CN113957081B

CN113957081B - Gene for regulating and controlling growth and development of tomato epidermal hair and application thereof

Info

Publication number: CN113957081B
Application number: CN202111243302.8A
Authority: CN
Inventors: 崔娜; 余兰; 丁琪; 杨芷; 黄竞楠; 张钰蛟; 邢红运; 范海延; 于洋; 孟祥南
Original assignee: Shenyang Agricultural University
Current assignee: Shenyang Agricultural University
Priority date: 2021-10-25
Filing date: 2021-10-25
Publication date: 2023-01-24
Anticipated expiration: 2041-10-25
Also published as: CN113957081A

Abstract

The invention relates to the technical field of plant genetic engineering, in particular to a gene for regulating and controlling the growth and development of tomato epidermal hair and application thereof, wherein the gene is an SlTOR gene, and the nucleotide sequence of the SlTOR gene is shown in SEQ ID NO. 1. Therefore, the invention constructs an SlTOR silent strain and applies TOR inhibitor and activator outside the seedling stage, and observes the change of tomato glandular hair density, finds that the tomato glandular hair density is obviously increased after the TOR inhibitor is treated, the transcription expression of terpene synthase genes (SlTPSs) and synthesis precursor pathway genes is obviously up-regulated, the silent strain result and the inhibitor treatment result trend are the same, and the activator treatment trend is opposite, which shows that SlTOR influences the synthesis of tomato glandular hair density. The method has important guiding significance and application value for reducing the use of harmful pesticides and selecting and breeding tomatoes in actual production.

Description

Gene for regulating and controlling growth and development of tomato epidermal hair and application thereof

Technical Field

The invention relates to the technical field of plant genetic engineering, in particular to a gene for regulating and controlling the growth and development of tomato epidermal hair and application thereof.

Background

Tomato (A)Solonum lycopersicum) The tomato is one of vegetable crops widely cultivated in the world, is delicious in taste and high in nutritional value, is popular with the public, has important market economic value, and is easily affected by biotic and abiotic stresses in actual production, so that the yield is reduced. The epidermal hair is a specialized hairy protrusion structure which originates in epidermal cells and is widely distributed on the surface of terrestrial plants and can be divided into glandular hair and non-glandular hair. Although not essential to plant growth, the coat serves as a first physical and chemical barrier to plant exposure to the external environment, and serves multiple biological functions, such as protection against insect and pathogen attack, reduction of water loss, and increased tolerance to abiotic stresses, including extremes of temperature and ultraviolet radiation; researches find that the epidermal hair can form a protective layer for resisting ultraviolet rays, so that the damage of the ultraviolet rays to plants is effectively reduced. Researches on birch find that frost and ozone increase in spring can cause leaf surface damage of the birch and influence the early growth of the birch; frost increases the density of epidermal hair, improves the structural adaptation of birch to low temperatures, and enhances the freezing resistance. Glandular hairs are important epidermal hair types in the epidermal hairs, can specifically synthesize, secrete or store a large amount of and various secondary metabolites, including acyl sugar, terpenoids, alkaloid, flavonoid and the like, and can protect plants from being damaged by herbivores, parasites, fungal infection and the like; in addition, some plant glandular hair secreted secondary metabolites have important commercial value, such as artemisinin secreted by artemisia apiacea epidermal hair to treat malaria.

Tomato epidermal hair is a typical multicellular structure that is divided into two major classes of 8 types: glandular type epidermal hair (glandular hair,

a model,

The model is,

Type and VII) and non-glandular epidermal hairs (non-glandular hairs, type II, type III, type V and type VIII). Glandular hairs are the main factories of various special secondary metabolites, which can synthesize, secrete or store various secondary metabolites including acylsugars, terpenoids, alkaloids, flavonoids, etc. Among these, terpenoids, in particular, represent the largest and most diverse group of plant volatile metabolites, which have a considerable role in tomato's resistance to pests, such as: carnivore or parasites which are repellent or toxic to herbivores and which attract attack insects, and the like.

Defines the important role of tomato epidermal hair in resisting biotic and abiotic stress, and how to regulate the growth and development of tomato epidermal hair becomes a technical problem to be solved by the technical personnel in the field.

