Use of mitochondrial GPAT1 gene and mitochondrial GPAT2 gene in regulating plant secondary cell wall thickness

Abstract

The present application relates to the technical field of genetic engineering, and particularly relates to application of mitochondrial GPAT1 gene and mitochondrial GPAT2 gene in regulating plant secondary cell wall thickness. The present application firstly finds that the mitochondrial GPAT1 gene and the mitochondrial GPAT2 gene have the effect of regulating the plant secondary cell wall thickness, and the effect is embodied in that the mitochondrial GPAT1 gene and the mitochondrial GPAT2 gene regulate the formation of the secondary cell wall by negatively regulating the lignin and cellulose biosynthesis related genes. The results of the specific embodiments of the present application show that when the mitochondrial GPAT1 gene and the mitochondrial GPAT2 gene are deleted, the lignin and cellulose synthesis genes are up-regulated, a large amount of lignin and cellulose is accumulated in the cell wall of the Arabidopsis thaliana double mutant gpat1gpat2 seedling, the cell wall becomes thicker and harder, and the cotyledon texture is harder.

Description

TECHNICAL FIELD

[0001] The present application relates to the technical field of genetic engineering, and particularly relates to application of mitochondrial GPAT1 gene and mitochondrial GPAT2 gene in regulating thickness of plant secondary cell wall. BACKGROUND

[0002] Secondary cell wall (SCW) is located between primary cell wall and plasma membrane, and is composed of lignin, cellulose and hemicellulose (Wang et al. 2023). The biosynthesis of lignin in plants starts from the phenylpropanoid metabolic pathway. Lignin is a natural macromolecular phenolic polymer with complex structure, which is located between cellulose and hemicellulose in the cell wall, and is an important component of the cell wall. It is the second most abundant high molecular polymer, accounting for about 30% of the content of organic carbon. Lignin plays an important role in plant growth and development. On the one hand, it can enhance the mechanical strength of plants, provide support for plant growth and development, and promote the transportation of minerals and water in the vascular bundle; on the other hand, lignin is involved in the response of plants to biological and abiotic stresses, which helps to improve the stress resistance of plants. Cellulose is the main component of plant cell wall, accounting for about 33% of the total weight of plants, and its economic value can be summarized as food, fiber, paper and bioenergy, which are closely related to human life. Therefore, cellulose is the most economically valuable cell wall component. Its structure is very simple, but the synthesis mechanism is extremely complex. Current research suggests that cellulose is composed of unbranched β-1,4-glucan chains, each chain containing up to 15,000 glucose residues. The glucan chains are combined by hydrogen bonds and van der Waals forces and arranged in parallel, and each 36 glucan chains form a microfibril, which further arranges and deposits to form the high-order structure of cellulose. Hemicellulose is the second most abundant polysaccharide in nature, and plays an irreplaceable role in plant growth and development. Plants with high hemicellulose content not only have strong resistance to lodging, but also can greatly improve the utilization rate of renewable energy.

[0003] The network formation of secondary cell wall is extremely complex and is regulated by multiple parties. It has been reported that transcription factors Tier 1, 2 and 3 containing MYB or NAC transcription factors play a crucial role in secondary cell wall biosynthesis (Zhong and Ye, 2009; Kumar et al. 2016; Ohtani and Demura, 2019). Ubiquitination is also involved in the formation of secondary cell wall. The subunit DDB1a of E3 ubiquitin ligase complex (Beris et al. 2016), cotton GhHUB2 (Feng et al. 2018), rice OsFBK1 (Borah and Khurana, 2018) as a component of SCF E3 ligase complex, have been reported to directly or indirectly regulate secondary cell wall synthesis. So far, the regulation of GPAT on secondary cell wall development is still blank. SUMMARY

[0004] The purpose of the present application is to provide the application of mitochondrial GPAT1 gene and mitochondrial GPAT2 gene in regulating the thickness of plant secondary cell wall, so as to solve the problems existing in the prior art. It is found for the first time that mitochondrial GPAT1 gene and mitochondrial GPAT2 gene have the function of regulating the thickness of plant secondary cell wall.

[0005] In order to achieve the above purpose, the present application provides the following scheme:

[0006] The present application provides the application of mitochondrial GPAT1 gene and mitochondrial GPAT2 gene in regulating the thickness of plant secondary cell wall.

