CN116606872A - Ssu TDX gene closely related to hemicellulose synthesis regulation and control and application thereof - Google Patents
Ssu TDX gene closely related to hemicellulose synthesis regulation and control and application thereof Download PDFInfo
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- 101150074323 TDX gene Proteins 0.000 title claims abstract description 25
- 229920002488 Hemicellulose Polymers 0.000 title claims abstract description 17
- 230000033228 biological regulation Effects 0.000 title claims abstract description 15
- 230000015572 biosynthetic process Effects 0.000 title claims abstract description 14
- 238000003786 synthesis reaction Methods 0.000 title claims abstract description 14
- 229920002522 Wood fibre Polymers 0.000 claims abstract description 35
- 108090000623 proteins and genes Proteins 0.000 claims abstract description 35
- 239000002025 wood fiber Substances 0.000 claims abstract description 35
- 230000009261 transgenic effect Effects 0.000 claims abstract description 12
- 238000011161 development Methods 0.000 claims abstract description 9
- 230000001105 regulatory effect Effects 0.000 claims abstract description 5
- 238000000034 method Methods 0.000 claims description 9
- 102000004169 proteins and genes Human genes 0.000 claims description 6
- 241000196324 Embryophyta Species 0.000 abstract description 13
- 241000219000 Populus Species 0.000 abstract description 13
- QMGVPVSNSZLJIA-UHFFFAOYSA-N Nux Vomica Natural products C1C2C3C4N(C=5C6=CC=CC=5)C(=O)CC3OCC=C2CN2C1C46CC2 QMGVPVSNSZLJIA-UHFFFAOYSA-N 0.000 abstract description 6
- 244000107975 Strychnos nux-vomica Species 0.000 abstract description 6
- 101100317563 Arabidopsis thaliana XTH15 gene Proteins 0.000 abstract description 2
- 101100317564 Arabidopsis thaliana XTH16 gene Proteins 0.000 abstract description 2
- 101100317565 Arabidopsis thaliana XTH17 gene Proteins 0.000 abstract description 2
- 239000000835 fiber Substances 0.000 description 8
- 241000894007 species Species 0.000 description 8
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- 230000014509 gene expression Effects 0.000 description 7
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- 239000000463 material Substances 0.000 description 6
- 238000012163 sequencing technique Methods 0.000 description 6
- 210000004027 cell Anatomy 0.000 description 5
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- 235000006508 Nelumbo nucifera Nutrition 0.000 description 4
- 235000006510 Nelumbo pentapetala Nutrition 0.000 description 4
- 238000004458 analytical method Methods 0.000 description 4
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- 238000011529 RT qPCR Methods 0.000 description 3
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- 229920005610 lignin Polymers 0.000 description 3
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- 241000589158 Agrobacterium Species 0.000 description 2
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- 102000004190 Enzymes Human genes 0.000 description 2
- 108010076504 Protein Sorting Signals Proteins 0.000 description 2
- 238000006243 chemical reaction Methods 0.000 description 2
- 239000011121 hardwood Substances 0.000 description 2
- 230000026731 phosphorylation Effects 0.000 description 2
- 238000006366 phosphorylation reaction Methods 0.000 description 2
- 239000000126 substance Substances 0.000 description 2
- 238000012360 testing method Methods 0.000 description 2
- 108010031061 xyloglucan - xyloglucosyltransferase Proteins 0.000 description 2
- 241000219194 Arabidopsis Species 0.000 description 1
- OKTJSMMVPCPJKN-UHFFFAOYSA-N Carbon Chemical compound [C] OKTJSMMVPCPJKN-UHFFFAOYSA-N 0.000 description 1
- 101150027068 DEGS1 gene Proteins 0.000 description 1
- 108091028043 Nucleic acid sequence Proteins 0.000 description 1
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- 241000610446 Schima Species 0.000 description 1
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- 241000422846 Sequoiadendron giganteum Species 0.000 description 1
- 241001122767 Theaceae Species 0.000 description 1
- 108700019146 Transgenes Proteins 0.000 description 1
- 229920002000 Xyloglucan Polymers 0.000 description 1
- 150000001413 amino acids Chemical class 0.000 description 1
- 239000004566 building material Substances 0.000 description 1
- 229910052799 carbon Inorganic materials 0.000 description 1
- 238000004737 colorimetric analysis Methods 0.000 description 1
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- 238000010079 rubber tapping Methods 0.000 description 1
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- 230000035939 shock Effects 0.000 description 1
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- 238000010257 thawing Methods 0.000 description 1
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- XLYOFNOQVPJJNP-UHFFFAOYSA-N water Substances O XLYOFNOQVPJJNP-UHFFFAOYSA-N 0.000 description 1
Classifications
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- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
- C12N9/00—Enzymes; Proenzymes; Compositions thereof; Processes for preparing, activating, inhibiting, separating or purifying enzymes
- C12N9/10—Transferases (2.)