Disclosure of Invention

In order to solve the above technical problems, the present invention provides, in a first aspect, a gene for regulating and controlling the growth and development of tomato epidermal hair, wherein the gene isSlTORThe nucleotide sequence of the gene is shown in SEQ ID NO. 1.

In the second aspect of the invention, the protein coded by the gene is provided, and the amino acid sequence of the protein is shown in SEQ ID NO. 2.

In a third aspect of the present invention, there is provided the use of the nucleic acid molecule expressed by the gene in the construction of a tomato plant with enhanced resistance to biotic and abiotic stress by increasing the glandular hair density of tomato.

In a fourth aspect of the invention, there is provided an expression cassette comprising said nucleic acid molecule.

In a fifth aspect of the invention, there is provided a plant expression vector comprising said nucleic acid molecule.

In a sixth aspect of the present invention, there is provided a recombinant bacterium containing the nucleic acid molecule.

In a seventh aspect of the invention, there is provided the use of a substance which reduces the protein content in the construction of a tomato plant having an increased tolerance to biotic and abiotic stress, wherein the increased tolerance to biotic and abiotic stress is achieved by increasing the glandular hair density of the tomato plant.

Further, the substance is AZD8055.

In an eighth aspect of the invention, there is provided a method of constructing a tomato plant with enhanced resistance to biotic and abiotic stress by silencing tomato plants of interest by gene editing techniquesSlTORThe gene or AZD8055 is applied to the target tomato plant to increase the glandular hair density of the obtained tomato plant, thereby enhancing the capability of resisting biotic and abiotic stress.

Compared with the prior art, the invention has the following beneficial effects:

the invention discovers a gene which is not reported in tomato and can regulate the growth and development of epidermal hairSlTORThe transcription expression of genes related to the synthesis of the tomato I and VI epidermal hair density and terpene compounds can be changed by applying a TOR inhibitor, an activator and constructing a silencing strain. The method has important guiding significance and application value for reducing the use of harmful pesticides and selecting and breeding tomatoes in actual production.

Drawings

FIG. 1 shows that SlTOR participates in the regulation of the number of tomato epidermal hairs;

a: AZD8055 and MHY1485 treat tomato epidermal hair phenotypic changes;

b: AZD8055 and MHY1485 treated tomatoes have total epidermal hair count, varying numbers of glandular and non-glandular hairs;

c: effect of AZD8055 and MHY1485 treatment on the variation in density of individual types of coat of tomato.

Con.: distilled water (added to the same amount of DMSO in experimental group) treated tomato plants; AZD8055: 5. mu m AZD8055 treated tomato plants; MHY1485: 20. μ m MHY1485 treated tomato plant;

wherein con is a control group; AZD8055 is a TOR inhibitor; MHY1485 is a TOR activator.

FIG. 2 isSlTORSilencing transcriptional expression of terpene synthesis related genes in the strain.

A：SlTORIn thatSlTORSilencing the level of transcriptional expression in the strain;

B：SlTORsilencing the transcriptional expression level of a terpene synthase gene in the strain;

C：SlTORsilencing transcriptional expression levels of terpene synthesis precursor-related genes in the strain.

TRV: 00: wild type tomato lines; TRV:SlTOR：SlTORand (4) silencing the strain.

Detailed Description

The invention is described in detail below with reference to the figures and the specific embodiments, but the invention should not be construed as being limited thereto. The technical means used in the following examples are conventional means well known to those skilled in the art, and materials, reagents and the like used in the following examples can be commercially available unless otherwise specified.

Example 1: tomatoSlTORGene cloning and vector double enzyme digestion.

