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Jun.  2021
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Research on Variation Pattern of Wood Properties of Red-heart Chinese Fir Plus Trees, a Featured Provenance from Rongshui of Guangxi

  • Corresponding author: HUANG Kai-yong, huangky73@163.com
  • Received Date: 2016-07-29
  • Objective To Measure and analyze the wood properties of red-heart Chinese fir (Cunninghamia lanceolata) plus trees, the provenance from Rongshui of Guangxi, and to investigate the wood properties variation pattern of the trees. Method Sixty 20-year-old plus trees of red-heart Chinese fir were chosen as research material. The wood properties such as wood volume, wood basic density, tissue proportion, tracheid properties and tracheid microfibril angle were assayed and the variation pattern of relevant wood properties was analyzed. Result The heartwood ratio, wood basic density, xylem ray proportion, tracheid proportion, tracheid length, tracheid width, and the length-width ratios of tracheid obeyed normal distribution. The variable coefficient of axially parenchyma proportion of red-heart Chinese fir plus trees from two plots in Rongshui was 35.08% and 44.97% respectively, showing a significant variation. The variable coefficient of tracheid proportion was 3.28% and 3.56% respectively, showing a small variation. There were highly significant differences (P < 0.01) in heartwood ratio, xylem ray proportion, tracheid length, tracheid width, tracheid microfibril angle and significant differences(P < 0.05)in parenchyma proportion between the plus trees from the two locations. There were 12 significant phenotype correlations among 10 wood properties data. Conclusion All the 10 characters assayed are continual quantitative. The data of tracheid length, tracheid width and length-width ratios of tracheid of early wood and late wood increase from heartwood to sapwood. The heartwood ratio and individual volume are both important economic characters. There are very significant positive correlation (P < 0.01) between heartwood ratio and xylem ray proportion, and significant positive correlation (P < 0.05) between the individual volume and axially parenchyma proportion. The volume of individual tree does not correlate significantly with the wood basic density, which makes it possible to the genetic improvement of volume combined with wood properties of red-heart Chinese fir.
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Research on Variation Pattern of Wood Properties of Red-heart Chinese Fir Plus Trees, a Featured Provenance from Rongshui of Guangxi

    Corresponding author: HUANG Kai-yong, huangky73@163.com
  • 1. Guangxi Forestry Research Institute, Key Laboratory of Central South Fast-growing Timber Cultivation of State Forestry Administration, Guangxi Key Laboratory of Superior Timber Trees Resource Cultivation, Nanning 530002, Guangxi, China
  • 2. Beijianghe State-owned Forest Farm of Guangxi, Rongshui 545300, Guangxi, China

Abstract:  Objective To Measure and analyze the wood properties of red-heart Chinese fir (Cunninghamia lanceolata) plus trees, the provenance from Rongshui of Guangxi, and to investigate the wood properties variation pattern of the trees. Method Sixty 20-year-old plus trees of red-heart Chinese fir were chosen as research material. The wood properties such as wood volume, wood basic density, tissue proportion, tracheid properties and tracheid microfibril angle were assayed and the variation pattern of relevant wood properties was analyzed. Result The heartwood ratio, wood basic density, xylem ray proportion, tracheid proportion, tracheid length, tracheid width, and the length-width ratios of tracheid obeyed normal distribution. The variable coefficient of axially parenchyma proportion of red-heart Chinese fir plus trees from two plots in Rongshui was 35.08% and 44.97% respectively, showing a significant variation. The variable coefficient of tracheid proportion was 3.28% and 3.56% respectively, showing a small variation. There were highly significant differences (P < 0.01) in heartwood ratio, xylem ray proportion, tracheid length, tracheid width, tracheid microfibril angle and significant differences(P < 0.05)in parenchyma proportion between the plus trees from the two locations. There were 12 significant phenotype correlations among 10 wood properties data. Conclusion All the 10 characters assayed are continual quantitative. The data of tracheid length, tracheid width and length-width ratios of tracheid of early wood and late wood increase from heartwood to sapwood. The heartwood ratio and individual volume are both important economic characters. There are very significant positive correlation (P < 0.01) between heartwood ratio and xylem ray proportion, and significant positive correlation (P < 0.05) between the individual volume and axially parenchyma proportion. The volume of individual tree does not correlate significantly with the wood basic density, which makes it possible to the genetic improvement of volume combined with wood properties of red-heart Chinese fir.

