KR102397314B1 - A Method for Controlling Plant Organ Abscission by Regulating Lignin Molecular Brace and a Mutant or Transgenic Plant Using the Same - Google Patents

Abstract

The present invention relates to a method for regulating plant detachment by regulating lignin formation in plants and to a mutant or transgenic plant accordingly, and more particularly, a gene encoding ROS-forming enzymes RbohD and/or RbohF is deleted or expressed This inhibition, characterized in that the lignin production is suppressed, mutant or transgenic plants with suppressed or delayed cleavage, and (a) a gene encoding ROS-forming enzymes RbohD and / or RbohF Deleting or suppressing the expression; and (b) selecting a plant in which lignin production is specifically suppressed by leaving cells at the detachment site; relates to a method for inhibiting detachment of a plant comprising; In the present invention, the mechanism of promoting precise processing of the cell wall, intercellular linkage, and cell structure throughout the dissociation site was confirmed, and the formation of escaped cell (SEC)-specific lignin was confirmed to be a major means for accurate organ isolation, There is an effect that can accurately induce the separation of organs within the plant.

Description

A method for controlling the detachment of plants by regulating the formation of lignin in plants, and a mutant or transgenic plant accordingly

The present invention relates to a method for regulating plant detachment by regulating lignin formation in plants and to a mutant or transgenic plant accordingly, and more particularly, a gene encoding ROS-forming enzymes RbohD and/or RbohF is deleted or expressed This inhibition, characterized in that the lignin production is suppressed, mutant or transgenic plants with suppressed or delayed cleavage, and (a) a gene encoding ROS-forming enzymes RbohD and / or RbohF Deleting or suppressing the expression; and (b) selecting a plant in which lignin production is inhibited specifically for deviated cells at the detachment site; a method for inhibiting detachment of a plant comprising a; Enhancing the expression of one or more enzymes selected from the group consisting of superoxide dismutase (SOD, superoxide dismutase) and NADPH oxidase (NOX); And (b) the step of selecting a plant in which the production of lignin is enhanced specifically for the cells leaving the detachment site; it relates to a method for promoting detachment of a plant comprising a.

In animal cells, cells move and exchange materials between cells through close interaction between cells to form tissues and organs. However, in plants, the movement of cells itself or the movement of substances between cells is restricted by the cell wall, which is a characteristic structure thereof. Plants continuously purify the cell wall despite the absence of such cell movement and physical obstacles in the movement of substances. This leads to cell division, expansion, differentiation, and development.

In order to overcome these mechanical limitations, plants must process and destroy the cell wall in a highly purified method during growth and development. It is essential to fine-tune the cell wall spatially and temporally. Tally requires degradation of the lamellae located between the two cell layers at the site of detachment, which increases the susceptibility of plants to infection. Therefore, the separation process must be limitedly performed in a limited area, and a resealing mechanism to protect the newly formed surface must operate. In other words, in the process of separation or detachment of plant organs such as leaves, fruits, and flowers, the cell wall must be accurately processed and destroyed to accurately remove these organs, preserve plant resources, and perfectly maintain the defense system. . However, it is not known how plants protect the cell surface of newly formed organs while at the same time induce the precise destruction and remodeling of the cell wall in space and time.

Accordingly, the present inventors have made intensive efforts to find out how the cell wall is precisely destroyed spatio-temporally and the surface of the remaining organs is protected at the detachment site of a plant. As a result, two adjacent cell types of Arabidopsis are completely different genes The present invention was completed by confirming that the expression pattern is shown, and by identifying that it is possible to induce precise detachment of the plant when regulating their activity to control cell wall treatment.

1. Martinez-Trujillo, M., Limones-Briones, V., Cabrera-Ponce, J. L. & Herrera-Estrella, L. Improving transformation efficiency of Arabidopsis thaliana by modifying the floral dip method. Plant Mol. Biol. Rep. 22, 63-70 (2004).

It is an object of the present invention to provide a novel mutant in which detachment is suppressed, a transgenic plant and a composition for inhibiting detachment of a novel plant.

Another object of the present invention is to provide a method for inhibiting detachment of a plant or inhibiting falling leaves or falling fruits.

Another object of the present invention is to provide a method for promoting detachment of plants.

In order to achieve the above object, the present invention is characterized in that the lignin production is suppressed by deletion or inhibition of the gene encoding the ROS-forming enzymes RbohD and / or RbohF, a mutant or transgenic plant with suppressed cleavage to provide.

The present invention also provides RNAi or antisense oligonucleotides that inhibit the expression of genes involved in lignin production; Or peroxidase (peroxidase), superoxide dismutase (SOD, superoxide dismutase), or a compound inhibiting NADPH oxidase (NOX); provides a composition for inhibiting detachment of plants comprising a.

The present invention also provides a method comprising: (a) deleting or inhibiting the expression of a gene encoding ROS-forming enzymes RbohD and/or RbohF; and (b) selecting a plant in which lignin production is inhibited specifically for deviated cells of the detachment site;

The present invention also comprises the steps of (a) enhancing the expression of one or more enzymes selected from the group consisting of peroxidase, superoxide dismutase (SOD) and NADPH oxidase (NOX); and (b) selecting a plant in which lignin production is inhibited specifically for deviated cells at the detachment site; provides a method for promoting detachment of plants, including.

In the present invention, the mechanism of promoting precise processing of the cell wall, intercellular linkage, and cell structure throughout the detachment site is confirmed, and the formation of secession cells (SEC)-specific lignin at the site of separation is correct for the separation of organs. As it was confirmed that it is a major means for this purpose, it was possible to accurately induce the separation of organs within the plant. Therefore, the present invention can increase crop production and improve crop quality by inducing precise desorption of organs in crop cultivation, and thus can be utilized in various ways for flower farmers and fruit farmers.

1 shows the characteristic cell activity and structure in residual cells (RECs) and SECs located in the remaining tissues of the detachment site. [Scale bars: 50 um (a, b, d), 20 um (e)]
(a) Location of plasma membrane proteins (GFP-SYP122) and cell wall proteins (QRT2-mCherry and XTH28-mCherry) in detachment site cells of the receptacle and detachment site cells of the separated sepal indicates
(b) Through RNA-seq analysis of residual cells (RECs) and escaped cells (SECs), the detachment site of the receptacle remaining after the organ has been separated (RECs, left) and the detachment site of the separated sepal Cell type-specific genes (right, cutoff values: P < 0.05 and log ₂ fold change > 2) were identified in cells (SECs, middle). In the scanning electron micrograph on the left, RECs, SECs, and nectaries were marked in red, blue, and yellow, respectively.
(c) Shows the results of ontology analysis of genes expressed in RECs and SECs using REVIGO.
(d) Shows promoter-GFP analysis of lignin biosynthesis enzymes.
(e) Visualization of lignin at the detachment site through UV autofluorescence, and the number in parentheses indicates the developmental stage of the flower.
2 shows RNAseq analysis of cell type specific transcripts.
(a) Scatter plots (upper right) and Spearman's rank correlation coefficients (lower left) in pairwise comparisons of biological replicates from RNA-seq analysis
(b) The minus-average (MA) plot shows differentially expressed genes in RECs and SECs. Genes showing significantly differential expression patterns (cut-off values: P _adj < 0.05 and log ₂ fold change > 2) are shown in red.
(c) The heat map shows the reproducibility of biological clones, and cell type-specific genes are indicated (cut-off values: P _adj < 0.05 and log ₂ fold change > 2).
Figure 3 shows the expression pattern of the detachment-related gene.
(a) Shows the expression patterns of previously known thalli-related genes (* indicates P _adj > 0.05).
Gene expression patterns by cell type in (b) transcription factor families and (c) MAPK cascade are shown (genes with P _adj < 0.05 and log ₂ fold change > 2 are shown).
(d) Shows the expression pattern of the receptor kinase gene by cell type ( P _adj < 0.05 and log ₂ fold change > 2 as one cell type, log ₂ fold change < 2 as another cell type, for each cell type The top 20 genes were shown, and the same color scale was used for all heatmaps).
Figure 4 confirmed the RNA-seq results through Promoter-GFP analysis of stage 16 flowers from transgenic plants, results consistent with RNA-seq in 17 genes, and RNA-seq in 4 genes (shown in blue) Inconsistent results were confirmed with (Scale bars: 100 um)
5 is an observation of lignin accumulation in SECs. [Scale bars: 20 um (c, d), 100 um (e)]
Genes encoding (a) monolignol biosynthetic enzyme family and (b) lignin polymerase exhibit cell type-specific expression patterns (cut-off values: P _adj < 0.05, log ₂ fold change > 2).
After treatment with PA, a lignin biosynthesis inhibitor, (c) autofluorescence from lignin, and (d) lignin staining with phloroglucinol-HCl was observed. Monolignol was treated externally for 6 hours.
(e) Lignin was stained with phloroglucinol-HCl at the detachment site of Arabidopsis cauline leaf and Ginkgo leaf.
6 confirms that ROS generation by RbohD and RbohF, which are NADPH oxidases, is required for SEC-specific lignin formation. [Scale bars: 20 um (a, b, d), 5 mm (e)]
(a) The inhibitory effect of lignin accumulation by DPI (diphenyleneiodonium), DDC (diethyldithiocarbamic acid) or SHAM (salicylhydroxamic acid) in the presence of monolignol and PA was confirmed.
(b) The accumulation of superoxide and H ₂ O ₂ was detected with NBT (nitroblue tetrazolium) or DAB (3,3-diaminobenzidine) in the flowers of stage 16, respectively.
(c) 2',7'-dichlorofluorescin (DCF) fluorescence was used to quantify H ₂ O ₂ at sepals and petals detachment sites. Three independent experiments were performed for the experimental group of n=9, and the error bar indicates the standard deviation.
(d) Wild-type, rbohD /F , proRbohF :: GFP - Lignin autofluorescence at the sepal detachment site of rbohD /F supplemented with RbohF.
(e) wild-type and It shows the development of flowers and siliques in rbohD /F , and the indicated numbers indicate the positions of the flowers.
7 is an analysis of ROS-metabolizing enzymes associated with ROS accumulation in SECs. [Scale bars: 50 um (a), 100 um (b)].
(a) The effects of ROS enzyme inhibitors, DPI, DDC, and SHAM, in SECs were confirmed by DCF fluorescence at the detachment sites of petals and sepals.
(b) Promoter-GUS analysis for four genes encoding NADPH oxidase in stage 16 flowers.
8 is a result confirming that the SEC-specific lignin brace having a honeycomb structure plays an important role in cell surface integrity and accurate cell wall processing. [*, P <0.05; **, P < 0.001. Auto F., autofluorescence. St., stage. CW, cell wall. Scale bars: 5 um (a, b), 20 um (c, d), 10 um (e), 1 um (f), 0.5 um (f-1, -2, -3), 0.1 mm (g)]
(a) Shows the honeycomb structure of lignin at the detachment site of the sepal, and the plasma membrane protein mCherry-SYP122 visualizes the cell shape.
(b) Indicating the location of the cell wall enzyme PLL18-mCherry at the sepal detachment site, the arrow shows that the location of PLL18 in the first layer of SECs is restricted by lignin.
(c) shows the location of pectinase (At4g18180)-mCherry at the sepal detachment site of wild-type and rbohD /F .
(d) Shows the detachment site located between the petal-receptacle in wild-type and rbohD /F , and the arrow indicates the detachment layer.
(e) After detachment of sepals from wild-type and stage 16 flowers of rbohD /F , detachment surfaces from receptacles and sepals were observed with a scanning electron microscope. Arrows indicate the remaining SECs placed on the RECs of the receptacle.
(f) Transmission electron microscopy of RECs in rbohD / F flowers after stage 16, with arrows indicating defective cutin structures on the surface of remaining SECs.
(g) wild-type and rbohD /F flowers showing pathogen experiments (Pathogen assays), the flowers were exposed to Pst DC3000 for 10 minutes, and the bacterial cfu (colony forming unit) was measured at two different sites and the average (n = 12) It is expressed as ± standard deviation. Four independent experiments were performed, with each measurement point representing 120 flowers.
9 is a scanning electron microscope observation of the lignin structure after cell wall degradation for 2 weeks (a, b) or 3 weeks (c). [Scale bars: 50 um (a), 5 um (b, c)]
10 shows the location of QRT2, a cell wall enzyme, fused with mCherry at the sepal detachment site according to the presence or absence of PA. Arrows indicate mCherry signals according to impaired lignin accumulation in non-deassociated cells. [Auto F., autofluorescence. Scale bars: 20 um]
11 shows the expression patterns according to cell types involved in the biosynthesis and transport of suberin (a) and cutin and/or wax (b). [ P _adj < 0.05, log ₂ fold change > 2, all heatmaps use the same color scale]
Figure 12 confirms the process of transdifferentiation of non-epithelial cells, RECs, into epidermal cells having a protective cutin coating. The development of the protective layer at the detachment site was visualized by (a) toluidine blue permeability, (b) scanning electron microscopy, (c) transmission electron microscopy, and (d) expression patterns of genes regulating epidermal cell specialization according to cell types. will be. Images were observed in transgenic plants expressing wild-type or proQRT2 :: spGFP - CDEF1 . [ P _adj < 0.05, log ₂ fold change > 2. Scale bars: 100 um (a), 5 um (b), 1 um (c)]

Unless defined otherwise, all technical and scientific terms used herein have the same meaning as commonly understood by one of ordinary skill in the art to which this invention belongs. In general, the nomenclature used herein is those well known and commonly used in the art.

In the present invention, it was confirmed that two adjacent cells have independent cell types at the detachment site of Arabidopsis , and the detachment process can be accurately induced by adjusting their activity to control cell wall processing. Specifically, one of the two adjacent cell types forms a honeycomb structure of lignin to localize cell wall disruption and serve as mechanical braces to spatially limit cell detachment, and It was confirmed that other cell types undergo transdifferentiation into epidermal cells producing a protective cutinoid coating. On the other hand, it was confirmed that the loss of the lignin brace causes a defect in the cutinoid surface having a protective action of the newly generated epidermis, and when a defect in the protective barrier occurs, the plant becomes susceptible to pathogen infection. In the present invention, it was confirmed how the above two adjacent cell types mutually regulate their physiological state and elaborate differential cell structures to induce precise detachment while maintaining the integrity of the cell surface, and through the regulation of these mechanisms It was confirmed that the detachment of plants can be artificially controlled.

In the present invention, secession site cells of calyx and separated organs are classified into residual cells (RECs, residuum cells) and secession cells (SECs, secession cells), and nlsGFP- induced by the detachment site-specific promoter Quartet2 (QRT2) - RECs and SECs were isolated from transgenic plants with GUS constructs using a fluorescence-activated cell isolation assay, and their transcripts were analyzed by RNA-sequencing. As a result, RECs and SECs showed different expression profiles. In RECs, genes related to metabolism, biosynthesis, and organ development were mainly expressed, and in SECs, genes related to stimulation response and aging were mainly expressed. .

Since RECs are non-epithelial cells in a state capable of converting into surface cells with protective functions, lignin biosynthesis genes were expected to be expressed, but rather, lignin biosynthesis enzymes and laccases involved in lignin polymerization in SECs were abundant. confirmed to be

As a result of confirming the mechanism for SECs-specific lignin formation, when salicylhydroxamic acid, a peroxidase inhibitor, was used, lignin accumulation was inhibited in SECs in the presence of monolignol and PA. In addition, diethyldithiocarbamate was used to inhibit ROS-metabolizing enzyme superoxide dismutase (SOD), and DPI (diphenyleneiodonium) was used to inhibit NADPH oxidase (NOX). , inhibited lignin formation in SECs.

H ₂ O ₂ formation in SECs is a result of the sequential reaction of NOX and SOD, and in the present invention, it was confirmed that 4 genes out of 10 genes encoding NOX protein were expressed in the desorption site of Arabidopsis thaliana, that is, RbohD and RbohF was expressed in both SECs and RECs, and RbohE and RbohH were expressed in vasculature and nectary cells. On the other hand, lignin formation disappeared in the rbohD / F mutant and was restored when RbohF was introduced again into the rbohD / F mutant, where lignin deficiency in the rbohD /F mutant delayed dissociation. These results suggest that the response by RbohD/F provides ROS for lignin polymerization in the SEC required for proper flower organ detachment.

As a result of observing the lignin structure in SECs, 2-3 layers of lignin in a hexagonal structure with pillars were identified. The lignin firmly supported each layer of the SEC, but was not observed in the upper part of the first layer of the SEC. In addition, the cell wall enzyme Pectate Lyase-Like18 (PLL18) was observed restricted to only the first layer of SECs. Thus, it can be seen that lignin blocks the diffusion of cell wall enzymes beyond the first layer of SEC, acts as a "brace" to hold the separate layers of cells together, and forms a mechanical barrier to enzyme diffusion. there was. Furthermore, when the mechanical barrier of lignin is defective, it can be seen that the uninhibited diffusion of enzymes that hydrolyze the cell wall dilutes the enzyme concentration at the dissociation site, delaying organ dissociation and also causing organ dissociation at the ectopic site. there was.

On the other hand, genes related to suberin biosynthesis were abundantly expressed in RECs, and genes encoding biosynthetic enzymes and transporters for cutin and/or wax were also expressed very abundantly. It protects plants from stress. As a result of observing the RECs at the detachment site with a transmission electron microscope, it was confirmed that the material with high electron density was located on the outside of the residual cell wall. Since it was deposited in the , it was found that the material was cutin.

In the present invention, a molecular mechanism that promotes precise processing of the cell wall, intercellular linkage, and cellular structure throughout the detachment site was identified. SEC-specific lignins aggregate individual cells into clusters for precise organ dissociation and act as mechanical scaffolds to limit cell wall enzymes to precise locations. In the process of organ dissociation, the process of improving the protective function of the permeable and fragile protective layer on the new epidermis surface, and the spatiotemporal and spatially accurate removal of the flower organ are very important for protecting the reproductive organs from environmental disasters. In addition, since manipulating the separation of periods is an early stage of crop cultivation and can contribute to the production of superior crops, the present invention may also aid in the development of strategies to improve crop and fruit production.

Accordingly, in one aspect, the present invention relates to a mutant or transgenic plant in which cleavage is inhibited or delayed.

In the present invention, the mutant or transgenic plant with suppressed or delayed cleavage may be characterized in that the gene encoding ROS-forming enzymes RbohD and/or RbohF is deleted or the expression is inhibited so that lignin production is suppressed, RbohD The gene encoding may be represented by the nucleotide sequence of SEQ ID NO: 1, and the gene encoding RbohF may be represented by the nucleotide sequence of SEQ ID NO: 2, but is not limited thereto.

In the present invention, the mutant or transgenic plant may be characterized in that the production of lignin is inhibited specifically for cells leaving the detachment site.

In the present invention, the term "plant" refers to an organism other than animals constituting the biological system, which has a cell wall and chlorophyll, so that it performs autotrophic photosynthesis. Plants that can be used in the present invention can be broad, such as classes of higher plants applicable to transformation techniques, generally including both monocots and dicotyledons, and certain lower plants, such as algae, which include polyploids, diploids and haploids. Plants of various ploidy levels, including

In the present invention, the term “mutation” refers to a change in the genetic properties of an organism due to insertion, deletion, or substitution of DNA, which is the intrinsic genetic material of an organism, and “transformation” refers to a change in the genetic material of an organism in DNA. It means that the trait of an individual or cell is genetically changed by

In the present invention, the term "plant" refers to a body of a plant type, the mutant of the present invention means a mutated plant, and the transgenic plant means a transformed plant, and the plant is the whole plant or part of a plant. , seeds or plant cells. In the present invention, the mutant is produced by selecting a naturally occurring mutation, processing Ethyl methanesulfonate (EMS) or X-ray, or using gene scissors technology, and the transgenic plant is produced by inserting T-DNA using Agrobacteria. It may be manufactured, but is not limited thereto.

In another aspect, the present invention relates to a composition for inhibiting detachment of plants.

In the present invention, the composition comprises RNAi or antisense oligonucleotides that inhibit the expression of genes involved in lignin production; Or a compound that inhibits peroxidase, superoxide dismutase (SOD), or NADPH oxidase (NOX); may be characterized as comprising.

In the present invention, the compound that inhibits peroxidase is salicylhydoxamic acid, and the compound that inhibits superoxide dismutase (SOD) is diethyldithiocarbamate (diethyldithiocarbamate), and the NADPH oxidase (NOX) inhibitory compound may be characterized as diphenylene iodonium (DPI, diphenyleneiodonium), but is not limited thereto.

In the present invention, the RNAi, antisense oligonucleotide, or the compound is characterized in that it inhibits the detachment of the plant by inhibiting the production of lignin specifically in the leaving cell at the detachment site, and thus, ROS for lignin polymerization in SECs. Those skilled in the art will be able to use the inhibitory or inhibitory substances with appropriate modifications.

In another aspect, the present invention relates to a method of inhibiting detachment of a plant or inhibiting falling leaves or falling fruits.

The method comprises the steps of (a) deleting or inhibiting the expression of a gene encoding ROS-forming enzymes RbohD and/or RbohF; And (b) it may include the step of selecting a plant in which lignin production is suppressed specifically for cells leaving the detachment site.

In the present invention, the gene encoding RbohD may be represented by the nucleotide sequence of SEQ ID NO: 1, and the gene encoding RbohF may be represented by the nucleotide sequence of SEQ ID NO: 2, but is not limited thereto.

In another aspect, the present invention relates to a method for promoting detachment of a plant.

The method comprises the steps of: (a) enhancing the expression of one or more enzymes selected from the group consisting of peroxidase, superoxide dismutase (SOD) and NADPH oxidase (NOX); And (b) it may include the step of selecting a plant in which the production of lignin is enhanced specifically for cells leaving the detachment site.

In the present invention, the expression of the enzyme may be characterized in that the gene encoding the enzyme is introduced into the plant by T-DNA insertion mutation to promote it, but is not limited thereto.

In the present invention, the gene encoding the peroxidase is selected from the group consisting of SEQ ID NOs: 3 to 11, and the gene encoding the superoxide dismutase is selected from the group consisting of SEQ ID NOs: 12 to 18, the NADPH The gene encoding oxidase may be characterized as represented by SEQ ID NO: 1 or SEQ ID NO: 2, but is not limited thereto.

Hereinafter, the present invention will be described in more detail through examples. These examples are only for illustrating the present invention, and it will be apparent to those of ordinary skill in the art that the scope of the present invention is not to be construed as being limited by these examples.

Example 1. Experimental method

1-1. Plants and growing conditions

Arabidopsis, Arabidopsis The accession Col - 0 was used as a wild type, and the rbohD /F double mutation was obtained from the Arabidopsis Biological Resource Center, a T-DNA insertion line, SALK_070610 ( rbohD , AT5G47910), SALK_059888 ( rbohF , AT1G64060) Produced by crossbreeding. Arabidopsis thaliana was grown with a photoperiod (110 μmol/m ² s) of 16/8 hours (light/dark) in soil in a growth chamber at 22°C±2°C.

1-2. transgenic plants

The plasmid was prepared using Gateway Cloning Technology (Invitrogen), and then introduced into plants using the floral dipping method (see Non-Patent Document 1). The primer sequence for constructing the promoter::GFP fusion is as follows.