RNA extraction is carried out on the sample according to the ultra-pure RNA extraction kit of the biological technology limited company in the Kangwei century. cDNA Synthesis tomato leaf cDNA was obtained by reverse transcription using FastKing cDNA first strand synthesis kit (Tiangen Biochemical technology Co., ltd.). Search for tomato at NCBI website (https:// www. NCBI. Nlm. Nih. Gov /)SlTORCDS region of (1), designSlTORThe silent vector primers were as follows: SEQ ID No.3 TOR (+): 5 'CCTCCATGGGATCCGACCGGAATTTCCCTCCTATACC-3' and SEQ ID NO.4 TOR (-): 5' CGTGAGCTCGGTACCGGATATATGATAATTTGC-; using tomato cDNA as template and primers SEQ ID NO.3 and SEQ ID NO.4 as primers, carrying out PCR amplificationSlTORThe size of the target fragment of cDNA of (1) is 379 bp. Use ofBamHI andKpni, carrying out double enzyme digestion on the pTRV2 vector, carrying out enzyme digestion at 37 ℃ for 3 h, and carrying out gel recovery by using a tiangen gel recovery kit to obtain a target fragment and a vector linearized fragment.

Example 2: plant expression vector construction

Connecting the recovered target fragment to a pTRV2 vector by using an One Step Seamless Cloning Mix kit of Congwort century Biotechnology Limited, adding all the connection products into competent cells containing 50 mu L DH5 alpha escherichia coli, gently sucking and uniformly mixing, placing on ice for incubation for 30 min, thermally shocking at 42 ℃ for 90 s, and incubating on ice for 5 min; then adding 500 mu L of LB liquid culture medium without antibiotics, placing the mixture on a shaker at 37 ℃ and 170 rpm for shake culture for 1 h;5000 Centrifuging for 2 min at rpm, discarding 400 muL supernatant, utilizing the remaining 100 muL to resuspend precipitate, and coating the precipitate with 50 mg ^. mL ^-1 On an LB solid medium plate of kan, the cultivation is carried out for 24 hours by shaking and inverting at 37 ℃. Picking single colony, placing positive colony in the medium with final concentration of 50 mg ^. mL ^-1 Culturing in LB liquid culture medium of kan at 200 rpm and 37 deg.C for 24 h with shaking, extracting plasmid with Tiangen plasmid small extraction kit, sequencing the obtained plasmid by Jinzhi Biotechnology Limited, and obtaining the plasmid with correct sequencing, i.e. pTRV-SlTORAnd (3) a carrier.

Example 3: establishment of plant transient expression system

(1) EHA105 Agrobacterium competent preparation: EHA105 original Agrobacterium contained 50 mg ^. mL ^-1 The YEP solid medium of (1) was streaked and cultured at 28 ℃ for 2 d. Single colony was picked and inoculated into a medium containing 50 mg ^. mL ^-1 In YEP liquid medium from Rif, shake overnight at 18 rpm and 28 ℃, and subculture to OD 2 d ₆₀₀ =0.5; 1.5 mL of bacterial liquid is sucked into a centrifuge tube, ice bath is carried out for 10 min, centrifugation is carried out for 2 min at 5000 rpm, and supernatant is discarded; suspending the thallus with 1.5 mL of 0.5M NaCl solution, carrying out ice bath for 20 min, centrifuging at 5000 rpm for 2 min, and removing the supernatant; then 100 muL of 20 mM Ca was added ₂ And (3) suspending the thallus by using a Cl solution, and quickly freezing for 5 min by using liquid nitrogen to obtain the agrobacterium tumefaciens competence.

(2) And (3) agrobacterium transformation: and adding 8 mu L of the successfully sequenced recombinant plasmid into 50 mu L of EHA105 agrobacterium infected state in an ice water mixture, sucking and blending uniformly by a gun, ice-bathing for 30 min, quickly freezing for 5 min by liquid nitrogen, and carrying out water bath for 5 min at 37 ℃. Then, 50 mg of 800 muL antibiotic-containing solution was added ^. mL ^-1 Rif YEP liquid medium, 28 ℃,170 rpm shaking culture 6h. 100 mu L of the solution is sucked and coated on the surface of the soilHas a weight of 50 mg ^. mL ^-1 Rif，50 mg ^. mL ^-1 The cells were cultured in an inverted state at 28 ℃ for 48 hours on a YEP solid medium plate of Kan. The plaque is picked up for PCR single colony identification, and the identified positive plaque is added into YEP liquid culture medium containing Rif and Kan, and is cultured for 48 h at 28 ℃ and 200 rpm in a shaking way.

