林木木材形成机制及材性改良研究进展

doi:10. 3969 /j. issn. 2096-4900. 2019. 02.007

温带林业研究

2019, Vol. 2

Issue (2): 40-47 DOI: 10. 3969 /j. issn. 2096-4900. 2019. 02.007

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林木木材形成机制及材性改良研究进展

李少锋

中国林业科学研究院华北林业实验中心，北京 102300

Wood Formation Mechanism and Properties Improvement in Forest Trees

LI Shao-feng

1.Harbin Research Institute of Forestry Machinery，State Forestry Administration，Harbin 150086，China；2.Research Center of Cold Temperate Forestry，CAF，Harbin 150086，China

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摘要我国森林覆盖率低，木材供求矛盾突出，迫切需要营造速生优质人工林，加速木材品质性状遗传改良进程。木材形成机理的研究可以为培育速生优质林木提供理论指导和原创技术，林木的次生生长包括维管组织形成、次生细胞壁形成及木质化、细胞程序化死亡、以及心材形成等过程。由于木本植物生长周期长、转基因材料较难获取以及缺乏有效的基因突变体库，对林木木材形成的机制研究相对滞后。随着近20 年研究技术的发展和多学科交叉研究的应用，材性相关基因的功能及其调控机制等得到了很大程度的揭示，例如发现了木聚糖乙酰化修饰存在乙酰化和去乙酰化过程，揭示了赤霉素调控纤维素合成的信号转导通路，解析了ILA1-IIP4-NAC 调控次生壁形成的信号转导通路，建立了木质素合成的新型系统生物学模型，从而有助于在木质素生物合成中提升木材应用性能。本文介绍了木材形成机理及转录调控网络、材性遗传改良的研究进展，并论述了次生生长与抗逆性状的平衡机制研究情况。这些研究进展不仅使人们对次生细胞壁的合成机理有了更深入的认识，也为木材形成的机制应用于材性遗传改良奠定了重要的理论基础，同时还对未来基因功能研究可能运用到的技术和方法进行了展望。

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关键词 ：木材形成, 速生优质林木, 调控机制, 材性改良

收稿日期: 2019-04-01

S781.7

基金资助:国家自然科学基金（31770705）和转基因生物新品种培育科技重大专项（2018ZX08020002）资助项目。

通讯作者: 李少锋 E-mail: lishaofeng2009@163.com

作者简介: 李少锋（1981—），男，博士，副研究员，主要研究方向：林木分子生物学及分子育种。

引用本文:

李少锋. 林木木材形成机制及材性改良研究进展[J]. 温带林业研究, 2019, 2(2): 40-47.
WANG Xiao-hong，SHEN Fang-yuan，ZHOU Xin，YE Tian-yi，Cong Ri-zheng，YANG Lan，HUANG Yan. Wood Formation Mechanism and Properties Improvement in Forest Trees. 温带林业研究, 2019, 2(2): 40-47.