  • 杉木(Cunninghamia ianceolata (Lamb.)Hook.)是我国特有的用材树种,分布于我国秦岭和长江流域以南各省区, 其生长快,材性好,用途多,栽培面积大,是我国最重要的用材和商品材树种之一。自20世纪50年代,我国开始以速生、高产为目标的杉木遗传育种工作取得了显著成效,针对材性的改良已成为现阶段杉木育种的重要目标。1976年中国林科院主持的全国杉木地理种源试验,收集全国62个种源的杉木在南方省区66个试验点进行测定研究。因种质好,速生性强,生产力高,适应性广等特点,确定广西融水杉木种源为全国优良种源[1]

    融水杉木主要包括糠杉和黄枝杉2种类别。本研究踏查选取广西融水种源60株20年生的杉木优树,其叶片较尖而稍硬,先端锐尖,叶表面有光泽,嫩枝和新叶为浅绿色,无白色蜡粉,属杉木的黄枝杉类别[2]。黄枝杉因心材比例大且色红而坚实,又被素称红心杉。目前,红心杉木良种选育、苗木培育等相关研究已在江西、广东陆续开展[3-4],但关于红心杉木材质性状变异规律的研究仍无报道。本研究通过测定红心杉木优树胸径、红心率、基本密度、组织比量、管胞性状、微纤丝角等材质指标,深入分析红心杉材质性状的变异规律,为开展广西融水种源红心杉的遗传多样性、木材产量与质量的综合遗传改良等研究工作打下良好的基础, 具有重要的理论价值和生产实践应用意义。

1.   材料
  • 运用5株优势木对比法(国标GB10018),在广西融水县永乐和下洞两乡分别踏查选取20年生红心杉优树30株,共60株。红心杉优树生长地的地理及林分情况见表 1。在红心杉优树1.3 m处,用直径5 mm的生长锥从北朝南方向取由树皮至髓心的完整木芯2根,其中,1根用于基本密度、组织比量和微纤丝角测定,另1根用于管胞形态测定。把每根木芯(从髓心至树皮)平均切成三部分,即心部(H)、中部(M)和边部(S)。

    地区
    Area
    纬度(N)
    Latitude/(°)
    经度(E)
    Longitude/(°)
    平均海拔
    Average Elevation/m
    林分组成
    Stand Composition
    郁闭度
    Canopy Density
    林分密度
    Stand Density/(tree·hm-2)
    永乐Yongle 25.03 109.09 395 纯林 0.7 2 200
    下洞Xiadong 25.51 109.17 235 纯林 0.7 2 800

    Table 1.  Geographical and stand situation of plus trees growth of Red-heart Chinese Fir

2.   方法
  • 按广西林业勘测设计院编制的杉木立木蓄积公式[5]计算材积(V):

    式中: D为胸径;H为树高。

  • 将每根木芯分别按心部(H)、中部(M)和边部(S)采用排水法测定生材体积(V),然后把木样置于鼓风干燥箱烘至绝干,称质量(W)。根据公式ρ=W/V计算基本密度(ρ)。具体方法依照《GB-1933-2009木材密度测定方法》进行。

  • 将每根木芯的心部(H)、中部(M)和边部(S)分别按早晚材取样,即H-早、H-晚、M-早、M-晚、S-早、S-晚,对6个部分样品进行测定。采用富兰克林离析法[6]对木材管胞进行离析。利用Nikon 80i显微成像系统对每一试件随机测定100根完整管胞长度、宽度和腔径。

  • 采用徒手切片法对上述每根木芯的心部(H)、中部(M)和边部(S)进行横切面切取,然后对切片进行蕃红染色、脱水、透明、树胶封片处理,制成玻片。采用普通光学显微镜, 用0.5网形目镜尺,每个试样测定60次以上,计算各组织的百分比量。