(3) Transforming tomato with the recombinant plasmid: taking out and shake culturing to OD ₆₀₀ 20 mL of the bacterial liquid with the speed of 1.0-1.2 is put in a centrifuge tube, centrifuged at 5000 rpm for 10 min, and the supernatant is discarded. With 2 mL of the padding liquor (containing 10 mmol) ^. L ^-1 MES and 10 mmol ^. L ^-1 MgCl ₂ ^. 6H ₂ O) resuspension and washing of the thalli, centrifugation at 5000 rpm for 10 min, and discarding of the supernatant. With a padding liquid (containing 10 mmol) ^. L ^-1 MES、10 mmol ^. L ^-1 MgCl2 ^. 6H2O and 200 mmol.L ^-1 Acetosyringone) suspension of the cells, the bacterial solution OD ₆₀₀ =0.5, left in the dark at 28 ℃ for 3 h. Soaking tomato seeds with 0.2-0.4 cm of germination roots in a bacterial liquid, vacuumizing until the bacterial liquid is soaked in the tomato roots, and sowing the seeds in vermiculite: culturing in soil with matrix soil = 1; and (3) extracting RNA from tomato seedling leaves to carry out qPCR identification and transformation on successful strains.

Example 4: tomato plant treatment with TOR inhibitor AZD8055 and activator MHY1485

Spraying 5 mu M AZD8055 on the surface of the leaf of the first-leaf and first-heart tomato seedling, culturing for 2 days, culturing for 20 days, taking the fourth-leaf tomato leaf to extract RNA for related gene transcription expression analysis, using 0.5cm below the first leaf for epidermal hair density analysis, and extracting epidermal hair secondary metabolites 6 cm above the cotyledon.

(1) Epidermal hair is a hairy protrusion structure widely distributed on the surface of a plant, is used as a key barrier on the surface of the plant, and plays an important defense role in the aspect of stress response of the plant. Tomato epidermal hairs are mainly divided into 8 types, 2 types, glandular hairs and non-glandular hairs. The effect of SlTOR on tomato epidermal hair density was analyzed by a bulk microscope and it was found that inhibitor AZD8055 treatment increased tomato epidermal hair density compared to the control, with a particularly marked increase in glandular hairs, especially type I and VI glandular hairs, whereas the opposite result was seen with activator treatment (fig. 1).

(2) Construction ofSlTORThe silent strain functional identification discovers that qRT-PCR detects the transcriptional expression of SlTOR on terpene synthesis related genes, and the tomato plant terpene synthase gene in the silent strainSlTPS5、SlTPS9、SlTPS12、SlTPS20Are all obviously adjusted upwards; the expression level of MVA pathway and MEP pathway related gene transcription is also significantly up-regulated (fig. 2), and the variation trend is consistent with the AZD8055 processing structure. Again, the description is givenSlTORThe transcription expression of the genes related to the synthesis of the tomato terpenoid is reduced and improved.

In conclusion, TOR can influence the ability of tomato to respond to external biotic and abiotic stresses by influencing the density of tomato epidermal hairs.

While preferred embodiments of the present invention have been described, additional variations and modifications in those embodiments may occur to those skilled in the art once they learn of the basic inventive concepts. Therefore, it is intended that the appended claims be interpreted as including the preferred embodiment and all changes and modifications that fall within the scope of the invention.

It will be apparent to those skilled in the art that various changes and modifications may be made in the present invention without departing from the spirit and scope of the invention. Thus, if such modifications and variations of the present invention fall within the scope of the claims of the present invention and their equivalents, the present invention is also intended to include such modifications and variations.