- C12N9/1048—Glycosyltransferases (2.4)
- C12N9/1051—Hexosyltransferases (2.4.1)
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- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
- C12N15/00—Mutation or genetic engineering; DNA or RNA concerning genetic engineering, vectors, e.g. plasmids, or their isolation, preparation or purification; Use of hosts therefor
- C12N15/09—Recombinant DNA-technology
- C12N15/63—Introduction of foreign genetic material using vectors; Vectors; Use of hosts therefor; Regulation of expression
- C12N15/79—Vectors or expression systems specially adapted for eukaryotic hosts
- C12N15/82—Vectors or expression systems specially adapted for eukaryotic hosts for plant cells, e.g. plant artificial chromosomes (PACs)
- C12N15/8241—Phenotypically and genetically modified plants via recombinant DNA technology
- C12N15/8242—Phenotypically and genetically modified plants via recombinant DNA technology with non-agronomic quality (output) traits, e.g. for industrial processing; Value added, non-agronomic traits
- C12N15/8243—Phenotypically and genetically modified plants via recombinant DNA technology with non-agronomic quality (output) traits, e.g. for industrial processing; Value added, non-agronomic traits involving biosynthetic or metabolic pathways, i.e. metabolic engineering, e.g. nicotine, caffeine
- C12N15/8245—Phenotypically and genetically modified plants via recombinant DNA technology with non-agronomic quality (output) traits, e.g. for industrial processing; Value added, non-agronomic traits involving biosynthetic or metabolic pathways, i.e. metabolic engineering, e.g. nicotine, caffeine involving modified carbohydrate or sugar alcohol metabolism, e.g. starch biosynthesis
- C12N15/8246—Non-starch polysaccharides, e.g. cellulose, fructans, levans
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- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
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- C12N15/00—Mutation or genetic engineering; DNA or RNA concerning genetic engineering, vectors, e.g. plasmids, or their isolation, preparation or purification; Use of hosts therefor
- C12N15/09—Recombinant DNA-technology
- C12N15/63—Introduction of foreign genetic material using vectors; Vectors; Use of hosts therefor; Regulation of expression
- C12N15/79—Vectors or expression systems specially adapted for eukaryotic hosts
- C12N15/82—Vectors or expression systems specially adapted for eukaryotic hosts for plant cells, e.g. plant artificial chromosomes (PACs)
- C12N15/8241—Phenotypically and genetically modified plants via recombinant DNA technology
- C12N15/8261—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield
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- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12Y—ENZYMES
- C12Y204/00—Glycosyltransferases (2.4)
- C12Y204/01—Hexosyltransferases (2.4.1)
- C12Y204/01207—Xyloglucan:xyloglucosyl transferase (2.4.1.207)
-
- Y—GENERAL TAGGING OF NEW TECHNOLOGICAL DEVELOPMENTS; GENERAL TAGGING OF CROSS-SECTIONAL TECHNOLOGIES SPANNING OVER SEVERAL SECTIONS OF THE IPC; TECHNICAL SUBJECTS COVERED BY FORMER USPC CROSS-REFERENCE ART COLLECTIONS [XRACs] AND DIGESTS
- Y02—TECHNOLOGIES OR APPLICATIONS FOR MITIGATION OR ADAPTATION AGAINST CLIMATE CHANGE
- Y02A—TECHNOLOGIES FOR ADAPTATION TO CLIMATE CHANGE
- Y02A40/00—Adaptation technologies in agriculture, forestry, livestock or agroalimentary production
- Y02A40/10—Adaptation technologies in agriculture, forestry, livestock or agroalimentary production in agriculture
- Y02A40/146—Genetically Modified [GMO] plants, e.g. transgenic plants
Abstract
An Ssu TDX gene closely related to hemicellulose synthesis regulation and control and application thereof belong to the field of biotechnology. The invention provides Ssu TDX genes closely related to hemicellulose synthesis regulation, the genes comprise 3 serial repeated units Ssu XTH15, ssu XTH16 and Ssu XTH17, CDS sequences of the 3 serial repeated units are respectively shown in SEQ ID NO. 1-3, and the invention provides application of the Ssu TDX genes. The Ssu TDX gene found in the nux vomica is closely related to hemicellulose synthesis regulation, the Ssu TDX gene is over-expressed in poplar, the length of wood fiber of transgenic poplar is longer than that of wild type contrast, the Ssu TDX gene is verified to have the function of regulating the growth and development of plant wood fiber, and candidate genes are provided for directional cultivation of nux vomica and other plant long wood fiber length varieties.