AGI Number Gene Name Promoter Size Primer Name Primer Sequence AT3G07970
QRT2
2297
prQRT2-attB4 GTATAGAAAAGTTGCTTAGGGATGCTAGAGAGGAATCTCG prQRT2-attB1r TTTGTACAAACTTGCTTTTTTTTGTTGGAGTGGGACTTC
At4g34230 CAD5
2400
prAtCAD5-attB4 GTATAGAAAAGTTGCTGGCATTGTGATTCACTGG prAtCAD5-attB1r TTTGTACAAACTTGCTTTTTGATGATTCTTTCTTCTT At3g09220
LAC7
2017
prLAC7-attB4 GTATAGAAAAGTTGCTAGAAAGTATGCAAGATCCTGGAGTT prLAC7-attB1r GTATAGAAAAGTTGCTAGAAAGTATGCAAGATCCTGGAGTT At5g48100
LAC15
3006
prAtLAC15-attB4 GTATAGAAAAGTTGCTCGTATGCATGTTTCAAATT prAtLAC15-attB1r TTTGTACAAACTTGCTTTGGAAGAGTTTTAGTAAATT AT1G35140
EXL1
2581
prEXL1-attB4 GTATAGAAAAGTTGCTAGTAGTTGGAGAAAATTGG prEXL1-attB1r TTTGTACAAACTTGCTTAATTTCTGTAATGTAGAAGC AT4G16190
RDL2
2957
prRDL2-attB4 GTATAGAAAAGTTGCTGATGAGTTTACCTGCCTGATGCAAG prRDL2-attB1r TTTGTACAAACTTGCGCTTCTTGGTTTCCAAGAGATAAGGAAG AT3G55090
ABCG16
1799
prABCG16-attB4 GTATAGAAAAGTTGCTACGCCATAATGTAAATTTTTCCGAAAC prABCG16 attB1r TTTGTACAAACTTGCTTAGAACAAAAGATGGTAAAGTGAGAAAT AT5G54230
MYB49
2624
prMyb49-attB4 GTATAGAAAAGTTGCTTATGTTGCTAGTGCTTGCTACCACACTT prMyb49-attB1r TTTGTACAAACTTGCCTTTTTCAAATGTTATATGTG AT4G28490
HAESA
1597
prHAESA-attB4 GTATAGAAAAGTTGCTCAGAAAGGATTAAACTAAAACTCTGTCA prHAESA-attB1r TTTGTACAAACTTGCTTTTTTGGAAAAGGAATCGTTATTC AT3G24670
PLL22
2529
prPLL22-attB4 GTATAGAAAAGTTGCTTCTTTTGGGGAATTGTTGTTTTGCC prPLL22-attB1r TTTGTACAAACTTGCTGTTGGATATATCAAAGCTCTCTCTCCTATT

AGI Number Gene Name Promoter Size Primer Name Primer Sequence AT3G48920
MYB45
2977
prMyb45-attB4 GTATAGAAAAGTTGCTATTTAATAACTTTTCCTGTGTA prMyb45-attB1r TTTGTACAAACTTGCTTTTATTTTTCCAAAGTTTT AT1G68765
IDA
1426
prIDA-attB4 GTATAGAAAAGTTGCTTTTTCAATTTTGTTATTGCATCTTAT prIDA-attB1r TTTGTACAAACTTGCTTGGTAGTCAATGTTTTTTTTCTTC AT2G37130
PER21
2858
prPER21-attB4 GTATAGAAAAGTTGCTAGGTATGTATTACTCTTGAAAACACACA prPER21-attB1r TTTGTACAAACTTGCTGCTCTCTCTCTCTCTATTCTCTCTTAT AT4G25810
XTR6
2318
prXTR6 attB4 GTATAGAAAAGTTGCTCATAAGAGGCTAACCGC prXTR6 attB1r TTTGTACAAACTTGCTGCTAAGTGATTGTTTCTT AT2G39350
ABCG1
3085
prABCG1 attB4 GTATAGAAAAGTTGCTCAGTTAGAAGTTAGTAGGCAAAAAGTCA prABCG1 attB1r TTTGTACAAACTTGCAGATACAATAGAAAAACTTTTAAAGAGTGA AT5G05340
PER52
2641
prPER52-attB4 GTATAGAAAAGTTGCTGACTAAGATACGTCACACGTGGAT prPER52-attB1r TTTGTACAAACTTGCTGTTAGGTTAAAACAAAGACTTGAATATA AT1G71380
CEL3 2907
prCEL3-attB4 GTATAGAAAAGTTGCTAGCCAAGATGTATTAAACACCACC prCEL3-attB1r TTTGTACAAACTTGCCTTGAAGACAGAGAGCTAAATTGCTTT AT2G47190
MYB2
2546
prMYB2-attB4 GTATAGAAAAGTTGCTTTTCCTATTCTCGCCAAGCAGT prMYB2-attB1r TTTGTACAAACTTGCTTCTTATAAAGAAACTTTAAATGAGG AT4G37520 PER50 2849 prPER50-attB4 GTATAGAAAAGTTGCTTGCTTCTTTAGACTGTTACTATCAAACAAAAAT prPER50-attB1r TTTGTACAAACTTGCTTAAAAAAAAATATTTTCACTCAATCTCTCTA AT1G62770
PMEi-3
2506
prPMEi-3-attB4 GTATAGAAAAGTTGCTATGAGTCTTCGAAAGATTTC prPMEi-3-attB1r TTTGTACAAACTTGCGGTGTTTTGGTTTGTTTG

Final Construct AGI Number Primer Name Primer Sequence prQRT2::
QRT2-mCherry
AT3G07970
QRT2-attB1 GTACAAAAAAGCAGGCTTAACCATGTATGAAAAGATCATAATCTTA QRT2-attB2 GTACAAGAAAGCTGGGTAAGTGCAAAAGAGGAGAAACAT prQRT2::
XTH28-mCherry
At1g14720
XTH28-attB1 GTACAAAAAAGCAGGCTTAACCATGGGTTTTATAACTCGATTTTTAGTTT XTH28-attB2 GTACAAGAAAGCTGGGTATATCGACTCAGTTCGAGCTAATCGGCTC pH7_prQRT2::
PLL18-mCherry
At3g27400
PLL18-attB1 GTACAAAAAAGCAGGCTTAACCATGGTTTCGTATTCTAATAATCATTTTGCATACGC PLL18-attB2 GTACAAGAAAGCTGGGTAACAGCGACGGCCTCGCCGAC pH7_prQRT2::
Pectinase-mCherry
at4g18180
Pectina-attB1 GTACAAAAAAGCAGGCTTAACCATGGAGAATTTTCGTTACCTGAAATTT Pectina-attB2 GTACAAGAAAGCTGGGTAGTGACATGGAGGTGGGATTTGAGTCC

1-3. Microscopic and histological analysis

Low magnification images were obtained using a Zeiss Discovery V12 stereo microscope. Confocal scanning laser microscopy analysis was performed with excitation and detection settings as follows: GFP 488 nm, 500-545 nm; Cherry 594 nm, 570-670 nm. For scanning electron microscopy of the detachment site in the flower organ, sepals and petals were removed before fixation, and critical point drying was performed using EM CPD 300 (Leica), and the tissue was placed on a steel stub and coated with osmium. After that, it was observed with a Hitachi S-4800 (Tokyo, Japan). For transmission electron microscopy, the tissue was prepared by Choi. et al. (An ABCG/WBC-type ABC transporter is essential for transport of sporopollenin precursors for exine formation in developing pollen. Plant J. 65, 181-193 (2011)) treated as described in the literature, and JEM- Analysis was performed using a 100CX-1 transmission electron microscope (JEOL, Japan). Tissue sections for optical microscopy were prepared using Technovit 7100 resin (Heraeus Kulzer). Embedded samples were cut into 2-μm sections using an RM2255 microtome (Leica) equipped with a TC65 tungsten carbide disposable blade (Leica) and observed using a Zeiss Axiovert 40 inverted microscope. To visualize the peroxide and H ₂ O ₂ accumulation, pollen was stained with nitroblue tetrazolium (NBT) and 3,3-diaminobenzidine (DAB), respectively. Briefly, inflorescences were vacuum infiltrated with NBT solution (0.05% in PBS) or DAB solution (0.1% in distilled water, pH 5.8, adjusted with KOH) and incubated in NBT and DAB for 3.5 h in the dark for 30 min. processed. After staining, inflorescences were fixed and dechlorinated in 3:1:1 ethanol:lactic acid:glycerol (bleach) at 70°C until complete removal of chlorophyll, followed by visualization using a Zeiss Discovery V12 stereo microscope.

For permeability analysis, flowers were treated with a solution (0.025 %) of toluidine blue O (Fisher Scientific) in distilled water for 5 min and washed with distilled water for 2 min. Images were imaged using an AxioCam HRC camera installed on a Zeiss Discovery V12 stereo microscope.

1-4. Observation of lignin structure

After flower removal, lignin autofluorescence was performed with some modifications as indicated in the literature of Malamy, JE & Benfey, P. N (Organization and cell differentiation in lateral roots of Arabidopsis thaliana. Development 124, 33-44 (1997)) as standard. Detection was performed using the GFP filter setting, with the reaction time increased to a maximum of 40 min at 57°C. When lignin was visualized with mCherry-labeled cell wall protein, 0.1% (w/v) berberine hemisulphate (Sigma) was used for lignin staining (Kurihara, D., Mizuta, Y., Sato, Y. & Higashiyama). , T. ClearSee: a rapid optical clearing reagent for whole-plant fluorescence imaging. Development 142, 4168-4179 (2015)). Alternatively, lignin was stained with phloroglucinol-HCl [1% (w/v) phloroglucinol in 6N HCl] and observed under a dissecting microscope.

In order to visualize lignin at the detachment site of the sepal, a conventionally known method for enzymatic degradation of plant cell walls was used (Schreiber, L. Chemical composition of Casparian strips isolated from Clivia miniata Reg roots: Evidence for lignin. Planta 199, 596). -601 (1996)) was used with some modifications, and the method was partially modified to be suitable for Arabidopsis inflorescences. In summary, carbon adhesive tape was attached to the sepal of stage 16 and reacted by lightly stirring at 60 ° C with an enzyme solution ((0.5% macerozyme, 0.5% cellulose RS, 200 μM cycloheximide in sodium acetate buffer, pH 5.0), The enzyme solution was replaced every 3 days, and after 2-3 weeks, the lignin structure of the sepal detachment site without other cellular structures was visualized with a scanning electron microscope.

1-5. pharmacological test

For the analysis of inhibition of lignin biosynthesis using piperonylic acid (PA), a partially modified excised pistil-feeding assay was performed. Briefly, flower buds were cut under the peduncle and seeded in a solidification medium containing 5 mM CaCl ₂ , 5 mM KCl, 1.6 mM HBO ₃ , and 5% sucrose (pH 7.5) in 24-well plates. After incubation until reaching floral stage 16 in a growth chamber for 3 days with or without 150 μM PA, respectively, monolignol (coniferyl alcohol and cyanyl alcohol in PBS, 30 μM each) was added with 100 μM diphenyleneiodonium (DPI) , was treated with 250 μM DDC (diethyldithiocarbamic acid) or 150 μM SHAM (salicylhydroxamic acid) for 6 hours.

For the ROS inhibition assay, calyx and petals in stage 16 were pre-incubated in PBS (pH 7.4) for 15 minutes, and treated with 125 μM DDC, 20 μM DPI or 500 μM SHAM at room temperature for 2 hours. Next, 10 μM 2',7'-DCF-DA (2',7'-dichlorofluorescin diacetate, Sigma-Aldrich) was added and incubated in the dark for 20 minutes. Samples were washed 3 times with PBS to remove excess DCF-DA and immediately visualized using an Olympus FluoView FV1200 confocal laser scanning microscope. H ₂ O ₂ production was quantified using Image J software.

1-6. Fluorescence-activated cell sorting and RNA extraction

Arabidopsis protoplasts were prepared by partially modifying the method of Bargmann, BO & Birnbaum, KD (Fluorescence activated cell sorting of plant protoplasts. J. Vis . Exp . 36 , 1673, doi:10.3791/1673 (2010)). For this purpose, flower organs including receptacles, stamens, sepals, and petals from about 300 plants were collected from flowers of stage 16, a stage in which each flower organ is attached but can be easily separated. did

Plant material in enzyme solution (1.2% Cellulose R-10, 0.5% Maceizayrin R-10, 400 mM mannitol, 0.1 % BSA, 10 mM CaCl ₂ , 4 mM MES, 20 mM KCl, 2.5 mM beta-mercaptoethanol, 40-U/ ml RNase inhibitor, and 10ug/ml actinomycin D, pH 5.7), vacuum infiltrated at 100mmHg for 30 minutes, and incubated with stirring (80-100 rpm) for 4 hours in the dark at 22°C. Protoplasts were diluted with an equal volume of cold W5 solution (1.154 mM NaCl, 125 mM CaCl ₂ , 5 mM KCl and 2 mM MES, pH 5.7), filtered through 2 sheets of 20 μm nylon mesh, and centrifuged at 1500 rpm for 5 min. and concentrated. The supernatant was aspirated and the pellet was dissolved in 1 ml of W5 solution. Protoplasts were sorted using a FACSAria III cell sorter (BD Biosciences) equipped with a 100 μm nozzle at a speed of 2,000 μm with a system pressure of up to 20 psi. Approximately 3,000 GFP-positive cells per sample were collected in 500 μL RLT buffer (Qiagen, RNeasy Micro Kit). Total RNA of protoplasts was extracted using the RNeasy™ micro kit (QIAGEN) according to the manufacturer's method. RNA was eluted in 14 μl distilled water, and RNA quality was checked using a Bioanalyzer Pico chip (Agilent Technologies).

1-7. RNA sequencing

Using SMART-Seq V4 Ultra Low Input RNA Kit (Clontech), cDNA was generated in 18 amplification cycles, and RNA-seq library was generated using TruSeq RNA Sample Prep Kit v2 (Illumina) according to the manufacturer's method. Libraries were qualitatively and quantitatively validated and then loaded into a flow cell for cluster generation using cBot and TruSeq PE Cluster Kit v3 (Illumina). Sequencing was performed on a HiSeq2000 using TruSeq SBS Kit v3 (pair-ended 200 cycles, Illumina) to generate 100-bp pair-ended reads. Illumina HiSeq generated raw images using HiSeq Control Software v2.2 for system control, and generated base calling using Real Time Analysis v1.18-integrated primary analysis software. Base call (BCL) binary files were converted to FASTQ format using Illumina package bcl2fastq (v1.8.4). Library preparation and RNA sequencing were performed by Macrogen Inc. (Korea).

1-8. RNA- seq analysis

Quality control and preprocessing of metagene data was performed using FastQC (v0.11.2; http://www.bioinformatics.babraham.ac.uk/projects/fastqc/). Trimomatic (v0.32.3) 37 was used to filter out adapters and low-quality sequences, and alignment was performed using Arabidopsis TAIR10 genome (ENSEMBL annotation) and TopHat 2.0.9 (ref38). PCR duplication was eliminated using Picard (v1.119, http://broadinstitute.github.io/picard/), and only mapped sequences were calculated using HTSeq v0.6.1. Differential gene expression analysis was performed using DESeq2. P values were adjusted through the Benjamini-Hochberg method, and differentially expressed genes were considered significant when the false discovery rate (FDR) was less than 0.05.

1-9. Gene Ontology (GO) Analysis

Gene ontology analysis was performed by inputting Arabidopsis genome (TAIR10 assembly) into Singular Enrichment Analysis of AgriGo. Hypergeometric tests were performed with Yekutieli as a multi-test adjustment method, and P values were adjusted using basic parameters. Gene ontology terms with FDR <0.05 were considered significantly enriched. In order to summarize and visualize the gene ontology terms, the results were applied as a treemap of REVIGO. Using the SimRel semantic similarity scale, terms were clustered with a specific similarity cutoff of 0.4 (“small REVIGO set”), and manually corrected to clarify the meaning of representative terms. The tree map shown in Fig. 1c was generated using modified representative terms.

1-10. Pathogen infection analysis

Pseudomonas syringae pv. Tomato DC3000 (Pst DC3000) was incubated overnight at 28 °C in King's B medium containing rifampicin. Bacteria were collected, washed, and diluted to an OD ₆₀₀ of 0.5. For infection analysis, flowers in stages 16 and 17 were immersed in a suspension of Pst DC3000 (OD ₆₀₀ = 0.5 × 10 ^{- 9} ) for 10 min and washed 3 times with distilled water. Bacterial colony forming units (cfu) were scored in zones 1 and 2 of flower pieces (Fig. 8g) rinsed for 1 h. After detachment, 10 flowers were used in each experiment 3 times, and 4 independent experiments were performed. Statistical analysis was performed by one-way analysis of variance (ANOVA) using Tukey's Honestly Significant Difference procedure in the SAS statistical software package (SAS, Inc., Cary, NC, v9.3).

1-11. genetic information

RbohD - AT5G47910 (SEQ ID NO: 1)

1 ATGAAAATGA GACGAGGCAA TTCAAGTAAC GACCATGAAC TTGGGATTCT

51 ACGAGGAGCT AACTCGGACA CCAACTCGGA CACGGAGAGC ATCGCTAGCG

101 ACCGTGGTGC CTTTAGCGGT CCGCTTGGCC GGCCTAAACG TGCGTCCAAG

151 AAAAACGCAA GATTCGCCGA CGATCTTCCC AAGAGAAGCA ATAGTGTTGC

201 TGGCGGCCGT GGTGATGACG ATGAGTACGT GGAGATCACG CTAGACATCA

251 GGGACGACTC GGTGGCCGTC CATAGTGTCC AACAAGCAGC TGGAGGTGGA

301 GGCCACCTGG AGGACCCGGA GCTAGCCCTT CTTACGAAGA AGACTCTCGA

351 GAGCAGCCTC AACAACACCA CCTCCTTATC TTTCTTCCGA AGCACCTCCT

401 CACGCATCAA GAACGCCTCC CGCGAGCTCC GCCGCGTGTT CTCTAGACGT

451 CCCTCCCCGG CCGTGCGGCG GTTTGACCGC ACGAGCTCCG CGGCCATCCA

501 CGCACTCAAA GGTCTCAAGT TCATTGCCAC CAAGACGGCC GCATGGCCGG

551 CCGTCGACCA ACGTTTCGAT AAACTCTCCG CTGATTCCAA CGGCCTCTTA

601 CTCTCTGCCA AGTTTTGGGA ATGCTTAGGA ATGAATAAGG AATCTAAAGA

651 CTTCGCTGAC CAGCTCTTTA GAGCATTAGC TCGCCGGAAT AACGTCTCCG

701 GCGATGCAAT CACAAAGGAA CAGCTTAGGA TATTCTGGGA ACAGATCTCA

751 GACGAAAGCT TTGATGCCAA ACTCCAAGTC TTTTTTGACA TGGTGGACAA

801 AGATGAAGAT GGGCGAGTAA CAGAAGAAGA GGTGGCTGAG ATTATTAGTC

851 TTAGTGCTTC TGCAAACAAG CTCTCAAATA TTCAAAAGCA AGCCAAAGAA

901 TATGCGGCAC TGATAATGGA AGAGTTGGAC CCAGACAATG CTGGGTTTAT

951 TATGATCGAA AACTTGGAAA TGTTGCTATT ACAAGCACCA AACCAGTCGG

1001 TGCGGATGGG AGACAGCAGG ATACTTAGTC AGATGTTAAG TCAGAAGCTT

1051 AGACCGGCAA AAGAGAGCAA CCCTTTAGTG AGATGGTCGG AGAAAATCAA

1101 ATATTTCATA CTTGACAATT GGCAGAGACT ATGGATAATG ATGTTATGGC

1151 TTGGCATCTG TGGTGGCCTC TTTACTTATA AATTCATTCA GTACAAGAAC

1201 AAAGCTGCCT ATGGTGTCAT GGGTTATTGC GTTTGTGTCG CCAAAGGAGG

1251 CGCCGAGACT CTCAAATTCA ACATGGCTCT CATATTGTTG CCTGTTTGTC

1301 GAAACACCAT CACTTGGCTT AGGAACAAGA CTAAGCTTGG TACTGTCGTT

1351 CCTTTTGATG ATAGTCTTAA CTTCCACAAG GTTATTGCAA GCGGGATAGT

1401 CGTCGGTGTT TTACTCCATG CGGGTGCCCA TTTAACGTGT GATTTTCCAC

1451 GTTTAATTGC CGCGGATGAG GACACCTATG AGCCGATGGA AAAATACTTT

1501 GGGGATCAAC CGACTAGCTA CTGGTGGTTT GTGAAAGGAG TGGAAGGATG

1551 GACTGGCATT GTGATGGTTG TGCTAATGGC TATAGCCTTT ACACTCGCGA

1601 CGCCTTGGTT CCGACGTAAC AAGCTTAACT TACCTAACTT CCTCAAGAAG

1651 CTTACCGGTT TCAACGCCTT TTGGTACACC CACCATTTGT TCATCATTGT

1701 TTATGCTCTT CTCATTGTCC ATGGTATCAA GCTCTACCTC ACAAAGATTT

1751 GGTATCAGAA AACGACATGG ATGTATCTTG CTGTACCCAT CCTTCTATAT

1801 GCATCGGAGA GGCTGCTCCG TGCTTTCAGA TCAAGCATCA AACCGGTTAA

1851 GATGATCAAG GTGGCTGTTT ACCCCGGGAA CGTGTTGTCT CTACACATGA

1901 CGAAGCCACA AGGATTCAAA TACAAAAGTG GACAGTTCAT GTTGGTGAAC

1951 TGCCGAGCCG TATCTCCATT CGAATGGCAT CCTTTCTCAA TCACATCAGC

2001 TCCCGGAGAC GATTACCTGA GCGTACATAT CCGCACTCTC GGTGACTGGA

2051 CACGTAAGCT CAGGACCGTT TTCTCCGAGG TTTGCAAACC TCCTACCGCC

2101 GGTAAAAGCG GTCTTCTCCG AGCAGACGGA GGAGATGGAA ACCTCCCGTT

2151 CCCGAAGGTC CTTATCGACG GTCCATACGG TGCTCCCGCA CAAGACTACA

2201 AGAAATACGA CGTGGTACTC CTCGTAGGTC TCGGCATTGG AGCCACGCCT

2251 ATGATCAGTA TCCTTAAGGA CATCATCAAC AACATGAAAG GTCCTGACCG

2301 CGACAGCGAC ATTGAGAACA ATAACAGTAA CAACAATAGT AAAGGGTTTA

2351 AGACAAGGAA AGCTTATTTC TACTGGGTGA CTAGGGAACA AGGATCATTC

2401 GAGTGGTTCA AGGGAATAAT GGACGAGATT TCGGAGTTAG ACGAGGAAGG

2451 AATCATCGAG CTTCACAATT ATTGCACGAG TGTGTACGAG GAAGGTGATG

2501 CAAGAGTGGC TCTCATTGCC ATGCTTCAGT CGTTGCAACA CGCTAAGAAC

2551 GGTGTGGATG TTGTGTCGGG TACACGTGTC AAGTCCCACT TCGCTAAACC

2601 TAACTGGAGA CAAGTCTACA AGAAGATCGC TGTTCAACAT CCCGGCAAAA

2651 GAATAGGAGT CTTCTACTGT GGAATGCCAG GAATGATAAA GGAATTAAAA

2701 AATCTAGCTT TGGATTTTTC TCGAAAGACA ACTACCAAGT TTGACTTCCA

2751 CAAAGAGAAC TTCTAG

RbohF - at1g64060 (SEQ ID NO: 2)

