂种的平均生长均显著高于自由授粉家系且变异系数较低;杂种的WDi平均值显著高于自由授粉家系,且变异系数稍高,表明尾细桉具有杂种优势。
表 1 尾细桉杂种和母本自由授粉家系7.5生时所测性状的平均值
Table 1. Mean values of the traits measured for E. urophylla ×E. tereticornis hybrids and maternal open pollinated families
性状Trait 杂种Hybrid 自由授粉家系Open pollinated family 平均值Mean 变异系数CV /% 平均值Mean 变异系数CV /% 树高H /m 16.70 ± 4.22*** 25.3 15.08 ± 3.98 26.4 胸径D /cm 13.16 ± 3.89** 29.6 11.48 ± 3.85 33.5 材积V/(×10-2 m3) 14.50 ± 9.41*** 65.1 10.64 ± 8.63 81.1 钻刺木材密度WDi 0.39 ± 0.06** 15.8 0.37 ± 0.05 15.0 木材基本密度WD /(g.cm-3) 0.49 ± 0.05 9.5 0.48 ± 0.04 9.3 注:表中数据为平均值±标准差,杂种和自由授粉家系的WD平均值分别基于53个杂交组合的79株和9个自由授粉家系的14株计算得出。***表示P<0.001显著,**表示P<0.01显著。
Note: data in the table as mean ± standard deviation. WD means for hybrids and open pollinated families were calculated with 79 trees of 53 crosses and 14 trees of 9 families, respectively. *** significance at P<0.001, ** significance at P<0.01. -
WDi与WD的表型相关系数(rp)为0.52 (P<0.001),遗传相关系数(rg)为0.55(P<0.05)。两种方法测定的木材密度相关显著,表明Resistograph钻刺法具有较高的可靠性,WDi可作为木材密度间接测定的有效指标。
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表 2表明:对于生长性状H、D和V,母本间和父本间均差异极显著(P<0.001或0.01),表明母本和父本对生长性状的加性效应均显著;但母本×父本互作不显著,表明参试群体的生长受显性效应的影响较弱。
表 2 7.5年生H、D、V和WDi的方差分析
Table 2. Analysis of variance on H, D, V and WDi at age of 7.5 years
状Trait 变异来源Source of variation 自由度d.f. 均方Mean square F值F value 树高H 重复Replicate 3 58.9 3.67* 母本Female 9 44.0 2.74** 父本Male 9 62.9 3.92*** 母本×父本Female×male 37 18.0 1.12 误差Error 432 16.0 胸径D 重复Replicate 3 28.5 2.07 母本Female 9 44.4 3.23*** 父本Male 9 62.8 4.57*** 母本×父本Female × male 37 11.9 0.86 误差Error 432 13.7 材积V 重复Replicate 3 0.028 9 3.69* 母本Female 9 0.030 4 3.87*** 父本Male 9 0.033 7 4.29*** 母本×父本Female × male 37 0.007 8 0.99 误差Error 432 0.007 9 钻刺木材密度WDi 重复Replicate 3 0.080 8 27.63*** 母本Female 9 0.005 6 1.90 父本Male 9 0.010 4 3.55*** 母本×父本Female × male 37 0.004 8 1.64* 误差Error 432 0.002 9 注: ***表示P<0.001显著,**表示P<0.01显著,*表示P<0.05显著。
Note: *** significance at P<0.001, ** significance at P<0.01, * significance at P<0.05.对于WDi,父本间差异极显著(P<0.001),而母本间差异不显著,表明父本对木材密度的影响极显著且比母本更重要;母本×父本互作显著(P<0.05),表明木材密度也受显性效应的影响。此外,重复间的差异也极显著(P<0.001),表明小环境亦可显著影响木材密度。
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表 3表明:7.5年生生长性状D、H和V之间的表型相关系数(rp)为0.82~0.93,遗传相关系数(rg)为0.93~0.99,均极显著相关(P<0.001),表明3个生长性状具有相似的遗传基础,可能有共同的基因影响树高、胸径和/或材积。
表 3 7.5年生H、D、V和WDi间的表型相关和遗传相关
Table 3. Phenotypic and genetic correlation coefficients between H, D, V and WDi at age of 7.5 years
性状Trait 树高H 胸径D 材积V 钻刺木材密度WDi 树高H 0.93±0.06*** 0.99±0.02*** 0.38±0.33 胸径D 0.82±0.02*** 0.97±0.03*** 0.38±0.32 材积V 0.93±0.01*** 0.92±0.01*** 0.32±0.34 钻刺木材密度WDi 0.36±0.04*** 0.36±0.04*** 0.35±0.04*** 注:表中数据为相关系数±标准误,斜对角线上方为遗传相关、下方为表型相关,***表示P<0.001显著。
Note: data in the table as correlation coefficient ± standard error. Phenotypic and genetic correlation coefficients were below and above the diagonal, respectively. *** significance at P<0.001.WDi与D、H、V间的rp和rg均较小,rp为0.35~0.36,极显著相关(P<0.001);rg为0.32~0.38,相关不显著。WDi与生长性状的不显著遗传相关表明,两类性状可能为独立遗传,育种改良中需要对这两类性状分别进行选择,以获得速生且木材密度较大的优良杂种。
