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在一个林木育种计划中,及时精确地估算主要经济性状的遗传参数对育种策略的制定和实施起着至关重要的作用[1]。遗传参数中遗传效应方差通常分为加性和非加性两部分,非加性遗传效应又可进一步分为显性和上位效应[2]。加性遗传效应(或一般配合力,GCA)被认为是性状在代际遗传中可以固定的遗传效应,是育种值大小的二分之一,在林木育种中占有相对重要的位置。因此,亲本选择、育种群体构建及相应的育种评价主要是基于加性遗传模型。显性效应(特殊配合力SCA的4倍)是等位基因间的相互作用,被认为是能遗传而不能被固定的遗传效应,是产生杂种优势的主要部分。早期遗传分析中忽视了非加性遗传效应尤其是显性效应的重要性,将其归入环境效应中,从而导致方差组分和育种值估计的偏差[3-5]。
双列交配设计可同时估计加性和显性效应方差,提供丰富的遗传信息,同时还可为高世代育种提供大量谱系清楚的供选子代,因此,国内外很多针叶树均开展了基于双列交配设计的配合力研究[6-8]。早期研究中,将GCA作为最重要的遗传参数用于育种策略的制定,而对SCA的关注较少。Carson利用辐射松(Pinus radiata)半双列杂交子代多点试验结果,报道了SCA相对于GCA的重要性在不同环境下变化较大,比值最高可达98%[9]。Wu等[7]在对辐射松不连续半双列交配设计的10个地点测定林的联合分析时发现,SCA方差近等于GCA方差。近期对火炬松(P.taeda)双列交配设计的多点子代测定研究中也证实树高受到更显著的显性效应控制[10]。因此,在特定的改良计划中,除了对GCA的利用,对SCA的研究也是必不可少的。
日本落叶松(Larix kaempferi (Lamb.) Carr.)具有早期速生、成林快、易于栽培、适应性广等特点,在我国的温带(吉林、辽宁、山东、河南等)及中北亚热带(湖北、湖南、四川、甘肃等)高山区得到迅速推广应用[11-12]。1965年在辽宁大孤家林场营建我国最早的日本落叶松种子园,随后陆续开展人工杂交和子代测定工作。已有研究多集中在对自由授粉家系的生长、形质及材性等遗传变异方面[12-15],但对全同胞子代表现的研究比较缺乏。本研究以来自13个母本和23个父本包括6×6和4×4两组全双列交配及部分随机交配的子代测定林为对象,对16年生和26年生的生长数据进行区组效应和空间环境效应校正后,研究自交子代的生长表现、正反交效应和GCA与SCA的相对重要性,为合理制定和调整日本落叶松遗传改良策略提供依据。
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育种种子园于1965年建于辽宁省清原县大孤家林场(42.72° N,124.88° E),建园无性系由选自辽宁地区30~40年生早期引种的日本落叶松人工林优树嫁接而成,包含12个小区,株行距为4 m×4 m。1986—1987年在种子园开展控制授粉进行人工制种,共获得来自13个母本和23个父本的杂交组合66个,其中,包括6×6和4×4两组全双列交配组合(组1和组2),以及19个随机交配组合,交配设计见表 1。1988年在圃地育苗,1990年以2年生苗在该林场内造林。试验林采用随机完全区组设计,5次重复,10株双列小区,株行距为2 m×2 m。土壤为山地棕壤,海拔约460 m。
表 1 交配设计表
Table 1. Mating design table
母本
Maternal
parents父本Paternal parents RC13 RC19 RC4 RC5 RC503 RC82 RF27 Y11 Y16 R1 Y33 Y5 Y13 RC2 RC34 RC35 RC40 RC402 RC404 RC41 RC61 RC8 RC81 RC13 ⊗ × × × × × × RC19 × ⊗ × × × × RC4 × × ⊗ × × RC5 × × × ⊗ × × × × × RC503 × × × × × × RC82 × × × × × × × × × RF27 ⊗ × × Y11 × × × Y16 × × ⊗ × R1 × × × ⊗ Y55 × Y10 × × × × × × × RC25 × × 注:×-交配;⊗-自交。
Notes: × and ⊗ indicate crossing and selfing, respectively. -
分别在林龄16年和26年对测定林进行每木生长调查。树高(H/m)采用激光测高仪(Vertex Ⅲ,Haglof Company Group,Sweden)测定,胸径(DBH/cm)采用测径尺测定,并计算材积(V/dm3):
$ V = H \times DB{H^2} $
(1) -
在估计遗传参数之前,通过混合效应模型(2)矫正表型性状中的环境变异。
$ y = X\beta + Zr + e $
(2) 式(2)中: y是表型观测值向量,β是固定效应向量(均值),r是随机的区组效应向量服从r~N(0, $I\hat \sigma _r^2$),I是单位矩阵,$\hat \sigma _r^2$是区组效应方差,X和Z分别是固定和随机效应的关联矩阵,e是随机残差向量,服从r~N(0, R),R是残差的方差-协方差结构(3)。
$ R = \hat \sigma _\xi ^2\sum\limits_c {\left( {{\rho _c}} \right)} \otimes \sum\limits_r {\left( {{\rho _r}} \right)} + I\hat \sigma _\eta ^2 $
(3) $\hat \sigma _\xi ^2$是空间残差方差,$\hat \sigma _\eta ^2$是独立残差方差,∑r(ρr)是具有维度r×r的行模型的相关矩阵,ρr是行方向上的自相关参数;∑c(ρc)是具有维度c×c的列模型的相关矩阵,ρc是列方向上的自相关参数,而且有:
$ \sum\limits_r = \left[ {\begin{array}{*{20}{c}} 1&{}&{}&{}&{}\\ {{\rho _r}}&1&{}&{}&{}\\ {\rho _r^2}&{{\rho _r}}&1&{}&{}\\ \vdots & \vdots &{ \vdots }& \ddots &{}\\ {\rho _r^{r - 1}}&{\rho _r^{r - 2}}&{\rho _r^{r - 3}}& \cdots &1 \end{array}} \right] $
$ \sum\limits_c = \left[ {\begin{array}{*{20}{c}} 1&{}&{}&{}&{}\\ {{\rho _c}}&1&{}&{}&{}\\ {\rho _c^2}&{{\rho _c}}&1&{}&{}\\ \vdots & \vdots & \vdots & \ddots &{}\\ {\rho _c^{c - 1}}&{\rho _c^{c - 2}}&{\rho _c^{c - 3}}& \cdots &1 \end{array}} \right] $
利用R软件(版本3.1.2)ASReml-R包(版本3.0)[16-17]拟合模型。最终通过从表型数据中减去随机的区组和空间位置效应,获得矫正表型数据用于后续分析。
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采用单株线性随机效应模型(4)和限制性最大似然估计方法(REML)估计随机效应的方差分量。
$ y = {Z_1}a + {Z_2}s + {Z_3}r + {Z_4}p + e $
(4) 式(4)中:y是矫正后的表型值向量,a、s、r、p、e分别是随机的加性、特殊配合力、正反交、小区、残差效应向量,Z1~Z4分别是对应效应的关联矩阵。
利用似然比检验(LRT)各方差组分的统计显著性。利用泰勒级数展开法(Taylor series expansion)计算遗传参数的标准误[16]。
遗传和表型变异系数(CVG和CVP)分别按照公式(5)和(6)计算。
$ C{V_G} = {{\hat \sigma }_a}/\bar X \times 100\% $
(5) $ C{V_P} = {{\hat \sigma }_P}/\bar X \times 100\% $
(6) 式(5)~(6)中:${\hat \sigma }$a、${\hat \sigma }$p、${\bar X}$分别为加性方差平方根、总方差平方根、总体均值。
狭义和广义单株遗传力(${\hat h}$i2和${\hat H}$i2)以及广义家系遗传力(f2)参照Weng等[18]中的公式进行计算:
$ \hat h_i^2 = \frac{{\hat \sigma _a^2}}{{0.5\hat \sigma _a^2 + \hat \sigma _s^2 + \hat \sigma _r^2 + \hat \sigma _p^2 + \hat \sigma _e^2}} $
(7) $ \hat H_i^2 = \frac{{\hat \sigma _a^2 + \hat \sigma _d^2}}{{0.5\hat \sigma _a^2 + \hat \sigma _s^2 + \hat \sigma _r^2 + \hat \sigma _p^2 + \hat \sigma _e^2}} $
(8) $ \hat H_f^2 = \frac{{0.5\hat \sigma _a^2 + \hat \sigma _s^2}}{{0.5\hat \sigma _a^2 + \hat \sigma _s^2 + \hat \sigma _r^2/{n_r} + \hat \sigma _p^2/{n_b} + \hat \sigma _e^2/{n_b}{n_i}}} $
(9) 式(7)~(9)中:$ \sigma _d^2 = 4 \times \sigma _s^2, \sigma _a^2, \sigma _d^2, \sigma _s^2, \sigma _r^2, \sigma _p^2$和$\hat { \sigma } _ { e } ^ { 2 }$分别是加性方差、显性方差、特殊配合力方差、正反交效应方差、小区方差和残差方差的估计值。nr、nb和ni分别是正反交、区组和调和小区株树。
根据亲本育种值排名选择前10个家系,并采用基于BLUP估计的育种值和遗传值(育种值+特殊配合力)计算遗传增益[19-20]。
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测定林在16年生和26年生时的保存率分别为44.7%和39.8%,组1、组2与随机交配组合个体存活表现较为一致。对两组全双列交配中自交与对应亲本的异交组合保存率均值进行t检验,未发现显著性差异(表 2)。同样地,组1中自交子代与对应亲本的异交子代的各生长性状也未有明显差异,组2中自交子代的生长表现则均优于对应亲本的异交子代,而且26年生时达到极显著水平(表 2)。
表 2 自交与对应亲本的异交子代的存活率和生长性状的差异分析
Table 2. Statistics of two-tailed t-test for means comparison between selfing and non-selfing progeny from the same parents for survival and growth traits
性状