1 ATGAAACCGT TCTCAAAGAA CGATCGGCGA CGGTGGTCAT TTGATTCAGT

51 TTCCGCCGGA AAAACCGCCG TCGGAAGTGC ATCAACTTCA CCGGGAACTG

101 AATACTCCAT TAACGGTGAT CAAGAGTTCG TTGAAGTCAC AATCGATCTT

151 CAAGACGATG ACACAATCGT TCTTCGTAGC GTCGAGCCAG CAACCGCCAT

201 TAATGTCATC GGAGATATCT CCGACGACAA CACCGGAATA ATGACTCCGG

251 TTTCGATTTC GAGATCTCCG ACGATGAAAC GAACTTCATC TAATCGGTTC

301 CGACAATTCT CACAAGAGCT TAAAGCCGAA GCTGTGGCGA AAGCGAAACA

351 GTTATCTCAG GAGTTGAAAC GATTCTCATG GTCTCGTTCT TTCTCAGGTA

401 ACTTAACCAC TACTAGTACC GCCGCTAATC AAAGCGGCGG TGCTGGTGGT

451 GGTTTGGTGA ACTCGGCTTT AGAAGCGCGA GCGTTGCGAA AGCAACGTGC

501 TCAGTTAGAT CGGACTCGGT CTAGTGCTCA AAGAGCTCTT CGTGGTTTGA

551 GATTCATTAG CAATAAGCAA AAGAACGTTG ATGGTTGGAA CGATGTTCAA

601 TCAAATTTCG AAAAATTCGA AAAAAATGGT TACATCTATC GCTCCGATTT

651 CGCTCAATGC ATAGGAATGA AAGATTCGAA AGATTTGCA TTGGAACTGT

701 TCGATGCATT GAGTAGAAGA AGAAGATTAA AAGTAGAGAA AATCAATCAC

751 GATGAGCTTT ATGAGTATTG GTCACAAATC AACGACGAGA GTTTTGATTC

801 TCGTCTCCAG ATCTTCTTCG ACATAGTGGA CAAGAATGAA GATGGGAGAA

851 TTACAGAAGA GGAAGTAAAA GAGATAATAA TGTTGAGTGC ATCTGCAAAT

901 AAGCTATCAA GATTAAAGGA ACAAGCAGAG GAATATGCAG CTTTGATTAT

951 GGAAGAGTTA GATCCTGAAA GACTTGGCTA CATAGAGCTA TGGCAACTAG

1001 AGACTTTGCT TCTACAAAAA GACACATACC TCAATTACAG TCAAGCATTG

1051 AGCTATACGA GCCAAGCATT GAGCCAAAAC CTTCAAGGGT TAAGGGGAAA

1101 GAGTCGAATA CATAGAATGA GTTCGGATTT CGTCTACATT ATGCAAGAGA

1151 ATTGGAAAAG GATATGGGTT TTATCCTTAT GGATCATGAT CATGATCGGA

1201 TTATTCTTGT GGAAATTCTT CCAATACAAG CAAAAAGATG CATTTCATGT

1251 GATGGGATAT TGTTTACTCA CAGCCAAAGG AGCAGCTGAA ACACTTAAAT

1301 TCAACATGGC TCTAATACTT TTCCCAGTTT GCAGAAACAC CATTACTTGG

1351 CTTAGATCCA CAAGACTCTC TTACTTCGTT CCTTTTGATG ATAATATCAA

1401 CTTCCACAAG ACAATTGCTG GAGCCATTGT AGTAGCTGTG ATCCTTCATA

1451 TTGGAGACCA TCTTGCTTGT GATTTCCCTA GAATTGTTAG AGCCACCGAA

1501 TACGATTACA ATCGGTATCT GTTTCATTAC TTTCAAACAA AACAGCCAAC

1551 ATACTTCGAC CTCGTTAAGG GACCTGAAGG AATCACTGGG ATTTTAATGG

1601 TCATTTTGAT GATTATTTCA TTCACATTAG CAACAAGATG GTTTAGGCGT

1651 AACCTAGTCA AGCTTCCTAA GCCATTTGAT CGACTAACCG GTTTTAACGC

1701 CTTTTGGTAT TCGCATCATT TGTTCGTCAT TGTTTATATC TTGCTTATTC

1751 TTCATGGTAT CTTCCTCTAT TTCGCCAAGC CTTGGTATGT TCGTACGACA

1801 TGGATGTATC TTGCAGTACC AGTTTTACTC TATGGTGGAG AAAGAACACT

1851 TAGGTACTTC CGTTCTGGTT CTTATTCGGT TCGACTGCTT AAGGTTGCTA

1901 TATATCCTGG TAATGTTCTA ACGCTACAAA TGTCGAAACC AACTCAATTT

1951 CGTTACAAAA GCGGACAATA CATGTTTGTC CAATGTCCTG CGGTTTCGCC

2001 ATTCGAGTGG CATCCATTCT CAATTACTTC CGCACCTGAA GATGATTATA

2051 TCAGCATTCA CATTAGACAA CTTGGTGATT GGACTCAAGA ACTCAAAAGA

2101 GTATTCTCTG AAGTTTGTGA GCCACCGGTT GGCGGTAAAA GCGGACTTCT

2151 CAGAGCCGAC GAAACAACAA AGAAAAGTTT GCCAAAGCTA TTGATAGATG

2201 GACCGTACGG TGCACCAGCA CAAGATTATA GGAAATATGA TGTTCTCTTA

2251 TTAGTTGGTC TTGGCATTGG TGCAACTCCA TTTATCAGTA TCTTGAAAGA

2301 TTTGCTTAAC AACATTGTTA AAATGGAAGA GCATGCGGAT TCGATCTCGG

2351 ATTTCAGTAG ATCATCAGAA TACAGCACAG GAAGCAACGG TGACACGCCA

2401 AGACGAAAGA GAATACTAAA AACCACAAAT GCTTATTTCT ACTGGGTCAC

2451 AAGAGAACAA GGCTCTTTTG ATTGGTTCAA AGGTGTCATG AACGAAGTTG

2501 CAGAACTTGA CCAACGGGGT GTGATAGAGA TGCATAACTA TTTAACAAGT

2551 GTGTATGAAG AAGGTGATGC TCGTTCTGCT CTCATTACAA TGGTTCAAGC

2601 TCTTAATCAT GCCAAAAATG GTGTCGACAT TGTCTCTGGC ACTAGGGTCA

2651 GAACACACTT TGCAAGACCT AATTGGAAGA AGGTTCTCAC AAAGCTAAGT

2701 TCCAAGCATT GCAATGCAAG AATAGGAGTG TTTTATTGCG GAGTACCGGT

2751 TTTAGGGAAG GAGCTTAGCA AACTATGCAA CACATTCAAT CAAAAAGGTT

2801 CAACCAAGTT TGAATTTCAC AAGGAGCATT TCTAA

## Peroxidase ( 73 genes A gene group that is frequently expressed in the SEC, Figure 5 reference)

PER03 - at1g05260 (SEQ ID NO: 3)

1 ATGAATTGCT TGATAGCTAT AGCTCTTTCA GTCTCTTTCT TTCTTGTGGG

51 AATTGTTGGA CCGATCCAAG CTCAATTGCA GATGAATTTC TATGCCAATT

101 CTTGTCCTAA TGCTGAAAAG ATTGTTCAAG ATTTTGTTTC AAACCACGTT

151 TCTAATGCTC CTTCTCTTGC TGCTGCTCTC ATTAGAATGC ATTTCCATGA

201 CTGTTTTGTC CGAGGTTGTG ATGGATCAGT GCTTATAAAC TCAACGTCAG

251 GAAACGCAGA GAGAGACGCG ACTCCTAACC TAACGGTTCG AGGGTTTGGC

301 TTCATCGACG CAATTAAATC TGTGCTTGAA GCTCAATGCC CTGGAATTGT

351 CTCTTGCGCT GATATTATCG CTCTAGCTTC TCGCGACGCT GTCGTTTTCA

401 CCGGAGGACC GAATTGGAGT GTACCGACCG GAAGAAGAGA CGGGAGGATA

451 TCAAACGCAG CGGAGGCATT AGCCAACATT CCTCCTCCAA CCAGTAATAT

501 CACCAATCTT CAGACACTCT TTGCAAACCA AGGACTTGAT CTTAAGGACC

551 TCGTTTTACT ATCCGGGGCT CACACTATTG GTGTATCTCA CTGCTCGTCT

601 TTCACAAACC GTCTCTACAA TTTTACGGGT CGTGGAGGCC AAGATCCGGC

651 TTTGGACAGC GAGTACGCAG CCAATCTCAA GTCTAGAAAA TGTCCTAGCC

701 TCAACGATAA CAAGACCATC GTAGAGATGG ATCCAGGGAG CCGCAAAACA

751 TTTGATCTAA GTTATTACCA GCTCGTACTC AAGCGTAGAG GTCTGTTTCA

801 ATCAGACTCT GCTCTCACCA CTAACCCCAC AACACTTTCA AACATAAACC

851 GGATCTTGAC GGGTTCGGTG GGGAGTTTCT TCTCTGAGTT TGCCAAGTCA

901 ATGGAGAAAA TGGGTCGGAT CAATGTCAAG ACTGGTTCAG CTGGAGTGGT

951 TAGGAGGCAA TGTTCCGTTG CAAATAGTTA A

PER09 - at1g44970 (SEQ ID NO: 4)

1 ATGGCTATCT CAAAGCTTAT TCCTACTCTT GTGCTCTTTG TTTTGTTCTC

51 TTTTGATGTT AGTGTTGCTC ATCCAGGTCT AGGATTTGGA TGGGGAAGTA

101 ATAGTCCCAT TGGAGGATCT TTTTACTCAA ATCTTTACCC ACAATTTTAC

151 CAGTTCTCTT GTCCACAAGC TGATGAGATT GTTATGACGG TGCTCGAAAA

201 AGCCATAGCT AAAGAACCAA GAATGGCAGC ATCTTTACTC AGACTTCACT

251 TCCACGACTG CTTCGTTCAG GGCTGTGATG CATCAATCTT GTTGGATGAT

301 AGTGCAACCA TAAGAAGTGA AAAGAATGCT GGACCAAACA AGAACTCCGT

351 TAGAGGGTTT CAAGTAATCG ACGAGATCAA AGCCAAACTT GAGCAAGCTT

401 GTCCTCAAAC TGTCTCCTGC GCCGACATTC TAGCTCTAGC TGCTCGTGGC

451 TCAACTATAC TGAGTGGTGG ACCATGG GAGCTTCCAC TAGGGAGGAG

501 AGACTCGAGG ACCGCTAGCC TTAATGGCGC AAACACGAAC ATTCCTGCAC

551 CTAACTCAAC TATTCAAAAT CTCTTAACCA TGTTCCAACG TAAAGGTTTG

601 AACGAAGAAG ACCTTGTTTC CCTATCAGGA GGACATACGA TTGGAGTAGC

651 GAGGTGCACA ACGTTTAAAC AAAGACTATA CAATCAAAAC GGTAATAACC

701 AACCAGACGA GACGCTAGAG AGGTCTTACT ACTATGGTCT CAGGTCAATT

751 TGTCCACCAA CAGGCGGTGA CAACAATATT TCACCTCTAG ACTTAGCCTC

801 TCCTGCAAGA TTCGACAACA CATACTTCAA GCTCCTTCTA TGGGGCAAAG

851 GCTTATTGAC ATCAGATGAA GTGCTTCTCA CTGGCAATGT AGGCAAGACT

901 GGTGCATTAG TGAAAGCTTA TGCCGAAGAC GAGAGACTTT TCTTCCAACA

951 ATTCGCAAAG TCGATGGTTA ACATGGGAAA CATTCAGCCT CTTACTGGTT

1001 TTAATGGTGA GATCAGGAAG AGTTGTCATG TGATTAACTA A

PER22 - at2g38380 (SEQ ID NO: 5)

1 ATGGGGTTTT CTCCTTCATT TTCTTGCAGT GCTATTGGAG CCCTAATTTT

51 GGGTTGCCTT CTGCTTCAAG CATCAAACTC TAATGCTCAG TTGAGGCCTG

101 ATTTCTACTT CGGGACGTGC CCATTTGTTT TTGATATCAT TGGAAATATC

151 ATCGTCGATG AACTGCAGAC TGATCCTCGT ATTGCCGCGA GCCTCCTTCG

201 TCTTCACTTT CACGACTGCT TTGTTCGTGG TTGTGATGCA TCGATCTTGC

251 TTGACAATTC CACATCATTC CGGACTGAGA AAGATGCTGC TCCGAACGCA

301 AATTCGGCTC GAGGGTTTAA TGTCATAGAT AGAATGAAAG TAGCCCTTGA

351 GAGAGCTTGC CCAGGAAGAG TCTCTTGTGC AGATATTCTC ACCATCGCCT

401 CTCAAATATC AGTGCTTTTG TCGGGAGGTC CATGGTGGCC GGTTCCGTTG

451 GGGAGGAGAG ACAGTGTAGA AGCTTTCTTC GCTCTGGCTA ATACAGCTCT

501 GCCCTCTCCG TTTTTCAATC TTACTCAACT TAAAACAGCC TTTGCTGACG

551 TTGGCCTAAA CCGCACTTCC GATCTAGTTG CTCTTTCCGG TGGTCACACA

601 TTTGGAAGAG CACAATGCCA GTTCGTGACG CCTCGTCTCT ACAACTTCAA

651 TGGTACAAAC AGTCCAGACC CAAGTCTGAA CCCAACTTAC CTTGTCGAAC

701 TCCGTCGATT GTGTCCTCAA AACGGAAACG GCACCGTTCT GGTCAACTTC

751 GATGTCGTGA CTCCAGATGC TTTCGATAGT CAATACTACA CCAACCTTCG

801 TAATGGGAAA GGTCTTATTC AGAGTGACCA GGAACTCTTT TCGACTCCAG

851 GAGCCGATAC GATCCCTTTA GTAAACCAAT ACAGCAGCGA CATGTCCGTG

901 TTCTTTCGAG CATTCATTGA CGCAATGATT AGGATGGGAA ATCTTAGACC

951 TTTGACTGGA ACTCAAGGAG AGATTAGACA AAATTGTAGG GTTGTGAATC

1001 CACGAATTAG GGTTGTGGAG AACGACGATG GTGTTGTGAG TTCTATCTGA

PER34 - at3g49120 (SEQ ID NO: 6)

1 ATGCATTTCT CTTCGTCTTC AACATCGTCC ACTTGGACAA TCTTAATCAC

51 ATTGGGATGT CTTATGCTTC ATGCATCTTT GTCCGCTGCT CAACTCACCC

101 CTACCTTCTA CGATAGGTCA TGTCCTAATG TCACTAACAT CGTACGAGAA

151 ACCATTGTAA ATGAGTTAAG GTCGGACCCT CGTATCGCTG CGAGCATCCT

201 TCGTCTTCAC TTCCACGACT GCTTTGTTAA TGGTTGTGAC GCATCCATCT

251 TGTTAGACAA CACGACATCA TTTCGAACAG AGAAAGATGC GTTTGGAAAC

301 GCAAATTCGG CTCGGGGATT TCCAGTGATT GATAGAATGA AAGCTGCGGT

351 GGAGAGGGCA TGCCCAAGAA CCGTTTCATG CGCAGATATG CTCACCATTG

401 CAGCTCAACA ATCTGTCACT TTGGCAGGAG GTCCTTCTTG GAGGGTTCCT

451 TTGGGAAGGA GAGACAGTTT ACAAGCATTC CTGGAACTCG CTAATGCAAA

501 TCTTCCAGCT CCATTCTTTA CACTTCCACA ACTTAAAGCC AGCTTCAGAA

551 ATGTTGGTCT CGATCGTCCT TCTGATCTCG TTGCTCTCTC CGGTGGTCAC

601 ACATTTGGTA AAAATCAATG TCAGTTTATT CTTGACAGAT TATACAATTT

651 CAGCAACACA GGTTTACCCG ACCCTACACT CAACACTACT TACCTCCAAA

701 CTCTTCGTGG ACTATGCCCC CTTAATGGCA ATCGAAGTGC CTTGGTAGAT

751 TTTGATCTAC GTACGCCTAC GGTTTTCGAC AACAAATACT ACGTGAATCT

801 CAAAGAGCGA AAAGGTCTTA TCCAGAGCGA CCAAGAGTTG TTCTCTAGCC

851 CCAATGCCAC TGACACAATC CCCTTGGTGA GAGCATATGC TGATGGCACA

901 CAAACATTCT TCAATGCATT TGTGGAGGCA ATGAATAGGA TGGGAAACAT

951 TACACCAACT ACAGGAACTC AAGGACAAAT CAGATTGAAC TGTAGAGTTG

1001 TGAACTCCAA CTCTCTGCTC CATGATGTGG TGGATATCGT TGACTTTGTT

1051 AGCTCTATGT GA

PER47 - at4g33420 (SEQ ID NO: 7)

1 ATGTTGACTA GATTTAAGAA ACAAAATAAT AAGATGGTTA GGGCAAATAT

51 AGTGAGCATG GTTCTGTTGA TGCATGCCAT CGTTGGGTTT CCTTTTCATG

101 CGAGGGGATT AAGTATGACT TACTACATGA TGAGCTGTCC TTTCGCTGAA

151 CAGATTGTGA AGAACAGTGT TAACAATGCT CTTCAAGCCG ATCCCACTTT

201 AGCCGCTGGT CTTATCCGTA TGTTGTTCCA CGACTGTTTC ATCGAGGGAT

251 GTGATGCGTC AATTCTGCTA GATTCAACGA AAGACAACAC TGCGGAAAAG

301 GATTCGCCTG CGAATCTGAG TCTACGTGGC TACGAGATCA TCGATGATGC

351 AAAGGAGAAA ATCGAGAATA GATGTCCAGG AGTTGTATCA TGCGCAGATA

401 TAGTTGCCAT GGCTGCTAGA GATGCTGTCT TTTGGGCTGG TGGTCCATAT

451 TATGACATAC CGAAAGGAAG GTTTGATGGT AAAAGATCAA AGATAGAAGA

501 TACAAGAAAT CTACCTTCAC CTTTTCTTAA TGCCTCTCAA CTCATTCAAA

551 CTTTTGGCCA ACGTGGTTTC ACTCCACAAG ATGTTGTTGC TCTCTCTGGA

601 GCACATACCC TAGGAGTTGC ACGATGCTCC TCCTTCAAGG CTAGACTTAC

651 CGTCCCAGAC TCTTCATTAG ACTCGACTTT TGCAAACACT CTCTCTAAAA

701 CATGCAGCGC GGGTGACAAT GCAGAGCAAC CCTTTGATGC TACGCGCAAC

751 GATTTCGACA ATGCTTACTT CAATGCGCTT CAGATGAAAT CAGGAGTCCT

801 CTTTTCAGAC CAAACTTTAT TCAACACACC AAGGACCAGG AATCTTGTTA

851 ACGGCTATGC ACTTAACCAA GCCAAGTTCT TCTTTGATTT CCAACAGGCC

901 ATGCGCAAGA TGAGCAATCT TGATGTTAAA CTTGGCTCTC AAGGTGAAGT

951 CCGTCAAAAT TGTCGGAGTA TTAATTAG

PER50 - at4g37520 (SEQ ID NO: 8)

1 ATGGTTGTGG TGAACAAAAC GAATCTCTTA TTGCTTCTTC TCTCTCTATG

51 TCTCACCCTC GACCTCTCCT CCGCACAACT CCGCCGTAAC TTCTACGCCG

101 GAAGTTGTCC TAACGTCGAA CAAATCGTTA GAAACGCCGT ACAGAAAAAA

151 GTCCAACAAA CTTTCACCAC CATTCCCGCC ACTCTACGCC TCTATTTCCA

201 TGATTGTTTC GTCAATGGGT GTGATGCATC AGTGATGATA GCGTCAACAA

251 ATAATAATAA GGCGGAGAAG GATCATGAGG AAAATTTATC GTTGGCCGGA

301 GATGGATTCG ACACCGTCAT TAAAGCTAAA GAAGCTCTCG ATGCCGTCCC

351 AAATTGTCGT AACAAAGTTT CATGCGCTGA TATTCTCACG ATGGCCACTC

401 GTGACGTCGT TAATCTTGCG GGTGGACCAC AATACGACGT GGAATTGGGA

451 AGGCTAGATG GACTATCATC AACAGCGGCT AGCGTCGGAG GGAAGTTGCC

501 GCATCCGACC GACGATGTCA ATAAACTCAC TTCACTCTTC GCCAAAAACG

551 GCCTTAGCCT TAACGACATG ATCGCTCTCT CCGGGGCACA CACATTAGGA

601 TTCGCGCATT GCACAAAAGT GTTCAATCGG ATATACACTT TCAACAAGAC

651 AACTAAAGTG GATCCCACGG TTAACAAAGA TTACGTGACA GAGCTAAAAG

701 CGTCGTGTCC TCGAAACATA GATCCAAGAG TGGCTATTAA CATGGACCCA

751 ACAACACCAA GACAATTCGA CAACGTTTAC TACAAAAACT TGCAACAAGG

801 GAAAGGATTG TTCACGTCGG ATCAAGTCTT ATTCACGGAT CGTCGGTCAA

851 AGCCAACCGT TGACTTATGG GCTAATAATG GACAGTTGTT TAATCAAGCT

901 TTTATTAACT CGATGATCAA GCTTGGTCGT GTTGGTGTTA AAACTGGTAG

951 TAATGGTAAT ATCCGTCGTG ATTGTGGAGC TTTTAATTGA

PER51 - at4g37530 (SEQ ID NO: 9)

1 ATGGTTGTGA TGAACAAAAC GAATCTCTTG TTGCTTATTC TCTCTCTTTT

51 TCTCGCCATC AACCTCTCCT CCGCTCAACT CCGCGGCGAC TTCTACGCCG

101 GAACTTGTCC GAACGTCGAG CAAATCGTCA GAAACGCCGT ACAGAAAAAA

151 ATCCAACAAA CTTTCACCAC CATTCCCGCC ACTCTACGCC TCTATTTCCA

201 TGATTGTTTC GTCAATGGAT GTGATGCATC AGTGATGATA GCGTCGACAA

251 ATACTAATAA GGCGGAGAAG GATCATGAGG ATAATTTATC GTTGGCCGGA

301 GATGGATTCG ACACCGTCAT TAAAGCTAAA GAAGCTGTTG ACGCCGTCCC

351 AAATTGTCGT AACAAAGTTT CTTGCGCTGA TATTCTCACG ATGGCCACTC

401 GTGACGTCGT TAATCTTGCG GGTGGACCAC AATACGCCGT AGAATTGGGA

451 AGGCGAGATG GACTTTCGTC GTCGGCTTCT AGCGTCACAG GGAAGCTACC

501 CAAACCGACC TTTGATCTCA ACCAACTCAA TGCACTTTTC GCAGAAAACG

551 GTCTTAGCCC TAATGACATG ATCGCTCTCT CCGGGGCACA CACATTAGGA

601 TTCGCGCATT GCACAAAAGT TTTCAATCGG CTATACAATT TCAACAAGAC

651 AAATAACGTG GATCCCACGA TTAACAAAGA TTACGTGACA GAGCTAAAAG

701 CGTCGTGTCC TCAAAACATA GATCCAAGAG TGGCTATTAA CATGGATCCC

751 AATACACCAA GACAATTCGA CAACGTTTAC TACAAAAACT TGCAACAAGG

801 GAAAGGATTG TTCACGTCTG ATCAAGTCTT ATTCACCGAT AGTCGGTCAA

851 AGCCAACCGT TGACTTATGG GCTAATAATG GACAGTTGTT TAATCAAGCT

901 TTTATTAGCT CGATGATCAA GCTTGGTCGT GTTGGTGTTA AAACTGGTAG

951 TAATGGTAAT ATTCGTCGTG ATTGCGGAGC TTTCAATTAA

PER52 - at5g05340 (SEQ ID NO: 10)

1 ATGGCTTCAA ATAAACTTAT TTCGATTCTA GTTCTCGTAG TTACACTTTT

51 ACTCCAAGGT GATAACAATT ACGTCGTCGA AGCACAACTC ACGACTAACT

101 TCTACTCAAC CTCTTGCCCC AACCTCCTCT CCACCGTCCA AACCGCCGTT

151 AAATCTGCCG TTAACAGCGA GGCTCGAATG GGTGCATCTA TCCTCCGTCT

201 CTTCTTCCAC GATTGCTTTG TCAACGGATG CGACGGTTCG ATTCTACTAG

251 ACGATACGTC AAGCTTCACA GGAGAACAAA ACGCGGCTCC AAACCGCAAT

301 TCAGCTCGTG GGTTTAATGT GATCGACAAC ATCAAATCAG CGGTTGAGAA

351 AGCATGTCCC GGGGTTGTGT CTTGTGCTGA TATCTTAGCC ATTGCTGCTA

401 GAGACTCCGT CGTAGCCCTT GGAGGACCGA ACTGGAATGT GAAAGTAGGA

451 AGAAGAGATG CAAGAACGGC GAGTCAAGCG GCAGCGAATA GTAACATTCC

501 GGCACCGACT TCGAGTCTGA GCCAACTCAT TAGTAGTTTC AGTGCTGTTG

551 GACTCTCCAC CAGAGATATG GTTGCTCTCT CTGGCGCACA CACGATCGGA

601 CAATCCCGTT GCACGAACTT CCGAGCGAGA ATCTACAACG AGACAAACAT

651 CAATGCCGCC TTCGCCACGA CACGTCAACG AACTTGCCCT AGAGCCTCCG

701 GTTCCGGCGA CGGAAACCTA GCTCCACTAG ACGTCACCAC GGCAGCTTCT

751 TTCGACAACA ACTACTTCAA GAATCTCATG ACTCAGAGAG GGCTCCTCCA

801 TTCCGACCAA GTGCTCTTCA ACGGCGGCTC TACTGACTCC ATCGTCCGTG

851 GATACAGCAA CAATCCGTCG AGCTTTAACT CTGACTTCAC GGCGGCGATG

901 ATAAAAATGG GTGATATCAG CCCGTTGACT GGTAGTAGCG GTGAAATCCG

951 CAAGGTTTGC GGGAGGACCA ACTGA

PER62 - at5g39580 (SEQ ID NO: 11)