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两对照无性系中,DH32-29的平均单株材积和平均钻刺木材密度均高于广林-9,故以DH32-29作为选择的标准,共选出V和WDi平均值均高于对照的杂交组合14个(表 4),其平均单株材积为标准的103.7%~156.7%,平均WDi为标准的106.9%~124.2%。来自母本E.u02、E.u03、E.u04、E.u06、E.u07和E.u09的中选组合各2个,来自父本E.t11、E.t17和E.t18的中选组合分别为3、3、4个,为较好的杂交亲本。
表 4 选出的14个杂种的V和WDi平均值
Table 4. Means of V and WDi for the 14 crosses selected out
杂种编号Cross no. 亲本组合Parental cross 材积V 钻刺木材密度WDi 排名Rank 平均值Mean /m3 杂种/对照Cross/CK /% 排名Rank 平均值Mean /m3 杂种/对照Cross/CK /% 45 E.u07 × E.t11 2 0.252 7 a 156.7 10 0.41 abc 114.8 28 E.u04 × E.t18 3 0.245 8 ab 152.4 5 0.43 abc 118.4 23 E.u03 × E.t18 5 0.233 7 abc 145.0 26 0.39 abc 109.3 15 E.u02 × E.t17 6 0.227 3 abc 141.0 18 0.40 abc 111.2 52 E.u07 × E.t18 7 0.222 9 abc 138.2 6 0.43 abc 118.3 60 E.u09 × E.t18 8 0.194 4 abc 120.6 20 0.40 abc 110.6 37 E.u06 × E.t11 10 0.183 9 abc 114.1 1 0.45 a 124.2 30 E.u05 × E.t11 11 0.183 8 abc 114.0 13 0.41 abc 112.7 21 E.u03 × E.t15 12 0.178 7 abc 110.8 17 0.40 abc 111.9 43 E.u06 × E.t17 14 0.173 8 abc 107.8 8 0.42 abc 115.3 59 E.u09 × E.t17 15 0.173 6 abc 107.6 33 0.39 abc 106.9 26 E.u04 × E.t16 16 0.170 2 abc 105.5 30 0.39 abc 107.0 17 E.u02 × E.t19 17 0.169 4 abc 105.1 14 0.41 abc 112.6 54 E.u08 × E.t12 18 0.167 2 abc 103.7 19 0.40 abc 111.1 注:对照为无性系DH32-29,平均值后相同的小写字母表示不同杂种间的差异不显著(基于Duncan法的多重比较,α= 0.01)。
Note: CK is represented by clone DH32-29. The same letters following means indicate non-significant difference between crosses (Duncan’s multiple comparison, α= 0.01).以对照无性系DH32-29最大单株的V (0.273 2 m3)和广林-9最高单株的WDi(0.441)为标准,评选出17株杂种单株,单株的V为标准的101.3%~149.1%,WDi为标准的100.1%~118.8% (具体数据未列示)。
尾叶桉×细叶桉木材密度与生长的联合选择
Combined Selection of Wood Density and Growth in Eucalyptus urophylla × E. tereticornis Hybrids
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摘要:
目的 评估Resistograph钻刺法间接测定尾叶桉×细叶桉木材密度的可靠性, 检测杂交亲本对子代表型的效应以及生长与木材密度的相关, 评选速生、优质的尾细桉杂种。 方法 基于10株尾叶桉与10株细叶桉不完全析因交配产生的56个杂交组合的7.5年生试验林, 利用79株分析容积法与Resistograph钻刺法测定的木材密度的相关, 通过方差分析检测亲本对杂种生长和木材密度的效应, 结合多重比较和独立淘汰法进行材积和木材密度的联合选择。 结果 容积法与Resistograph钻刺法测定的木材密度的表型相关系数为0.52(P < 0.001), 遗传相关系数为0.55(P < 0.05);树高、胸径及材积的母本间和父本间均呈极显著差异(P < 0.001或0.01), 但母本×父本互作的效应不显著; 对钻刺木材密度, 父本间呈极显著差异(P < 0.001), 母本×父本互作显著(P < 0.05), 但母本间差异不显著; 树高、胸径和材积间的表型相关和遗传相关均极显著(P < 0.001), 其与钻刺木材密度的表型相关极显著(P < 0.001), 但遗传相关不显著; 评选出速生、木材密度较高的杂交组合14个、单株17株。 结论 Resistograph钻刺法是一种间接测定尾细桉木材密度的简便、经济和可靠的方法; 母本和父本选择以及母本与父本的组配对培育速生、材质优良的尾细桉杂种均较重要; 尾细桉生长与木材密度的遗传相关不显著, 需要对这两类性状分别进行选择; 评选的尾细桉杂交组合和单株为培育速生、优质的桉树良种提供了有用的材料。 -
关键词:
- 尾叶桉×细叶桉杂种
- / Resistograph钻刺法
- / 木材密度
- / 生长
- / 联合选择
