树脂铸型法研究毛竹材细胞壁的纹孔特征
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摘要
【目的】研究毛竹材细胞壁纹孔的形态、结构和分布排列等特征,揭示维管束组织细胞间的纹孔通道,为竹子生长发育机制的深入研究和竹材高效利用提供理论依据。【方法】毛竹竹条经树脂渗透、树脂固化、组织消化和清洗干燥等处理后,获得单个细胞铸型和空间范围内多个细胞连接的立体铸型网络。结合毛竹切片扫描电镜图像,研究导管、薄壁细胞和纤维细胞的纹孔特征以及细胞间纹孔对的连接和分布特点。【结果】毛竹后生木质导管纹孔具明显的纹孔缘结构,并呈较高的选择性分布,即靠近维管束几何中心位置一侧的导管壁上往往密集分布大量纹孔,而另一侧纹孔则十分稀少或基本没有。导管铸型上,纹孔口和纹孔膜分别为0. 9～2. 7μm和1. 1～3. 8μm;维管束内存在小导管且相互间以具缘纹孔对连接;维管束内薄壁细胞纹孔比基本组织中的数量多、孔径大,紧邻木质部导管的薄壁细胞,其纹孔分布尤为密集;纤维细胞上纹孔稀疏分布,纹孔小、纹孔道狭长,纤维状管胞上分布有具缘纹孔;维管束内细胞间由纹孔对进行横向和纵向连接,并进一步形成流体移动的空间网络。【结论】树脂铸型法可直接展现纹孔立体结构,客观反映特定流体在竹材内部的扩散或渗透路径,适用于毛竹材细胞壁纹孔特征的研究。
【Objective】To observe the pit morphology and distribution in moso bamboo,resin casting,a very proper way to obtain three-dimensional duplicate of pits,was applied.【Method】 A two-component resin BS was used to complete the following procedures,infiltration of resin,resin curing,corrosion of bamboo tissue and dry cleaning in order. Resin casts of different types of individual cell and cells within certain scope served to reveal characteristics of pits in cell wall,pit pairs between cells and channels in vascular bundle. Moreover,a combination with observing bamboo sections was able to make verification of result getting from two ways.【Result】Pits in metaxylem vessels of moso bamboo presented a high selectivity of distribution,that is,large scale of pits were centered in single side of cell wall near to the geometric center of vascular bundle,though pits in the other side turned to be the opposite. And clear pit border structure was found. Diameter of pit aperture and pit membrane of vessel resin casts were 0. 9-2. 7 μm and 1. 1-3. 8 μm respectively. There were some smaller metaxylem vessel elements scattered or arranged adjacent connecting with bordered pit. As for parenchyma cells within the vascular bundle,pits were more and larger than those in the ground tissue. And parenchyma cells adjacent to metaxylem showed particularity. Pits in fiber cells were in sparse distribution with small openings,long and narrow cavities and suspect pit borders. Cells in vascular bundle connected each other through the pit and formatted space networks further.【Conclusion】Resin casting,which is capable of showing the pit stereo structure and reflecting diffusion permeation of specific fluid inside bamboo both directly and objectively,is a suitable method for investigating pits in moso bamboo.
【Objective】To observe the pit morphology and distribution in moso bamboo,resin casting,a very proper way to obtain three-dimensional duplicate of pits,was applied.【Method】 A two-component resin BS was used to complete the following procedures,infiltration of resin,resin curing,corrosion of bamboo tissue and dry cleaning in order. Resin casts of different types of individual cell and cells within certain scope served to reveal characteristics of pits in cell wall,pit pairs between cells and channels in vascular bundle. Moreover,a combination with observing bamboo sections was able to make verification of result getting from two ways.【Result】Pits in metaxylem vessels of moso bamboo presented a high selectivity of distribution,that is,large scale of pits were centered in single side of cell wall near to the geometric center of vascular bundle,though pits in the other side turned to be the opposite. And clear pit border structure was found. Diameter of pit aperture and pit membrane of vessel resin casts were 0. 9-2. 7 μm and 1. 1-3. 8 μm respectively. There were some smaller metaxylem vessel elements scattered or arranged adjacent connecting with bordered pit. As for parenchyma cells within the vascular bundle,pits were more and larger than those in the ground tissue. And parenchyma cells adjacent to metaxylem showed particularity. Pits in fiber cells were in sparse distribution with small openings,long and narrow cavities and suspect pit borders. Cells in vascular bundle connected each other through the pit and formatted space networks further.【Conclusion】Resin casting,which is capable of showing the pit stereo structure and reflecting diffusion permeation of specific fluid inside bamboo both directly and objectively,is a suitable method for investigating pits in moso bamboo.