1 ATGGGCTTGG TCCGATCATT TGCTTTGGTC ATAGTCTTCC TTAGTTGTCT

51 CATCGCCGTT TATGGCCAAG GTACGAGGAT CGGGTTCTAC TCGACTACGT

101 GCCCTAATGC CGAGACAATT GTTCGGACCA CTGTGGCATC TCACTTTGGT

151 TCAGATCCAA AGGTTGCACC CGGGTTACTG AGATGCCACA ACCATGATTG

201 CTTTGTCCAA GGTTGTGATG GTTCGGTGCT TTTATCGGGA CCTAACTCTG

251 AAAGAACCGC TGGCGCAAAC GTTAACCTCC ATGGTTTTGA GGTCATTGAC

301 GATGCCAAGA GGCAGCTCGA GGCAGCATGT CCTGGTGTTG TCTCTTGTGC

351 TGATATCTTG GCCTTAGCGG CTCGTGATTC GGTTTCTCTC ACAAACGGAC

401 AAAGTTGGCA AGTTCCAACA GGACGTAGAG ATGGAAGAGT TTCCTTGGCA

451 TCGAACGTTA ACAATCTTCC TTCTCCAAGT GACTCTCTAG CCATTCAACA

501 AAGGAAATTC TCCGCTTTTC GCCTCAACAC TCGCGATCTC GTCACTCTTG

551 TTGGAGGAGG ACACACGATC GGAACAGCTG CATGTGGGTT TATCACGAAC

601 AGGATATTCA ACTCGAGCGG AAACACAGCA GATCCAACAA TGGACCAAAC

651 ATTTGTACCA CAACTTCAAA GACTTTGTCC CCAAAACGGC GACGGATCAG

701 CACGTGTTGA TCTTGACACC GGAGTGGAA ACACTTTTGA CACATCTTAT

751 TTCATCAATC TCAGCCGTAA CAGAGGAATT CTTCAATCCG ATCACGTTCT

801 TTGGACTAGT CCGGCCACAA GATCCATAGT GCAAGAGTTT ATGGCACCTA

851 GAGGCAACTT CAATGTCCAG TTCGCAAGGT CAATGGTTAA AATGAGTAAT

901 ATTGGTGTGA AGACGGGGAC AAATGGAGAA ATTCGTAGGG TTTGCTCTGC

951 GGTTAATTAA

## Superoxide Dismutase (Genes encoding seven SODs known to be present in Arabidopsis)

CSD1 - at1g08830 (SEQ ID NO: 12)

1 ATGGCGAAAG GAGTTGCAGT TTTGAACAGC AGTGAGGGTG TTACGGGGAC

51 TATCTTTTTC ACCCAGGAAG GCGATGGTGT GACCACTGTG AGTGGAACAG

101 TTTCTGGCCT TAAGCCTGGT CTTCATGGTT TCCATGTCCA TGCTCTTGGT

151 GACACCACTA ACGGTTGCAT GTCTACTGGT CCACATTTCA ACCCCGATGG

201 TAAAACACAC GGTGCCCCTG AGGATGCTAA TCGACATGCT GGTGATCTAG

251 GAAACATCAC TGTTGGAGAT GATGGAACTG CCACCTTCAC AATCACTGAT

301 TGCCAGATTC CTCTTACTGG ACCAAACTCT ATTGTTGGTA GGGCTGTTGT

351 TGTCCATGCA GACCCTGATG ACCTCGGAAA GGGAGGCCAT GAACTCAGCC

401 TGGCTACTGG AAACGCAGGC GGCCGTGTTG CTTGCGGCAT CATTGGTCTC

451 CAGGGCTAA

CSD2 - at2g28190 (SEQ ID NO: 13)

1 ATGGCTGCCA CCAACACAAT CCTCGCATTC TCATCTCCTT CTCGTCTTCT

51 CATTCCTCCT TCCTCCAATC CTTCAACTCT CCGTTCCTCT TTCCGCGGCG

101 TCTCTCTCAA CAACAACAAT CTCCACCGTC TCCAATCTGT TTCCTTCGCC

151 GTTAAAGCTC CGTCGAAAGC GTTGACAGTT GTTTCCGCGG CGAAGAAGGC

201 TGTTGCAGTG CTTAAAGGTA CTTCTGATGT CGAAGGAGTT GTTACTTTGA

251 CCCAAGATGA CTCAGGTCCT ACAACTGTGA ATGTTCGTAT CACTGGTCTC

301 ACTCCAGGGC CTCATGGATT TCATCTCCAT GAGTTTGGTG ATACAACTAA

351 TGGATGTATC TCAACAGGAC CACATTTCAA CCCTAACAAC ATGACACACG

401 GAGCTCCAGA AGATGAGTGC CGTCATGCGG GTGACCTGGG AAACATAAAT

451 GCCAATGCCG ATGGCGTGGC AGAAACAACA ATAGTGGACA ATCAGATTCC

501 TCTGACTGGT CCTAATTCTG TTGTTGGAAG AGCCTTTGTG GTTCACGAGC

551 TTAAGGATGA CCTCGGAAAG GGTGGCCATG AGCTTAGTCT GACCACTGGA

601 AACGCAGGCG GGAGATTGGC ATGTGGTGTG ATTGGCTTGA CGCCGCTCTA

651 A

CSD3 - at5g18100 (SEQ ID NO: 14)

1 ATGGAAGCTC CTAGAGGAAA TCTGAGAGCG GTGGCTCTTA TCGCCGGCGA

51 TAACAACGTC CGAGGCTGTC TTCAATTCGT ACAAGACATT TCCGGAACTA

101 CTCATGTCAC CGGGAAGATC TCAGGACTCT CTCCTGGCTT CCATGGATTT

151 CATATTCACT CTTTTGGTGA TACCACAAAT GGCTGCATCT CTACTGGACC

201 TCACTTCAAT CCATTAAATC GAGTTCACGG GCCACCGAAC GAGGAAGAGC

251 GTCATGCTGG CGATTTGGGT AACATTCTTG CAGGATCAAA CGGTGTTGCT

301 GAAATCTTAA TCAAAGACAA ACATATACCG CTTAGTGGGC AGTATTCCAT

351 ACTCGGGAGG GCGGTTGTTG TGCATGCGGA TCCTGATGAC CTTGGGAAAG

401 GAGGGCACAA GCTTAGCAAA TCAACTGGAA ACGCAGGCTC GAGAGTTGGA

451 TGTGGTATCA TCGGACTTCA ATCATCTGCG GATGCTAAAC TATAG

FSD1 - at4g25100 (SEQ ID NO: 15)

1 ATGGCTGCTT CAAGTGCTGT CACCGCAAAC TACGTCCTCA AGCCACCTCC

51 ATTCGCACTG GATGCTTTGG AGCCGCATAT GAGCAAACAA ACTCTGGAGT

101 TTCACTGGGG AAAACATCAC AGAGCTTACG TGGACAACCT CAAGAAACAG

151 GTTCTTGGAA CCGAGCTTGA AGGCAAGCCC TTAGAGCACA TTATCCACAG

201 CACTTACAAC AATGGTGATC TCCTCCCTGC TTTCAACAAC GCTGCTCAGG

251 CGTGGAACCA CGAGTTCTTC TGGGAGTCAA TGAAACCAGG TGGTGGAGGA

301 AAACCATCAG GAGAGCTTCT TGCTTTGCTT GAAAGAGATT TCACTTCTTA

351 TGAGAAGTTC TATGAAGAGT TCAATGCTGC TGCAGCCACT CAGTTTGGAG

401 CTGGCTGGGC CTGGCTTGCT TATTCAAATG AAAAACTCAA AGTAGTGAAA

451 ACTCCCAATG CTGTGAATCC CCTTGTGCTC GGCTCTTTCC CATTGCTTAC

501 CATTGATGTC TGGGAGCATG CTTACTACCT TGACTTCCAG AACCGAAGAC

551 CAGATTACAT AAAGACATTC ATGACCAATC TTGTGTCTTG GGAAGCTGTA

601 AGTGCCAGAC TTGAGGCCGC CAAGGCTGCT TCTGCTTAA

FSD2 - at5g51100 (SEQ ID NO: 16)

1 ATGATGAATG TTGCAGTGAC AGCCACTCCC TCGTCTCTCT TGTACTCTCC

51 TCTGCTTCTT CCTTCTCAAG GGCCAAACCG GCGAATGCAA TGGAAAAGAA

101 ACGGAAAGAG ACGGTTAGGG ACAAAGGTGG CTGTTTCCGG TGTTATCACA

151 GCTGGATTTG AGCTGAAGCC ACCTCCATAT CCTCTTGATG CTCTGGAACC

201 GCATATGAGC CGGGAAACCT TGGATTATCA CTGGGGCAAA CATCACAAAA

251 CTTATGTAGA GAACCTGAAC AAGCAAATCT TAGGCACGGA TCTAGATGCA

301 TTATCCTTGG AAGAAGTTGT GCTTCTTTCA TACAACAAAG GCAATATGCT

351 TCCTGCTTTC AACAACGCTG CACAGGCTTG GAACCACGAG TTCTTCTGGG

401 AGTCTATCCA ACCTGGAGGT GGAGGAAAGC CAACTGGAGA GCTCCTCAGA

451 TTAATAGAAA GAGATTTTGG GTCTTTCGAA GAGTTTTTGG AAAGGTTCAA

501 GTCGGCTGCA GCTTCGAATT TTGGTTCGGG TTGGACATGG CTTGCATATA

551 AGGCGAATAG ACTTGACGTT GCAAATGCCG TTAATCCTCT CCCAAAGGAG

601 GAAGACAAGA AACTTGTTAT AGTGAAGACG CCCAATGCAG TAAATCCGCT

651 CGTATGGGAT TATTCTCCAC TTCTCACCAT TGATACCTGG GAGCACGCTT

701 ACTATCTGGA TTTTGAGAAC CGAAGAGCTG AATACATAAA TACATTCATG

751 GAAAAGCTTG TGTCATGGGA AACTGTAAGC ACAAGGTTGG AATCCGCAAT

801 TGCTCGAGCA GTGCAAAGAG AACAAGAAGG AACAGAGACA GAAGATGAAG

851 AGAATCCAGA TGATGAAGTA CCAGAGGTCT ATTTAGATAG TGACATCGAT

901 GTATCTGAGG TTGACTAA

FSD3 - at5g23310 (SEQ ID NO: 17)

1 ATGAGTTCTT GTGTTGTGAC GACAAGCTGT TTCTATACAA TTTCAGATTC

51 TAGTATACGT TTGAAATCCC CCAAGCTCCT CAATCTGAGT AACCAGCAGA

101 GAAGACGCTC TCTTAGGTCT CGAGGTGGTT TAAAGGTTGA AGCTTACTAC

151 GGTCTAAAGA CACCTCCTTA TCCACTTGAT GCTTTGGAGC CGTATATGAG

201 TAGAAGAACA CTAGAAGTGC ATTGGGGAAA ACACCATCGA GGTTATGTAG

251 ATAATCTGAA TAAACAGTTA GGGAAAGATG ATAGACTCTA TGGATACACC

301 ATGGAAGAGC TTATCAAGGC TACATACAAC AACGGGAATC CTTTACCCGA

351 GTTCAACAAC GCTGCACAGG TCTATAACCA TGATTTCTTC TGGGAGTCGA

401 TGCAACCTGG TGGTGGAGAC ACGCCTCAAA AGGGTGTTCT TGAGCAGATT

451 GATAAGGATT TTGGTTCTTT CACAAATTTT AGAGAAAAGT TCACTAATGC

501 AGCTCTTACT CAGTTTGGTT CTGGATGGGT CTGGCTTGTC TTAAAGAGGG

551 AAGAGAGAAG GCTTGAGGTG GTCAAAACCT CAAACGCCAT TAACCCACTC

601 GTGTGGGACG ATATTCCAAT CATCTGCGTG GATGTGTGGG AGCACTCTTA

651 TTATCTGGAC TACAAGAACG ACAGGGCTAA GTATATAAAC ACATTTCTGA

701 ACCACTTGGT GTCGTGGAAC GCTGCCATGA GTCGGATGGC CCGTGCAGAA

751 GCGTTTGTGA ATCTTGGTGA ACCCAACATC CCAATCGCTT AA

MSD1 - at3g10920 (SEQ ID NO: 18)

1 ATGGCGATTC GTTGTGTAGC GAGTAGAAAA ACCCTAGCCG GCTTGAAGGA

51 GACATCATCG AGGCTATTGA GGATCAGAGG GATTCAGACT TTTACGCTTC

101 CTGATCTTCC TTACGATTAT GGCGCATTGG AACCGGCCAT TAGTGGAGAG

151 ATCATGCAGA TTCATCACCA GAAGCATCAC CAGGCTTATG TTACTAATTA

201 CAATAATGCT CTTGAGCAGC TTGATCAAGC TGTGAACAAG GGAGATGCTT

251 CCACTGTTGT TAAGTTGCAG AGCGCCATCA AATTCAACGG CGGAGGTCAT

301 GTCAACCATT CGATTTTCTG GAAGAACCTT GCTCCTTCCA GTGAAGGTGG

351 TGGAGAGCCA CCAAAAGGAT CTCTTGGTAG TGCCATTGAC GCTCACTTTG

401 GCTCCCTTGA AGGTCTGGTG AAAAAGATGA GTGCTGAGGG TGCTGCAGTG

451 CAAGGCTCAG GATGGGTGTG GCTCGGACTA GACAAAGAAC TGAAGAAGCT

501 AGTTGTTGAC ACAACTGCCA ATCAGGATCC ATTAGTGACA AAAGGAGGAA

551 GCTTGGTACC TCTGGTGGGT ATAGATGTTT GGGAGCACGC CTACTACTTG

601 CAGTACAAAA ATGTGAGGCC TGAGTATCTG AAGAATGTAT GGAAAGTGAT

651 CAACTGGAAA TATGCAAGCG AGGTTTATGA GAAGGAAAAC AACTGA

## Genes related to lignin production (a group of genes highly expressed in SEC among related genes, see FIG. 5)

PAL1 - at2g37040 (SEQ ID NO: 19)

1 ATGGAGATTA ACGGGGCACA CAAGAGCAAC GGAGGAGGAG TGGACGCTAT

51 GTTATGCGGC GGAGACATCA AGACAAAGAA CATGGTGATC AACGCGGAGG

101 ATCCTCTCAA CTGGGGAGCT GCAGCGGAGC AAATGAAAGG TAGCCATTTG

151 GATGAAGTGA AGGAATGGT TGCTGAGTTT AGGAAGCCAG TTGTGAATCT

201 TGGTGGTGAG ACTCTGACCA TTGGACAAGT GGCTGCGATC TCAACTATTG

251 GTAACAGTGT GAAGGTGGAG CTATCGGAGA CAGCTAGAGC CGGTGTGAAT

301 GCTAGTAGTG ATTGGGTTAT GGAGAGTATG AACAAAGGCA CTGATAGTTA

351 TGGTGTTACT ACTGGTTTTG GTGCTACTTC TCATCGGAGA ACCAAAAACG

401 GTGTCGCACT TCAGAAGGAA CTTATTAGAT TCCTTAACGC CGGAATATTC

451 GGAAGCACGA AAGAAACAAG CCACACATTG CCACACTCCG CCACAAGAGC

501 CGCCATGCTT GTACGAATCA ACACTCTCCT CCAAGGATTT TCCGGTATCC

551 GATTTGAGAT TCTCGAAGCA ATTACCAGTT TCCTCAACAA CAACATCACT

601 CCATCTCTCC CCCTCCGTGG TACAATCACC GCCTCCGGAG ATCTCGTTCC

651 TCTCTCCTAC ATCGCCGGAC TTCTCACCGG TCGTCCCAAT TCCAAAGCTA

701 CTGGTCCCAA CGGTGAAGCT TTAACAGCAG AGGAAGCTTT CAAATTAGCA

751 GGAATCAGCT CCGGATTCTT TGATCTCCAG CCTAAGGAAG GTCTCGCGCT

801 AGTCAATGGC ACGGCGGTTG GATCTGGAAT GGCGTCAATG GTGTTATTCG

851 AAACGAATGT TCTCTCTGTT TTGGCTGAGA TTTTGTCGGC GGTTTTCGCA

901 GAGGTGATGA GTGGTAAGCC TGAGTTCACC GATCATCTCA CTCACAGACT

951 TAAACATCAT CCCGGTCAAA TCGAAGCGGC GGCGATAATG GAGCATATCC

1001 TCGACGGAAG CTCGTACATG AAATTAGCTC AGAAGCTTCA CGAGATGGAT

1051 CCGTTACAGA AACCTAAACA AGATCGTTAC GCTCTTCGTA CTTCTCCTCA

1101 ATGGTTAGGT CCTCAAATCG AAGTGATCCG TTACGCAACG AAATCGATCG

1151 AGCGTGAGAT TAACTCCGTC AACGATAATC CGTTGATCGA TGTTTCGAGG

1201 AACAAGGCGA TTCACGGTGG TAACTTCCAA GGAACACCAA TCGGAGTTTC

1251 AATGGATAAC ACGAGATTGG CGATAGCAGC GATTGGTAAA CTCATGTTTG

1301 CTCAATTCTC AGAGCTTGTG AATGATTTCT ACAACAATGG TTTACCCTCG

1351 AATCTAACCG CTTCGAGGAA TCCAAGTTTG GATTATGGAT TCAAGGGAGC

1401 TGAGATTGCA ATGGCTTCTT ATTGTTCAGA GCTTCAATAC TTAGCTAATC

1451 CTGTGACTAG CCATGTTCAA TCAGCAGAGC AACATAACCA AGATGTCAAC

1501 TCTTTGGGAC TAATCTCGTC TCGCAAAACT TCTGAAGCTG TTGATATTCT

1551 CAAGCTTATG TCAACAACGT TCCTCGTTGC GATTTGTCAA GCTGTGGATT

1601 TGAGACATTT GGAGGAGAAT TTGAGACAGA CTGTGAAGAA CACTGTCTCT

1651 CAAGTGGCGA AGAAAGTTCT TACTACTGGA GTCAATGGTG AGCTTCATCC

1701 TTCTCGCTTC TGCGAAAAGG ATTTACTCAA AGTTGTAGAC CGTGAACAAG

1751 TCTACACATA CGCGGATGAT CCTTGTAGCG CAACGTACCC GTTGATTCAG

1801 AAGCTGAGAC AAGTTATTGT TGACCATGCT TTGATCAATG GTGAGAGTGA

1851 GAAGAATGCA GTGACTTCAA TCTTCCATAA GATTGGAGCT TTCGAGGAGG

1901 AGCTTAAGGC AGTGCTACCG AAAGAAGTGG AAGCAGCAAG AGCAGCCTAC

1951 GATAACGGAA CATCGGCTAT CCCGAACAGG ATCAAGGAAT GTAGGTCGTA

2001 TCCATTGTAT AGATTCGTGA GGGAAGAGCT TGGAACAGAG CTTTTGACCG

2051 GAGAGAAAGT GACGTCGCCT GGAGAAGAGT TCGACAAGGT TTTCACGGCG

2101 ATTTGTGAAG GTAAAATCAT TGATCCGATG ATGGAATGTC TCAACGAGTG

2151 GAACGGAGCT CCCATTCCAA TATGTTAA

C4H - at2g30490 (SEQ ID NO: 20)

1 ATGGACCTCC TCTTGCTGGA GAAGTCTTTA ATCGCCGTCT TCGTGGCGGT

51 GATTCTCGCC ACGGTGATTT CAAAGCTCCG CGGCAAGAAA TTGAAGCTAC

101 CTCCAGGTCC TATACCAATT CCGATCTTCG GAAACTGGCT TCAAGTCGGA

151 GATGATCTCA ACCACCGTAA TCTCGTCGAT TACGCTAAGA AATTCGGCGA

201 TCTCTTCCTC CTCCGTATGG GTCAGCGAAA CCTAGTCGTC GTCTCCTCAC

251 CGGATCTAAC AAAGGAAGTG CTCCTCACTC AAGGCGTTGA GTTTGGATCC

301 AGAACGAGAA ACGTCGTGTT CGACATTTTC ACCGGGAAAG GTCAAGATAT

351 GGTGTTCACT GTTTACGGCG AGCATTGGAG GAAGATGAGA AGAATCATGA

401 CGGTTCCTTT CTTCACCAAC AAAGTTGTTC AACAGAATCG TGAAGGTTGG

451 GAGTTTGAAG CAGCTAGTGT TGTTGAAGAT GTTAAGAAGA ATCCAGATTC

501 TGCTACGAAA GGAATCGTGT TGAGGAAACG TTTGCAATTG ATGATGTATA

551 ACAATATGTT CCGTATCATG TTCGATAGAA GATTTGAGAG TGAGGATGAT

601 CCTCTTTTCC TTAGGCTTAA GGCTTTGAAT GGTGAGAGAA GTCGATTAGC

651 TCAGAGCTTT GAGTATAACT ATGGAGATTT CATTCCTATC CTTAGACCAT

701 TCCTCAGAGG CTATTTGAAG ATTTGTCAAG ATGTGAAAGA TCGAAGAATC

751 GCTCTTTTCA AGAAGTACTT TGTTGATGAG AGGAAGCAAA TTGCGAGTTC

801 TAAGCCTACA GGTAGTGAAG GATTGAAATG TGCCATTGAT CACATCCTTG

851 AAGCTGAGCA GAAGGGAGAA ATCAACGAGG ACAATGTTCT TTACATCGTC

901 GAGAACATCA ATGTCGCCGC GATTGAGACA ACATTGTGGT CTATCGAGTG

951 GGGAATTGCA GAGCTAGTGA ACCATCCTGA AATCCAGAGT AAGCTAAGGA

1001 ACGAACTCGA CACAGTTCTT GGACCGGGTG TGCAAGTCAC CGAGCCTGAT

1051 CTTCACAAAC TTCCATACCT TCAAGCTGTG GTTAAGGAGA CTCTTCGTCT

1101 GAGAATGGCG ATTCCTCTCC TCGTGCCTCA CATGAACCTC CATGATGCGA

1151 AGCTCGCTGG CTACGATATC CCAGCAGAAA GCAAAATCCT TGTTAATGCT

1201 TGGTGGCTAG CAAACAACCC CAACAGCTGG AAGAAGCCTG AAGAGTTTAG

1251 ACCAGAGAGG TTCTTTGAAG AAGAATCGCA CGTGGAAGCT AACGGTAATG

1301 ACTTCAGGTA TGTGCCATTT GGTGTTGGAC GTCGAAGCTG TCCCGGGATT

1351 ATATTGGCAT TGCCTATTTT GGGGATCACC ATTGGTAGGA TGGTCCAGAA

1401 CTTCGAGCTT CTTCCTCCTC CAGGACAGTC TAAAGTGGAT ACTAGTGAGA

1451 AAGGTGGACA ATTCAGCTTG CACATCCTTA ACCACTCCAT AATCGTTATG

1501 AAACCAAGGA ACTGTTAA

4CL1 - at1g51680 (SEQ ID NO: 21)