Abstract:Objective To evaluate the reliability of Resistograph penetration method on indirect measurement of wood density of Eucalyptus urophylla×E. tereticornis hybrids, detect the parental effects on cross phenotypic traits and the correlations between growth and wood density, and select fast-growing and high wood-quality Eucalyptus urophylla×E. tereticornis hybrids. Method In a 7.5-year-old trial stand with fifty six hybrid combinations derived from an incomplete factorial mating among 10 E.urophylla (female) and 10 E.tereticornis(male) parents, 79 trees were sampled to determine the correlation in wood density between volumetric and Resistograph measurements. The effects of parents on hybrid growth and wood density were detected by variance analysis. Multiple comparison and independent selection were performed to make joint selection of stand volume and wood density. Result The phenotypic and genetic correlation coefficients between volumetric and Resistograph measurements were 0.52 (P < 0.001) and 0.55 (P < 0.05), respectively. The differences in the traits of height, DBH and stock volume, were highly significant (P < 0.001 or 0.01) among either females or males, but no significant difference was observed for female×male interactions. For ResistograPh-based wood density, however, the differences were highly significant (P < 0.001) among the males and significant (P < 0.05) for female×male interactions, but insignificant among the females. The growth traits of height, DBH and stock volume were highly significantly correlated in both phenotypic and genetic terms (P < 0.001), while their phenotypic and genetic correlations with wood density were highly significant (P < 0.001) and insignificant, respectively. A total of 14 hybrids and 17 individual trees were selected out. Conclusion Resistograph method is convenient, economic and reliable for indirect measurement of wood density of E. urophylla×E. tereticornis hybrids. Female and male parents selecting and parental pairing are important for breeding fast-grow and high wood-quality hybrids. Since there is no significant genetic correlation between growth and wood density, it indicates the necessity of selecting separately against the two traits. The hybrids and trees selected out are of valuable plant materials for further cultivating fast-grow and high wood-quality eucalypt hybrids. -
表 1 尾细桉杂种和母本自由授粉家系7.5生时所测性状的平均值
Table 1. Mean values of the traits measured for E. urophylla ×E. tereticornis hybrids and maternal open pollinated families
性状Trait 杂种Hybrid 自由授粉家系Open pollinated family 平均值Mean 变异系数CV /% 平均值Mean 变异系数CV /% 树高H /m 16.70 ± 4.22*** 25.3 15.08 ± 3.98 26.4 胸径D /cm 13.16 ± 3.89** 29.6 11.48 ± 3.85 33.5 材积V/(×10-2 m3) 14.50 ± 9.41*** 65.1 10.64 ± 8.63 81.1 钻刺木材密度WDi 0.39 ± 0.06** 15.8 0.37 ± 0.05 15.0 木材基本密度WD /(g.cm-3) 0.49 ± 0.05 9.5 0.48 ± 0.04 9.3 注:表中数据为平均值±标准差,杂种和自由授粉家系的WD平均值分别基于53个杂交组合的79株和9个自由授粉家系的14株计算得出。***表示P<0.001显著,**表示P<0.01显著。