  • 采用徒手切片法对上述每根木芯的心部(H)、中部(M)和边部(S)进行弦切面切取。采用硝酸-铬酸法离析切片,采用Nikon 80i显微成像系统测定次生壁S2层的微纤丝与管胞主轴的夹角,微纤丝角测定100次。

  • 统计各优树的基本密度、木射线比量、轴向薄壁细胞比量、管胞比量、管胞长、管胞宽、管胞长宽比、微纤丝角数据,取平均值。百分率性状数据经反正弦(arcsin $ \sqrt x $) 转换后统计,采用SPSS19.软件进行数据分析。

3.   结果与分析
  • 对60株红心杉优树的材积、红心率、基本密度、木射线比量、轴向薄壁细胞比量、管胞比量、管胞长、管胞宽、管胞长宽比、微纤丝角等10个材性表型性状数据频度分布的分析表明:60株红心杉优树所有10个性状均具有连续性数量性状(图 1)。对60株红心杉优树的10个表型性状数据进行正态分布检验,从表 2看出:材积、红心率、基本密度、轴向薄壁细胞比量、管胞比量性状偏度数据为正值,呈正偏态分布,其余性状数据呈负偏态分布。材积、轴向薄壁细胞比量、微纤丝角性状P值<0.05,表明这3个性状数据不服从正态分布,其余性状数据符合正态分布。

    Figure 1.  Histogram of 10 wood property characters number of Red-heart Chinese Fir from Rongshui

    统计量
    Statistics
    单株材积
    Individual Volume
    红心率
    Heartwood ratio
    基本密度
    Wood basic density
    木射线比量
    Xylem ray proportion
    轴向薄壁细胞比量
    Axially parenchyma proportion
    偏度Skewness 0.821 0.136 0.026 -0.273 0.643
    峰度Kurtosis -0.048 -0.626 -0.311 -0.553 -0.355
    W检验hapiro-Wilk 0.925 0.982 0.990 0.969 0.948
    P值P-value 0.002 0.560 0.907 0.135 0.020
    统计量
    Statistics
    管胞比量
    Tracheid properties
    管胞长
    Tracheid length
    管胞宽
    Tracheid width
    管胞长宽比
    Length-width ratios
    of tracheid
    微纤丝角
    Tracheid microfibril
    angle
    偏度Skewness 0.138 -0.095 -0.134 -0.087 -0.342
    峰度Kurtosis -0.742 -0.931 -0.390 -0.619 -0.986
    W检验Shapiro-Wilk 0.980 0.972 0.989 0.989 0.960
    P值P-value 0.448 0.207 0.881 0.895 0.049

    Table 2.  Normal distribution test of 10 wood property characters number of Red-heart Chinese Fir from Rongshui

  • 对60株红心杉优树心部(H)、中部(M)和边部(S)的早、晚材管胞长度、管胞宽度、管胞长宽比等进行测定,结果(图 2)表明:红心杉优树早、晚材的管胞长度、管胞宽度和管胞长宽比都呈从木材心部至边部逐渐增加的趋势,心部、中部及边部晚材的管胞长度和管胞长宽比均比早材的大,而心部、中部及边部晚材的管胞宽度均比早材的小。

    Figure 2.  Radial variation of Tracheid morphology of Red-heart Chinese Fir from Rongshui

  • 对融水2地区各30株红心杉优树材质性状变异的分析(表 3)表明:红心杉木材品质在融水2地区具有不同程度的差异。经T-检验,融水2地区优树间红心率、木射线比量、管胞长、管胞宽、微纤丝角等性状差异极显著(P<0.01),轴向薄壁细胞比量差异显著(P<0.05),单株材积和基本密度差异显著(P<0.1)。2地区红心杉优树轴向薄壁细胞比量变异系数分别为35.08%和44.97%,变异较大;管胞比量变异系数分别为3.28%和3.56%,变异较小。材积和红心率作为重要的经济性状,2地区红心杉优树单株材积和红心率平均值分别为0.684、0.594 m3和46.809%、55.042%。