Traits组1 Group 1 组2 Group 2 t值t-value p值p-value t值t-value p值p-value Sur16 0.69 0.52 0.87 0.41 H16 -0.27 0.80 -1.83 0.16 DBH16 0.68 0.53 -1.94 0.12 V16 0.50 0.64 -2.11 0.12 Sur26 0.93 0.40 0.61 0.56 H26 -0.04 0.97 -3.20 0.01 DBH26 0.92 0.41 -4.72 < 0.01 V26 1.01 0.36 -4.86 < 0.01 注:Sur-存活率;H-树高;DBH-胸径;V-材积指数。下同。
Notes: Sur, H, DBH and V present survival, height, diameter at breast height and volume index, respectively. The following 16 and 26 indicate the tree ages. The same bellow.16年生和26年生各生长性状的表型统计值及变异系数列于表 3。整体上,26年生树高、胸径和材积的表型变异系数分别为6.22%、11.33%和25.92%,胸径的变异系数高于树高,同性状26年生时的变异系数大于16年生,树高的加性遗传变异系数极低(0.01%)。组1、组2与整体的变异趋势基本一致,只是组2各性状的加性遗传变异系数均 < 0.01%。
表 3 存活率和生长性状的描述性统计
Table 3. Descriptive statistics for survival and growth traits
性状Traits 均值
Mean标准差
SD极小值
Min极大值
Max加性遗传变异
系数CVG/%表型变异系
数CVP/%组1
Group 1Sur16/% 45.04 15.68 9.09 75.00 NA NA H16/m 17.52 0.49 16.68 18.32 0.97 5.80 DBH16/cm 15.47 0.68 14.07 16.30 2.56 9.67 V16/dm3 43.01 4.68 34.16 51.44 6.56 24.04 Sur26/% 38.81 14.60 9.09 75.00 NA NA H26/m 22.33 0.87 20.43 23.37 0.52 6.53 DBH26/cm 20.02 1.25 17.59 22.04 4.28 11.85 V26/dm3 93.41 14.81 64.06 117.74 9.00 28.37 组2
Group 2Sur16/% 44.98 9.51 25.00 62.00 NA NA H16/m 17.79 0.43 16.93 19.02 < 0.01 5.43 DBH16/cm 15.46 0.80 12.82 16.93 < 0.01 9.56 V16/dm3 43.96 5.30 28.17 54.61 < 0.01 21.44 Sur26/% 40.36 10.18 16.67 58.00 NA NA H26/m 22.95 0.77 20.19 24.87 < 0.01 7.29 DBH26/cm 20.32 1.30 14.93 23.00 < 0.01 11.85 V26/dm3 98.60 12.93 52.67 126.28 0.01 27.57 全部Total Sur16/% 44.72 11.04 9.09 75.00 NA NA H16/m 17.73 0.48 16.68 19.02 < 0.01 5.70 DBH16/cm 15.58 0.86 12.82 18.29 3.38 9.49 V16/dm3 44.40 5.71 28.17 62.27 7.88 23.34 Sur26/% 39.82 10.59 9.09 75.00 NA NA H26/m 22.88 0.94 20.19 25.87 < 0.01 6.22 DBH26/cm 20.43 1.39 14.93 24.85 4.97 11.33 V26/dm3 99.62 15.98 52.67 161.16 9.49 25.92 注:NA-缺失。
Notes: NA presents not available. -
各性状的方差分量和遗传力估计值见表 4。似然比检验(LRT)结果显示:胸径和材积的加性效应作用极显著,而显性效应则随着年龄而变化,16年生时的显性效应极显著,且大于加性效应($ \sigma _d^2/ \sigma _a^2$=1.33~1.34),至26年生时显性效应消失(方差分量均 < 0.001);相对于胸径和材积,树高的各方差分量较小,16年生时加性效应显著,而26年生时显性效应显著,且大于加性效应($ \sigma _d^2/ \sigma _a^2$=6.36)。组1中的结果与整体基本一致,只是16年生时的显性效应低于加性效应($\hat \sigma _d^2/\hat \sigma _a^2$=0.59~0.85)。树高的各方差分量均 < 0.001,加性和显性效应可忽略不计。组2材料各性状未发现显著的加性和显性,说明其变异主要来源于环境因素,遗传变异极小,也不再进行后续的分析。各性状的正反交效应不显著,暗示交配方向对子代的生长没有影响。在整体材料中,小区对所有性状均有显著作用,表明家系在不同重复间有显著的差异。16年生时,胸径和材积的单株狭义遗传力较低,分别为0.070和0.074,树高遗传力更低为0.028;至26年生时,胸径和材积的单株狭义遗传力分别增加至0.130和0.101,树高则降至0.006。而组1中的胸径和材积受中度遗传控制,16年生时单株狭义遗传力分别为0.127和0.114,26年生时分别增加至0.192和0.134。受SCA的影响,整体材料在26年生时胸径和材积的单株广义遗传力高于单株狭义遗传力,分别0.164和0.173;家系水平上广义遗传力较高,分别为0.572和0.546。26年生时因SCA的作用减弱,胸径和材积的广义遗传力分别降至0.130和0.101;家系遗传力略有下降,分别为0.561和0.501。组1几乎贡献了整体材料的全部遗传变异,因此其遗传力和整体有相同的变化趋势,仅在数值上较高。