1 ATGGCGCCAC AAGAACAAGC AGTTTCTCAG GTGATGGAGA AACAGAGCAA

51 CAACAACAAC AGTGACGTCA TTTTCCGATC AAAGTTACCG GATATTTACA

101 TCCCGAACCA CCTATCTCTC CACGACTACA TCTTCCAAAA CATCTCCGAA

151 TTCGCCACTA AGCCTTGCCT AATCAACGGA CCAACCGGCC ACGTGTACAC

201 TTACTCCGAC GTCCACGTCA TCTCCCGCCA AATCGCCGCC AATTTTCACA

251 AACTCGGCGT TAACCAAAAC GACGTCGTCA TGCTCCTCCT CCCAAACTGT

301 CCCGAATTCG TCCTCTCTTT CCTCGCCGCC TCCTTCCGCG GCGCAACCGC

351 CACCGCCGCA AACCCTTTCT TCACTCCGGC GGAGATAGCT AAACAAGCCA

401 AAGCCTCCAA CACCAAACTC ATAATCACCG AAGCTCGTTA CGTCGACAAA

451 ATCAAACCAC TTCAAAACGA CGACGGAGTA GTCATCGTCT GCATCGACGA

501 CAACGAATCC GTGCCAATCC CTGAAGGCTG CCTCCGCTTC ACCGAGTTGA

551 CTCAGTCGAC AACCGAGGCA TCAGAAGTCA TCGACTCGGT GGAGATTTCA

601 CCGGACGACG TGGTGGCACT ACCTTACTCC TCTGGCACGA CGGGATTACC

651 AAAAGGAGTG ATGCTGACTC ACAAGGGACT AGTCACGAGC GTTGCTCAGC

701 AAGTCGACGG CGAGAACCCG AATCTTTATT TCCACAGCGA TGACGTCATA

751 CTCTGTGTTT TGCCCATGTT TCATATCTAC GCTTTGAACT CGATCATGTT

801 GTGTGGTCTT AGAGTTGGTG CGGCGATTCT GATAATGCCG AAGTTTGAGA

851 TCAATCTGCT ATTGGAGCTG ATCCAGAGGT GTAAAGTGAC GGTGGCTCCG

901 ATGGTTCCGC CGATTGTGTT GGCCATTGCG AAGTCTTCGG AGACGGAGAA

951 GTATGATTTG AGCTCGATAA GAGTGGTGAA ATCTGGTGCT GCTCCTCTTG

1001 GTAAAGAACT TGAAGATGCC GTTAATGCCA AGTTTCCTAA TGCCAAACTC

1051 GGTCAGGGAT ACGGAATGAC GGAAGCAGGT CCAGTGCTAG CAATGTCGTT

1101 AGGTTTTGCA AAGGAACCTT TTCCGGTTAA GTCAGGAGCT TGTGGTACTG

1151 TTGTAAGAAA TGCTGAGATG AAAATAGTTG ATCCAGACAC CGGAGATTCT

1201 CTTTCGAGGA ATCAACCCGG TGAGATTTGT ATTCGTGGTC ACCAGATCAT

1251 GAAAGGTTAC CTCAACAATC CGGCAGCTAC AGCAGAGACC ATTGATAAAG

1301 ACGGTTGGCT TCATACTGGA GATATTGGAT TGATCGATGA CGATGACGAG

1351 CTTTTCATCG TTGATCGATT GAAAGAACTT ATCAAGTATA AAGGTTTTCA

1401 GGTAGCTCCG GCTGAGCTAG AGGCTTTGCT CATCGGTCAT CCTGACATTA

1451 CTGATGTTGC TGTTGTCGCA ATGAAAGAAG AAGCAGCTGG TGAAGTTCCT

1501 GTTGCATTTG TGGTGAAATC GAAGGATTCG GAGTTATCAG AAGATGATGT

1551 GAAGCAATTC GTGTCGAAAC AGGTTGTGTT TTACAAGAGA ATCAACAAAG

1601 TGTTCTTCAC TGAATCCATT CCTAAAGCTC CATCAGGGAA GATATTGAGG

1651 AAAGATCTGA GGGCAAAACT AGCAAATGGA TTGTGA

HCT - at5g48930 (SEQ ID NO: 22)

1 ATGAAAATTA ACATCAGAGA TTCCACCATG GTCCGGCCTG CCACCGAGAC

51 ACCAATCACT AATCTTTGGA ACTCCAACGT CGACCTTGTC ATCCCCAGAT

101 TCCATACCCC TAGTGTCTAC TTCTACAGAC CCACCGGCGC TTCCAATTTC

151 TTTGACCCTC AGGTCATGAA GGAAGCTCTT TCCAAAGCCC TTGTCCCTTT

201 TTACCCTATG GCTGGTCGCT TGAAGAGAGA CGATGATGGT CGTATTGAGA

251 TCGATTGTAA CGGTGCTGGT GTTCTCTTCG TTGTGGCTGA TACTCCTTCT

301 GTTATCGATG ATTTTGGTGA TTTTGCTCCT ACCCTTAATC TCCGTCAGCT

351 TATTCCCGAA GTTGATCACT CCGCTGGCAT TCACTCTTTC CCGCTTCTCG

401 TTTTGCAGGT GACTTTCTTT AAATGTGGGG GAGCTTCACT TGGGGTTGGG

451 ATGCAACATC ACGCGGCAGA TGGTTTCTCT GGTCTTCATT TTATCAACAC

501 ATGGTCTGAT ATGGCTCGTG GTCTTGACCT AACCATTCCA CCTTTCATTG

551 ATCGAACACT CCTCCGAGCT AGGGACCCGC CACAGCCTGC TTTTCATCAT

601 GTTGAATATC AGCCTGCACC AAGTATGAAG ATACCTCTTG ATCCGTCTAA

651 ATCAGGACCT GAGAATAACCA CTGTCTCTAT ATTCAAATTA ACACGAGACC

701 AGCTTGTTGC TCTTAAGGCG AAATCCAAGG AGGATGGGAA CACTGTCAGC

751 TACAGCTCAT ACGAGATGTT GGCAGGGCAT GTGTGGAGAT CAGTGGGAAA

801 GGCGCGAGGG CTTCCAAACG ACCAAGAGAC GAAACTGTAC ATTGCAACTG

851 ATGGAAGGTC TAGACTACGT CCGCAGCTGC CTCCTGGTTA CTTTGGGAAT

901 GTGATATTCA CTGCAACACC ATTGGCTGTT GCAGGGGATT TGTTATCTAA

951 GCCAACATGG TATGCTGCAG GACAGATTCA TGATTTCTTG GTTCGTATGG

1001 ATGATAACTA TCTGAGGTCA GCTCTTGACT ACCTGGAGAT GCAGCCTGAT

1051 CTGTCAGCCC TTGTCCGCGG TGCACATACC TACAAGTGCC CAAATTTGGG

1101 AATCACAAGC TGGGTTAGAT TACCTATTTA TGATGCAGAC TTTGGTTGGG

1151 GTCGTCCTAT CTTTATGGGA CCTGGTGGAA TTCCATACGA GGGTTTGTCT

1201 TTTGTGCTAC CAAGTCCTAC TAATGATGGC AGCTTATCCG TTGCCATTGC

1251 CCTCCAATCT GAACACATGA AACTGTTTGA GAAGTTTTTG TTTGAGATAT

1301 GA

CCoAOMT6 - at1g67980 (SEQ ID NO: 23)

1 ATGGCCAATG AAATTCCTAC CAAAGGGATT CTCAAGAGCG AAGCTCTAAA

51 ACAGTACATC ATGGAAACGT CAGCGTATCC AAGAGAGCAT GAATTGCTCA

101 AGGAACTTCG AAAAGCTACT GTCCAAAAAT ACGGAAATTT AAGCGAGATG

151 GAGGTTCCGG TTGATGAGGG TCATTTCCTA TCGATGCTTG TAAAGATCAT

201 GAATGCGAAA AACACTATCG AGATCGGTGT TTTCACCGGT TACTCGCTTC

251 TCACTACAGC TCTTGCTTTA CCTGAAGATG GCCGTATTAC CGCGATAGAC

301 ATTGATAAAG AAGCTTACGA AGTTGGACTA GAGTTTATTA AGAAGGCTGG

351 TGTTGATCAC AAGATTAATT TTATCCATTC CGATGGTCTT AAGGCCTTAG

401 ACCAATTGGT GAACGACAAA TGTGAGTTTG ATTTTGCATT TGCGGATGCT

451 GACAAATCAA GCTACGTCAA CTTCCATGAG AGGCTTCTAA AATTGGTGAA

501 GGTTGGGGGA ATCATTGCGT TCGACAACAC CTTGTGGTTT GGTTTTGTGG

551 CGGAGGATGA AGATGGAGTT CCGGAGCATA TGAGAGAATA TAGAGCAGCT

601 CTTATAGAAT TCAATAAGAA ATTGGCTTTG GATCCCCGAG TCGAGGTCTC

651 TCAGATTTCC ATTGGAGATG GTATCACGCT CTGTAGACGT CTTGTTTGA

CCR - like1 - at1g76470 (SEQ ID NO: 24)

1 ATGGCTGTAA AGCAAAAGGT CTGTGTTACT GGTGCAGGAG GGTTTATTGC

51 TTCATGGCTC GTCAAGTTTC TTCTCTCTCG TGGCTACACT GTCCATGGAA

101 CCGTTCGAGA TCCTTGTGAT GAAAAGAATG ACCATTTGAG GAAACTGGAT

151 AATGCTTCAA AAAATCTGAA ACTTTTCAAG GCAGATTTGT TTGATGATGA

201 AGGTCTTTTC TCAGCCATTG ATGGTTGCTC TGGCGTTTTC CATATCGCTT

251 CTCCTGTGCC TTTTGAAGGA GTTCCTCTCA CAGAGGAAGA ACTGATTAAA

301 CCAGCTCTCA CTGGCACCAA GAATGTTCTA GAAGCATGCA CAGAAACCAA

351 AGTGCAGAAA GTTGTGGTTG TATCATCAAT CGCTGCAGTT GTTTACAATC

401 CAAAATGGCC TCAAGATGTT GCTAAGGACG AGGATTGTTG GTCAGACACT

451 CAATACCTTC ACTCACTTGA GGGTTATTGG CGTTATTACT ATCTTGCCAA

501 AACATTAACG GAACGTGAAG CTTTAGAGTG GAGTAAGAGA AACTTTGCTG

551 ATGTTGTTAC GCTCTGTCCG TCTGTCATTA TAGGCCCTAG GCTTCAATCC

601 ACGCTTAACT CTAGTAGTTT GGGACTTCTC AAGTTCATAA AAGGTGGTAT

651 AAAGTCCTTG TTGAGCGATG AACTGTATCT TGTGGATGTG CGTGATGTTG

701 CGGATGCGCT TTTGTTGGTG TATGAAAATC GAGAAGCAAC AGGAAGATAC

751 ATATGTAATT CTCATTCACT TTACACTGAT TCTCTTATGG AGAAGCTCAA

801 GAATATGTAT CCTAAACGCA ACTTCCCTGA AAGTTTCACG GAGGTAAAAG

851 AGAAGGAAGT GAGGCCTCTA AGTGCAGAGA AGTTGAAAAA CTTGGGGTGG

901 AAGTTTAGGC CATTAGAAGA AACCATTGAT GATTCTGTTG TGAGCTTTGA

951 AGCAGCTGGT GATCTTCCAA AGGCATAA

F5H2 - at5g04330 (SEQ ID NO: 25)

1 ATGCTTACTC TAATGACTCT CATCGTTCTT GTCCCTCTTC TTCTTTTTCT

51 CTTCCCCCAC CTCCTTTTAC GGCGGCAGAT GCTTCTGAAA CCCTACCCTC

101 CTGGCCCCAA AGGCTTACCC GTCATTGGAA ATATTCTCAT GATGAACCAG

151 TTTAACCACC GGGGTCTGGC CAAGCTCAGC CGTATATACG GTGGGTTGCT

201 CCATCTCCGC CTCGGATTTT CCCACATTTT CGTCGTTTCT TCTCCGGACA

251 TTGCGCGTCA AGTCCTCCAA GTCCAAGACC ACGTTTTCTC AAACCGTCCC

301 ACCACAATCG CAATCCGATA CTTGACCTAC GGAGGATCCG ATCTTGCATT

351 CTGCAATTAC GGCCCGTTTT GGCGTCGGAT GAGAAAACTC TACGTCATGA

401 TGCTCTTTAG CCGTAAACGG GCCGAGTCGT GGGTCTCTGT TGATGAAGAG

451 GTCCACAAAT CGGTCCGTTT AGTGGCTTCC AACGTCGGAA AACCACTAAA

501 CATATGTAAA CTAGCATTTT CCCTATCAAG AGACATAACG TTCCGAGCAG

551 CTTTCGGTTC CTCCTCATCC ACATCTGATG AAAGCCGTTT GGACGAGTTC

601 CTTGAGATCA TTCAAGAGTT CTCTAAGCTC TTTGGTGAGT TTAACGTAGC

651 GGATTATGTC CCGTCGTGGC TCAGTTGGAT TGACCCGCAA GGGATAAACG

701 GGCGAGTTGA GAAAGCCCGA AAATCTCTAG ACGGTTTCAT TGAGTCCGTC

751 ATCGATGATC ACTTGCACAA GAAGAAGAGA GAACATGATA ACGTTGACGA

801 AGAGACTGAT ATGGTAGATC AATTACTTGC GTTCTACGAA GAAGAAGTCA

851 AAGTCAACAA CTCAGTCACC AAAATTAATC TCGATAACAT AAAAGGCATC

901 ATCATGGATG TGATGTTCGG AGGAACGGAG ACGGTGGCAT TAGCAATCGA

951 ATGGGTGCTA ACCGAGATAC TCCGGAGCCC CGAGAACATG AAACGGGTCC

1001 AGGACGAGCT AACGAGTGTG GTCGGGCTTG ACAGATGGCG CGTGGAGGAC

1051 ACGCACCTCG AGAAGCTCAC TTTCCTCAAG TGTATCCTCA AGGAGACCCT

1101 CCGGCTCCAC CCGCCTTTCC CTCTCCTCCT CCACGAGACG GTGAAGGACA

1151 CTGAGATTTC CGGTTACTTC ATCCCCAAGG GTTCGCGTGT GATGGTCAAT

1201 ACCTACGCTC TAGGGCGTGA CCCGAATTCT TGGTCCGACC CGGAAAGTTT

1251 TAACCCGGGT AGGTTTTTAA ACCCGATTGC TCCGGATCTG AAGGGGAATA

1301 ATTTCGAATT TGTTCCATTC GGGTCGGGTC GGAGATCGTG CCCGGGTATG

1351 CAACTCGGGT TGTACGCATT TGAGCTTGCG GTGGCTCATC TTCTACACTG

1401 CTTCACGTGG AGTTTACCGG ACGGTATGAA TCCCGGTGAC GTCGACACCG

1451 TTGAAGGACC GGGTCTCACC GTTCCTAAGG CAATTCCTTT GGTGGCGGTG

1501 CCGACCACGC GCCTCCTCTG CCCCATCGTC GTCTCGTGA

COMT - like5 - at1g33030 (SEQ ID NO: 26)

1 ATGGAAGAAC AAAACCTCTC CTCGTACGCA ATGATCTTAT CAAGCTCTTC

51 TGTTCTTCCC ATGGTTCTTA AGACAGCCAT TGATCTTGGT CTCTTTGACA

101 TATTAGCTGA ATCTGGTCCT TCCTCTGCTT CTCAGATCTT TTCTTTGTTG

151 TCTAATGAGA CAAAGAAACA TCATGATTCA AGCTTGGTCA ATCGGATTCT

201 GAGATTTCTT GCGAGTTACT CTATACTCAC ATGCTCTGTT TCTACTGAAC

251 ACGGTGAGCC ATTTGCAATC TATGGCTTAG CTCCTGTTGC CAAGTACTTC

301 ACCAAGAACC AAAATGGAGG TGGATCGTTG GCTCCGATGG TGAATCTGTT

351 TCAGGACAAG GTCGTGACTG ATATGTGGTA CAACCTTAAA GATTCTGTTC

401 TTGAAGGAGG GCTTCCATTC AACAACACTC ACGGTTCAAG CGCGGTTGAA

451 CTGGTAGGCA GTGATTCAAG ATTCAGAGAA GTGTTCCAAA GCTCCATGAA

501 AGGTTTCAAT GAAGTATTCA TAGAAGAATT TCTAAAGAAC TACAATGGTT

551 TTGATGGTGT GAAGTCTTTG GTTGATGTTG GTGGTGGAGA TGGCTCTCTT

601 CTCAGTAGAA TCATCTCTAA GCATACTCAC ATTATCAAAG CTATTAACTT

651 TGATTTGCCC ACTGTTATCA ACACTTCATT ACCTTCTCCA GGAATTGAGC

701 ATGTTGCTGG AGATATGTTT ACAAACACTC CTAAAGGAGA AGCTATTTTC

751 ATGAAGTGGA TGCTCCATAG CTGGGACGAT GACCATTGCG TGAAGATACT

801 TTCCAACTGT TATCAGTCGT TACCGTCAAA TGGGAAGGTG ATTGTAGTCG

851 ATATGGTAAT CCCAGAGTTC CCAGGAGACA CTCTCTTAGA CAGAAGCTTG

901 TTTCAGTTTG AGTTGTTCAT GATGAACATG AACCCATCAG GGAAAGAACG

951 AACAAAGAAA GAATTCGAGA TCTTAGCTCG TCTCGCCGGA TTTCTCTAATG

1001 TCCAAGTTCC ATTCACATCT CTCTGTTTCT CAGTTCTTGA GTTCCACAAG

1051 AACAAGTGA

CAD4 - at3g19450 (SEQ ID NO: 27)

1 ATGGGAAGTG TAGAAGCAGG AGAAAAGAAG GCTTTAGGAT GGGCTGCAAG

51 AGATCCTTCT GGTGTTCTCT CTCCTTACTC CTACACTTTA AGGAGCACAG

101 GAGCAGATGA TGTGTATATA AAGGTTATAT GCTGTGGAAT ATGCCATACT

151 GATATTCACC AAATTAAAAA TGATCTTGGA ATGTCTAATT ATCCTATGGT

201 TCCTGGGCAC GAGGTAGTAG GTGAGGTGTT GGAAGTGGGA TCAGATGTGA

251 GCAAGTTCAC CGTAGGAGAT GTAGTCGGAG TCGGCGTGGT CGTTGGATGC

301 TGCGGTAGCT GCAAACCATG CAGCTCAGAA CTTGAGCAGT ATTGTAACAA

351 GAGAATCTGG AGTTACAACG ATGTCTACAC CGATGGAAAA CCTACTCAAG

401 GTGGCTTCGC CGACACCATG ATCGTCAATC AAAAGTTTGT GGTGAAGATT

451 CCAGAAGGAA TGGCGGTGGA GCAGGCTGCA CCGCTACTCT GCGCGGGAGT

501 AACGGTTTAT AGTCCGTTGA GCCACTTCGG TCTAATGGCG AGTGGGCTTA

551 AAGGAGGGAT TCTAGGGTTA GGTGGAGTAG GGCACATGGG TGTGAAGATA

601 GCTAAAGCAA TGGGACACCA TGTGACTGTG ATAAGCTCAT CAGATAAGAA

651 AAAAGAAGAA GCTATCGAAC ATTTAGGAGC TGACGACTAT GTGGTGAGCT

701 CTGATCCAGC AGAGATGCAA AGACTCGCAG ATTCCCTTGA CTACATCATT

751 GACACCGTAC CGGTGTTTCA CCCTCTCGAC CCTTACTTGG CTTGTCTTAA

801 ACTCGACGGG AAATTGATTC TCATGGGTGT TATCAACACT CCTCTTCAGT

851 TCGTGACCCC TCTCGTCATT CTTGGGAGGA AAGTGATATC AGGGAGCTTT

901 ATAGGGAGCA TTAAAGAGAC AGAAGAAGTG TTGGCTTTCT GCAAAGAGAA

951 AGGGCTTACG TCGACGATTG AAACTGTGAA GATCGATGAG CTTAACATTG

1001 CGTTTGAGAG GCTTCGCAAG AACGATGTTC GTTACAGATT TGTGGTTGAT

1051 GTTGCAGGAA GTAATCTGGT GGAGGAGGCT GCAACAACTA CAAACTAA

CAD5 - at4g34230 (SEQ ID NO: 28)

1 ATGGGAATAA TGGAGGCAGA GAGGAAAACA ACAGGCTGGG CTGCCAGAGA

51 CCCATCTGGG ATCCTCTCTC CTTACACTTA CACTCTTAGA GAGACTGGAC

101 CAGAGGATGT GAACATAAGA ATCATTTGCT GTGGAATCTG CCACACCGAT

151 CTTCATCAAA CTAAAAATGA TCTTGGCATG TCTAATTACC CCATGGTTCC

201 TGGGCATGAA GTGGTAGGGG AAGTAGTGGA GGTGGGATCA GATGTGAGCA

251 AGTTCACCGT AGGGGACATA GTTGGAGTTG GTTGCCTCGT TGGATGTTGC

301 GGAGGTTGTA GCCCCTGCGA GAGAGATCTG GAACAGTATT GTCCAAAGAA

351 GATTTGGAGC TACAATGATG TTTACATCAA TGGTCAACCT ACACAAGGCG

401 GCTTCGCTAA AGCCACCGTC GTTCACCAAA AGTTTGTGGT CAAGATTCCA

451 GAAGGAATGG CGGTTGAGCA GGCTGCGCCG CTACTGTGCG CTGGTGTGAC

501 TGTGTACAGT CCACTGAGCC ACTTTGGTCT GAAACAACCA GGCCTAAGAG

551 GAGGTATACT AGGGTTAGGT GGAGTCGGTC ACATGGGTGT GAAAATAGCC

601 AAAGCAATGG GTCACCATGT GACTGTCATA AGCTCATCAA ACAAGAAGAG

651 AGAAGAGGCA TTGCAAGATC TTGGAGCTGA TGATTACGTG ATCGGATCCG

701 ACCAAGCGAA GATGAGCGAA TTGGCTGATT CGTTGGATTA CGTAATTGAC

751 ACGGTGCCTG TTCATCATGC ACTTGAGCCA TATTTGTCTC TGCTTAAGCT

801 TGATGGTAAA CTCATTCTCA TGGGAGTTAT CAACAATCCA TTACAGTTTC

851 TCACTCCTCT GCTTATGCTT GGGAGGAAAG TGATAACGGG GAGCTTCATA

901 GGGAGCATGA AGGAGACAGA GGAGATGCTT GAGTTCTGTA AAGAAAAGGG

951 TTTGAGTTCG ATTATCGAAG TTGTGAAGAT GGATTATGTT AACACTGCGT

1001 TTGAGAGACT CGAGAAGAAC GATGTGCGTT ATAGGTTCGT CGTTGATGTC

1051 GAAGGAAGCA ATCTCGACGC TTGA

CAD8 - at4g37990 (SEQ ID NO: 29)

1 ATGGGAAAGG TTCTTCAGAA AGAGGCGTTC GGATTAGCCG CGAAAGACAA

51 TTCCGGAGTT CTCTCGCCTT TCAGTTTCAC TAGAAGGGAA ACAGGAGAAA

101 AGGATGTGAG GTTCAAAGTG TTATTTTGTG GAATTTGTCA CTCTGATTTG

151 CATATGGTCA AGAACGAGTG GGGAATGTCT ACTTATCCTC TTGTCCCAGG

201 GCATGAGATC GTGGGCGTGG TGACTGAAGT CGGAGCCAAA GTGACTAAAT

251 TCAAAACCGG AGAAAAAGTC GGAGTTGGGT GTTTGGTCAG CTCGTGCGGG

301 TCATGTGACA GCTGCACCGA AGGTATGGAA AACTATTGTC CAAAATCAAT

351 CCAAACGTAC GGATTCCCGT ACTACGACAA CACCATAACA TACGGTGGTT

401 ACTCCGACCA CATGGTTTGC GAGGAAGGTT TCGTCATCCG TATTCCAGAC

451 AATCTCCCCT TGGACGCCGC CGCACCGCTC CTCTGTGCCG GTATCACGGT

501 CTATTCCCCT ATGAAGTATC ACGGGCTCGA CAAACCCGGT ATGCACATCG

551 GTGTGGTAGG ATTAGGCGGT TTAGGTCATG TAGGAGTGAA ATTTGCCAAG

601 GCTATGGGTA CTAAGGTTAC GGTTATTAGT ACTTCGGAGA AAAAGAGAGA

651 TGAGGCGATT AATCGGCTTG GTGCGGATGC TTTCTTGGTG AGCCGTGACC

701 CAAAACAGAT TAAGGATGCA ATGGGTACTA TGGATGGTAT AATTGATACC

751 GTCTCTGCGA CTCATTCACT TCTTCCGTTG CTCGGTTTGC TGAAGCATAA

801 GGGAAAACTT GTTATGGTTG GTGCACCCGA GAAGCCACTC GAGCTACCTG

851 TCATGCCTCT CATCTTTGAG AGGAAGATGG TAATGGGAAG TATGATAGGA

901 GGGATAAAAG AGACCCAGGA AATGATAGAT ATGGCCGGGA AACACAACAT

951 CACTGCGGAT ATTGAGCTTA TCTCTGCCGA TTATGTCAAC ACCGCCATGG

1001 AACGGCTAGA GAAAGCCGAC GTTAGGTACC GCTTTGTGAT TGATGTTGCC

1051 AACACATTGA AGCCTAATCC TAATTTATAA

Example 2: tally Confirmation of gene expression patterns according to cell types in the site

By confirming the location of the cell wall hydrolase at the detachment site of Arabidopsis of stage 16, we tried to analyze the mechanism that spatially regulates cell wall decomposition in the process of detachment of external organs from flowers. Two other cell wall hydrolyzing proteins, QRT2 and XTH28, were located inside the cell in REC and secreted into the apoptotic cell wall region at the detachment site (SEC) of the dissociating organ (Fig. 1a, b). From this, it was found that the two adjacent cell types have different regulatory mechanisms that regulate the same cell wall degrading enzyme in the process of organ dissociation.