Sequence listing

<110> Shenyang agriculture university

<120> gene for regulating and controlling growth and development of tomato epidermal hair and application thereof

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<170> SIPOSequenceListing 1.0

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gctcgtttta tggatcatct atatgaacgc gttaccacat ctcttgatag taatgaagtt 240

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aatgcatcaa aagtggcaaa attctccaac tacatgcgcg ttgcttttga aacaaagcgt 360

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gttattgaaa agcgtgagac acgatggcgt gttcagtggt attatcgaat gtttgaggct 720

acccaagatg gattggggag aaatgcgcct gttcatagta tacatggatc tcttctcgca 780

gtcggggagt tgctaaggaa cacaggggag ttcatgatgt caagatacag ggaggttgct 840

gaaattgttc taagatacct ggagcaccga gatcgcctag ttcgcctcag cataacttca 900

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gctctggggg agatggcagg tgctctggat ggtgaactta tcaactattt gccgacaata 1080

acctctcact tgcgtgatgc gatcgctccc cgtagaggca gaccctcact tgaggctcta 1140

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ctttgttgct gcaaactaat agcaaattct tttttggcta tgtcttctac ccagtttagt 1620

cctagtagaa tcaatcgcgc tagtggaaag cgacgtcgac ttgttgaaga gattgtgcaa 1680

aaacttctca ttgccgctgt tgctgatgct gatgttactg ttcggcattc tatattttcc 1740

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atatttgcca ctttaaatga tgaggatttt gaagttcgtg agtatgcaat ttcactagct 1860

ggcagattat ctgaaaagaa tccagcatat gttctaccag cccttcgtcg ccatcttatt 1920

cagctgttga cttacctaga acaaagcagt gcagacaata agtgtaaaga agagagtgca 1980

aagttattgg gttgcttgat tcgcaattgt gaacgcttag ttcttccata cgtttctcct 2040

atacacaagg ctcttgtcgc aaaactctgt gagggcacag gggtcaatgc aaatagtggc 2100

attattagtg gagtcctagt gactgttgga gatctcgcaa gagtgggtgg ctttgccatg 2160

cgacagtata tttcagaact tatgccatta attgttgaag ctctattgga tggggctgct 2220

gttaccaaac gtgaagtggc tgtttcaaca cttggtcaag ttgtacagag tacaggatat 2280

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ttagatcccc atgtgcacaa gcgcaatcag caaagcttac ctggatccca tggtgaagtt 2460

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ctctggccat cctttgcaac atctgaggat tattattcca ctgttgctat caactcactc 2580

atgcggattc tcagggatcc atctctgtca agttaccacc agaaagtagt tggatctctt 2640

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ggagagagtg acctctatga ggttgggacc gacatgcaga agcagctcag aaaccatcag 3540

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tgggcagagt ggatgagaca tttcagcatt gaacttttga aagaatcacc tagtccagca 3660

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gcaggttttg ttagttgctg gtcacaactt aacgaggcca gtcaaaggca gctcgtacgg 3780

agtttagaaa tggcgttttc ttctccaaat atcccacctg aaattcttgc tacacttctg 3840

aacttggcgg agtttatgga acatgatgag agaccccttc ctatagatat ccgtctgctg 3900

ggtgctctag cggagaagtg tcgagcattt gcaaaggccc tacactacaa ggaaatggaa 3960

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cggcttgatg atggtgatga aaccaaactc cgagtcttgg gaaatactgc ttccagcggt 4440