表 4 生长性状方差组分及遗传力估计
Table 4. Estimates of variance components, heritability, and standard errors (in parentheses) for growth traits
组别
Group性状
Traits加性方差
${\hat \sigma }$a(SE)2显性方差
${\hat \sigma }$d(SE)2正反交效
应方差
${\hat \sigma }$r(SE)2小区方差
${\hat \sigma }$p(SE)2残差方差
${\hat \sigma }$e(SE)2显性/加
性方差
${\hat \sigma }$d2/a2单株狭义
遗传力
${\hat \sigma }$i(SE)2单株广义
遗传力
${\hat \sigma }$i(SE)2家系广义
遗传力
${\hat \sigma }$f(SE)2组1
Group 1H16 < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)0.382
(0.066)**0.638
(0.046)NA < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)DBH16 0.277
(0.299)**0.164
(0.329)**0.03
(0.057)0.119
(0.083)**1.856
(0.165)0.59 0.127
(0.083)0.202
(0.078)0.642
(0.357)V16 12.233
(15.224)**10.34
(19.26)**0.614
(2.851)12.571
(4.75)**85.48
(7.811)0.85 0.114
(0.084)0.21
(0.078)0.593
(0.337)H26 < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)0.143
(0.087)**1.885
(0.146)NA < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)DBH26 1.031
(0.886)**< 0.001
(< 0.001)< 0.001
(< 0.001)0.478
(0.229)**4.367
(0.444)< 0.01 0.192
(0.099)0.192
(0.109)0.621
(0.349)V26 89.286
(84.68)**< 0.001
(< 0.001)< 0.001
(< 0.001)47.629
(28.112)*574.56
(52.833)< 0.01 0.134
(0.075)0.134
(0.094)0.539
(0.315)组2
Group 2H16 < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)0.144
(0.098)**0.789
(0.133)NA < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)DBH16 < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)2.187
(0.306)NA < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)V16 < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)88.85
(12.442)NA < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)H26 < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)0.016
(0.364)2.784
(0.594)NA < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)DBH26 < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)5.794
(0.905)NA < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)V26 < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)739.077
(115.425)< 0.01 < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)全部
CombinedH16 0.029
(0.063)*< 0.001
(< 0.001)< 0.001
(< 0.001)0.356
(0.049)**0.661
(0.041)< 0.01 0.028
(0.034)0.028
(0.036)0.128
(0.091)DBH16 0.157
(0.173)**0.21
(0.288)**0.013
(0.043)0.084
(0.061)*2.013
(0.124)1.34 0.07
(0.046)0.164
(0.04)0.572
(0.229)V16 7.951
(8.926)**10.578
(15.038)**< 0.001
(< 0.001)9.343
(3.29)**90.961
(5.751)1.33 0.074
(0.05)0.173
(0.043)0.546
(0.218)H26 0.014
(0.103)0.087
(0.28)*< 0.001
(< 0.001)0.159
(0.076)*1.937
(0.122)6.36 0.006
(0.028)0.047
(0.025)0.182
