木材定量解剖研究新进展

doi:10.11707/j.1001-7488.LYKX20220849

摘要/Abstract

摘要：

木材解剖学是研究木材（次生木质部）中各类细胞形态特征、排列、比量、结构和功能的一门科学。在木材解剖学框架内，开展木材构造特征定量化研究，实现木材解剖信息的定量提取、挖掘、整合与应用，科学构建木材构造特征与树种生理功能、气候环境、木材性质和加工工艺等之间的相互关系，可进一步推动木材学、林学、植物学、古生物学、古气候学、考古学和物候学等学科的发展。当前，在系统科学的时代背景下，梳理木材定量解剖研究的发展脉络，厘清木材构造特征的结构性和整体性，从系统论视角探讨木材定量解剖研究的涌现性质，有利于促进木材解剖学的跨学科交叉融合，开创木材解剖学发展新格局。本研究首先介绍木材解剖学的重要发展阶段，针对木材构造特征的共性和多样性，从样品制备、信息采集和数据分析等方面总结木材定量解剖研究的主要方法，然后分别从木材构造特征与树木系统进化、木材构造特征与气候环境变化、木材构造特征与木材性质及加工利用关系3方面归纳近10年来木材定量解剖研究取得的最新进展，最后针对当前存在的问题与不足，提出未来发展展望：1）木材定量解剖研究方法的持续性革新迭代，推动木材定量解剖高通量测试与分析平台的研发，为木材解剖学的广泛应用提供基础；2）木材信息资源及其共享体系的全球化构建完善，增强木材定量解剖数据的可靠性和共享性，为发展基于数据驱动的木材科学研究新范式提供重要基础；3）木材定量解剖研究体系的多学科交叉融合，从底层逻辑和层级架构视角比较木材解剖学与植物解剖学、生态学、木材加工利用等领域的关联与异同，促进多学科融合创新发展。通过进一步开展木材定量解剖研究，加强木材解剖学的跨学科交叉融合，从木材解剖学角度推动基于数据驱动的木材科学研究新范式的构建与发展，能够为林木培育、森林经营、树木分类、森林碳汇、气候变化、木材生产及可持续利用等研究提供理论基础和科学依据。

关键词: 次生木质部, 木材解剖学, 定量解剖, 构造特征, 高通量测试

Abstract:

Wood anatomy studies about the morphological characteristics, cell arrangement, tissue proportion, structure and function of various cells in secondary xylem, the wood. In the frame of wood anatomy research, carrying out quantitative wood anatomy (QWA) can realize the quantitative extraction, mining, integration and application of wood anatomical traits information. It can further be beneficial for scientifically constructing the relationships between wood anatomical traits and physiological functions of tree species, climate and environment, wood properties and processing technology. QWA can also advance the developments of other research fields, including wood science, forestry, botany, paleontology, palaeoclimatology, archaeology and phenology. Therefore, in the era of systematic science, we should find out the developmental characteristics of quantitative anatomy of wood, clarify the structural frame and integrity of wood anatomical traits, and discuss the emerging nature of quantitative wood anatomy of wood from the perspective of scientific system theory. This would enhance the multi-disciplinary integration and update the quantitative wood anatomy. Hence, in this review, we firstly introduced the important development stages of wood anatomy, and then described main methods for quantitative wood anatomy studies on basis of sample preparation, information collection and data analysis. Furthermore, new research progress on QWA over the decade was summarized from three aspects; wood anatomy and tree system evolution, wood anatomy and climate and environment change, wood anatomy and wood properties, processing and utilization. Finally, to solve the problems and shortcomings of QWA, three prospects for future development are proposed. 1) Innovate the methods for QWA, and build the high flux testing and analysis platform for QWA. This could provide a basis for the wide application of wood anatomy. 2) Improve wood information resources and build the information sharing system, enhance the reliability and sharing of QWA information. This can provide an important basis for the development of a new paradigm of wood science research driven by data. 3) Frame the QWA research from the perspective of systematic development theory. Comparatively investigate the similarities and differences between wood anatomy and other fields, such as plant anatomy, ecology, wood processing and utilization by the view of underlying logic and hierarchy architecture. Promote and innovate the multidisciplinary development. QWA promotes multi-disciplinary integration, and provides theoretical and scientific basis for research on forest cultivation, forest management, tree classification, forest carbon sink, climate change, wood production and sustainable utilization.