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[1] 2020 年中国国内木材缺口将达到2 亿立方[R].[2018-06-01].http ：//www.forestry.gov.cn/xdly/5198/20180601/085824756459828.html. [2] Schrader J，Nilsson J，Mellerowicz E，et al. A highresolution transcript profile across the wood-forming meristem of poplar identifies potential regulators of cambial stem cell identity[J]. The Plant Cell，2004，16（9）：2278-2292. [3] Li L，Zhou Y，Cheng X，et al. Combinatorial modification of multiple lignin traits in trees through multigene cotransformation[J]. Proceedings of the National Academy of Sciences，2003，100（8）：4939-4944. [4] Wei J H，Wang Y Z，Wang H Z，et al. Pulping performance of transgenic poplar with depressed Caffeoyl-CoA O-methyltransferase[J]. Chinese Science Bulletin，2008，53（22）：3535. [5] Wang J P，Matthews M L，Williams C M，et al.Improving wood properties for wood utilization through multi-omics integration in lignin biosynthesis[J]. Nature Communications，2018，9（1）：1579. [6] Peng L，Kawagoe Y，Hogan P，et al. Sitosterol-bglucoside as primer for cellulose synthesis in plants[J].Science，2002，2959（5552）：147-150. [7] Gardiner J C，Taylor N G，Turner S R，et al. Control of cellulose synthase complex localization in developing xylem[J]. Plant Cell，2003，15（8）：1740-1748. [8] Lu S，Li L G，Yi X P，et al. Differential expression of three eucalyptus secondary cell wall-related cellulose synthase genes in response to tension stress[J]. Journal of Experimental Botany，2008，59（3）：681-695. [9] Ranik M，Myburg A A. Six new cellulose synthase genes from Eucalyptus are associated with primary and secondary cell wall biosynthesis[J]. Tree Physiology，2006，26（5）：545-556. [10] Bhandari S，Fujino T，Thammanagowda S，et al. Xylemspecific and tension stress responsive coexpression of KORRIGAN endoglucanase and three secondary wall associated cellulose synthase genes in aspen trees[J].Planta，2006，224（4）：828-837. [11] Samuga A，Joshi C P. Cloning and characterization of cellulose synthase-like gene，PtrCSLD2 from developing xylem of aspen trees[J]. Physiologia Plantarum，2004，120（4）：631-641. [12] Liu L，Shang-Guan K，Zhang B，et al. Brittle Culm1，a COBRA-like protein，functions in cellulose assembly through binding cellulose microfibrils[J]. PLoS Genet，2013，9（8）：e1003704. [13] Zhang D，Xu Z，Cao S，et al. An uncanonical CCCHtandem zinc-finger protein represses secondary wall synthesis and controls mechanical strength in rice[J]. Molecular plant，2018，11（1）：163-174. [14] Kulkarni N，Shendye A，Rao M. Molecular and biotechnological aspects of xylanases[J]. FEMS microbiology reviews，1999，23（4）：411-456. [15] Scheller H V，Ulvskov P . Hemicelluloses [J]. Annual review of plant biology，2010，61（1）：263-289. [16] 秦丽霞，张德静，李龙，等. 参与植物细胞壁半纤维素木聚糖合成的糖基转移酶[J]. 植物生理学报，2011，47（9）：831-839. [17] Liepman A H，Wilkerson C G，Keegstra K. Expression of cellulose synthase-like（Csl）genes in insect cells reveals that CslA family members encode mannan synthases[J].Proceedings of the National Academy of Sciences，2005，102（6）：2221-2226. [18] Hamann T，Osborne E，Youngs H L，et al. Global e x p r e s s i o n analysi s of CESA and CSL genes in Arabidopsis[J]. Cellulose，2004，11（3-4）：279-286. [19]Gao Y，He C，Zhang D，et al. Two trichome birefringencelike proteins mediate xylan acetylation，which is essential for leaf blight resistance in rice[J]. Plant physiology，2017，173（1）：470-481. [20] Zhang B，Zhang L，Li F，et al. Control of secondary cellwall patterning involves xylan deacetylation by a GDSL esterase[J]. Nature plants，2017，3（3）：17017. [21] Yu Q，McKeand S E，Nelson C D，et al. Differences in wood density and growth of fertilized and nonfertilized loblolly pine associated with a mutant gene，cad-n1[J].Canadian Journal of Forest Research，2005，35（7）：1723-1730. [22] Groover A T，Mansfield S D，DiFazio S P，et al.The Populus homeobox gene ARBORKNOX1 reveals overlapping mechanisms regulating the shoot apical meristem and the vascular cambium[J]. Plant molecular biology，2006，61（6）：917-932. [23] Canam T，Mak S W Y，Mansfield S D. Spatial and temporal expression profiling of cell-wall invertase genes during early development in hybrid poplar[J]. Tree physiology，2008，28（7）：1059-1067. [24] Coleman H D，Cánovas F M，Man H，et al. Enhanced expression of glutamine synthetase（GS1a）confers altered fibre and wood chemistry in field grown hybrid poplar（Populus tremula×alba）（717-1B4）[J]. Plantbiotechnology journal，2012，10（7）：883-889. [25] Derba-Maceluch M，Awano T，Takahashi J，et al.Suppression of xylan endotransglycosylase PtxtXyn10A affects cellulose microfibril angle in secondary wall in aspen wood[J]. New Phytologist，2015，205（2）：666-681. [26] Szechyńska-Hebda M，Czarnocka W，Hebda M，et al.PAD4，LSD1 and EDS1 regulate drought tolerance，plant biomass production，and cell wall properties[J]. Plant cell reports，2016，35（3）：527-539 [27] 王明庥，黄敏仁. 