(0.113)DBH26 0.735
(0.479)**< 0.001
(< 0.001)< 0.001
(< 0.001)0.166
(0.162)*5.098
(0.346)< 0.01 0.13
(0.051)0.13
(0.065)0.561
(0.324)V26 70.653
(52.565)**< 0.001
(< 0.001)< 0.001
(< 0.001)12.532
(19.864)*654.455
(43.092)< 0.01 0.101
(0.045)0.101
(0.055)0.501
(0.31)注:*和**分别表示在0.05和0.01水平显著;NA-缺失;SE-标准误。
Note: * and ** present significance at 0.05 and 0.01 level, respectively; NA presents not available; SE indicates standard error. -
根据亲本育种值选择排名前10的组合,然后分别根据育种值和遗传效应值(育种值+SCA效应值)估算遗传增益,结果见表 5。仅根据育种值进行选择,16年生时胸径和材积的遗传增益分别为2.76%和7.12%;当联合育种值和SCA效应值进行选择时,遗传增益分别为3.21%和8.04%,比单纯育种值选择遗传增益分别提高16.30%和12.92%。26年生时SCA消失,根据育种值和遗传值进行选择的效果一致,胸径和材积的遗传增益分别为6.31%和11.00%(表 5)。
表 5 各生长性状根据育种值和遗传值选择的遗传增益
Table 5. Estimates of genetic gains from selection based on breeding values and genetic values
性状
Traits均值
Mean育种值
Breeding value遗传值
Genetic value育种值遗传增益/%
Genetic gains with breeding value遗传值遗传增益/%
Genetic gains with genetic valueH16/m 17.73 0.12 0.12 0.68 0.68 DBH16/cm 15.58 0.43 0.50 2.76 3.21 V16/dm3 44.40 3.16 3.57 7.12 8.04 H26/m 22.88 0.05 0.08 0.22 0.35 DBH26/cm 20.43 1.29 1.29 6.31 6.31 V26/dm3 99.62 10.96 10.96 11.00 11.00
日本落叶松全双列交配生长性状的遗传分析
Genetic Analysis of Larix kaempferi Growth Traits in Full-diallel Crosses
-
摘要:
目的 通过全双列交配设计开展配合力分析,研究SCA(特殊配合力)对重要性状的相对贡献,旨在为日本落叶松的育种策略制定和育种群体管理提供重要遗传参数信息。 方法 本研究以13个母本和23个父本获得的6×6和4×4两组全双列交配设计的日本落叶松子代测定林为对象,利用空间模型对16年生和26年生的生长表型数据进行校正,采用单株模型对其配合力和正反交效应分析,研究加性效应、SCA效应的相对重要性及其在落叶松育种中的应用。 结果 自交子代在保存率性状上未表现出明显的自交衰退,生长性状在一些亲本中甚至优于相应异交后代;生长性状的正反交效应不明显,暗示在今后的育种中无需考虑交配方向。16年生时胸径和材积加性效应和显性效应显著,且显性效应大于加性效应,显性和加性效应方差比值分别为1.34和1.33。此时按亲本育种值排名选择前10个家系,当联合育种值和SCA进行选择时遗传增益分别为3.21%和8.04%,比单纯育种值选择的遗传增益(分别为2.76%和7.12%)分别提高16.30%和12.92%;26年生时胸径和材积的加性效应显著而显性效应消失。16年生时胸径和材积的单株狭义遗传力、单株广义遗传力和平均家系广义遗传力的变幅分别为0.070~0.074、0.164~0.173和0.546~0.572;26年生时胸径和材积的单株狭义遗传力分别增至0.13和0.10。 结论 胸径和材积在早期的显性效应显著,通过对SCA的利用可获得更高的遗传增益,即选择一般配合力高的亲本通过控制授粉配制高特殊配合力组合的种子,并从子代中选择优良单株进行无性扩繁,可获得最大化的遗传增益。 Abstract:Objective The analysis of combining ability was conducted through full-diallel mating design to study the relative contribution of special combining ability (SCA) to economically important traits, aiming to provide important genetic parameter information for the development of Japanese larch (Larix kaempferi) breeding strategies and the management of breeding populations. Method The progeny trail mainly consists of two full-diallels (6×6 and 4×4, respectively) derived from 13 parent trees used as female and 23 used as male. The phenotypic data for growth traits measured at age 16 and age 26 were adjusted using first