Cell type-specific expression profile analysis was conducted to elucidate the functions of these two cell types in the process of organ detachment. The cells remaining in the detachment site and the detachment site of the separated organ were named as residual cells (RECs, residuum cells) and secession cells (SECs, secession cells), respectively, and Quartet2 ( QRT2 ) , a specific promoter for the detachment site. RECs and SECs were isolated from transgenic plants having the nlsGFP-GUS construct induced by fluorescence-activated cell isolation assay, and their transcripts were analyzed by RNA-sequencing (FIG. 2).

As a result, it was found that RECs and SECs exhibit different gene expression profiles (Fig. 1b, Fig. 2, Fig. 3), and 21 genes showing these differences in expression patterns were expressed by promoter-reporter analysis. It could be reconfirmed (FIG. 4). Through gene ontology analysis, it was found that genes preferentially expressed in RECs are mainly related to metabolism, biosynthesis, and organ development, and genes preferentially expressed in SECs are mainly related to stimuli response and aging (Fig. 1c). Therefore, it was found that RECs and SECs each have their own cell activity profile, and thus play a unique role according to cell type in cell wall processing.

Example 3: SECs Specific lignin formation

RECs are non-epithelial cells in a state that can be converted into surface cells with protective functions, and suberin and lignin are components of the cell protective layer and are formed by protective epithelial cells after organ detachment. Therefore, it was expected that the lignin biosynthesis gene would be expressed in RECs, but as a result of the analysis, it was rather confirmed that the lignin biosynthesis enzyme and laccases involved in lignin polymerization were abundantly expressed in SECs (Fig. 1d, Fig. 5a, b). . To reconfirm this, as a result of observing the location of lignin through cell wall autofluorescence and tissue staining, cell wall autofluorescence was not confirmed in RECs, but strong fluorescence was observed in SECs of isolated flower organs (Fig. 1e). Autofluorescence and phloroglucinol staining were rapidly reduced when flowers were grown in a medium containing piperonylic acid (PA), a lignin biosynthesis inhibitor, and recovered again when monolignol lignin precursor was added (Fig. 5c, d). From these results, it can be seen that lignin is synthesized in SECs, not RECs, and is not located in the protective layer. SECs-specific lignin formation showed the same pattern in various plants (Fig. 5e).

Example 4: SECs Identification of mechanisms for specific lignin formation

The lignin polymerization reaction involves the oxidation of monolignol mediated by peroxidase and/or laccase. In order to confirm the mechanism for SECs-specific lignin formation, salicylhydroxamic acid, a peroxidase inhibitor, was used. As a result, lignin accumulation was inhibited in SECs in the presence of monolignol and PA (FIG. 6a).

Focusing on the fact that peroxidase oxidizes monolignol using H ₂ O ₂ during lignin polymerization, DDC (diethyldithiocarbamate) is used to inhibit ROS-metabolizing enzyme superoxide dismutase (SOD) and , NADPH oxidase (NOX) was inhibited using diphenyleneiodonium (DPI), and both inhibitors inhibited lignin formation in SECs (FIG. 6a).

Accumulation of H ₂ O ₂ in SECs and accumulation of superoxide in RECs indicates that there is a cell type-specific ROS distribution at the site of dissociation, which may play an important role in regulating the determination of cell status. H ₂ O ₂ formation in SECs was a result of the sequential reaction of NOX and SOD ( FIGS. 6c and 7a ), and it was confirmed that 4 genes out of 10 genes encoding NOX proteins were expressed at the detachment site of Arabidopsis thaliana. That is, RbohD and RbohF were expressed in both SECs and RECs, and RbohE and RbohH were expressed in vasculature and nectary cells (Fig. 6b). Lignin formation disappeared in the rbohD /F mutant (Fig. 6d) and was restored when RbohF was introduced into the rbohD /F mutant (Fig. 6d). Lignin deficiency in the rbohD /F mutant delayed dissociation (Fig. 6e). From these results, it was found that the reaction by RbohD/F provides ROS for lignin polymerization in SECs required for proper flower organ detachment.

Example 5: Mechanical barrier function of lignin to cell wall enzyme diffusion

Lignin did not exhibit a function to protect the organs from being detached, so it was hypothesized that its role was to help the organs dissociate. To confirm this, as a result of observing the lignin structure of the SEC, it was possible to confirm 2-3 layers of hexagonal structures with ligninized columns only at the cell edges. as expected (Fig. 8a; Fig. 9). Lignin was shown to firmly support each layer of SEC, but was not observed on top of the first layer of SEC (Fig. 8b), and the cell wall enzyme Pectate Lyase-Like18 (PLL18) was restricted to only the first layer of SEC. In its presence (Figure 8b), lignin prevents diffusion of cell wall enzymes beyond the first layer of SEC, acts as a "brace" to hold the separate layers of cells together, and provides a mechanical barrier to enzyme diffusion. was found to be formed. To confirm the role of lignin, when PA, a lignin biosynthesis inhibitor, was treated on the calyx, it was found that the cell wall enzyme QRT2 was no longer limited to the first SEC layer (FIG. 10). On the other hand, it was confirmed that the diffusion of pectinase was not controlled in the calyx of the rbohD /F mutant without a lignin barrier (Fig. 8c). Combining the above results, it was found that lignin functions as a molecular brace in the cells of the detachment site.

Based on these results, it was expected that the uninhibited diffusion of enzymes that hydrolyze the cell wall would dilute the enzyme concentration at the dissociation site, thereby delaying organ dissociation and also causing organ dissociation at the ectopic site. In order to confirm this, as a result of observing the detachment layer of the rbohD /F mutant, it was confirmed that both sides of the detachment site in the rbohD /F mutant had irregular surfaces that were not observed in the wild type (FIGS. 8d, e). The irregular surface of the rbohD /F mutant is due to the lack of lignin in SECs and RECs to control the diffusion of cell wall degrading enzymes. From these results, the lignin brace only limits the diffusion of cell wall enzymes Rather, it can be seen that mechanical constraints support SECs through precise separation.

Example 6: RECs outside the cell wall cutie Check the distribution pattern

Cell type-specific RNA-seq analysis revealed that RECs were rich in genes related to suberin biosynthesis (Fig. 11a), and suberin was judged to be a major component of the protective layer of the new cell surface . In addition, it was confirmed that genes encoding biosynthetic enzymes and transporters for cutin and/or wax, including ABCG12 / CER5, ABCG11 / WBC11 and ABCG32 / PEC115-17, were highly abundant in RECs (Fig. 11b). . Cutin and suberin are glycerol lipid polymers bound to the cell wall that regulate the flow of gas, water and solutes, and serve to protect plants from environmental stress. It is deposited on the inside of the primary cell wall. As a result of observing the RECs at the detachment site with a transmission electron microscope, it was confirmed that the material with high electron density was located outside the residual cell wall ( FIG. 8f ), and it was confirmed that it was cutin quality.

In order to observe the characteristics of materials with high electron density located outside the residual cell wall, the cell wall changes of RECs were investigated at different time points after desorption. Immediately after flower organ isolation, RECs appeared to have a thin primary cell wall that was permeable to toluidine blue, indicating a defect in the cuticle layer (Fig. 12a-c, left). However, stage 16 flowers start accumulating electron-dense substances on the REC surface within one day, and later penetrate between the cells and become thick enough to cover the entire surface, reducing cell permeability to toluidine blue (Fig. 12a). -c, medium). Materials with high electron density were degraded in plants ectopic expressing cutinase epidermal degeneration factor 1 (CDEF1) (Fig. 12a-c, right). CDEF1 overexpressing plants showed much higher permeability of toluidine blue compared to RECs of wild-type plants (Fig. 12a, right). As a result, it was confirmed that the material with high electron density was cutin.

Since the cell surface cutin formation is a crucial feature of epidermal cells, the above results indicate that non-epidermal RECs undergo transdifferentiation to become epidermal cells during the detachment process, which in RECs is ACR4 , ALE2 , ATML1 , DEK1 , And it is consistent with the fact that genes related to epidermal specification, such as PDF2 , are expressed (FIG. 12d). It is known that the characteristics of epidermal cells are newly acquired in the early embryogenesis stage and maintained throughout the plant life cycle, and the detachment process is considered to be the only period in which the characteristics of epidermal cells are newly specialized after embryogenesis.

We investigated whether SECs remaining on the surface of RECs in rbohD /F mutants lacking lignin affect cutin formation. Unlike the cell wall of RECs (Fig. 8f, 8f-1), the cutinoid structure on the surface of the remaining SECs was found to be defective (Fig. 8f-2). Also, there was no accumulation of cutin at the cell wall junction between RECs and SECs (Fig. 8f-3).

Example 7: rbohD /F Confirmation of pathogen susceptibility due to defects in cutin structure in mutants

Cutin is the first line of defense against biological stress. Therefore, it was confirmed how the defect in the cutin structure in the rbohD/F mutant works to counteract the bacterial pathogen. The fragment containing the exit site in the flower of stage 16 of the rbohD /F mutant was wild-type ( P <0.001; Fig. 8g, zone 1) or stage 16 and 17 ( P >0.5; Fig. 8g, zone 2) of rbohD /F . Pseudomonas syringae compared to other parts of the mutant flower pv . It reacted sensitively to tomato DC3000 ( Pst DC3000). Corresponding to the toluidine blue staining results, the region containing the detachment site of wild-type flowers of stage 16 was more sensitive to pathogens compared to wild-type flowers of stage 17 ( P < 0.05) ( FIG. 12A ). In the rbohD /F mutant flower, the region containing the stage 17 exit site was less sensitive to Pst DC3000 ( P < 0.001) comparable to wild-type ( P >0.5; FIG. 8g, zone 1). Therefore, it was found that the modified cutin structure of the new epidermal surface of the reproductive organ forms an incomplete and permeable barrier by the remaining SECs due to the lack of lignin, and is sensitive to pathogens.

As the specific parts of the present invention have been described in detail above, for those of ordinary skill in the art, it is clear that these specific descriptions are only preferred embodiments, and the scope of the present invention is not limited thereby. will be. Accordingly, it is intended that the substantial scope of the present invention be defined by the claims and their equivalents.