黑杨派新无性系木材性状遗传改良[J].南京林业大学学报，1989，13（3）：9-16. [28] Hertzberg M，Aspeborg H，Schrader J，et al. A transcriptional roadmap to wood formation[J]. Proceedings of the National Academy of Sciences，2001，98（25）：14732-14737. [29] Thumma B R，Nolan M F，Evans R，et al. Polymorphisms in Cinnamoyl CoA Reductase（CCR）are associated with variation in microfibril angle in Eucalyptus spp[J].Genetics，2005，171（3）：1257-1265. [30] Spokevicius A V，Southerton S G ，MacMillan C P，et al.β-tubulin affects cellulose microfibril orientation in plant secondary fibre cell walls[J]. The Plant Journal，2007，51（4）：717-726. [31] Zhong R Q，Burk D H，Morrison III W H，et al. A kinesinlike protein is essential for oriented deposition of cellulose microfibrils and cell wall strength[J]. Plant cell，2002，14（12）：3101-3117. [32] Wang X D，Wu X J. Phenotypic characterization，genetic analysis and gene-mapping for a brittle mutant in rice[J].Journal of integrative plant biology，2008，50（3）：319-328. [33] Li S F，Zhang Y X，Ding C J，et al. Proline-rich protein gene PdPRP regulates secondary cell wall formation in poplar[J]. Jounal of plant physiology，2019，233 ：58-72. [34] Li S，Huang Q，Zhang B，et al. Small GTP-binding protein PdRanBP regulates vascular tissue development in poplar[J]. BMC genetics，2016，17（1）：96. [35] Li S，Su X，Zhang B，et al. Molecular cloning and functional analysis of the Populus deltoides remorin gene PdREM[J]. Tree physiology，2013，33（10）：1111-1121. [36] 李少锋，苏晓华，张冰玉，等. 美洲黑杨（Populus deltoides）材性相关候选基因PdCYTOB 的功能鉴定[J].植物学报，2011，46（6）：642-651. [37] Zhang Q，Xie Z，Zhang R，et al. Blue light regulates secondary cell wall thickening via MYC2 /MYC4 activation of the NST1-directed transcriptional network in Arabidopsis[J]. The Plant cell，2018，30（10）：2512-2528. [38] Savidge R A，Barnett J R，Napier R. Cell and molecular biology of wood formation[M]. BIOS，2000. [39] Silvertown J，Charlesworth D. Introduction to plant population biology[M]. John Wiley & Sons，2009. [40] Buchanan B B，Gruissem W，Jones R L. Biochemistry and molecular biology of plants[M]. John Wiley & Sons，2015. [41] Huang D，Wang S，Zhang B，et al. A gibberellinmediated DELLA-NAC signaling cascade regulates cellulose synthesis in rice[J]. The Plant Cell，2015，27（6）：1681-1696. [42] Xu C，Shen Y，He F，et al. Auxin-mediated Aux/IAAARF-HB signaling cascade regulates secondary xylem development in Populus[J]. New Phytologist，2019，222（2）：752-767. [43] Shu W，Zhou H，Jiang C，et al. The auxin receptor TIR1 homolog（Pag FBL 1）regulates adventitious rooting through interactions with Aux/IAA 28 in Populus[J]. Plant biotechnology journal，2019，17（2）：338-349. [44] Zhang J，Xie M，Tuskan GA，et al. Recent advances in the transcriptional regulation of secondary cell wall biosynthesis in the woody plants[J]. Frontiers in Plant Science，2018，9：1535. [45] Zhong R，Mccarthy R L，Lee C，et al. Dissection of the transcriptional program regulating secondary wall biosynthesis during wood formation in Poplar[J]. Plant physiology，2011，157（3）：1452-1468. [46] Chai G，Qi G，Cao Y，et al. Poplar PdC3H17 and PdC3H18 are direct targets of PdMYB3 and PdMYB21， and positively regulate secondary wall formation in Arabidopsis and poplar[J]. New Phytologist，2014，203 （2）：520-534. [47] Zhao Y，Sun J，Xu P，et al. Intron-mediated alternative splicing of wood-associated NAC transcription factor 1B regulates cell wall thickening during fiber development in Populus species[J]. Plant physiology，2014，164（2）：765-776. [48] Li Q，Lin Y C，Sun Y H，et al. Splice variant of the SND1 transcription factor is a dominant negative of SND1 members and their regulation in Populus trichocarpa[J]. Proceedings of the National Academy of Sciences，2012，109（36）：14699-14704. [49] Wang H，Avci U，Nakashima J，et al. Mutation of WRKY transcription factors initiates pith secondary wall formation and increases stem biomass in dicotyledonous plants[J].Proceedings of the National Academy of Sciences，2010， 107（51）：22338-22343. [50] Yu Y，Hu R，Wang H，et al. MlWRKY12，a novel Miscanthus transcription factor，participates in pith secondary cell