order autoregression model, and then were used in individual mixed models to conduct combining ability and reciprocal effect analysis. The importance of dominance related to additive effect was investigated and its application in Japanese larch breeding was discussed. Result The self-crossed offspring showed no significant self-depression in the survival, and even outperformed the corresponding non-selfed individuals in some parents in terms of growth. The reciprocal effects of the growth traits were not notable, indicating that the mating direction needs not to be considered in future's breeding. At age 16, the additive and dominance effect of DBH and volume were significant, and the dominant effect was greater than the additive effect. At this age, the genetic gains were 3.21% and 8.04% when selected for the top 10 families based on combining breeding value and SCA, which increased by 16.30% and 12.92% compared with selection on breeding value solely (2.76% and 7.12%, respectively). The additive effects of DBH and volume were significant at age 26 while the dominant effects were disappeared. The narrow-sense heritability, broad sense heredity and broad family heritability of the average family were 0.070-0.074, 0.164-0.173, and 0.546-0.572 for DBH and volume at age 16, and the narrow-sense heredity increased to 0.13 and 0.10 for this two traits, respectively. Conclusion The SCA effects are significant for DBH and volume at age 16. More genetic gain can be captured by the utilization of SCA through producing improved seeds from mating those parent pairs with high GCA and high SCA effects. Further, genetic gain can be maximized through vegetative propagation of the trees developed from those seeds. -
Key words:
- tree genetic improvement
- / SCA
- / full-sib
- / genetic parameters
- / Larix kaempferi
-
表 1 交配设计表
Table 1. Mating design table
母本
Maternal
parents父本Paternal parents RC13 RC19 RC4 RC5 RC503 RC82 RF27 Y11 Y16 R1 Y33 Y5 Y13 RC2 RC34 RC35 RC40 RC402 RC404 RC41 RC61 RC8 RC81 RC13 ⊗ × × × × × × RC19 × ⊗ × × × × RC4 × × ⊗ × × RC5 × × × ⊗ × × × × × RC503 × × × × × × RC82 × × × × × × × × × RF27 ⊗ × × Y11 × × × Y16 × × ⊗ × R1 × × × ⊗ Y55 × Y10 × × × × × × × RC25 × × 注:×-交配;⊗-自交。
Notes: × and ⊗ indicate crossing and selfing, respectively.表 2 自交与对应亲本的异交子代的存活率和生长性状的差异分析
Table 2. Statistics of two-tailed t-test for means comparison between selfing and non-selfing progeny from the same parents for survival and growth traits
性状