[0007] As an additional scheme, the sequence number of the mitochondrial GPAT1 gene in TAIR (The Arabidopsis Information Resource) is AT1G06520; the sequence number of the mitochondrial GPAT2 gene in TAIR is AT1G02390.

[0008] The present application provides a method for increasing the thickness of plant secondary cell wall, comprising the step of inhibiting the expression amount of mitochondrial GPAT1 gene and mitochondrial GPAT2 gene in the plant.

[0009] The present application provides the application of mitochondrial GPAT1 gene and mitochondrial GPAT2 gene in cultivating transgenic plants with thick secondary cell wall.

[0010] As an additional scheme, the present application further provides a method for cultivating transgenic plants with thick secondary cell wall, comprising the steps of inhibiting the expression amount of mitochondrial GPAT1 gene and mitochondrial GPAT2 gene in the plant, and obtaining transgenic plants with thick secondary cell wall.

[0011] The application provides applications of mitochondrial GPAT1 genes and mitochondrial GPAT2 genes in regulating lignin content of plants.

[0012] The application provides a method for increasing lignin content of plants, which comprises the step of inhibiting expression amount of mitochondrial GPAT1 genes and mitochondrial GPAT2 genes in the plants.

[0013] The application provides applications of mitochondrial GPAT1 genes and mitochondrial GPAT2 genes in cultivating transgenic plants with high lignin content.

[0014] As an additional solution, the application further provides a method for cultivating transgenic plants with high lignin content, which comprises the steps of inhibiting expression amount of mitochondrial GPAT1 genes and mitochondrial GPAT2 genes in the plants, and obtaining the transgenic plants with high lignin content.

[0015] The application provides applications of mitochondrial GPAT1 genes and mitochondrial GPAT2 genes in regulating cellulose content of plants.

[0016] The application provides a method for increasing cellulose content of plants, which comprises the step of inhibiting expression amount of mitochondrial GPAT1 genes and mitochondrial GPAT2 genes in the plants.

[0017] The application provides applications of mitochondrial GPAT1 genes and mitochondrial GPAT2 genes in cultivating transgenic plants with high cellulose content.

[0018] As an additional solution, the application further provides a method for cultivating transgenic plants with high cellulose content, which comprises the steps of inhibiting expression amount of mitochondrial GPAT1 genes and mitochondrial GPAT2 genes in the plants, and obtaining the transgenic plants with high lignin content.

[0019] Preferably, the plants are Arabidopsis thaliana.

[0020] As an additional solution, the application provides applications of mitochondrial GPAT1 genes and mitochondrial GPAT2 genes in regulating drought resistance of plants.

[0021] Further preferably, the plants are Arabidopsis thaliana.

[0022] As an additional solution, the application provides a method for increasing drought resistance of plants, which comprises the step of inhibiting expression amount of mitochondrial GPAT1 genes and mitochondrial GPAT2 genes in the plants.

[0023] Further preferably, the plants are Arabidopsis thaliana.

[0024] As an additional solution, the present application provides the use of mitochondrial GPAT1 gene and mitochondrial GPAT2 gene in cultivating drought-resistant transgenic plants.

[0025] Further preferably, the plant is Arabidopsis thaliana.

[0026] As an additional solution, the present application provides a method for cultivating drought-resistant transgenic plants, comprising the step of inhibiting the expression amount of mitochondrial GPAT1 gene and mitochondrial GPAT2 gene in a plant to obtain the drought-resistant transgenic plant.

[0027] Further preferably, the plant is Arabidopsis thaliana.

[0028] As an additional solution, the present application provides the use of mitochondrial GPAT1 gene and mitochondrial GPAT2 gene in cultivating drought-resistant transgenic plants.

[0029] Further preferably, the plant is Arabidopsis thaliana.