wall formation and promotes flowering[J].Plant science，2013，212 ：1-9. [51] Yang L，Zhao X，Yang F，et al. PtrWRKY19，a novel WRKY transcription factor，contributes to the regulation of pith secondary wall formation in Populus trichocarpa[J].Scientific reports，2016，6 ：18643. [5 2] Chen H，Wang J P，Liu H，et al. Hierarchical transcription-factor and chromatin binding network for wood formation in Populus trichocarpa[J]. The Plant Cell，2019 ：tpc. 00620.2018. [53] Rogers L，Dubos C，Surman C，et al. Comparison of lignin deposition in three ectopic lignification mutants[J].New Phytol，2005，168 ：123-140. [54] Bedon F，Grima-Pettenati J，Mackay J. Conifer R2R3- MYB transcription factors ：sequence analyses and gene expression in wood-forming tissues of white spruce（Picea glauca）[J]. BMC Plant Biology，2007，7（1）：17. [55] Wilkins O，Nahal H，Foong J，et al. Expansion and diversification of the Populus R2R3-MYB family of transcription factors[J]. Plant Physiology，2009，149（2）：981-993. [56] Tang X，Zhuang Y，Qi G，et al. Poplar PdMYB221 is involved in the direct and indirect regulation of secondary wall biosynthesis during wood formation[J]. Scientific reports，2015，5 ：12240. [57] Yang L，Zhao X，Ran L，et al. PtoMYB156 is involved in negative regulation of phenylpropanoid metabolism and secondary cell wall biosynthesis during wood formation in poplar[J]. Scientific reports，2017，7 ：41209. [58] Goicoechea M，Lacombe E，Legay S，et al. EgMYB2，a new transcriptional activator from Eucalyptus xylem，regulates secondary cell wall formation and lignin biosynthesis[J]. The Plant Journal，2005，43（4）：553-567. [59] Gómez-Maldonado J，Avila C，de la Torre F，et al.Functional interactions between a glutamine synthetase promoter and MYB proteins[J]. The Plant Journal，2004，39（4）：513-526. [60] Mccarthy R L，Zhong R ，Fowler S ， et al. The poplar MYB transcription factors，PtrMYB3 and PtrMYB20，are involved in the regulation of secondary wall biosynthesis[J].Plant & Cell Physiology，2010，51（6）：1084-1090. [61] Lu S，Li Q，Wei H，et al. Ptr-miR397a is a negative regulator of laccase genes affecting lignin content in Populus trichocarpa[J]. Proceedings of the National Academy of Sciences，2013，110（26）：10848-10853. [62] Gerber L，Zhang B，Roach M，et al. Deficient sucrose synthase activity in developing wood does not specifically affect cellulose biosynthesis，but causes an overall decrease in cell wall polymers[J]. New Phytologist，2014，203（4）：1220-1230. [63] Guo H，Wang Y，Wang L，et al. Expression of the MYB transcription factor gene BplMYB46 affects abiotic stress tolerance and secondary cell wall deposition in Betula platyphylla[J]. Plant biotechnology journal，2017，15（1）：107-121. [64] Hu P，Zhang K，Yang C. BpNAC012 positively regulates abiotic stress responses and secondary wall biosynthesis[J].Plant physiology，2019，179（2）：700-717. [65] Li S，Lin Y C J，Wang P，et al. Histone acetylation cooperating with AREB1 transcription factor regulates drought response and tolerance in Populus trichocarpa[J].The Plant Cell，2018 ：tpc. 00437.2018. [66] Xu C，Fu X，Liu R，et al. PtoMYB170 positively regulates lignin deposition during wood formation in poplar and confers drought tolerance in transgenic Arabidopsis[J]. Treephysiology，2017，37（12）：1713-1726. [67] Park M Y，Kang J，Kim S Y. Overexpression of AtMYB52 confers ABA hypersensitivity and drought tolerance[J].Molecules and cells，2011，31（5）：447-454. [68] Jeong C，Lee W，Truong H，et al. Dual role of SND1 facilitates effcient communication between abiotic stress signalling and normal growth in Arabidopsis [J]. Scientific Reports，2018，8（1）：10114. [69] Schreiber T，Prange A，Hoppe T，et al. Split-TALE ：A TALE-based two-component system for synthetic biology applications in planta[J]. Plant physiology，2019，179（3）：1001-1012. [70] Xie M，Muchero W，Bryan A C，et al. A 5-enolpyruvylshikimate 3-phosphate synthase functions as a transcriptional repressor in Populus[J]. The Plant Cell，2018，30（7）：1645-1660. [71] Lu S，Li Q，Wei H，et al. Ptr-miR397a is a negative regulator of laccase genes affecting lignin content in Populus trichocarpa[J]. Proceedings of the National Academy of Sciences，2013，110（26）：10848-10853. [72] Zhang B，Zhou Y. Plant cell wall formation and regulation[J]. Scientia Sinica，2015，45（6）：544-556. [73] Wang L，Xue Y，Xing J，et al. E x p l o r i n g the spatiotemporal organization of membrane proteins in livingplant cells[J]. Annual review of plant biology，2018，69（1）：525-551.