Traits组1 Group 1 组2 Group 2 t值t-value p值p-value t值t-value p值p-value Sur16 0.69 0.52 0.87 0.41 H16 -0.27 0.80 -1.83 0.16 DBH16 0.68 0.53 -1.94 0.12 V16 0.50 0.64 -2.11 0.12 Sur26 0.93 0.40 0.61 0.56 H26 -0.04 0.97 -3.20 0.01 DBH26 0.92 0.41 -4.72 < 0.01 V26 1.01 0.36 -4.86 < 0.01 注:Sur-存活率;H-树高;DBH-胸径;V-材积指数。下同。
Notes: Sur, H, DBH and V present survival, height, diameter at breast height and volume index, respectively. The following 16 and 26 indicate the tree ages. The same bellow.表 3 存活率和生长性状的描述性统计
Table 3. Descriptive statistics for survival and growth traits
性状Traits 均值
Mean标准差
SD极小值
Min极大值
Max加性遗传变异
系数CVG/%表型变异系
数CVP/%组1
Group 1Sur16/% 45.04 15.68 9.09 75.00 NA NA H16/m 17.52 0.49 16.68 18.32 0.97 5.80 DBH16/cm 15.47 0.68 14.07 16.30 2.56 9.67 V16/dm3 43.01 4.68 34.16 51.44 6.56 24.04 Sur26/% 38.81 14.60 9.09 75.00 NA NA H26/m 22.33 0.87 20.43 23.37 0.52 6.53 DBH26/cm 20.02 1.25 17.59 22.04 4.28 11.85 V26/dm3 93.41 14.81 64.06 117.74 9.00 28.37 组2
Group 2Sur16/% 44.98 9.51 25.00 62.00 NA NA H16/m 17.79 0.43 16.93 19.02 < 0.01 5.43 DBH16/cm 15.46 0.80 12.82 16.93 < 0.01 9.56 V16/dm3 43.96 5.30 28.17 54.61 < 0.01 21.44 Sur26/% 40.36 10.18 16.67 58.00 NA NA H26/m 22.95 0.77 20.19 24.87 < 0.01 7.29 DBH26/cm 20.32 1.30 14.93 23.00 < 0.01 11.85 V26/dm3 98.60 12.93 52.67 126.28 0.01 27.57 全部Total Sur16/% 44.72 11.04 9.09 75.00 NA NA H16/m 17.73 0.48 16.68 19.02 < 0.01 5.70 DBH16/cm 15.58 0.86 12.82 18.29 3.38 9.49 V16/dm3 44.40 5.71 28.17 62.27 7.88 23.34 Sur26/% 39.82 10.59 9.09 75.00 NA NA H26/m 22.88 0.94 20.19 25.87 < 0.01 6.22 DBH26/cm 20.43 1.39 14.93 24.85 4.97 11.33 V26/dm3 99.62 15.98 52.67 161.16 9.49 25.92 注:NA-缺失。
Notes: NA presents not available.表 4 生长性状方差组分及遗传力估计
Table 4. Estimates of variance components, heritability, and standard errors (in parentheses) for growth traits
组别
Group性状
Traits加性方差
${\hat \sigma }$a(SE)2显性方差
${\hat \sigma }$d(SE)2正反交效
应方差
${\hat \sigma }$r(SE)2小区方差
${\hat \sigma }$p(SE)2残差方差
${\hat \sigma }$e(SE)2显性/加
性方差
${\hat \sigma }$d2/a2单株狭义
遗传力
${\hat \sigma }$i(SE)2单株广义
遗传力
${\hat \sigma }$i(SE)2家系广义
遗传力
${\hat \sigma }$f(SE)2组1
Group 1H16 < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)0.382
(0.066)**0.638
(0.046)NA < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)DBH16 0.277
(0.299)**0.164
(0.329)**0.03
(0.057)0.119
(0.083)**1.856
(0.165)0.59 0.127
(0.083)0.202
(0.078)0.642
(0.357)V16 12.233
(15.224)**10.34
(19.26)**0.614
(2.851)12.571
(4.75)**85.48
(7.811)0.85 0.114
(0.084)0.21
(0.078)0.593
(0.337)H26 < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)0.143
(0.087)**1.885
(0.146)NA < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)DBH26 1.031