Key words: secondary xylem, wood anatomy, quantitative wood anatomy, anatomical traits, high flux testing

中图分类号:

S781

郭娟,焦立超,何拓,马灵玉,姜笑梅,殷亚方. 木材定量解剖研究新进展[J]. 林业科学, 2025, 61(2): 204-218.

Juan Guo,Lichao Jiao,Tuo He,Lingyu Ma,Xiaomei Jiang,Yafang Yin. New Research Progress on Quantitative Wood Anatomy[J]. Scientia Silvae Sinicae, 2025, 61(2): 204-218.

图/表 3

表1

木材构造特征的定量测试分析方法①"

主要构造特征 Main anatomical traits	定义 Definition	定量测试分析方法 Methods for quantitative analysis	推荐样本量 Recommended sampling number	参考文献 Reference
组织比量（轴向管胞、木纤维、导管、木射线、轴向薄壁组织） Percentages of tracheids, fibers, vessels, ray parenchyma, axial parenchyma	一定区域内，不同类型组织的面积占比 Tissue area as a percentage of measured transverse section area	尺寸建议不低于5 mm生长锥等样品；LM观测 At least the 5 mm diameter core is suggested; LM technique	≥20个视野 At least 20 fields of appropriate size	—
细胞长度（轴向管胞、纤维细胞、导管分子） Tracheid length, fibre length, vessel element length	完整细胞从一端到另一端的长度，包含细胞锐端 Measure the whole length of each cell from one tail end to the other	组织离析；LM观测 In a maceration; LM technique	≥25	IAWA Committee, 1989；2004
胞壁厚度（轴向管胞、纤维细胞、导管分子） Wall thickness of tracheid, fibre, vessel element	横切面上细胞双壁厚度 Total wall thickness measured as the double wall between 2 adjacent tracheids, fibers or vessels	横切面显微切片；区分早晚材；区分径向壁和弦向壁，建议测量径向壁；LM观测 Transverse sections; Measurement in earlywood and latewood respectively; Measured radially or tangentially, the radial measurement is suggested; LM technique	≥50	Scholz et al., 2013; IAWA Committee, 1989；2004
胞腔直径（纤维细胞、导管分子） Fiber lumen diameter, arithmic vessel diameter	细胞腔等效圆的算术直径 Fiber lumen diameter= arithmic diameter corresponding to equivalent circle diameter of fiber lumina; Arithmic vessel diameter= the simple average of the equivalent circle diameters	横切面显微切片；区分早晚材；LM观测Transverse sections; Measurement in earlywood and latewood respectively; LM technique	≥100	Scholz et al., 2013
导管分布密度 Vessels frequency	每平方毫米管孔数 Vessels per square millimetre	横切面显微切片；环孔材不计；LM观测Transverse sections; Not count vessel frequency for ring-porous woods; LM technique	≥5个视野 Count all the vessels in at least five fields of appropriate size (depending on vessel diameter and distribution)	IAWA Committee, 1989
导管腔弦向直径 Tangential diameter of vessel lumina	导管开口的最大距离，不包含壁厚 The tangential diameter of the vessel lumina, excluding the wall, is measured at the widest part of the opening	横切面显微切片；对环孔材和具有2类导管尺寸的散孔材，仅测量大导管；LM观测 Transverse sections; In ring-porous woods and woods with ‘vessels of two distinct diameter classes, wood not ring-porous’, only measure and record the larger size class; LM technique	≥25	IAWA Committee, 1989
复合导管比例 Vessel multiple fraction	复合导管数量占导管总数比例 Ratio of grouped vessels to total number of vessels	横切面显微切片；LM观测 Transverse sections; LM technique	≥50	Scholz et al., 2013
射线密度 Ray density	每毫米射线数 Rays per millmetre	尺寸建议不小于5 mm生长锥等样品；弦切面上沿一条垂直于射线高度的直线测定；LM观测 At least the 5 mm diameter core; The number of rays per linear unit is best determined from a tangential section along a line perpendicular to the ray’s axis; LM technique	10	IAWA Committee, 1989；von Arx et al., 2015
射线高度 Ray height	组成射线高度的细胞数量 Number of cells	弦切面测量；区分单列射线与纺锤型射线；纺锤型射线应包括两端延伸的单列部分；LM观测 Tangential sections; All features referring to ray height exclude rays containing intercellular canals (“fusiform rays”) unless otherwise specified; Rays that contain intercellular canals are called fusiform. Ensure that the entire ray is measured, including the uniseriate extensions at both ends of the ray; LM technique	≥25	IAWA Committee, 2004
管间纹孔直径 Intervessel pit size	导管间纹孔的直径 The diameter of pits between vessel elements	弦切面；观测导管重叠端壁上管间纹孔式；一般测定水平直径；LM观测 Surface views of intervessel pits are easiest to find in tangential sections; Intervessel pit size must be observed in overlapping end wall portions of vessel elements in a single vessel; The most widely used convention for determining pit size is to measure horizontal pit diameter; LM technique	10	IAWA Committee, 1989
管间纹孔膜厚度 Intervessel pit membrane thickness	导管间纹孔膜最厚处的厚度 Intervessel pit membrane thickness measured at its thickest point	TEM观测 TEM technique	≥25	Scholz et al., 2013
导管纹孔口面积 Pit aperture surface area	导管纹孔口所占据的面积 Pit aperture surface area	测量不同导管的纹孔；SEM测量 Pits from different vessels; SEM technique	≥50	Scholz et al., 2013
纹孔口直径 Diameter of outer pit aperture	纹孔最大开口处测量的纹孔口直径 Measure at the widest part of the opening	SEM测量 SEM technique	≥50	Scholz et al., 2013
纹孔室深度 Pit chamber depth	纹孔膜至纹孔口内侧的距离 Distance from the pit membrane to the inner pit aperture	TEM测量 TEM technique	≥25	Scholz et al., 2013