    地区
    Area
    单株材积
    Individual volume
    红心率
    Heartwood ratio
    基本密度
    Wood basic density
    木射线比量
    Xylem ray proportion
    轴向薄壁细胞比量
    Axially parenchyma
    roportion
    均值
    Mean/m3
    变异系数
    C·V/%
    均值
    Mean/%
    变异系数
    C·V/%
    均值Mean/
    (g·cm-3)
    变异系数
    C·V/%
    均值
    Mean/%
    变异系数
    C·V/%
    均值
    Mean/%
    变异系数
    C·V/%
    永乐Yongle 0.684±0.155 22.70 46.809±8.236 17.60 0.304±0.023 7.46 8.726±1.371 14.55 5.319±1.916 35.08
    下洞Xiadong 0.594±0.117 19.75 55.042±8.070 14.66 0.288±0.041 14.32 10.046±1.813 18.42 4.432±2.087 44.97
    T-检验T-test 2.545 -5.073** 1.952 -4.076** 2.524*
    地区
    Area
    管胞比量
    Tracheid proportion
    管胞长
    Tracheid length
    管胞宽
    Tracheid width
    管胞长宽比
    Length-width ratios
    of tracheid
    微纤丝角
    Tracheid microfibril
    angle
    均值
    Mean/%
    变异系数
    C·V/%
    均值
    Mean/μm
    变异系数
    C·V/%
    均值
    Mean/μm
    变异系数
    C·V/%
    均值
    Mean
    变异系数
    C·V/%
    均值
    Mean/(°)
    变异系数
    C·V/%
    永乐Yongle 85.955±2.891 3.28 2 502.7±233.1 9.31 40.076±1.804 4.50 66.224±5.673 8.57 7.771±0.576 7.38
    下洞Xiadong 85.522±3.046 3.56 2 296.7±246.1 10.71 38.965±1.887 4.84 64.250±6.466 10.97 6.820±0.543 7.96
    T-检验T-test 0.431 3.236** 2.897** 0.861 7.324**
    注:Δ表示0.1显著;*表示0.05显著;**表示0.01显著,下同。
    Note:Δ presents 0.1 significant difference level, * presents 0.05 significant difference level, **presents 0.01 significant difference level.The same below.

    Table 3.  Variation of 10 wood propertiesof Red-heart Chinese Fir from Rongshui

  • 运用Spearman方法对融水红心杉优树材质性状进行相关性分析,结果(表 4)表明:10个材质性状间存在12对表型显著相关,其中,红心率与木射线比量、木射线比量与轴向薄壁细胞比量、木射线比量与管胞比量、木射线比量与微纤丝角、轴向薄壁细胞比量与管胞比量、管胞长与管胞宽、管胞长与管胞长宽比、管胞长与微纤丝角、管胞宽与微纤丝角等9对性状极显著相关(P<0.01),单株材积与轴向薄壁细胞比量、基本密度与管胞宽等2对性状显著相关(P<0.05),单株材积与管胞比量显著相关(P<0.1)。

    红心率
    Heartwood
    ratio
    基本密度
    Wood Basic
    density
    木射线比量
    Xylem ray
    proportion
    轴向薄壁细胞
    比量Axially par-
    enchyma proportion
    管胞比量
    Tracheid
    proportion
    管胞长
    Tracheid
    length
    管胞宽
    Tracheid
    width
    管胞长宽比
    Length-width
    ratios of tracheid
    微纤丝角
    Tracheid mic-
    rofibril angle
    单株材积Individual volume 0.05 -0.18 0.16 0.29* -0.23 0.17 0.04 0.18 -0.14
    红心率Heartwood ratio -0.08 0.36** -0.01 -0.16 0.03 -0.19 0.20 -0.05
    基本密度Wood basic density -0.16 0.00 0.10 -0.14 -0.27* -0.01 0.16
    木射线比量Xylem ray proportion 0.33** -0.71** 0.00 -0.02 0.12 -0.34**
    轴向薄壁细胞比量
    Axially parenchyma proportion
    -0.86** 0.15 0.03 0.14 0.08
    管胞比量Tracheid proportion -0.07 -0.04 -0.09 0.09
    管胞长Tracheid length 0.57** 0.81** 0.42**
    管胞宽Tracheid width 0.08 0.35**
    管胞长宽比
    Length-width ratios of tracheid
    0.16