引文
鲍甫成,赵有科,吕建雄.2003.杉木和马尾松木材渗透性与微细结构的关系研究.北京林业大学学报,25(1):1-5.(Bao F C,Zhao Y K,LüJ X.2003.Relationship between permeability and fine structure of common Chinese fir and massion pine wood.Journal of Beijing Forestry University,25(1):1-5.[in Chinese])
刘波.2008.毛竹发育过程中细胞壁形成的研究.北京:中国林业科学研究院博士学位论文.(Liu B.2008.Formation of cell wall in developmental culms of Phyllostachys pubescens.Beijing:PhD thesis of Chinese Academy of Forestry.[in Chinese])
鈴木寧.1952.竹材の研究(第7報)竹材柔細胞腔の空気及び液体に対する閉鎖について.東京大学農学部演習林報告,43:151-157.(Suzuki Y.1952.Studies on bamboo(VII)-Stopping of pit pore in bamboo parenchyma cell and its effect on gas and liquid permeability into bamboo.Tokyo University Agriculture Department Exercise Forest Report,43:151-157.[in Japanese])
刘嵘.2017.毛竹材细胞壁的纹孔特征研究.北京:中国林业科学研究院博士学位论文.(Liu R.2017.Characteristics of pits in bamboo cell wall.Beijing:PhDthesis of Chinese Academy of Forestry.[in Chinese])
刘嵘,杨淑敏,李晖,等.2017.毛竹材导管分子的纹孔特征.南京林业大学学报:自然科学版,41(6):163-168.Liu R,Yang S M,Li H,et al.2017.Characteristics of pits in the vessel element of moso bamboo.Journal of Nanjing Forestry University:Natural Sciences Edition,41(6):163-168.[in Chinese])
刘主凰.2011.福建主要竹材纤维特性的研究.福州:福建农林大学硕士学位论文.(Liu Z H.2011.Study on fiber properties of chief bamboo species in Fujian province.Fuzhou:MS thesis of Fujian Agriculture and Forestry University.[in Chinese])
江泽慧.2007.世界竹藤.北京:中国林业出版社.(Jiang Z H.2007.Bamboo and rattan in the world.Beijing:China Forestry Publishing House.[in Chinese])
Carlquist S,Schneider E L.2010.Origins and nature of vessels in monocotyledons.12.Pit membrane microstructure diversity in tracheary elements of Astelia.Pacific Science,64(4):607-618.
Carlquist S.2012.Monocot xylem revisited:new information,new paradigms.The Botanical Review,78(2):87-153.
Cho D,Jin M K,Kim D.2013.Phenolic resin infiltration and carbonization of cellulose-based bamboo fibers.Materials Letters,104(3):24-27.
Choat B,Cobb A R,Jansen S.2008.Structure and function of bordered pits:new discoveries and impacts on whole-plant hydraulic function.New Phytologist,177(3):608-626.
Cosgrove D J.2005.Growth of the plant cell wall.Nature Reviews Molecular Cell Biology,6(11):850-861.
Dusotoit-Coucaud A,Brunel N,Tixier A,et al.2014.Hydrolase treatments help unravel the function of intervessel pits in xylem hydraulics.Physiologia Plantarum,150(3):388-396.
Fei B H,Gao Z M,Wang J,et al.2016,Biological,anatomical,and chemical characteristics of bamboo∥Yoon S K,Ryo F,Adya P S.Secondary xylem biology.Elsevier,Netherlands,283-306.
Fujii T.1993.Application of resin casting method to wood anatomy.Plant Morphology,5(1):3-18.
Liese W,Tang T K H.2015.Properties of the bamboo culm,bamboo.Springer,227-256.
Meyra A G,Kuz V A,Zarragoicoechea G J.2007.Geometrical and physicochemical considerations of the pit membrane in relation to air seeding:the pit membrane as a capillary valve.Tree Physiology,27(10):1401-1405.