<110> INSTITUTE FOR BASIC SCIENCE Daegu Gyeongbuk Institute of Science and Technology <120> A Method for Controlling Plant Organ Abscission by Regulating Lignin Molecular Brace and a Mutant or Transgenic Plant Using the Same <130> P17-B189 <160> 29 <170 > KoPatentIn 3.0 <210> 1 <211> 2766 <212> DNA <213> Arabidopsis sp. <400> 1 atgaaaatga gacgaggcaa ttcaagtaac gaccatgaac ttgggattct acgaggagct 60 aactcggaca ccaactcgga cacggagagc atcgctagcg accgtggtgc ctttagcggt 120 ccgcttggcc ggcctaaacg tgcgtccaag aaaaacgcaa gattcgccga cgatcttccc 180 aagagaagca atagtgttgc tggcggccgt ggtgatgacg atgagtacgt ggagatcacg 240 ctagacatca gggacgactc ggtggccgtc catagtgtcc aacaagcagc tggaggtgga 300 ggccacctgg aggacccgga gctagccctt cttacgaaga agactctcga gagcagcctc 360 aacaacacca cctccttatc tttcttccga agcacctcct cacgcatcaa gaacgcctcc 420 cgcgagctcc gccgcgtgtt ctctagacgt ccctccccgg ccgtgcggcg gtttgaccgc 480 acgagctccg cggccatcca cgcactcaaa ggtctcaagt tcattgccac caagacggcc 540 gcatggccgg ccgtcgacca acgtttcgat aaactctccg ctgattccaa cggcctctta 600 ctctctgcca agttttggga atgcttagga atgaataagg aatctaaaga cttcgctgac 660 cagctcttta gagcattagc tcgccggaat aacgtctccg gcgatgcaat cacaaaggaa 720 cagcttagga tattctggga acagatctca gacgaaagct ttgatgccaa actccaagtc 780 ttttttgaca tggtggacaa agatgaagat gggcgagtaa cagaagaaga ggtggctgag 840 attattagtc tt agtgcttc tgcaaacaag ctctcaaata ttcaaaagca agccaaagaa 900 tatgcggcac tgataatgga agagttggac ccagacaatg ctgggtttat tatgatcgaa 960 aacttggaaa tgttgctatt acaagcacca aaccagtcgg tgcggatggg agacagcagg 1020 atacttagtc agatgttaag tcagaagctt agaccggcaa aagagagcaa ccctttagtg 1080 agatggtcgg agaaaatcaa atatttcata cttgacaatt ggcagagact atggataatg 1140 atgttatggc ttggcatctg tggtggcctc tttacttata aattcattca gtacaagaac 1200 aaagctgcct atggtgtcat gggttattgc gtttgtgtcg ccaaaggagg cgccgagact 1260 ctcaaattca acatggctct catattgttg cctgtttgtc gaaacaccat cacttggctt 1320 aggaacaaga ctaagcttgg tactgtcgtt ccttttgatg atagtcttaa cttccacaag 1380 gttattgcaa gcgggatagt cgtcggtgtt ttactccatg cgggtgccca tttaacgtgt 1440 gattttccac gtttaattgc cgcggatgag gacacctatg agccgatgga aaaatacttt 1500 ggggatcaac cgactagcta ctggtggttt gtgaaaggag tggaaggatg gactggcatt 1560 gtgatggttg tgctaatggc tatagccttt acactcgcga cgccttggtt ccgacgtaac 1620 aagcttaact tacctaactt cctcaagaag cttaccggtt tcaacgcctt ttggtacacc 1680 caccatttgt tcatcattgt ttatgctctt ctcattgtcc atggtatcaa gctctacctc 1740 acaaagattt ggtatcagaa aacgacatgg atgtatcttg ctgtacccat ccttctatat 1800 gcatcggaga ggctgctccg tgctttcaga tcaagcatca aaccggttaa gatgatcaag 1860 gtggctgttt accccgggaa cgtgttgtct ctacacatga cgaagccaca aggattcaaa 1920 tacaaaagtg gacagttcat gttggtgaac tgccgagccg tatctccatt cgaatggcat 1980 cctttctcaa tcacatcagc tcccggagac gattacctga gcgtacatat ccgcactctc 2040 ggtgactgga cacgtaagct caggaccgtt ttctccgagg tttgcaaacc tcctaccgcc 2100 ggtaaaagcg gtcttctccg agcagacgga ggagatggaa acctcccgtt cccgaaggtc 2160 cttatcgacg gtccatacgg tgctcccgca caagactaca agaaatacga cgtggtactc 2220 ctcgtaggtc tcggcattgg agccacgcct atgatcagta tccttaagga catcatcaac 2280 aacatgaaag gtcctgaccg cgacagcgac attgagaaca ataacagtaa caacaatagt 2340 aaagggttta agacaaggaa agcttatttc tactgggtga ctagggaaca aggatcattc 2400 gagtggttca agggaataat ggacgagatt tcggagttag acgaggaagg aatcatcgag 2460 cttcacaatt attgcacgag tgtgtacgag gaaggtgatg caagagtggc tctcattgcc 2520 atgcttcagt cgttgcaaca cgcta agaac ggtgtggatg ttgtgtcggg tacacgtgtc 2580 aagtcccact tcgctaaacc taactggaga caagtctaca agaagatcgc tgttcaacat 2640 cccggcaaaa gaataggagt cttctactgt ggaatgccag gaatgataaa ggaattaaaa 2700 aatctagctt tggatttttc tcgaaagaca actaccaagt ttgacttcca caaagagaac 2760 ttctag 2766 <210> 2 <211> 2835 <212> DNA <213> Arabidopsis sp. <400> 2 atgaaaccgt tctcaaagaa cgatcggcga cggtggtcat ttgattcagt ttccgccgga 60 aaaaccgccg tcggaagtgc atcaacttca ccgggaactg aatactccat taacggtgat 120 caagagttcg ttgaagtcac aatcgatctt caagacgatg acacaatcgt tcttcgtagc 180 gtcgagccag caaccgccat taatgtcatc ggagatatct ccgacgacaa caccggaata 240 atgactccgg tttcgatttc gagatctccg acgatgaaac gaacttcatc taatcggttc 300 cgacaattct cacaagagct taaagccgaa gctgtggcga aagcgaaaca gttatctcag 360 gagttgaaac gattctcatg gtctcgttct ttctcaggta acttaaccac tactagtacc 420 gccgctaatc aaagcggcgg tgctggtggt ggtttggtga actcggcttt agaagcgcga 480 gcgttgcgaa agcaacgtgc tcagttagat cggactcggt ctagtgctca aagagctctt 540 cgtggtttga gattcattag caataagcaa aagaacgttg atggttggaa cgatgttcaa 600 tcaaatttcg aaaaattcga aaaaaatggt tacatctatc gctccgattt cgctcaatgc 660 ataggaatga aagattcgaa agaatttgca ttggaactgt tcgatgcatt gagtagaaga 720 agaagattaa aagtagagaa aatcaatcac gatgagcttt atgagtattg gtcacaaatc 780 aacgacgaga gttttgattc tcgtctccag atcttcttcg acatagtgga caagaatgaa 840 gatgggagaa tt acagaaga ggaagtaaaa gagataataa tgttgagtgc atctgcaaat 900 aagctatcaa gattaaagga acaagcagag gaatatgcag ctttgattat ggaagagtta 960 gatcctgaaa gacttggcta catagagcta tggcaactag agactttgct tctacaaaaa 1020 gacacatacc tcaattacag tcaagcattg agctatacga gccaagcatt gagccaaaac 1080 cttcaagggt taaggggaaa gagtcgaata catagaatga gttcggattt cgtctacatt 1140 atgcaagaga attggaaaag gatatgggtt ttatccttat ggatcatgat catgatcgga 1200 ttattcttgt ggaaattctt ccaatacaag caaaaagatg catttcatgt gatgggatat 1260 tgtttactca cagccaaagg agcagctgaa acacttaaat tcaacatggc tctaatactt 1320 ttcccagttt gcagaaacac cattacttgg cttagatcca caagactctc ttacttcgtt 1380 ccttttgatg ataatatcaa cttccacaag acaattgctg gagccattgt agtagctgtg 1440 atccttcata ttggagacca tcttgcttgt gatttcccta gaattgttag agccaccgaa 1500 tacgattaca atcggtatct gtttcattac tttcaaacaa aacagccaac atacttcgac 1560 ctcgttaagg gacctgaagg aatcactggg attttaatgg tcattttgat gattatttca 1620 ttcacattag caacaagatg gtttaggcgt aacctagtca agcttcctaa gccatttgat 1680 cgactaaccg gttttaacgc cttttggtat tcgcatcatt tgttcgtcat tgtttatatc 1740 ttgcttattc ttcatggtat cttcctctat ttcgccaagc cttggtatgt tcgtacgaca 1800 tggatgtatc ttgcagtacc agttttactc tatggtggag aaagaacact taggtacttc 1860 cgttctggtt cttattcggt tcgactgctt aaggttgcta tatatcctgg taatgttcta 1920 acgctacaaa tgtcgaaacc aactcaattt cgttacaaaa gcggacaata catgtttgtc 1980 caatgtcctg cggtttcgcc attcgagtgg catccattct caattacttc cgcacctgaa 2040 gatgattata tcagcattca cattagacaa cttggtgatt ggactcaaga actcaaaaga 2100 gtattctctg aagtttgtga gccaccggtt ggcggtaaaa gcggacttct cagagccgac 2160 gaaacaacaa agaaaagttt gccaaagcta ttgatagatg gaccgtacgg tgcaccagca 2220 caagattata ggaaatatga tgttctctta ttagttggtc ttggcattgg tgcaactcca 2280 tttatcagta tcttgaaaga tttgcttaac aacattgtta aaatggaaga gcatgcggat 2340 tcgatctcgg atttcagtag atcatcagaa tacagcacag gaagcaacgg tgacacgcca 2400 agacgaaaga gaatactaaa aaccacaaat gcttatttct actgggtcac aagagaacaa 2460 ggctcttttg attggttcaa aggtgtcatg aacgaagttg cagaacttga ccaacggggt 2520 gtgatagaga tgcataacta tttaa caagt gtgtatgaag aaggtgatgc tcgttctgct 2580 ctcattacaa tggttcaagc tcttaatcat gccaaaaatg gtgtcgacat tgtctctggc 2640 actagggtca gaacacactt tgcaagacct aattggaaga aggttctcac aaagctaagt 2700 tccaagcatt gcaatgcaag aataggagtg ttttattgcg gagtaccggt tttagggaag 2760 gagcttagca aactatgcaa cacattcaat caaaaaggtt caaccaagtt tgaatttcac 2820 aaggagcatt tctaa 2835 <210> 3 <211> 981 <212> DNA <213 > Arabidopsis sp. <400> 3 atgaattgct tgatagctat agctctttca gtctctttct ttcttgtggg aattgttgga 60 ccgatccaag ctcaattgca gatgaatttc tatgccaatt cttgtcctaa tgctgaaaag 120 attgttcaag attttgtttc aaaccacgtt tctaatgctc cttctcttgc tgctgctctc 180 attagaatgc atttccatga ctgttttgtc cgaggttgtg atggatcagt gcttataaac 240 tcaacgtcag gaaacgcaga gagagacgcg actcctaacc taacggttcg agggtttggc 300 ttcatcgacg caattaaatc tgtgcttgaa gctcaatgcc ctggaattgt ctcttgcgct 360 gatattatcg ctctagcttc tcgcgacgct gtcgttttca ccggaggacc gaattggagt 420 gtaccgaccg gaagaagaga cgggaggata tcaaacgcag cggaggcatt agccaacatt 480 cctcctccaa ccagtaatat caccaatctt cagacactct ttgcaaacca aggacttgat 540 cttaaggacc tcgttttact atccggggct cacactattg gtgtatctca ctgctcgtct 600 ttcacaaacc gtctctacaa ttttacgggt cgtggaggcc aagatccggc tttggacagc 660 gagtacgcag ccaatctcaa gtctagaaaa tgtcctagcc tcaacgataa caagaccatc 720 gtagagatgg atccagggag ccgcaaaaca tttgatctaa gttattacca gctcgtactc 780 aagcgtagag gtctgtttca atcagactct gctctcacca ctaaccccac aacactttca 840 aacataaacc gg atcttgac gggttcggtg gggagtttct tctctgagtt tgccaagtca 900 atggagaaaa tgggtcggat caatgtcaag actggttcag ctggagtggt taggaggcaa 960 tgttccgtt 213 Arabidopsis sp. 4 <400> 4 atggctatct caaagcttat tcctactctt gtgctctttg ttttgttctc ttttgatgtt 60 agtgttgctc atccaggtct aggatttgga tggggaagta atagtcccat tggaggatct 120 ttttactcaa atctttaccc acaattttac cagttctctt gtccacaagc tgatgagatt 180 gttatgacgg tgctcgaaaa agccatagct aaagaaccaa gaatggcagc atctttactc 240 agacttcact tccacgactg cttcgttcag ggctgtgatg catcaatctt gttggatgat 300 agtgcaacca taagaagtga aaagaatgct ggaccaaaca agaactccgt tagagggttt 360 caagtaatcg acgagatcaa agccaaactt gagcaagctt gtcctcaaac tgtctcctgc 420 gccgacattc tagctctagc tgctcgtggc tcaactatac tgagtggtgg accatcatgg 480 gagcttccac tagggaggag agactcgagg accgctagcc ttaatggcgc aaacacgaac 540 attcctgcac ctaactcaac tattcaaaat ctcttaacca tgttccaacg taaaggtttg 600 aacgaagaag accttgtttc cctatcagga ggacatacga ttggagtagc gaggtgcaca 660 acgtttaaac aaagactata caatcaaaac ggtaataacc aaccagacga gacgctagag 720 aggtcttact actatggtct caggtcaatt tgtccaccaa caggcggtga caacaatatt 780 tcacctctag acttagcctc tcctgcaaga ttcgacaaca catacttcaa gctccttcta 840 tggggcaaag gc ttattgac atcagatgaa gtgcttctca ctggcaatgt aggcaagact 900 ggtgcattag tgaaagctta tgccgaagac gagagacttt tcttccaaca attcgcaaag 960 tcgatggtta acatgggaaa cattcagcct cttactggtt ttaatggtga gatcaggaag 1020 agttgtcatg tgattaacta a 1041 <210> 5 <211> 1050 <212> DNA <213> Arabidopsis sp. <400> 5 atggggtttt ctccttcatt ttcttgcagt gctattggag ccctaatttt gggttgcctt 60 ctgcttcaag catcaaactc taatgctcag ttgaggcctg atttctactt cgggacgtgc 120 ccatttgttt ttgatatcat tggaaatatc atcgtcgatg aactgcagac tgatcctcgt 180 attgccgcga gcctccttcg tcttcacttt cacgactgct ttgttcgtgg ttgtgatgca 240 tcgatcttgc ttgacaattc cacatcattc cggactgaga aagatgctgc tccgaacgca 300 aattcggctc gagggtttaa tgtcatagat agaatgaaag tagcccttga gagagcttgc 360 ccaggaagag tctcttgtgc agatattctc accatcgcct ctcaaatatc agtgcttttg 420 tcgggaggtc catggtggcc ggttccgttg gggaggagag acagtgtaga agctttcttc 480 gctctggcta atacagctct gccctctccg tttttcaatc ttactcaact taaaacagcc 540 tttgctgacg ttggcctaaa ccgcacttcc gatctagttg ctctttccgg tggtcacaca 600 tttggaagag cacaatgcca gttcgtgacg cctcgtctct acaacttcaa tggtacaaac 660 agtccagacc caagtctgaa cccaacttac cttgtcgaac tccgtcgatt gtgtcctcaa 720 aacggaaacg gcaccgttct ggtcaacttc gatgtcgtga ctccagatgc tttcgatagt 780 caatactaca ccaaccttcg taatgggaaa ggtcttattc agagtgacca ggaactcttt 840 tcgactccag ga gccgatac gatcccttta gtaaaccaat acagcagcga catgtccgtg 900 ttctttcgag cattcattga cgcaatgatt aggatgggaa atcttagacc tttgactgga 960 actcaaggag agattagaca aaattgtagg gttgtgaatc cacgaattag ggttgtggag 1020 aacgacgatg gtgttgtgag ttctatctga 1050 <210> 6 <211> 1062 <212> DNA <213> Arabidopsis sp. <400> 6 atgcatttct cttcgtcttc aacatcgtcc acttggacaa tcttaatcac attgggatgt 60 cttatgcttc atgcatcttt gtccgctgct caactcaccc ctaccttcta cgataggtca 120 tgtcctaatg tcactaacat cgtacgagaa accattgtaa atgagttaag gtcggaccct 180 cgtatcgctg cgagcatcct tcgtcttcac ttccacgact gctttgttaa tggttgtgac 240 gcatccatct tgttagacaa cacgacatca tttcgaacag agaaagatgc gtttggaaac 300 gcaaattcgg ctcggggatt tccagtgatt gatagaatga aagctgcggt ggagagggca 360 tgcccaagaa ccgtttcatg cgcagatatg ctcaccattg cagctcaaca atctgtcact 420 ttggcaggag gtccttcttg gagggttcct ttgggaagga gagacagttt acaagcattc 480 ctggaactcg ctaatgcaaa tcttccagct ccattcttta cacttccaca acttaaagcc 540 agcttcagaa atgttggtct cgatcgtcct tctgatctcg ttgctctctc cggtggtcac 600 acatttggta aaaatcaatg tcagtttatt cttgacagat tatacaattt cagcaacaca 660 ggtttacccg accctacact caacactact tacctccaaa ctcttcgtgg actatgcccc 720 cttaatggca atcgaagtgc cttggtagat tttgatctac gtacgcctac ggttttcgac 780 aacaaatact acgtgaatct caaagagcga aaaggtctta tccagagcga ccaagagttg 840 ttctctagcc cc aatgccac tgacacaatc cccttggtga gagcatatgc tgatggcaca 900 caaacattct tcaatgcatt tgtggaggca atgaatagga tgggaaacat tacaccaact 960 acaggaactc aaggacaaat cagattgaac tgtagagttg tgaactccaa ctctctgctc 1020 catgatgtgg tggatatcgt tgactttgtt agctctatgt ga 1062 <210> 7 <211> 978 <212> DNA <213> Arabidopsis sp. <400> 7 atgttgacta gatttaagaa acaaaataat aagatggtta gggcaaatat agtgagcatg 60 gttctgttga tgcatgccat cgttgggttt ccttttcatg cgaggggatt aagtatgact 120 tactacatga tgagctgtcc tttcgctgaa cagattgtga agaacagtgt taacaatgct 180 cttcaagccg atcccacttt agccgctggt cttatccgta tgttgttcca cgactgtttc 240 atcgagggat gtgatgcgtc aattctgcta gattcaacga aagacaacac tgcggaaaag 300 gattcgcctg cgaatctgag tctacgtggc tacgagatca tcgatgatgc aaaggagaaa 360 atcgagaata gatgtccagg agttgtatca tgcgcagata tagttgccat ggctgctaga 420 gatgctgtct tttgggctgg tggtccatat tatgacatac cgaaaggaag gtttgatggt 480 aaaagatcaa agatagaaga tacaagaaat ctaccttcac cttttcttaa tgcctctcaa 540 ctcattcaaa cttttggcca acgtggtttc actccacaag atgttgttgc tctctctgga 600 gcacataccc taggagttgc acgatgctcc tccttcaagg ctagacttac cgtcccagac 660 tcttcattag actcgacttt tgcaaacact ctctctaaaa catgcagcgc gggtgacaat 720 gcagagcaac cctttgatgc tacgcgcaac gatttcgaca atgcttactt caatgcgctt 780 cagatgaaat caggagtcct cttttcagac caaactttat tcaacacacc aaggaccagg 840 aatcttgtta ac ggctatgc acttaaccaa gccaagttct tctttgattt ccaacaggcc 900 atgcgcaaga tgagcaatct tgatgttaaa cttggctctc aaggtgaagt ccgtcaaaat 960 tgtcggagta 213 ttaattag 978 <210> 8 <211> sp990 <400> 8 atggttgtgg tgaacaaaac gaatctctta ttgcttcttc tctctctatg tctcaccctc 60 gacctctcct ccgcacaact ccgccgtaac ttctacgccg gaagttgtcc taacgtcgaa 120 caaatcgtta gaaacgccgt acagaaaaaa gtccaacaaa ctttcaccac cattcccgcc 180 actctacgcc tctatttcca tgattgtttc gtcaatgggt gtgatgcatc agtgatgata 240 gcgtcaacaa ataataataa ggcggagaag gatcatgagg aaaatttatc gttggccgga 300 gatggattcg acaccgtcat taaagctaaa gaagctctcg atgccgtccc aaattgtcgt 360 aacaaagttt catgcgctga tattctcacg atggccactc gtgacgtcgt taatcttgcg 420 ggtggaccac aatacgacgt ggaattggga aggctagatg gactatcatc aacagcggct 480 agcgtcggag ggaagttgcc gcatccgacc gacgatgtca ataaactcac ttcactcttc 540 gccaaaaacg gccttagcct taacgacatg atcgctctct ccggggcaca cacattagga 600 ttcgcgcatt gcacaaaagt gttcaatcgg atatacactt tcaacaagac aactaaagtg 660 gatcccacgg ttaacaaaga ttacgtgaca gagctaaaag cgtcgtgtcc tcgaaacata 720 gatccaagag tggctattaa catggaccca acaacaccaa gacaattcga caacgtttac 780 tacaaaaact tgcaacaagg gaaaggattg ttcacgtcgg atcaagtctt attcacggat 840 cgtcggtcaa ag ccaaccgt tgacttatgg gctaataatg gacagttgtt taatcaagct 900 tttattaact cgatgatcaa gcttggtcgt gttggtgtta aaactggtag taatggtaat 960 atccgtcgtg attgtggido Arab psi 990. <400> 9 atggttgtga tgaacaaaac gaatctcttg ttgcttattc tctctctttt tctcgccatc 60 aacctctcct ccgctcaact ccgcggcgac ttctacgccg gaacttgtcc gaacgtcgag 120 caaatcgtca gaaacgccgt acagaaaaaa atccaacaaa ctttcaccac cattcccgcc 180 actctacgcc tctatttcca tgattgtttc gtcaatggat gtgatgcatc agtgatgata 240 gcgtcgacaa atactaataa ggcggagaag gatcatgagg ataatttatc gttggccgga 300 gatggattcg acaccgtcat taaagctaaa gaagctgttg acgccgtccc aaattgtcgt 360 aacaaagttt cttgcgctga tattctcacg atggccactc gtgacgtcgt taatcttgcg 420 ggtggaccac aatacgccgt agaattggga aggcgagatg gactttcgtc gtcggcttct 480 agcgtcacag ggaagctacc caaaccgacc tttgatctca accaactcaa tgcacttttc 540 gcagaaaacg gtcttagccc taatgacatg atcgctctct ccggggcaca cacattagga 600 ttcgcgcatt gcacaaaagt tttcaatcgg ctatacaatt tcaacaagac aaataacgtg 660 gatcccacga ttaacaaaga ttacgtgaca gagctaaaag cgtcgtgtcc tcaaaacata 720 gatccaagag tggctattaa catggatccc aatacaccaa gacaattcga caacgtttac 780 tacaaaaact tgcaacaagg gaaaggattg ttcacgtctg atcaagtctt attcaccgat 840 agtcggtcaa ag ccaaccgt tgacttatgg gctaataatg gacagttgtt taatcaagct 900 tttattagct cgatgatcaa gcttggtcgt gttggtgtta aaactggtag taatggtaat 960 attcgtcgtg attgcggagc ttttcaatta < 990 <210> DNA <212> Arabidopsis sp. <400> 10 atggcttcaa ataaacttat ttcgattcta gttctcgtag ttacactttt actccaaggt 60 gataacaatt acgtcgtcga agcacaactc acgactaact tctactcaac ctcttgcccc 120 aacctcctct ccaccgtcca aaccgccgtt aaatctgccg ttaacagcga ggctcgaatg 180 ggtgcatcta tcctccgtct cttcttccac gattgctttg tcaacggatg cgacggttcg 240 attctactag acgatacgtc aagcttcaca ggagaacaaa acgcggctcc aaaccgcaat 300 tcagctcgtg ggtttaatgt gatcgacaac atcaaatcag cggttgagaa agcatgtccc 360 ggggttgtgt cttgtgctga tatcttagcc attgctgcta gagactccgt cgtagccctt 420 ggaggaccga actggaatgt gaaagtagga agaagagatg caagaacggc gagtcaagcg 480 gcagcgaata gtaacattcc ggcaccgact tcgagtctga gccaactcat tagtagtttc 540 agtgctgttg gactctccac cagagatatg gttgctctct ctggcgcaca cacgatcgga 600 caatcccgtt gcacgaactt ccgagcgaga atctacaacg agacaaacat caatgccgcc 660 ttcgccacga cacgtcaacg aacttgccct agagcctccg gttccggcga cggaaaccta 720 gctccactag acgtcaccac ggcagcttct ttcgacaaca actacttcaa gaatctcatg 780 actcagagag ggctcctcca ttccgaccaa gtgctcttca acggcggctc tactgactcc 840 atcgtccgtg gatacagcaa caatccgtcg agctttaact ctgacttcac ggcggcgatg 900 ataaaaatgg gtgatatcag cccgttgact ggtagtagcg gtgaaatccg caaggtttgc 960 ggtgaggacca actga 975 <210> 11 <211> sp. <400> 11 atgggcttgg tccgatcatt tgctttggtc atagtcttcc ttagttgtct catcgccgtt 60 tatggccaag gtacgaggat cgggttctac tcgactacgt gccctaatgc cgagacaatt 120 gttcggacca ctgtggcatc tcactttggt tcagatccaa aggttgcacc cgggttactg 180 agaatgcaca accatgattg ctttgtccaa ggttgtgatg gttcggtgct tttatcggga 240 cctaactctg aaagaaccgc tggcgcaaac gttaacctcc atggttttga ggtcattgac 300 gatgccaaga ggcagctcga ggcagcatgt cctggtgttg tctcttgtgc tgatatcttg 360 gccttagcgg ctcgtgattc ggtttctctc acaaacggac aaagttggca agttccaaca 420 ggacgtagag atggaagagt ttccttggca tcgaacgtta acaatcttcc ttctccaagt 480 gactctctag ccattcaaca aaggaaattc tccgcttttc gcctcaacac tcgcgatctc 540 gtcactcttg ttggaggagg acacacgatc ggaacagctg catgtgggtt tatcacgaac 600 aggatattca actcgagcgg aaacacagca gatccaacaa tggaccaaac atttgtacca 660 caacttcaaa gactttgtcc ccaaaacggc gacggatcag cacgtgttga tcttgacacc 720 ggaagtggaa acacttttga cacatcttat ttcatcaatc tcagccgtaa cagaggaatt 780 cttcaatccg atcacgttct ttggactagt ccggccacaa gatccatagt gcaagagttt 840 atggcaccta g aggcaactt caatgtccag ttcgcaaggt caatggttaa aatgagtaat 900 attggtgtga agacggggac aaatggagaa attcgtaggg tttgctctgc ggttaattaa 960 960 <210> 12 <211> 459 <212> DNA <213> Arabidopsis sp. <400> 12 atggcgaaag gagttgcagt tttgaacagc agtgagggtg ttacggggac tatctttttc 60 acccaggaag gcgatggtgt gaccactgtg agtggaacag tttctggcct taagcctggt 120 cttcatggtt tccatgtcca tgctcttggt gacaccacta acggttgcat gtctactggt 180 ccacatttca accccgatgg taaaacacac ggtgcccctg aggatgctaa tcgacatgct 240 ggtgatctag gaaacatcac tgttggagat gatggaactg ccaccttcac aatcactgat 300 tgccagattc ctcttactgg accaaactct attgttggta gggctgttgt tgtccatgca 360 gaccctgatg acctcggaaa gggaggccat gaactcagcc tggctactgg aaacgcaggc 420 ggccgtgttg cttgcggcat cattggtctc cagggctaa 459 <210> 13 <211> 651 <212> DNA <213> Arabidopsis sp. <400> 13 atggctgcca ccaacacaat cctcgcattc tcatctcctt ctcgtcttct cattcctcct 60 tcctccaatc cttcaactct ccgttcctct ttccgcggcg tctctctcaa caacaacaat 120 ctccaccgtc tccaatctgt ttccttcgcc gttaaagctc cgtcgaaagc gttgacagtt 180 gtttccgcgg cgaagaaggc tgttgcagtg cttaaaggta cttctgatgt cgaaggagtt 240 gttactttga cccaagatga ctcaggtcct acaactgtga atgttcgtat cactggtctc 300 actccagggc ctcatggatt tcatctccat gagtttggtg atacaactaa tggatgtatc 360 tcaacaggac cacatttcaa ccctaacaac atgacacacg gagctccaga agatgagtgc 420 cgtcatgcgg gtgacctggg aaacataaat gccaatgccg atggcgtggc agaaacaaca 480 atagtggaca atcagattcc tctgactggt cctaattctg ttgttggaag agcctttgtg 540 gttcacgagc ttaaggatga cctcggaaag ggtggccatg agcttagtct gaccactgga 600 aacgcaggcg ggagattggc atgtggtgtg attggcttga cgccgctcta a 651 <210> 14 <211> 495 <212> DNA <213> Arabidopsis sp. <400> 14 atggaagctc ctagaggaaa tctgagagcg gtggctctta tcgccggcga taacaacgtc 60 cgaggctgtc ttcaattcgt acaagacatt tccggaacta ctcatgtcac cgggaagatc 120 tcaggactct ctcctggctt ccatggattt catattcact cttttggtga taccacaaat 180 ggctgcatct ctactggacc tcacttcaat ccattaaatc gagttcacgg gccaccgaac 240 gaggaagagc gtcatgctgg cgatttgggt aacattcttg caggatcaaa cggtgttgct 300 gaaatcttaa tcaaagacaa acatataccg cttagtgggc agtattccat actcgggagg 360 gcggttgttg tgcatgcgga tcctgatgac cttgggaaag gagggcacaa gcttagcaaa 420 tcaactggaa acgcaggctc gagagttgga tgtggtatca tcggacttca atcatctgcg 480 gatgctaaac tatag 495 <210> 15 <211> 639 <212> DNA <213> Arabidopsis sp. <400> 15 atggctgctt caagtgctgt caccgcaaac tacgtcctca agccacctcc attcgcactg 60 gatgctttgg agccgcatat gagcaaacaa actctggagt ttcactgggg aaaacatcac 120 agagcttacg tggacaacct caagaaacag gttcttggaa ccgagcttga aggcaagccc 180 ttagagcaca ttatccacag cacttacaac aatggtgatc tcctccctgc tttcaacaac 240 gctgctcagg cgtggaacca cgagttcttc tgggagtcaa tgaaaccagg tggtggagga 300 aaaccatcag gagagcttct tgctttgctt gaaagagatt tcacttctta tgagaagttc 360 tatgaagagt tcaatgctgc tgcagccact cagtttggag ctggctgggc ctggcttgct 420 tattcaaatg aaaaactcaa agtagtgaaa actcccaatg ctgtgaatcc ccttgtgctc 480 ggctctttcc cattgcttac cattgatgtc tgggagcatg cttactacct tgacttccag 540 aaccgaagac cagattacat aaagacattc atgaccaatc ttgtgtcttg ggaagctgta 600 agtgccagac ttgaggccgc caaggctgct tctgcttaa 639 <210> 16 <211> 918 <212> DNA <213> Arabidopsis sp. <400> 16 atgatgaatg ttgcagtgac agccactccc tcgtctctct tgtactctcc tctgcttctt 60 ccttctcaag ggccaaaccg gcgaatgcaa tggaaaagaa acggaaagag acggttaggg 120 acaaaggtgg ctgtttccgg tgttatcaca gctggatttg agctgaagcc acctccatat 