[0030] The present application discloses the following technical effects:

[0031] The present application first discovers that mitochondrial GPAT1 gene and mitochondrial GPAT2 gene have the function of regulating the thickness of plant secondary cell wall, which is embodied in that mitochondrial GPAT1 gene and mitochondrial GPAT2 gene regulate the formation of secondary cell wall by negatively regulating the biosynthesis of lignin and cellulose related genes. The results of specific embodiments of the present application show that when mitochondrial GPAT1 gene and mitochondrial GPAT2 gene are deleted, the synthesis genes of lignin and cellulose are up-regulated, and a large amount of lignin and cellulose accumulates in the cell wall of Arabidopsis thaliana double mutant gpat1 gpat2 seedlings, so that the cell wall becomes thicker and harder, and the cotyledon texture becomes harder. Therefore, mitochondrial GPAT1 gene and mitochondrial GPAT2 gene can be used for cultivating thick secondary cell wall transgenic plants, and provide a new way for the cultivation of new plant varieties. BRIEF DESCRIPTION OF DRAWINGS

[0032] In order to more clearly illustrate the technical solutions in the embodiments of the present application or the prior art, the drawings needed in the embodiments will be briefly introduced below. Obviously, the drawings in the following description are only some embodiments of the present application, and those skilled in the art can obtain other drawings according to these drawings without creative labor.

[0033] Figure 1Figure 6 shows that mitochondrial GPAT1 and GPAT2 genes regulate lignin synthesis. A, cotyledon and hypocotyl of wild-type Arabidopsis seedlings (right) and Arabidopsis double mutant gpat1 gpat2 seedlings (left) dyed with phloroglucinol; B, roots of wild-type Arabidopsis seedlings (right) and Arabidopsis double mutant gpat1 gpat2 seedlings (left) dyed with phloroglucinol; C, results of quantitative analysis of lignin content, *** indicates p<0.001; D, results of real-time quantitative PCR (qPCR) of lignin biosynthesis genes; all data are expressed as mean ± standard deviation of three biologically independent samples, three independent experiments were performed with similar results, statistical analysis was performed using one-way ANOVA, * indicates p<0.05, ** indicates p<0.01, *** indicates p<0.001, n.s. indicates no significant difference; WT is wild-type Arabidopsis, gpat1 gpat2 is Arabidopsis double mutant gpat1 gpat2;

[0034] Figure 2 Figure 7 shows a comparison of different seedling lignin extracts and cellulose extracts. A, comparison of lignin extracts; B, comparison of cellulose extracts, the lower right corner is the precipitate during cellulose extraction; Blank is a blank control, WT is wild-type Arabidopsis, gpat1 gpat2 is Arabidopsis double mutant gpat1 gpat2;

[0035] Figure 3 Figure 8 shows that mitochondrial GPAT1 and GPAT2 genes regulate cellulose synthesis. A, cotyledon and hypocotyl of wild-type Arabidopsis seedlings (right) and Arabidopsis double mutant gpat1 gpat2 seedlings (left) dyed with zinc chloride-iodine method, scale = 500 μm; B, results of cellulose content determination kit analysis of cellulose content, *** indicates p<0.05; C, qPCR results of lignin biosynthesis genes, all data are expressed as mean ± standard deviation of three biologically independent samples, three independent experiments were performed with similar results, statistical analysis was performed using one-way ANOVA, ** indicates p<0.01, *** indicates p<0.001, n.s. indicates no significant difference; WT is wild-type Arabidopsis, gpat1 gpat2 is Arabidopsis double mutant gpat1 gpat2;

[0036] Figure 4Fig. 6A and Fig. 6B are the staining images and thickness quantification of cotyledon cell walls of wild type Arabidopsis and Arabidopsis double mutant gpat1 gpat2 seedlings; Fig. 6A is the transmission electron microscopy images of the adaxial surface of wild type Arabidopsis and Arabidopsis double mutant gpat1 gpat2 cotyledons, CW is the cell wall, the black arrow indicates the cuticle, the scale bar = 200 nm; Fig. 6B is the thickness quantification of cell walls of wild type Arabidopsis and Arabidopsis double mutant gpat1 gpat2; *** indicates P < 0.001; WT is wild type Arabidopsis, gpat1 gpat2 is Arabidopsis double mutant gpat1 gpat2;

[0037] Figure 5 Fig. 5 is the investigation results of the sensitivity to water loss of wild type Arabidopsis, Arabidopsis double mutant gpat1 gpat2, Arabidopsis complementation line GPAT1 ::gpat1 gpat2 and Arabidopsis complementation line GPAT2 ::gpat1 gpat2, all data are expressed as the mean ± standard deviation of three biologically independent samples, statistical analysis was performed by one-way ANOVA, *** indicates p < 0.001; WT is wild type Arabidopsis, gpat1 is Arabidopsis mutant gpat1, gpat2 is Arabidopsis mutant gpat2, gpat1 gpat2 is Arabidopsis double mutant gpat1 gpat2, GPAT1 ::gpat1 gpat2 is Arabidopsis complementation line GPAT1 ::gpat1 gpat2, GPAT2 ::gpat1 gpat2 is Arabidopsis complementation line GPAT2 ::gpat1 gpat2. DETAILED DESCRIPTION

[0038] Various exemplary embodiments of the present application will now be described in detail with reference to the drawings. The detailed description is not intended to limit the present application, but to explain certain aspects, features and embodiments of the present application.