Nirmala C,Bisht M S,Bajwa H K,et al.2018.Bamboo:a rich source of natural antioxidants and its applications in the food and pharmaceutical industry.Trends in Food Science&Technology,77:91-99.
Stieber J.1981.A new method of examining vessels.Annals of Botany,48(3):411-414.
刘波.2008.毛竹发育过程中细胞壁形成的研究.北京:中国林业科学研究院博士学位论文.(Liu B.2008.Formation of cell wall in developmental culms of Phyllostachys pubescens.Beijing:PhD thesis of Chinese Academy of Forestry.[in Chinese])
鈴木寧.1952.竹材の研究(第7報)竹材柔細胞腔の空気及び液体に対する閉鎖について.東京大学農学部演習林報告,43:151-157.(Suzuki Y.1952.Studies on bamboo(VII)-Stopping of pit pore in bamboo parenchyma cell and its effect on gas and liquid permeability into bamboo.Tokyo University Agriculture Department Exercise Forest Report,43:151-157.[in Japanese])
刘嵘.2017.毛竹材细胞壁的纹孔特征研究.北京:中国林业科学研究院博士学位论文.(Liu R.2017.Characteristics of pits in bamboo cell wall.Beijing:PhDthesis of Chinese Academy of Forestry.[in Chinese])
刘嵘,杨淑敏,李晖,等.2017.毛竹材导管分子的纹孔特征.南京林业大学学报:自然科学版,41(6):163-168.Liu R,Yang S M,Li H,et al.2017.Characteristics of pits in the vessel element of moso bamboo.Journal of Nanjing Forestry University:Natural Sciences Edition,41(6):163-168.[in Chinese])
刘主凰.2011.福建主要竹材纤维特性的研究.福州:福建农林大学硕士学位论文.(Liu Z H.2011.Study on fiber properties of chief bamboo species in Fujian province.Fuzhou:MS thesis of Fujian Agriculture and Forestry University.[in Chinese])
江泽慧.2007.世界竹藤.北京:中国林业出版社.(Jiang Z H.2007.Bamboo and rattan in the world.Beijing:China Forestry Publishing House.[in Chinese])
Carlquist S,Schneider E L.2010.Origins and nature of vessels in monocotyledons.12.Pit membrane microstructure diversity in tracheary elements of Astelia.Pacific Science,64(4):607-618.
Carlquist S.2012.Monocot xylem revisited:new information,new paradigms.The Botanical Review,78(2):87-153.
Cho D,Jin M K,Kim D.2013.Phenolic resin infiltration and carbonization of cellulose-based bamboo fibers.Materials Letters,104(3):24-27.
Choat B,Cobb A R,Jansen S.2008.Structure and function of bordered pits:new discoveries and impacts on whole-plant hydraulic function.New Phytologist,177(3):608-626.
Cosgrove D J.2005.Growth of the plant cell wall.Nature Reviews Molecular Cell Biology,6(11):850-861.
Dusotoit-Coucaud A,Brunel N,Tixier A,et al.2014.Hydrolase treatments help unravel the function of intervessel pits in xylem hydraulics.Physiologia Plantarum,150(3):388-396.
Fei B H,Gao Z M,Wang J,et al.2016,Biological,anatomical,and chemical characteristics of bamboo∥Yoon S K,Ryo F,Adya P S.Secondary xylem biology.Elsevier,Netherlands,283-306.
Fujii T.1993.Application of resin casting method to wood anatomy.Plant Morphology,5(1):3-18.
Liese W,Tang T K H.2015.Properties of the bamboo culm,bamboo.Springer,227-256.
Meyra A G,Kuz V A,Zarragoicoechea G J.2007.Geometrical and physicochemical considerations of the pit membrane in relation to air seeding:the pit membrane as a capillary valve.Tree Physiology,27(10):1401-1405.
Nirmala C,Bisht M S,Bajwa H K,et al.2018.Bamboo:a rich source of natural antioxidants and its applications in the food and pharmaceutical industry.Trends in Food Science&Technology,77:91-99.
Stieber J.1981.A new method of examining vessels.Annals of Botany,48(3):411-414.