(0.886)**< 0.001
(< 0.001)< 0.001
(< 0.001)0.478
(0.229)**4.367
(0.444)< 0.01 0.192
(0.099)0.192
(0.109)0.621
(0.349)V26 89.286
(84.68)**< 0.001
(< 0.001)< 0.001
(< 0.001)47.629
(28.112)*574.56
(52.833)< 0.01 0.134
(0.075)0.134
(0.094)0.539
(0.315)组2
Group 2H16 < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)0.144
(0.098)**0.789
(0.133)NA < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)DBH16 < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)2.187
(0.306)NA < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)V16 < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)88.85
(12.442)NA < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)H26 < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)0.016
(0.364)2.784
(0.594)NA < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)DBH26 < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)5.794
(0.905)NA < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)V26 < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)739.077
(115.425)< 0.01 < 0.001
(< 0.001)< 0.001
(< 0.001)< 0.001
(< 0.001)全部
CombinedH16 0.029
(0.063)*< 0.001
(< 0.001)< 0.001
(< 0.001)0.356
(0.049)**0.661
(0.041)< 0.01 0.028
(0.034)0.028
(0.036)0.128
(0.091)DBH16 0.157
(0.173)**0.21
(0.288)**0.013
(0.043)0.084
(0.061)*2.013
(0.124)1.34 0.07
(0.046)0.164
(0.04)0.572
(0.229)V16 7.951
(8.926)**10.578
(15.038)**< 0.001
(< 0.001)9.343
(3.29)**90.961
(5.751)1.33 0.074
(0.05)0.173
(0.043)0.546
(0.218)H26 0.014
(0.103)0.087
(0.28)*< 0.001
(< 0.001)0.159
(0.076)*1.937
(0.122)6.36 0.006
(0.028)0.047
(0.025)0.182
(0.113)DBH26 0.735
(0.479)**< 0.001
(< 0.001)< 0.001
(< 0.001)0.166
(0.162)*5.098
(0.346)< 0.01 0.13
(0.051)0.13
(0.065)0.561
(0.324)V26 70.653
(52.565)**< 0.001
(< 0.001)< 0.001
(< 0.001)12.532
(19.864)*654.455
(43.092)< 0.01 0.101
(0.045)0.101
(0.055)0.501
(0.31)注:*和**分别表示在0.05和0.01水平显著;NA-缺失;SE-标准误。
Note: * and ** present significance at 0.05 and 0.01 level, respectively; NA presents not available; SE indicates standard error.表 5 各生长性状根据育种值和遗传值选择的遗传增益
Table 5. Estimates of genetic gains from selection based on breeding values and genetic values
性状
Traits均值
Mean育种值
Breeding value遗传值
Genetic value育种值遗传增益/%
Genetic gains with breeding value遗传值遗传增益/%
Genetic gains with genetic valueH16/m 17.73 0.12 0.12 0.68 0.68 DBH16/cm 15.58 0.43 0.50 2.76 3.21 V16/dm3 44.40 3.16 3.57 7.12 8.04 H26/m 22.88 0.05 0.08 0.22 0.35 DBH26/cm 20.43 1.29 1.29 6.31 6.31 V26/dm3 99.62 10.96 10.96 11.00 11.00 -
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