表1

表2

木材构造特征的信息采集方法①"

手段 Techniques	优点 Advantages	缺点 Limitations
iWood	宏观构造；最大图像采集分辨率为4 000像素×3 000像素；成像视场为6.35 mm×6.35 mm；光源配置有LED和UV；适用于木材表面图像的高清、快速采集 Macrostructure; The maximum image acquisition resolution is 4000 pixels × 3000 pixels; The field of view is 6.35 mm × 6.35 mm; Selection of LED and UV illumination; Suitable for high-resolution and fast-acquisition of imaging	局限于表面分析 Structural features of surfaces can be visualized
Xylophone	宏观构造；视野6485 μm（相机放大倍数2.9×），视野6331 μm（相机放大倍数3×）；光源配置有LED和UV；适用于木材表面图像的高清、快速采集 Macrostructure; Field of view is 6485 μm (camera zoom of 2.9×), field of view is 6331 μm (camera zoom of 3×); Selection of visible light or UV illumination; Suitable for high-resolution and fast-acquisition imaging.	局限于表面分析 Structural features of surfaces can be visualized
LM	微观构造；放大倍数×10~×1 000；分辨率极限200 nm Microstructure; The magnification of the microscope can range from ×10 to ×1 000; The best resolution for a light microscope is 200 nm	需要样品制备，耗时耗力；侧重2D图像采集，3D检测需要连续切片，结合逐层扫描显微图像合成系统 Extensive sample preparation; 2D image of wood, 3D analysis of wood structure is quite laborious and time-consuming as it requires serial sectioning
RLM	微观构造；适用于表面抛光不透明样品 Microstructure; RLM does not require sectioning and can be used on opaque wood samples	无法用于表面涂膜处理的样品；较难适用于腐朽、考古样品 RLM cannot be applied on artefacts whose surfaces are generally treated and coated because this hinders the identification of anatomical structures; RLM cannot be used on deteriorated old sample
3D-RLM	微观构造，无损检测；对样品表面粗糙度无要求，适用于木材、木制品及木炭样品；放大倍数×10~×5000 Macrostructure; The advantages of using the 3D-RLM technology are that fresh fracture planes of wood and charcoal can be directly observed under the microscope without further preparation or surface treatment; The 3D-technique with integrated polarized light illumination creates high-contrast images of uneven and black charcoal surfaces; Magnification up to 5000×	局限于表面分析 Structural features of surfaces can be visualized
CLSM	微观构造；横向极限分辨率0.2 μm，轴向极限分辨率0.6 μm Microstructure; The best resolution that can be obtained is ~0.2 μm laterally and ~0.6 μm axially, though in practice these values are not always achieved	侧重2D图像采集，3D图像需通过2D图像叠加模块实现 A “z-stack” of the images can be compiled to form a composite 3D image of a thick sample
全自动数字玻片扫描系统	微观构造；放大倍数20×，可高达43×或86×，分辨率0.087 μm/像素；一次可自动扫描高达12张切片；提供荧光扫描通道 Microstructure; Magnification of 20×, up to 43× or 86×; Resolution of 0.087 μm/pixel; Up to 12 slices can be automatically scanned at once; Selection of fluorescence scanning channel	设备昂贵 Expensive equipment