Description
Technical Field
The invention belongs to the technical field of biology, and particularly relates to an Ssu TDX gene closely related to hemicellulose synthesis regulation and control and application thereof.
Background
The tree lotus (Schima superba) is evergreen hardwood broad-leaved big tree of the genus Phyllostachys (Schima reinw.) of the family Theaceae, and the ninth nationwide forest resource checking data shows that the tree is ranked according to the important value of the nationwide tree species and is the first most precious high-quality broad-leaved afforestation tree species in the south of China, and the main promotion of Zhejiang is the first of ten carbon sink tree species. The trunk end is straight, and the wood is dense (air dry density is 0.697 g/cm) 3 ) Even structure, better machining comprehensive performance and huge market demand. The study shows that compared with other hardwood wood species, the method has the advantages that the content of the xylogen hemicellulose is lower (lower than the average 2.5% of the tree species), the anatomical structure of the wood fiber (wood fiber length, width, wall thickness and the like) is obviously better than that of other tree species by 1.6-3.3 times (Table 1), and the cell wall of the wood fiber has stronger plasticity, but the variation in the population is larger (the variation coefficient is as high as 10%).
Note that: fiber classification standard: fiber length: < 900 μm, short: 900-1600 μm, medium: > 1600um, long (CLAWA, 1937): fiber width: < 16um, fine: 16-25 μm, medium: 26-30 μm, coarse: > 30 μm, very coarse (Pashin et al 1964): wall thickness: very thin, cells are much thicker than cell walls; thin, cells have a larger wall thickness than cells; the thickness of the cell cavity is smaller than that of the cell wall; very thick, the cells are almost completely closed.
There is less intensive research on the mechanism of growth and development of wood lotus wood fibers. In model plants arabidopsis and poplar, genes associated with vascular bundle/wood fiber growth and development are mainly involved in synthesis and transcriptional regulation of cellulose, hemicellulose and lignin. What kind of substances and their regulation and control processes are limiting factors in the carpus littoralis are not known at present. The wood fiber anatomical structure has remarkable influence on the physical properties of the wood, and has important influence on the application of the wood, such as the paper-making material needs the materials with long wood fiber and strong wall thickness and toughness, the building material needs the materials with wide wood fiber and strong wall thickness and rigidity, and the like. The work of improving the morphology of the target material wood fiber anatomical structure by utilizing the regulation genes related to the length development of the wood lotus wood fiber has important scientific significance and application value.
Disclosure of Invention
Aiming at the problems existing in the prior art, the invention aims at providing a Ssu TDX gene closely related to hemicellulose synthesis regulation and control and a technical scheme of application thereof.
The invention is realized by the following technical scheme:
in a first aspect, the invention provides a Ssu TDX gene closely related to hemicellulose synthesis regulation, the gene comprises 3 serial repeated units Ssu XTH15, ssu XTH16 and Ssu XTH17, and CDS sequences of the 3 serial repeated units of the X XTH are shown in SEQ ID NO. 1-3 respectively.
Further, the SkuXTH 15, skuXTH 16 and SkuXTH 17 genes encode proteins as shown in SEQ ID NO. 4-6.
The second aspect of the invention provides application of the Ssu TDX gene in regulating and controlling growth and development of plant wood fibers.
The third aspect of the invention provides an application of the SsuTDX gene in improving the length of plant wood fibers.
In a fourth aspect, the present invention provides a method of increasing the length of a plant wood fiber, comprising the steps of: introducing the Ssu TDX gene of claim 1 into a target plant to obtain a transgenic plant; the length of the transgenic wood fiber is significantly increased compared to the wild type control of the target plant.
Experiments prove that the Ssu TDX gene found in the nux vomica is closely related to hemicellulose synthesis regulation, the Ssu TDX gene is over-expressed in poplar, the length of wood fiber of transgenic poplar is longer than that of wild control, the Ssu TDX gene is verified to have the function of regulating the growth and development of plant wood fiber, and candidate genes are provided for directional cultivation of the nux vomica and other long plant wood fiber length varieties.