[0039] It is to be understood that the terminology used herein is for the purpose of describing particular embodiments only and is not intended to be limiting of the present application. Additionally, for a range of values of a parameter, unless otherwise indicated, each intervening value by each intervening value, or any other stated value or implicitly supported value by every stated value or implicitly supported value within the stated range is expressly contemplated. The above description is intended to be illustrative, and not restrictive. Many other embodiments will be apparent to those of skill in the art upon reviewing the above description.

[0040] All technical and scientific terms used herein have, unless otherwise defined, the same meaning as commonly understood by one of ordinary skill in the art to which this application pertains. Although methods and materials similar or equivalent to those described herein can be used in the practice or testing of the present application, preferred methods and materials are described. All publications mentioned in this specification are herein incorporated by reference to disclose and describe the methods and / or materials in connection with which the publications are cited. In case of conflict, the content of the present specification will control.

[0041] Various modifications and changes can be made to the specific embodiments described herein without departing from the scope or spirit of the application. Other embodiments will be apparent to those of ordinary skill in the art from the description and examples provided herein. The description and examples are illustrative only.

[0042] As used herein, the terms "comprise", "comprising", "include", "including", "have", "having" and the like are open-ended and do not exclude additional, unrecited elements or method steps.

[0043] Example 1

[0044] 1. Plant material

[0045] Arabidopsis thaliana double mutant gpatl gpat2, Arabidopsis thaliana complemented line GPAT1 ::gpatl gpat2 and Arabidopsis thaliana complemented line GPAT2::gpatl gpat2 are disclosed in the document "Mitochondrial GPAT-derived LPA controls auxin-dependent embryonic and postembryonic development" and are committed to be released for 20 years.

[0046] 2. Plant material growth conditions

[0047] Seeds of the above plant material and wild type Arabidopsis thaliana were germinated on half-strength Murashige and Skoog (MS) medium and, after two days of vernalization, were transferred to a growth chamber with a light intensity of 160 μιηοΙ·ηι2·ηηη2for 6 days, a day-night cycle of 14h / 10h (14h of light and 10h of darkness) and a temperature of 23°C. After that, seedlings were transferred to soil under the same conditions for further cultivation. -2 -1

[0048] 3. Investigation method

[0049] ​​Lignin content was detected by phloroglucinol method: 6-day-old Arabidopsis seedlings were soaked in lignin acidified solution for 10 min, then equal amount of phloroglucinol staining solution (Mcclin, China) was added dropwise to the acidified Arabidopsis seedlings, and after standing, the staining of lignin in Arabidopsis seedlings was observed under a stereomicroscope. Then, the lignin content in Arabidopsis seedlings was investigated using a lignin content determination kit (Mcclin, China), and the experimental steps were performed according to the production instructions of the kit, with reagent without sample as blank control (Blank).

[0050] Cellulose content was detected by zinc chloride iodide method: 6-day-old Arabidopsis seedlings were soaked in cellulose staining solution (Yuan Ye, China) for 5 min, and the staining of cellulose in Arabidopsis seedlings was observed under a stereomicroscope. Then, the cellulose content in Arabidopsis seedlings was investigated using a cellulose content determination kit (Mcclin, China), and the experimental steps were performed according to the production instructions of the kit, with reagent without sample as blank control (Blank).

[0051] Cotyledon cell wall thickness was observed by transmission electron microscopy (TEM) (HITACH HT7800, Japan). Specifically, the cotyledons of wild-type Arabidopsis and Arabidopsis double mutant gpat1 gpat2 were fixed with 2.5% (w / v) glutaraldehyde in 0.2M PBS at 4°C for more than 2h. The samples were washed with 0.1M PBS for 3 times and dehydrated by a series of gradient ethanol. Then, the cotyledons were embedded with epoxy resin and ultrathin sectioned. Finally, the samples were placed on copper mesh, stained with uranyl acetate and lead citrate, and observed under a transmission electron microscope (80.0KV).