μCT	微观构造，三维检测；无损检测 Non-destructive, non-invasive technique that can be utilized to construct virtual 3D images of wood microstructure	体素与样品检测空间有关，如体素1 μm时，样品需为亚毫米尺寸 The resolution is inversely proportional to sample width and can be up to 1 μm; The resolution of 1 μm is only possible with submillimeter sample size
LATscan	微观构造，二维/三维检测；自动化检测；1 min内能采集上百张图像；利用木质素等自发荧光现象，可实现细胞构造与组分化学的分析整合 Microstructure; Generate high-quality 2D images and 3D reconstructions of wood plant samples; Automatic imaging during ablation; LATscan is a high-throughput imaging system that yields hundreds of images per minute; LATscan is a powerful technology to generate morphological and anatomical features while revealing differential fluorescent signals corresponding to cellulosic, lignified, and suberized cell walls	破坏性检测；激光烧灼可能产生热伪影；成本贵 Destructive technique; Material removal via laser ablation probably lends itself to thermal effects. The thermal effect would only get completely removed in the low-picosecond range and femtosecond pulsed range with a similar pulse energy. There could also be additional thermal effects due to thick samples as the outer edges of the sample would fall on the edges or outside the Gaussian beam profile along the y-direction; It may not be cost-effective compared with other traditional techniques (e.g. light microscopy)
SEM	超微构造；分辨率极限20 nm；聚焦深度高，比LM高300倍 Ultrastructure; SEM resolution up to 20 nm; High depth of focus, field depth is 300 times higher than that of LM	需要样品制备，耗时耗力；局限于表面分析 For SEM studies, an appropriate size of cube of wood is cut, ensuring that surfaces have a clean cut to make sure no fractures or cutting lines are seen in the final image. The cubes are finally mounted on sample stubs and then coated with gold in a high-vacuum evaporating unit; Examines only exposed surfaces of a sample
AFM	超微构造；x-y轴的分辨率极限10 nm，z轴的分辨率<0.1 nm；探针尖端可多款选配，实现木材细胞壁力学、化学或者热性能的原位分析 Ultrastructure; Ever decreasing AFM probe tip size (<10 nm) produce topographical images with high x-y resolution (of tens of nanometers) and ultrahigh z-resolution (<0.1 nm); The real beauty with AFM lies in the possibility to modify the probe tip to simultaneously image wood cell wall ultrastructure and measure cell wall properties (mechanical, chemical or thermal), enabling direct elucidation of underlying structure-property relations	需要样品制备，耗时耗力；局限于表面分析 Preparation of smooth samples may require ultra-microtoming with a diamond knife, although thicker samples can be used; Examines only exposed surfaces of a sample
TEM	超微构造；放大倍数×35~×700 000，分辨率极限0.5 ? Ultrastructure; Modern TEMs have a resolution of over 0.5 ? and a magnification ratio ranging from ×35 to ×700 000	需要样品制备，制样难度高，耗时耗力；2D图像采集 Extensive sample preparation; 2D image of wood