Drawings
Fiber morphology and chemical composition analysis of the wood charges S1-S3 of FIG. 1. In the figure: a-D fiber length, width, secondary wall thickness, and fiber aspect ratio. E cellulose, hemicellulose and lignin content. Error bars represent standard deviation. Different letters represent significant differences in the progeny of different half-siblings (p <0.01 by one-way anova).
FIG. 2SsuTDX gene structure schematic.
FIG. 3 alignment of amino acid sequences of Ssu TDX genes with model species.
FIG. 4 enzymatic Activity of SkuTDX protein.
FIG. 5 validates RNA-seq data by qRT-PCR. In the figure: the bar graph shows qRT-PCR results and the line graph shows transcriptome results.
FIG. 6 analysis of wood fiber length of SsuTDX transgenic poplar. In the figure: a:2 months old; b:3 months of age.
FIG. 7 effect of SsuTDX gene on Yang Shumu fiber length.
FIG. 8 effect of SsuTDX gene on Yang Shujing anatomy.
FIG. 9 effect of Ssu TDX gene on poplar stem cell wall thickness.
Detailed Description
The invention is further illustrated by the following examples.
Examples:
sample preparation
The test materials are from a free pollination family measuring forest of a Ou 12-year old Mulotus excellent tree of Fujian, and 3 materials with obvious variation of wood fiber anatomical morphology among individuals in one family are screened. The cortex (secondary xylem, secondary phloem and cambium) of the 3 half-sibling progenies (S1, S2 and S3, fig. 1) was collected, snap frozen with liquid nitrogen, stored at-80 ℃ for transcriptome sequencing and subsequent gene function studies.
(II) sequencing and transcriptome analysis
Using Yisi Easy Plant RNA Kit (Polysaccharides)&Polyphenolic-rich) was used to extract total RNAUltra TM RNA Library Prep Kit for/>(Beckman Coulter, beverly, USA) cDNA libraries were constructed and sequenced using Illumina Novaseq platform (IlluminaInc, USA) from Norway. The raw data obtained by sequencing contains a small amount of reads with sequencing adaptors or lower sequencing quality. In order to ensure the quality and reliability of data analysis, the original data needs to be filtered. Meanwhile, Q20, Q30 and GC content of clean data were calculated. All analyses that follow are high quality analyses based on clean data.
The wood lotus genome has been sequenced and assembled, so clean reads are aligned directly to the genome using HISAT2 v 2.0.5. And new transcripts were assembled by StringTie software. Gene expression levels were determined by FPKM, and identification criteria for differentially expressed genes (differential expressed genes, DEGs) were referenced to |log2 (FoldChange) | >0 and padj.ltoreq.0.05.
(III) differential Gene screening
The expression levels of the genes were compared with FPKM. Differential expression (FDR.ltoreq.0.01) was determined using cuffdiff software. And analyzing differential genes related to synthesis and regulation of cellulose, hemicellulose and lignin to obtain an Ssu TDX gene sequence closely related to synthesis and regulation of hemicellulose. The gene comprises 3 serial repeated units (SkuXTH 15, skuXTH 16 and SkuXTH 17) (figure 2), the CDS sequences of the three genes are 876 bp, 912 bp and 888bp respectively, the nucleotide sequences are shown as SEQ ID NO. 1-3, the encoding proteins respectively comprise amino acids 291, 300 aa and 295aa, and the amino acid sequences are shown as SEQ ID NO. 4-6.
The Ssu TDX gene is mainly expressed in phloem and xylem, has XET enzyme and XEH enzyme activities as measured by a colorimetry and a viscosity method, has the highest XET enzyme activity at pH 6.5 and has a hydrolytic activity of about 2 hours, and has specificity to xyloglucan. Wherein the membrane-spanning structure, signal peptide and phosphorylation site of the xylem protein are predicted as shown in table 2, the amino acid sequence of the Ssu TDX gene is compared with the model species as shown in FIG. 3, and the enzyme activity of the Ssu TDX protein is shown in FIG. 4.