[0052] Before water loss determination, Arabidopsis seedlings were grown at 23°C for 1 week, and the water loss rate was calculated, with the formula: water loss rate = (initial weight - weight at a certain time point) / initial weight x 100%.

[0053] 4. RNA extraction and transcription analysis

[0054] Total RNA was extracted from the cotyledons of wild-type Arabidopsis and Arabidopsis double mutant gpat1 gpat2 by using RNA-easy Isolation Reagent kit (Norgen, China). First-strand cDNA was synthesized using 1st Strand cDNA Synthesis Kit (Yikesheng, China). The cDNA was used as template for quantitative real-time PCR (qRT-PCR) using SYBR Green PCR Master Mix (TaKaRa, Japan). ​Quantitative real-time PCR (qPCR) was performed using SYBR Green Master Mix (no rox) (Yisheng, China). Relative quantification results were obtained using ACTIN2 as an internal control and compared via the cycle threshold method (2...). -ΔΔCt The average normalized transcription level was presented. Detection was performed using a Gentier 96E / 96R system (Tianlong, China). Primer sequences used for qPCR and gene accession numbers are shown in Table 1.

[0055] Table 1 Primers and gene accession numbers used in this embodiment

[0056]

[0057]

[0058]

[0059] 5. Data Analysis

[0060] Statistical analysis was performed using IBM SPSS Statistics 25 software. One-way ANOVA was used to compare the means. Quantitative analysis of cell wall thickness was performed using ImageJ software.

[0061] 6. Results and Analysis

[0062] 6.1 Mitochondrial GPAT1 and GPAT2 genes can regulate the formation of the secondary cell wall (SCW).

[0063] In this embodiment, phloroglucinol was used to stain wild-type Arabidopsis thaliana and Arabidopsis thaliana double mutant gpat1 gpat2 seedlings. Phloroglucinol is a lignin-specific dye; after staining, different degrees of lignification result in red or purplish-red hues, with deeper lignification producing darker colors. The results showed that the cotyledons and hypocotyls of the Arabidopsis thaliana double mutant gpat1 gpat2 seedlings were darker than those of wild-type Arabidopsis thaliana, while there was no significant difference in root color. Figure 1 (A and B in the original text). Furthermore, the total lignin content was detected using a kit, and it was found that the total lignin content of Arabidopsis double mutants gpat1 and gpat2 seedlings was significantly higher than that of wild-type Arabidopsis. Figure 1 C and Figure 2 (A) Therefore, we can conclude that the mitochondrial GPAT1 and mitochondrial GPAT2 genes have a negative regulatory effect on lignin content.

[0064] Subsequently, the cellulose content of wild-type Arabidopsis thaliana and Arabidopsis thaliana double mutant gpat1 gpat2 seedlings was analyzed using the zinc chloride-iodine method. The cotyledons and hypocotyls of the Arabidopsis thaliana double mutant gpat1 gpat2 seedlings were significantly darker in color than those of wild-type Arabidopsis thaliana, indicating that the cellulose content of the Arabidopsis thaliana double mutant gpat1 gpat2 seedlings was higher than that of wild-type Arabidopsis thaliana. Figure 3 (A) Furthermore, to clarify this conclusion, this embodiment analyzed the cellulose content of wild-type Arabidopsis thaliana and Arabidopsis thaliana double mutant gpat1 gpat2 seedlings, and similarly found that the cellulose content of Arabidopsis thaliana double mutant gpat1 gpat2 seedlings was higher than that of wild-type Arabidopsis thaliana (A). Figure 3 B and Figure 2 (B in the middle).

[0065] To analyze how the mitochondrial GPAT1 and GPAT2 genes regulate lignin and cellulose content, qPCR was performed on lignin and cellulose synthesis genes in the cotyledons of wild-type Arabidopsis and Arabidopsis double mutants gpat1 and gpat2 seedlings. For lignin synthesis, several genes from the PAL, C4H, 4CL, CCR, CAD, F5H, COM, and LAC families were selected to detect transcriptional levels. The results showed that almost all genes except LAC15 were upregulated, with LAC12 showing the highest upregulation, followed by PAL3. Figure 1 (D in the example). For cellulose synthesis, this example selected 9 genes from the CESA family. Among them, CESA4, CESA8, and CESA9 were significantly upregulated in Arabidopsis double mutant gpat1 gpat2 seedlings, while CESA1 and CESA6 were slightly downregulated. Figure 3 (C in the middle).