表2

表3

适用于图像信息的木材定量解剖分析软件①"

手段 Techniques	木材构造特征 Wood anatomical traits			对象筛选策略 Cell filtering	其他软件特征 Some other features	软件许可 License
手段 Techniques	细胞尺寸 Cell size	细胞壁厚度 Cell wall thickness	生长轮 Tree ring	对象筛选策略 Cell filtering	其他软件特征 Some other features	软件许可 License
AutoCellRow	可分析 Yes	自动分析 Automatic	自动识别 Automatic	尺寸、形状 Size, shape	多种图像格式的兼容输入能力；数据自动输出；基于Python语言开发的软件，具有较高的可扩展性 tiff., jpg., bmp., png. formats; Developed in Python programming language	免费 Free
AxioVision	可分析 Yes	手动分析 Manual	无法识别 No	尺寸、形状 Size, shape	自动化工作流模块；宏开发工具；适用于显微图像 Automating workflow and macro development tools; Microscope control	商业 Commercial
CellProfilter	可分析 Yes	手动分析 Manual	无法识别 No	尺寸、颜色、形状 Size, color, shape	批量分析超1 000张图像集合；自动化工作流模块 Batch analyzing large sets (>1000) of images; Automating workflow using modules	免费 Free
ImageJ	可分析 Yes	手动分析 Manual	无法识别 No	尺寸、形状 Size, shape	拥有大量插件；宏开发工具 Large collection of plugins by community; Macro development tools	免费 Free
Image-Pro Plus	可分析 Yes	手动分析 Manual	无法识别 No	尺寸、颜色、形状 Size, color, shape	较强的图像处理与分析能力；宏开发工具 Powerful image processing and analysis functions; Macro development tools	商业 Commercial
Image-warp	可分析 Yes	自动分析 Automatic	无法识别 No	尺寸、形状 Size, shape	自动的图像处理与分析能力 Automatic image processing morphometric method	商业 Commercial
NIE-Elements	可分析 Yes	手动分析 Manual	无法识别 No	尺寸、形状 Size, shape	宏开发工具；适用于显微图像 Macro development tools; Microscope control	商业 Commercial
ROXAS	可分析 Yes	自动分析 automatic	自动识别 automatic	尺寸、颜色、形状、背景 Size, color, shape, context	能分析细胞数高达1 000 000的大图像；数据自动输出；图像自动处理；可批量处理；在线库和定制配置 Analyzing large images (up to 1000,000 cells); Automatic image processing; Batch processing options; Online library and customization of tailored configurations	免费，但需要商业软件Image-Pro Plus (v6.1 或更高版本) Free, but requires commercial Image-Pro Plus (v6.1 or higher)
WinCELL	可分析 Yes	自动分析 Automatic	手动识别 Manual	尺寸、颜色、形状 Size, color, shape	数据自动输出；可批量处理；定制配置；区分显微图像和扫描图像录入选择 Large set of anatomical output parameters; Batch processing options; Customizing tailored configurations; Microscope and scanner control	商业 Commercial

表3

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