TABLE 2 prediction of transmembrane Structure, signal peptide and phosphorylation site of xylem XTH protein
(IV) quantitative analysis of fluorescence
To verify the SquTDX gene, the relative expression levels of S1-S3 material were determined, and RT-qPCR was performed using the xylem-specific SquACT gene as an internal control and a Applied Biosystems Q (USA) instrument. mu.L of the reaction system contained 10. Mu.L of 2X TB GreenPremix Ex TaqII (Tli RNaseH Plus) (RR 820A, taKaRa, japan), 2. Mu.L of diluted cDNA, 0.8. Mu.L of each of the front and rear primers (Table 3) (10 mM), 0.4. Mu. L ROX Reference Dye (50X) and 6. Mu.L of water. The reaction procedure was as follows: incubating at 95 ℃ for 30s; incubating for 5s at 95 ℃ and incubating for 30s at 60 ℃ and circulating for 40 times; and (5) collecting a dissolution curve. Average Ct values were calculated from values of 3 biological replicates and 3 technical replicates using 2 -ΔΔCt The relative expression level of the gene was calculated by the method. As shown in FIG. 5, the Ssu TDX gene expression level was consistent with the result of transcriptome sequencing, and the expression level was highest in S2.
(fifth) transgene verification
The full-length cDNA of Ssu TDX was cloned into the plant expression vector pCAMIA1300-GFP containing the 35S promoter. The method comprises the following specific steps: specific primers were designed, see Table 3, and Ssu TDX cDNA sequence was obtained using Mulotus S2cDNA as template. The pMD20-T vector was ligated by PCR tapping recovery, TOP10 was transformed by heat shock, and pCAMIA1300 was ligated by Gateway to construct the overexpression vector p35S: : ssuTDX. The recombinant vector is transferred into GV3101 agrobacterium competent by freeze thawing method. Agrobacterium-mediated leaf disc transformation of poplar. And (5) carrying out phenotypic character observation, wood fiber morphology, hemicellulose content and the like on the transgenic positive strain.
TABLE 3 primers related thereto
The test results show that:
SsuTDX can obviously promote the elongation development of Yang Shumu fiber, and the elongation rate can reach about 15%.
Evidence 1: as shown in FIG. 6, at2 months of soil cultivation of transgenic poplar, the wood fiber lengths of the SkuXTH 15-OE, skuXTH 16-OE and SkuXTH 17-OE lines were significantly greater than WT at 15 internodes, increased by 13.34%, 20.76% and 21.14%, respectively. When the transgenic poplar is cultivated in soil for 3 months, the wood fiber lengths of SkuXTH 15-OE, skuXTH 16-OE and SkuXTH 17-OE strains are extremely obviously larger than those of WT, and the wood fiber lengths are increased by 8.46%, 15.44% and 12.70% respectively; between 20 internodes, the wood fiber lengths of SkuXTH 15-OE, skuXTH 16-OE and SkuXTH 17-OE strains were significantly greater than WT, increased by 9.23%, 15.65% and 17.99%, respectively.
Evidence 2: as shown in FIG. 7, when the transgenic poplar was cultivated in soil for 90d, the length of wood fiber between 10 th, 15 th and 20 th nodes of SkuXTH 15&16&17-OE strain was significantly greater than WT, the thickness of xylem between 10 th nodes of SkuXTH 15&16&17-OE strain was significantly less than WT (FIG. 8), and the thickness of cell wall of SkuXTH 15&16&17-OE strain was significantly less than WT (FIG. 9).
In conclusion, the Ssu TDX gene found in the nux vomica is closely related to hemicellulose synthesis regulation, the Ssu TDX gene is overexpressed in poplar, the length of wood fiber of transgenic poplar is longer than that of wild type control, the Ssu TDX gene is verified to have the function of regulating the growth and development of plant wood fiber, and candidate genes are provided for directional cultivation of the nux vomica and other plant long wood fiber length varieties.
Claims (5)
1. SkuTDX gene closely related to hemicellulose synthesis regulation, which is characterized by comprising 3 serial repeated units SkuXTH 15, skuXTH 16 and SkuXTH 17, wherein CDS sequences of the 3 serial repeated units of the SkuXTH are respectively shown in SEQ ID NO. 1-3.
2. The SkuTDX gene according to claim 1, wherein the proteins encoded by SkuXTH 15, skuXTH 16 and SkuXTH 17 genes are shown in SEQ ID NO. 4-6.
3. Use of the Ssu TDX gene according to claim 1 for regulating growth and development of plant wood fiber.
4. Use of the Ssu TDX gene according to claim 1 for increasing the length of plant wood fiber.
5. A method for increasing the length of plant wood fibers comprising the steps of: introducing the Ssu TDX gene of claim 1 into a target plant to obtain a transgenic plant; compared with the wild type of the target plant, the length of the transgenic wood fiber is obviously improved.
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