[0066] To observe the epidermal structure more clearly, this example used transmission electron microscopy (TEM) to observe the cotyledons of 20-day-old wild-type Arabidopsis thaliana and Arabidopsis thaliana double mutant gpat1gpat2 seedlings. The results showed that the cell wall thickness of the cotyledons of Arabidopsis thaliana double mutant gpat1gpat2 seedlings was significantly higher than that of wild-type Arabidopsis thaliana. Figure 4 ).

[0067] These results indicate that the mitochondrial GPAT1 and GPAT2 genes play a negative regulatory role in lignin and cellulose synthesis. When the mitochondrial GPAT1 and GPAT2 genes are missing, several lignin and cellulose synthesis genes are upregulated, and large amounts of lignin and cellulose accumulate in the cell walls of Arabidopsis double mutant gpat1 gpat2 seedlings, resulting in thicker and harder cell walls and harder cotyledon texture. This result confirms the cause of the cell wall thickening.

[0068] 6.2 Mitochondrial GPAT1 gene and mitochondrial GPAT2 gene maintain plant water balance

[0069] To explore whether the deletion of mitochondrial GPAT1 gene and mitochondrial GPAT2 gene affects the response of plants to water stress, the water loss of seedlings was determined. The results showed that the water loss rate of Arabidopsis double mutant gpat1 gpat2 seedlings was slower than that of wild type Arabidopsis, Arabidopsis complement line GPAT1::gpat1 gpat2 and Arabidopsis complement line GPAT2::gpat1 gpat2 Figure 5 ), and Arabidopsis double mutant gpat1 gpat2 could adapt to more drought conditions. Therefore, mitochondrial GPAT1 gene and mitochondrial GPAT2 gene can be used to cultivate drought-tolerant plants.

[0070] From the above, changes in cell arm thickness can cause imbalance in plant water metabolism. The deletion of mitochondrial GPAT1 gene and mitochondrial GPAT2 gene promotes the synthesis of lignin and cellulose, and the increase in lignin and cellulose content leads to thickening of the cell wall, which can achieve the purpose of regulating the drought tolerance of plants.

[0071] The above-described embodiments only describe the preferred modes of the present application and do not limit the scope of the present application. Without departing from the design spirit of the present application, various modifications and improvements to the technical solutions of the present application made by those skilled in the art shall fall within the protection scope determined by the claims of the present application.

Claims

1. Use of knocking out mitochondrial GPAT1 gene and mitochondrial GPAT2 gene in improving secondary cell wall thickness in Arabidopsis thaliana.

2. A method of increasing secondary cell wall thickness in Arabidopsis thaliana, characterized by, comprising the step of knocking out mitochondrial GPAT1 gene and mitochondrial GPAT2 gene in said Arabidopsis thaliana.

3. Use of knocking out mitochondrial GPAT1 gene and mitochondrial GPAT2 gene in breeding transgenic Arabidopsis thaliana with thick secondary cell wall.

4. Use of knocking out mitochondrial GPAT1 gene and mitochondrial GPAT2 gene in improving lignin content in Arabidopsis thaliana.

5. A method of increasing the lignin content of Arabidopsis thaliana, characterized by, comprising the step of knocking out mitochondrial GPAT1 gene and mitochondrial GPAT2 gene in said Arabidopsis thaliana.

6. Use of knocking out mitochondrial GPAT1 gene and mitochondrial GPAT2 gene in breeding transgenic Arabidopsis thaliana with high lignin content.

7. Use of knocking out mitochondrial GPAT1 gene and mitochondrial GPAT2 gene in improving cellulose content in Arabidopsis thaliana.

8. A method of increasing cellulose content in Arabidopsis thaliana, characterized by, comprising the step of knocking out mitochondrial GPAT1 gene and mitochondrial GPAT2 gene in said Arabidopsis thaliana.

9. Use of knocking out mitochondrial GPAT1 gene and mitochondrial GPAT2 gene in breeding transgenic Arabidopsis thaliana with high cellulose content.

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