    Table 4.  Correlation analysis of 10 wood propertiesof Red-heart Chinese Fir from Rongshui

4.   讨论
  • 生长量直接决定林木个体和单位面积的木材产量, 是用材树种遗传改良重要的育种目标,而生长与材质性状的遗传相关性是目前林木遗传研究的热点,研究主要集中在生长性状与木材基本密度、管胞长度、管胞宽等材性性状的相关关系上[7-8]。木材密度是木材品质重要的因子之一, 它影响木材的力学强度, 直接或间接决定木材的产量和品质。木材基本密度作为针叶树材性的重要指标,在针叶材改良中具有重要地位[9-11]。有研究表明,杉木生长与材质性状间呈不显著负相关[12-15]。本研究中,融水红心杉优树单株材积与轴向薄壁细胞比量呈显著正相关,与管胞比量呈显著负相关,而与基本密度间呈不显著负相关,这使红心杉生长量与材性相结合的遗传改良成为可能。

    本研究采用生长锥取样测定分析融水红心杉优树材质性状变异规律,选取木质部心部、中部和边部3个样点进行材质性状测定, 基本反应了株内径向的变异趋势[16-17]。融水红心杉优树胸径处管胞长度从髓心到树皮表现为增加的趋势, 径向变异规律应属PashinⅡ型[18]。针叶树材主要由管胞、薄壁细胞、木射线组织构成,而管胞组织占针叶木材体积的90%以上[19]。有研究表明,管胞长宽比大于50的木材适宜制浆造纸,且愈大愈好[19];管胞长度保持在330 μm以上, 纸张的耐折度、撕裂指数、耐破指数等不再受其影响, 而管胞长度越长、管胞长宽比越大,纸张的各项物理力学性能均得到有效提高[21-22]。本研究中,融水红心杉优树的管胞比量分布在80.012%~91.937%,管胞长度分布在1 951~2 898 μm,远大于330 μm,管胞宽度分布在36~44 μm,管胞长宽比分布在52~79。融水红心杉木材不仅是优异的建筑材,也是良好的纸浆材。

    微纤丝角为细胞壁中纤维素链的螺旋卷索与纤维轴之间的夹角[23]。已有研究表明, 微纤丝角影响木材管胞的基本强度和特性[24], 与木材强度、硬度密切相关[25],是无节木材强度的重要决定因素,是影响木材综合性质的一个重要因子。本研究中,融水红心杉优树微纤丝角分布在5.879°~8.675°,且微纤丝角与木射线比量显著负相关,与管胞长、管胞宽显著正相关。

    近年来,融水种源杉木面临着基因资源逐渐丧失的困境,种质资源的保护已成当务之急。本研究通过对融水下洞、永乐2地区红心杉优树材质性状研究发现,2地区红心杉优树在材质性状上存在广泛而显著差异,本研究结果为开展融水红心杉种质资源的挖掘、保护和利用工作提供了坚实的理论依据和实践基础。

5.   小结
  • 通过分析广西融水种源红心杉优树材质性状的变异规律,发现测定的10个性状均是连续性数量性状,红心率、基本密度、木射线比量、管胞比量、管胞长、管胞宽、管胞长宽比等性状数据服从正态分布;早、晚材的管胞长度、管胞宽度和管胞长宽比等性状数据呈从心材至边材逐渐增加的规律;单株材积和红心率是重要的经济性状,这2个性状在两地区的表型平均值分别为0.684、0.594 m3和46.809%、55.042%;红心率等6个表型性状存在极显著或显著差异。10个材质性状间存在12对表型显著相关,其中,单株材积与轴向薄壁细胞比量呈显著正相关(P<0.05),红心率与木射线比量呈极显著正相关(P<0.01),而单株材积与基本密度相关不显著,这使红心杉生长量与材性相结合的遗传改良成为可能。

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