Note: data in the table as mean ± standard deviation. WD means for hybrids and open pollinated families were calculated with 79 trees of 53 crosses and 14 trees of 9 families, respectively. *** significance at P<0.001, ** significance at P<0.01.表 2 7.5年生H、D、V和WDi的方差分析
Table 2. Analysis of variance on H, D, V and WDi at age of 7.5 years
状Trait 变异来源Source of variation 自由度d.f. 均方Mean square F值F value 树高H 重复Replicate 3 58.9 3.67* 母本Female 9 44.0 2.74** 父本Male 9 62.9 3.92*** 母本×父本Female×male 37 18.0 1.12 误差Error 432 16.0 胸径D 重复Replicate 3 28.5 2.07 母本Female 9 44.4 3.23*** 父本Male 9 62.8 4.57*** 母本×父本Female × male 37 11.9 0.86 误差Error 432 13.7 材积V 重复Replicate 3 0.028 9 3.69* 母本Female 9 0.030 4 3.87*** 父本Male 9 0.033 7 4.29*** 母本×父本Female × male 37 0.007 8 0.99 误差Error 432 0.007 9 钻刺木材密度WDi 重复Replicate 3 0.080 8 27.63*** 母本Female 9 0.005 6 1.90 父本Male 9 0.010 4 3.55*** 母本×父本Female × male 37 0.004 8 1.64* 误差Error 432 0.002 9 注: ***表示P<0.001显著,**表示P<0.01显著,*表示P<0.05显著。
Note: *** significance at P<0.001, ** significance at P<0.01, * significance at P<0.05.表 3 7.5年生H、D、V和WDi间的表型相关和遗传相关
Table 3. Phenotypic and genetic correlation coefficients between H, D, V and WDi at age of 7.5 years
性状Trait 树高H 胸径D 材积V 钻刺木材密度WDi 树高H 0.93±0.06*** 0.99±0.02*** 0.38±0.33 胸径D 0.82±0.02*** 0.97±0.03*** 0.38±0.32 材积V 0.93±0.01*** 0.92±0.01*** 0.32±0.34 钻刺木材密度WDi 0.36±0.04*** 0.36±0.04*** 0.35±0.04*** 注:表中数据为相关系数±标准误,斜对角线上方为遗传相关、下方为表型相关,***表示P<0.001显著。
Note: data in the table as correlation coefficient ± standard error. Phenotypic and genetic correlation coefficients were below and above the diagonal, respectively. *** significance at P<0.001.表 4 选出的14个杂种的V和WDi平均值
Table 4. Means of V and WDi for the 14 crosses selected out
杂种编号Cross no. 亲本组合Parental cross 材积V 钻刺木材密度WDi 排名Rank 平均值Mean /m3 杂种/对照Cross/CK /% 排名Rank 平均值Mean /m3 杂种/对照Cross/CK /% 45 E.u07 × E.t11 2 0.252 7 a 156.7 10 0.41 abc 114.8 28 E.u04 × E.t18 3 0.245 8 ab 152.4 5 0.43 abc 118.4 23 E.u03 × E.t18 5 0.233 7 abc 145.0 26 0.39 abc 109.3 15 E.u02 × E.t17 6 0.227 3 abc 141.0 18 0.40 abc 111.2 52 E.u07 × E.t18 7 0.222 9 abc 138.2 6 0.43 abc 118.3 60 E.u09 × E.t18 8 0.194 4 abc 120.6 20 0.40 abc 110.6 37 E.u06 × E.t11 10 0.183 9 abc 114.1 1 0.45 a 124.2 30 E.u05 × E.t11 11 0.183 8 abc 114.0 13 0.41 abc 112.7 21 E.u03 × E.t15 12 0.178 7 abc 110.8 17 0.40 abc 111.9 43 E.u06 × E.t17 14 0.173 8 abc 107.8 8 0.42 abc 115.3 59 E.u09 × E.t17 15 0.173 6 abc 107.6 33 0.39 abc 106.9 26 E.u04 × E.t16 16 0.170 2 abc 105.5 30 0.39 abc 107.0 17 E.u02 × E.t19 17 0.169 4 abc 105.1 14 0.41 abc 112.6 54 E.u08 × E.t12 18 0.167 2 abc 103.7 19 0.40 abc 111.1 注:对照为无性系DH32-29,平均值后相同的小写字母表示不同杂种间的差异不显著(基于Duncan法的多重比较,α= 0.01)。
Note: CK is represented by clone DH32-29. The same letters following means indicate non-significant difference between crosses (Duncan’s multiple comparison, α= 0.01). -
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