180 cctcttgatg ctctggaacc gcatatgagc cgggaaacct tggattatca ctggggcaaa 240 catcacaaaa cttatgtaga gaacctgaac aagcaaatct taggcacgga tctagatgca 300 ttatccttgg aagaagttgt gcttctttca tacaacaaag gcaatatgct tcctgctttc 360 aacaacgctg cacaggcttg gaaccacgag ttcttctggg agtctatcca acctggaggt 420 ggaggaaagc caactggaga gctcctcaga ttaatagaaa gagattttgg gtctttcgaa 480 gagtttttgg aaaggttcaa gtcggctgca gcttcgaatt ttggttcggg ttggacatgg 540 cttgcatata aggcgaatag acttgacgtt gcaaatgccg ttaatcctct cccaaaggag 600 gaagacaaga aacttgttat agtgaagacg cccaatgcag taaatccgct cgtatgggat 660 tattctccac ttctcaccat tgatacctgg gagcacgctt actatctgga ttttgagaac 720 cgaagagctg aatacataaa tacattcatg gaaaagcttg tgtcatggga aactgtaagc 780 acaaggttgg aatccgcaat tgctcgagca gtgcaaagag aacaagaagg aacagagaca 840 gaagatgaag a gaatccaga tgatgaagta ccagaggtct atttagatag tgacatcgat 900 gtatctgagg ttgactaa 918 <210> 17 <211> 792 <212> DNA <213> Arabidopsis sp. <400> 17 atgagttctt gtgttgtgac gacaagctgt ttctatacaa tttcagattc tagtatacgt 60 ttgaaatccc ccaagctcct caatctgagt aaccagcaga gaagacgctc tcttaggtct 120 cgaggtggtt taaaggttga agcttactac ggtctaaaga cacctcctta tccacttgat 180 gctttggagc cgtatatgag tagaagaaca ctagaagtgc attggggaaa acaccatcga 240 ggttatgtag ataatctgaa taaacagtta gggaaagatg atagactcta tggatacacc 300 atggaagagc ttatcaaggc tacatacaac aacgggaatc ctttacccga gttcaacaac 360 gctgcacagg tctataacca tgatttcttc tgggagtcga tgcaacctgg tggtggagac 420 acgcctcaaa agggtgttct tgagcagatt gataaggatt ttggttcttt cacaaatttt 480 agagaaaagt tcactaatgc agctcttact cagtttggtt ctggatgggt ctggcttgtc 540 ttaaagaggg aagagagaag gcttgaggtg gtcaaaacct caaacgccat taacccactc 600 gtgtgggacg atattccaat catctgcgtg gatgtgtggg agcactctta ttatctggac 660 tacaagaacg acagggctaa gtatataaac acatttctga accacttggt gtcgtggaac 720 gctgccatga gtcggatggc ccgtgcagaa gcgtttgtga atcttggtga acccaacatc 780 ccaatcgctt aa 792 <210 > 18 <211> 696 <212> DNA <213> Arabidopsis sp. <400> 18 atggcgattc gttgtgtagc gagtagaaaa accctagccg gcttgaagga gacatcatcg 60 aggctattga ggatcagagg gattcagact tttacgcttc ctgatcttcc ttacgattat 120 ggcgcattgg aaccggccat tagtggagag atcatgcaga ttcatcacca gaagcatcac 180 caggcttatg ttactaatta caataatgct cttgagcagc ttgatcaagc tgtgaacaag 240 ggagatgctt ccactgttgt taagttgcag agcgccatca aattcaacgg cggaggtcat 300 gtcaaccatt cgattttctg gaagaacctt gctccttcca gtgaaggtgg tggagagcca 360 ccaaaaggat ctcttggtag tgccattgac gctcactttg gctcccttga aggtctggtg 420 aaaaagatga gtgctgaggg tgctgcagtg caaggctcag gatgggtgtg gctcggacta 480 gacaaagaac tgaagaagct agttgttgac acaactgcca atcaggatcc attagtgaca 540 aaaggaggaa gcttggtacc tctggtgggt atagatgttt gggagcacgc ctactacttg 600 cagtacaaaa atgtgaggcc tgagtatctg aagaatgtat ggaaagtgat caactggaaa 660 tatgcaagcg aggtttatga gaaggaaaac aactga 696 <210> 19 <211> 2178 <212> DNA <213 > Arabidopsis sp. <400> 19 atggagatta acggggcaca caagagcaac ggaggaggag tggacgctat gttatgcggc 60 ggagacatca agacaaagaa catggtgatc aacgcggagg atcctctcaa ctggggagct 120 gcagcggagc aaatgaaagg tagccatttg gatgaagtga agagaatggt tgctgagttt 180 aggaagccag ttgtgaatct tggtggtgag actctgacca ttggacaagt ggctgcgatc 240 tcaactattg gtaacagtgt gaaggtggag ctatcggaga cagctagagc cggtgtgaat 300 gctagtagtg attgggttat ggagagtatg aacaaaggca ctgatagtta tggtgttact 360 actggttttg gtgctacttc tcatcggaga accaaaaacg gtgtcgcact tcagaaggaa 420 cttattagat tccttaacgc cggaatattc ggaagcacga aagaaacaag ccacacattg 480 ccacactccg ccacaagagc cgccatgctt gtacgaatca acactctcct ccaaggattt 540 tccggtatcc gatttgagat tctcgaagca attaccagtt tcctcaacaa caacatcact 600 ccatctctcc ccctccgtgg tacaatcacc gcctccggag atctcgttcc tctctcctac 660 atcgccggac ttctcaccgg tcgtcccaat tccaaagcta ctggtcccaa cggtgaagct 720 ttaacagcag aggaagcttt caaattagca ggaatcagct ccggattctt tgatctccag 780 cctaaggaag gtctcgcgct agtcaatggc acggcggttg gatctggaat ggcgtcaatg 840 gtgttattcg a aacgaatgt tctctctgtt ttggctgaga ttttgtcggc ggttttcgca 900 gaggtgatga gtggtaagcc tgagttcacc gatcatctca ctcacagact taaacatcat 960 cccggtcaaa tcgaagcggc ggcgataatg gagcatatcc tcgacggaag ctcgtacatg 1020 aaattagctc agaagcttca cgagatggat ccgttacaga aacctaaaca agatcgttac 1080 gctcttcgta cttctcctca atggttaggt cctcaaatcg aagtgatccg ttacgcaacg 1140 aaatcgatcg agcgtgagat taactccgtc aacgataatc cgttgatcga tgtttcgagg 1200 aacaaggcga ttcacggtgg taacttccaa ggaacaccaa tcggagtttc aatggataac 1260 acgagattgg cgatagcagc gattggtaaa ctcatgtttg ctcaattctc agagcttgtg 1320 aatgatttct acaacaatgg tttaccctcg aatctaaccg cttcgaggaa tccaagtttg 1380 gattatggat tcaagggagc tgagattgca atggcttctt attgttcaga gcttcaatac 1440 ttagctaatc ctgtgactag ccatgttcaa tcagcagagc aacataacca agatgtcaac 1500 tctttgggac taatctcgtc tcgcaaaact tctgaagctg ttgatattct caagcttatg 1560 tcaacaacgt tcctcgttgc gatttgtcaa gctgtggatt tgagacattt ggaggagaat 1620 ttgagacaga ctgtgaagaa cactgtctct caagtggcga agaaagttct tactactgga 1680 gtcaatggtg agcttcatc c ttctcgcttc tgcgaaaagg atttactcaa agttgtagac 1740 cgtgaacaag tctacacata cgcggatgat ccttgtagcg caacgtaccc gttgattcag 1800 aagctgagac aagttattgt tgaccatgct ttgatcaatg gtgagagtga gaagaatgca 1860 gtgacttcaa tcttccataa gattggagct ttcgaggagg agcttaaggc agtgctaccg 1920 aaagaagtgg aagcagcaag agcagcctac gataacggaa catcggctat cccgaacagg 1980 atcaaggaat gtaggtcgta tccattgtat agattcgtga gggaagagct tggaacagag 2040 cttttgaccg gagagaaagt gacgtcgcct ggagaagagt tcgacaaggt tttcacggcg 2100 atttgtgaag gtaaaatcat tgatccgatg atggaatgtc tcaacgagtg gaacggagct 2160 cccattccaa tatgttaa 2178 <210> 20 <211> 1518 <212> DNA <213> Arabidopsis sp. <400> 20 atggacctcc tcttgctgga gaagtcttta atcgccgtct tcgtggcggt gattctcgcc 60 acggtgattt caaagctccg cggcaagaaa ttgaagctac ctccaggtcc tataccaatt 120 ccgatcttcg gaaactggct tcaagtcgga gatgatctca accaccgtaa tctcgtcgat 180 tacgctaaga aattcggcga tctcttcctc ctccgtatgg gtcagcgaaa cctagtcgtc 240 gtctcctcac cggatctaac aaaggaagtg ctcctcactc aaggcgttga gtttggatcc 300 agaacgagaa acgtcgtgtt cgacattttc accgggaaag gtcaagatat ggtgttcact 360 gtttacggcg agcattggag gaagatgaga agaatcatga cggttccttt cttcaccaac 420 aaagttgttc aacagaatcg tgaaggttgg gagtttgaag cagctagtgt tgttgaagat 480 gttaagaaga atccagattc tgctacgaaa ggaatcgtgt tgaggaaacg tttgcaattg 540 atgatgtata acaatatgtt ccgtatcatg ttcgatagaa gatttgagag tgaggatgat 600 cctcttttcc ttaggcttaa ggctttgaat ggtgagagaa gtcgattagc tcagagcttt 660 gagtataact atggagattt cattcctatc cttagaccat tcctcagagg ctatttgaag 720 atttgtcaag atgtgaaaga tcgaagaatc gctcttttca agaagtactt tgttgatgag 780 aggaagcaaa ttgcgagttc taagcctaca ggtagtgaag gattgaaatg tgccattgat 840 cacatccttg a agctgagca gaagggagaa atcaacgagg acaatgttct ttacatcgtc 900 gagaacatca atgtcgccgc gattgagaca acattgtggt ctatcgagtg gggaattgca 960 gagctagtga accatcctga aatccagagt aagctaagga acgaactcga cacagttctt 1020 ggaccgggtg tgcaagtcac cgagcctgat cttcacaaac ttccatacct tcaagctgtg 1080 gttaaggaga ctcttcgtct gagaatggcg attcctctcc tcgtgcctca catgaacctc 1140 catgatgcga agctcgctgg ctacgatatc ccagcagaaa gcaaaatcct tgttaatgct 1200 tggtggctag caaacaaccc caacagctgg aagaagcctg aagagtttag accagagagg 1260 ttctttgaag aagaatcgca cgtggaagct aacggtaatg acttcaggta tgtgccattt 1320 ggtgttggac gtcgaagctg tcccgggatt atattggcat tgcctatttt ggggatcacc 1380 attggtagga tggtccagaa cttcgagctt cttcctcctc caggacagtc taaagtggat 1440 actagtgaga aaggtggaca attcagcttg cacatcctta accactccat aatcgttatg 1500 aaaccaagga actgttaa 1518 <210> 21 <211> 1686 <212> DNA <213> Arabidopsis sp. <400> 21 atggcgccac aagaacaagc agtttctcag gtgatggaga aacagagcaa caacaacaac 60 agtgacgtca ttttccgatc aaagttaccg gatatttaca tcccgaacca cctatctctc 120 cacgactaca tcttccaaaa catctccgaa ttcgccacta agccttgcct aatcaacgga 180 ccaaccggcc acgtgtacac ttactccgac gtccacgtca tctcccgcca aatcgccgcc 240 aattttcaca aactcggcgt taaccaaaac gacgtcgtca tgctcctcct cccaaactgt 300 cccgaattcg tcctctcttt cctcgccgcc tccttccgcg gcgcaaccgc caccgccgca 360 aaccctttct tcactccggc ggagatagct aaacaagcca aagcctccaa caccaaactc 420 ataatcaccg aagctcgtta cgtcgacaaa atcaaaccac ttcaaaacga cgacggagta 480 gtcatcgtct gcatcgacga caacgaatcc gtgccaatcc ctgaaggctg cctccgcttc 540 accgagttga ctcagtcgac aaccgaggca tcagaagtca tcgactcggt ggagatttca 600 ccggacgacg tggtggcact accttactcc tctggcacga cgggattacc aaaaggagtg 660 atgctgactc acaagggact agtcacgagc gttgctcagc aagtcgacgg cgagaacccg 720 aatctttatt tccacagcga tgacgtcata ctctgtgttt tgcccatgtt tcatatctac 780 gctttgaact cgatcatgtt gtgtggtctt agagttggtg cggcgattct gataatgccg 840 aagtttgaga t caatctgct attggagctg atccagaggt gtaaagtgac ggtggctccg 900 atggttccgc cgattgtgtt ggccattgcg aagtcttcgg agacggagaa gtatgatttg 960 agctcgataa gagtggtgaa atctggtgct gctcctcttg gtaaagaact tgaagatgcc 1020 gttaatgcca agtttcctaa tgccaaactc ggtcagggat acggaatgac ggaagcaggt 1080 ccagtgctag caatgtcgtt aggttttgca aaggaacctt ttccggttaa gtcaggagct 1140 tgtggtactg ttgtaagaaa tgctgagatg aaaatagttg atccagacac cggagattct 1200 ctttcgagga atcaacccgg tgagatttgt attcgtggtc accagatcat gaaaggttac 1260 ctcaacaatc cggcagctac agcagagacc attgataaag acggttggct tcatactgga 1320 gatattggat tgatcgatga cgatgacgag cttttcatcg ttgatcgatt gaaagaactt 1380 atcaagtata aaggttttca ggtagctccg gctgagctag aggctttgct catcggtcat 1440 cctgacatta ctgatgttgc tgttgtcgca atgaaagaag aagcagctgg tgaagttcct 1500 gttgcatttg tggtgaaatc gaaggattcg gagttatcag aagatgatgt gaagcaattc 1560 gtgtcgaaac aggttgtgtt ttacaagaga atcaacaaag tgttcttcac tgaatccatt 1620 cctaaagctc catcagggaa gatattgagg aaagatctga gggcaaaact agcaaatgga 1680 ttgtga 1686 < 210 > 22 <211> 1302 <212> DNA <213> Arabidopsis sp. <400> 22 atgaaaatta acatcagaga ttccaccatg gtccggcctg ccaccgagac accaatcact 60 aatctttgga actccaacgt cgaccttgtc atccccagat tccatacccc tagtgtctac 120 ttctacagac ccaccggcgc ttccaatttc tttgaccctc aggtcatgaa ggaagctctt 180 tccaaagccc ttgtcccttt ttaccctatg gctggtcgct tgaagagaga cgatgatggt 240 cgtattgaga tcgattgtaa cggtgctggt gttctcttcg ttgtggctga tactccttct 300 gttatcgatg attttggtga ttttgctcct acccttaatc tccgtcagct tattcccgaa 360 gttgatcact ccgctggcat tcactctttc ccgcttctcg ttttgcaggt gactttcttt 420 aaatgtgggg gagcttcact tggggttggg atgcaacatc acgcggcaga tggtttctct 480 ggtcttcatt ttatcaacac atggtctgat atggctcgtg gtcttgacct aaccattcca 540 cctttcattg atcgaacact cctccgagct agggacccgc cacagcctgc ttttcatcat 600 gttgaatatc agcctgcacc aagtatgaag atacctcttg atccgtctaa atcaggacct 660 gagaatacca ctgtctctat attcaaatta acacgagacc agcttgttgc tcttaaggcg 720 aaatccaagg aggatgggaa cactgtcagc tacagctcat acgagatgtt ggcagggcat 780 gtgtggagat cagtgggaaa ggcgcgaggg cttccaaacg accaagagac gaaactgtac 840 attgcaactg a tggaaggtc tagactacgt ccgcagctgc ctcctggtta ctttgggaat 900 gtgatattca ctgcaacacc attggctgtt gcaggggatt tgttatctaa gccaacatgg 960 tatgctgcag gacagattca tgatttcttg gttcgtatgg atgataacta tctgaggtca 1020 gctcttgact acctggagat gcagcctgat ctgtcagccc ttgtccgcgg tgcacatacc 1080 tacaagtgcc caaatttggg aatcacaagc tgggttagat tacctattta tgatgcagac 1140 tttggttggg gtcgtcctat ctttatggga cctggtggaa ttccatacga gggtttgtct 1200 tttgtgctac caagtcctac taatgatggc agcttatccg ttgccattgc cctccaatct 1260 gaacacatga aactgtttga gaagtttttg tttgagatat ga 1302 <210> 23 <211> 699 <212> DNA <213> Arabidopsis sp. <400> 23 atggccaatg aaattcctac caaagggatt ctcaagagcg aagctctaaa acagtacatc 60 atggaaacgt cagcgtatcc aagagagcat gaattgctca aggaacttcg aaaagctact 120 gtccaaaaat acggaaattt aagcgagatg gaggttccgg ttgatgaggg tcatttccta 180 tcgatgcttg taaagatcat gaatgcgaaa aacactatcg agatcggtgt tttcaccggt 240 tactcgcttc tcactacagc tcttgcttta cctgaagatg gccgtattac cgcgatagac 300 attgataaag aagcttacga agttggacta gagtttatta agaaggctgg tgttgatcac 360 aagattaatt ttatccattc cgatggtctt aaggccttag accaattggt gaacgacaaa 420 tgtgagtttg attttgcatt tgcggatgct gacaaatcaa gctacgtcaa cttccatgag 480 aggcttctaa aattggtgaa ggttggggga atcattgcgt tcgacaacac cttgtggttt 540 ggttttgtgg cggaggatga agatggagtt ccggagcata tgagagaata tagagcagct 600 cttatagaat tcaataagaa attggctttg gatccccgag tcgaggtctc tcagatttcc 660 attggagatg gtatcacgct ctgtagacgt cttgtttga 699 <210> 24 <211> 978 <212 > DNA <213> Arabidopsis sp. <400> 24 atggctgtaa agcaaaaggt ctgtgttact ggtgcaggag ggtttattgc ttcatggctc 60 gtcaagtttc ttctctctcg tggctacact gtccatggaa ccgttcgaga tccttgtgat 120 gaaaagaatg accatttgag gaaactggat aatgcttcaa aaaatctgaa acttttcaag 180 gcagatttgt ttgatgatga aggtcttttc tcagccattg atggttgctc tggcgttttc 240 catatcgctt ctcctgtgcc ttttgaagga gttcctctca cagaggaaga actgattaaa 300 ccagctctca ctggcaccaa gaatgttcta gaagcatgca cagaaaccaa agtgcagaaa 360 gttgtggttg tatcatcaat cgctgcagtt gtttacaatc caaaatggcc tcaagatgtt 420 gctaaggacg aggattgttg gtcagacact caataccttc actcacttga gggttattgg 480 cgttattact atcttgccaa aacattaacg gaacgtgaag ctttagagtg gagtaagaga 540 aactttgctg atgttgttac gctctgtccg tctgtcatta taggccctag gcttcaatcc 600 acgcttaact ctagtagttt gggacttctc aagttcataa aaggtggtat aaagtccttg 660 ttgagcgatg aactgtatct tgtggatgtg cgtgatgttg cggatgcgct tttgttggtg 720 tatgaaaatc gagaagcaac aggaagatac atatgtaatt ctcattcact ttacactgat 780 tctcttatgg agaagctcaa gaatatgtat cctaaacgca acttccctga aagtttcacg 840 gaggtaaaag a gaaggaagt gaggcctcta agtgcagaga agttgaaaaa cttggggtgg 900 aagtttaggc cattagaaga aaccattgat gattctgttg tgagctttga agcagctggt 960 gatctttccaa aggcataa 978 <210> 25 <211> Arabidopsis <210> 25 <211> 1539 <400> 25 atgcttactc taatgactct catcgttctt gtccctcttc ttctttttct cttcccccac 60 ctccttttac ggcggcagat gcttctgaaa ccctaccctc ctggccccaa aggcttaccc 120 gtcattggaa atattctcat gatgaaccag tttaaccacc ggggtctggc caagctcagc 180 cgtatatacg gtgggttgct ccatctccgc ctcggatttt cccacatttt cgtcgtttct 240 tctccggaca ttgcgcgtca agtcctccaa gtccaagacc acgttttctc aaaccgtccc 300 accacaatcg caatccgata cttgacctac ggaggatccg atcttgcatt ctgcaattac 360 ggcccgtttt ggcgtcggat gagaaaactc tacgtcatga tgctctttag ccgtaaacgg 420 gccgagtcgt gggtctctgt tgatgaagag gtccacaaat cggtccgttt agtggcttcc 480 aacgtcggaa aaccactaaa catatgtaaa ctagcatttt ccctatcaag agacataacg 540 ttccgagcag ctttcggttc ctcctcatcc acatctgatg aaagccgttt ggacgagttc 600 cttgagatca ttcaagagtt ctctaagctc tttggtgagt ttaacgtagc ggattatgtc 660 ccgtcgtggc tcagttggat tgacccgcaa gggataaacg ggcgagttga gaaagcccga 720 aaatctctag acggtttcat tgagtccgtc atcgatgatc acttgcacaa gaagaagaga 780 gaacatgata acgttgacga agagactgat atggtagatc aattacttgc gttctacgaa 840 gaagaagtca a agtcaacaa ctcagtcacc aaaattaatc tcgataacat aaaaggcatc 900 atcatggatg tgatgttcgg aggaacggag acggtggcat tagcaatcga atgggtgcta 960 accgagatac tccggagccc cgagaacatg aaacgggtcc aggacgagct aacgagtgtg 1020 gtcgggcttg acagatggcg cgtggaggac acgcacctcg agaagctcac tttcctcaag 1080 tgtatcctca aggagaccct ccggctccac ccgcctttcc ctctcctcct ccacgagacg 1140 gtgaaggaca ctgagatttc cggttacttc atccccaagg gttcgcgtgt gatggtcaat 1200 acctacgctc tagggcgtga cccgaattct tggtccgacc cggaaagttt taacccgggt 1260 aggtttttaa acccgattgc tccggatctg aaggggaata atttcgaatt tgttccattc 1320 gggtcgggtc ggagatcgtg cccgggtatg caactcgggt tgtacgcatt tgagcttgcg 1380 gtggctcatc ttctacactg cttcacgtgg agtttaccgg acggtatgaa tcccggtgac 1440 gtcgacaccg ttgaaggacc gggtctcacc gttcctaagg caattccttt ggtggcggtg 1500 ccgaccacgc gcctcctctg ccccatcgtc gtctcgtga 1539 <210> 26 <211> 1050 <212> DNA <213> Arabidopsis sp. <400> 26 atggggtttt ctccttcatt ttcttgcagt gctattggag ccctaatttt gggttgcctt 60 ctgcttcaag catcaaactc taatgctcag ttgaggcctg atttctactt cgggacgtgc 120 ccatttgttt ttgatatcat tggaaatatc atcgtcgatg aactgcagac tgatcctcgt 180 attgccgcga gcctccttcg tcttcacttt cacgactgct ttgttcgtgg ttgtgatgca 240 tcgatcttgc ttgacaattc cacatcattc cggactgaga aagatgctgc tccgaacgca 300 aattcggctc gagggtttaa tgtcatagat agaatgaaag tagcccttga gagagcttgc 360 ccaggaagag tctcttgtgc agatattctc accatcgcct ctcaaatatc agtgcttttg 420 tcgggaggtc catggtggcc ggttccgttg gggaggagag acagtgtaga agctttcttc 480 gctctggcta atacagctct gccctctccg tttttcaatc ttactcaact taaaacagcc 540 tttgctgacg ttggcctaaa ccgcacttcc gatctagttg ctctttccgg tggtcacaca 600 tttggaagag cacaatgcca gttcgtgacg cctcgtctct acaacttcaa tggtacaaac 660 agtccagacc caagtctgaa cccaacttac cttgtcgaac tccgtcgatt gtgtcctcaa 720 aacggaaacg gcaccgttct ggtcaacttc gatgtcgtga ctccagatgc tttcgatagt 780 caatactaca ccaaccttcg taatgggaaa ggtcttattc agagtgacca ggaactcttt 840 tcgactccag g agccgatac gatcccttta gtaaaccaat acagcagcga catgtccgtg 900 ttctttcgag cattcattga cgcaatgatt aggatgggaa atcttagacc tttgactgga 960 actcaaggag agattagaca aaattgtagg gttgtgaatc cacgaattag ggttgtggag 1020 aacgacgatg gtgttgtgag ttctatctga 1050 <210> 27 <211> 1098 <212> DNA <213> Arabidopsis sp. <400> 27 atgggaagtg tagaagcagg agaaaagaag gctttaggat gggctgcaag agatccttct 60 ggtgttctct ctccttactc ctacacttta aggagcacag gagcagatga tgtgtatata 120 aaggttatat gctgtggaat atgccatact gatattcacc aaattaaaaa tgatcttgga 180 atgtctaatt atcctatggt tcctgggcac gaggtagtag gtgaggtgtt ggaagtggga 240 tcagatgtga gcaagttcac cgtaggagat gtagtcggag tcggcgtggt cgttggatgc 300 tgcggtagct gcaaaccatg cagctcagaa cttgagcagt attgtaacaa gagaatctgg 360 agttacaacg atgtctacac cgatggaaaa cctactcaag gtggcttcgc cgacaccatg 420 atcgtcaatc aaaagtttgt ggtgaagatt ccagaaggaa tggcggtgga gcaggctgca 480 ccgctactct gcgcgggagt aacggtttat agtccgttga gccacttcgg tctaatggcg 540 agtgggctta aaggagggat tctagggtta ggtggagtag ggcacatggg tgtgaagata 600 gctaaagcaa tgggacacca tgtgactgtg ataagctcat cagataagaa aaaagaagaa 660 gctatcgaac atttaggagc tgacgactat gtggtgagct ctgatccagc agagatgcaa 720 agactcgcag attcccttga ctacatcatt gacaccgtac cggtgtttca ccctctcgac 780 ccttacttgg cttgtcttaa actcgacggg aaattgattc tcatgggtgt tatcaacact 840 cctcttcagt t cgtgacccc tctcgtcatt cttgggagga aagtgatatc agggagcttt 900 atagggagca ttaaagagac agaagaagtg ttggctttct gcaaagagaa agggcttacg 960 tcgacgattg aaactgtgaa gatcgatgag cttaacattg cgtttgagag gcttcgcaag 1020 aacgatgttc gttacagatt tgtggttgat gttgcaggaa gtaatctggt ggaggaggct 1080 gcaacaacta caaactaa 1098 <210> 28 <211> 1074 <212> DNA <213> Arabidopsis sp. <400> 28 atgggaataa tggaggcaga gaggaaaaca acaggctggg ctgccagaga cccatctggg 60 atcctctctc cttacactta cactcttaga gagactggac cagaggatgt gaacataaga 120 atcatttgct gtggaatctg ccacaccgat cttcatcaaa ctaaaaatga tcttggcatg 180 tctaattacc ccatggttcc tgggcatgaa gtggtagggg aagtagtgga ggtgggatca 240 gatgtgagca agttcaccgt aggggacata gttggagttg gttgcctcgt tggatgttgc 300 ggaggttgta gcccctgcga gagagatctg gaacagtatt gtccaaagaa gatttggagc 360 tacaatgatg tttacatcaa tggtcaacct acacaaggcg gcttcgctaa agccaccgtc 420 gttcaccaaa agtttgtggt caagattcca gaaggaatgg cggttgagca ggctgcgccg 480 ctactgtgcg ctggtgtgac tgtgtacagt ccactgagcc actttggtct gaaacaacca 540 ggcctaagag gaggtatact agggttaggt ggagtcggtc acatgggtgt gaaaatagcc 600 aaagcaatgg gtcaccatgt gactgtcata agctcatcaa acaagaagag agaagaggca 660 ttgcaagatc ttggagctga tgattacgtg atcggatccg accaagcgaa gatgagcgaa 720 ttggctgatt cgttggatta cgtaattgac acggtgcctg ttcatcatgc acttgagcca 780 tatttgtctc tgcttaagct tgatggtaaa ctcattctca tgggagttat caacaatcca 840 ttacagtttc t cactcctct gcttatgctt gggaggaaag tgataacggg gagcttcata 900 gggagcatga aggagacaga ggagatgctt gagttctgta aagaaaaggg tttgagttcg 960 attatcgaag ttgtgaagat ggattatgtt aacactgcgt ttgagagact cgagaagaac 1020 gatgtgcgtt ataggttcgt cgttgatgtc gaaggaagca atctcgacgc ttga 1074 <210> 29 <211> 1080 <212> DNA <213> Arabidopsis sp. <400> 29 atgggaaagg ttcttcagaa agaggcgttc ggattagccg cgaaagacaa ttccggagtt 60 ctctcgcctt tcagtttcac tagaagggaa acaggagaaa aggatgtgag gttcaaagtg 120 ttattttgtg gaatttgtca ctctgatttg catatggtca agaacgagtg gggaatgtct 180 acttatcctc ttgtcccagg gcatgagatc gtgggcgtgg tgactgaagt cggagccaaa 240 gtgactaaat tcaaaaccgg agaaaaagtc ggagttgggt gtttggtcag ctcgtgcggg 300 tcatgtgaca gctgcaccga aggtatggaa aactattgtc caaaatcaat ccaaacgtac 360 ggattcccgt actacgacaa caccataaca tacggtggtt actccgacca catggtttgc 420 gaggaaggtt tcgtcatccg tattccagac aatctcccct tggacgccgc cgcaccgctc 480 ctctgtgccg gtatcacggt ctattcccct atgaagtatc acgggctcga caaacccggt 540 atgcacatcg gtgtggtagg attaggcggt ttaggtcatg taggagtgaa atttgccaag 600 gctatgggta ctaaggttac ggttattagt acttcggaga aaaagagaga tgaggcgatt 660 aatcggcttg gtgcggatgc tttcttggtg agccgtgacc caaaacagat taaggatgca 720 atgggtacta tggatggtat aattgatacc gtctctgcga ctcattcact tcttccgttg 780 ctcggtttgc tgaagcataa gggaaaactt gttatggttg gtgcacccga gaagccactc 840 gagctacctg t catgcctct catctttgag aggaagatgg taatgggaag tatgatagga 900 gggataaaag agacccagga aatgatagat atggccggga aacacaacat cactgcggat 960 attgagctta tctctgccga ttatgtcaac accgtgccatgg tattgtcaac accgtgtga 1080 cctagt tagtga

Claims (19)

delete delete delete delete delete delete delete delete delete delete delete A method for inhibiting detachment of a plant or inhibiting falling or falling fruit comprising the steps of:
(a) deletion or suppression of expression of genes encoding ROS-forming enzymes RbohD and RbohF; and
(b) obtaining a plant in which lignin production is suppressed specifically for cells leaving the detachment site. The method according to claim 12, wherein the gene encoding RbohD is represented by the nucleotide sequence of SEQ ID NO: 1. The method according to claim 12, wherein the gene encoding RbohF is represented by the nucleotide sequence of SEQ ID NO:2. 13. The method of claim 12,
A method further comprising inhibiting the activity of one or more enzymes selected from the group consisting of peroxidase, superoxide dismutase (SOD), and NADPH oxidase (NOX). The method according to claim 12, wherein step (a) is performed by treating RNAi or antisense oligonucleotides of RbohD and RbohF.
delete delete delete

Images