gatggaagta gtaatggcac cttttatagg gccgttcttc tagttcgtcg agggaagtac 4500

gacgaagcac gtgaatatgt tgaaagagca aggaaatgtt tggcaaccga gcttgctgca 4560

ctggttcttg agagctatga acgtgcttac agcaacatgg ttcgcgttca acagctttct 4620

gaattagaag aggtcattga gtactgtact cttcctccta cgggaaaccc tgttgctgaa 4680

ggaagaagag cccttgttcg caatatgtgg aatgagcgca taaagggtgc aaaaagaaat 4740

gtcgaggtct ggcaagcact tttagctgtg agggcacttg tattgcctcc tacagaagat 4800

atagaaacat ggatcaagtt tgcatcactc tgccggaaaa atggcagaat tagccaagcg 4860

agatctacgt tgattaaact cttacagttt gatcctgaaa cgactcctgc aactgtgcgg 4920

tatcacggcc cccctcaggt gatgctagca tacttaaagt accaatggtc acttggtgag 4980

gatcataagc gaaaggaagc ctttgctagg ttgcaggacc ttgctatgga cctctcgaga 5040

acagctactc tgcaaccagt aatgcagaat gcattagttg cttcttctgg tgcgccactt 5100

gttgctcgta tatacctcag actcggcact tggaagtggg cactttctcc tggtttggat 5160

gatgattcta tacaagaaat ccttagtgca tttagaaatg ctactcactg tgcaacgaag 5220

tggggaaaag catggcatac ttgggcactt ttcaatacag cagtgatgtc tcattacact 5280

ctaagaggtt ttgcaaatat tgctgcacag tttgttgttg ctgccgtaac tggttatttt 5340

cactctatag cttgcggagc acatgctaag ggtgttgatg atagtttaca ggatattctt 5400

cgtcttctta ctttgtggtt caaccatgga gctacttctg aggtccaaat ggcattacag 5460

aaaggattca cacatgttaa catcaacaca tggttggttg ttttacctca gattatcgca 5520

cggatacatt caaataacca tgctgtcaga gaactgatac agtccttgtt agttcgaatt 5580

ggacagagtc atccacaggc tcttatgtat ccacttcttg tggcatgtaa gtcgattagc 5640

aatttgcgca gagctgcagc tcaagaggtg gttgataaag ttaggcagca tagtggcgta 5700

ctcgttgatc aggcccaact tgtctcaaag gagcttatca gggttgcaat actttggcat 5760

gaaatgtggc atgaggcatt ggaagaggcc agtcgtctgt attttggtga acacaacatt 5820

gagggcatgc tgaaggtgtt agagcctctg catgaaatgc ttgaggaagg agcgatgcgg 5880

aataatacta ctatgaagga gaaagcattc atccaggcat accgtattga gttgttggag 5940

gcctatgaat gctgtatgaa gtatcggaga acaggaaaag atgctgaatt aatacaggct 6000

tgggatctct attaccatgt attcaggcgg atagataagc agcttcaaac actcactacg 6060

ctggatttac agtccgtttc ccccgagttg ctagaatgtc gagatttaga gctagctgtt 6120

cctggaactt atcgagcaga tacaccagtg gtgacaattg catcatttgc accccaactt 6180

gttgtaatta catccaaaca acgacctcgt aaattgacaa tccatgggag tgatggaaaa 6240

gactatgctt tcttgctcaa agggcatgaa gatttacgcc aagatgaacg tgtcatgcag 6300

ttgtttggtc tggttaatac tttgctggag aactcgagaa agactgcaga gaaagattta 6360

tcaattcaac gatatgatgt cattccattg tcccctaata gtggactgat agaatgggtt 6420

ccaaattgtg acaccttgca ccaacttatt cgagaatata gggatgctcg gaagatcacc 6480

ctaaatcaag agcataaatt gatgctgagt tttgcaccgg attatgataa tttgccactt 6540

attgctaagg tggaggtgtt tgaatatgca ctgcaaaata cagaagggaa tgacttatca 6600

agggttcttt ggttaaagag tcgcacttct gaggtctggc tggacagaag aacaaattat 6660

acaagaagtt tggctgtcat gagcatggtt ggatacttac ttggtttggg cgatcgacat 6720

cctagtaacc tcatgcttca ccgatttagt gggaagattc tgcatattga ttttggagat 6780

tgttttgaag cttcaatgaa tagggagaag tttccagaga aggttccgtt tcgcctcact 6840

aggatgcttg taaaagcaat ggaggtcagt ggtatagagg gaaatttccg gtcaacatgt 6900

gagaatgtaa tgcaagttct ccgactgcat aaagatagtg ttatggctat gatggaggcc 6960

tttgttcatg acccacttat aaattggcgt cttttcaatt tcaatgaagt tccgcaaatg 7020

tccacacttg ccagtgcaca tgtgcctcct gttgtgaata gtgaggattc ctcttcagat 7080

agagagcttc ttcaaccaca aaggggtgca agggagagag aactgcttca ggcggtcaat 7140

caattaggtg atgccaatga agttctaaat gaacgtgctg tggctgtcat ggctcgaatg 7200

agtaataaac tcactggacg tgattttgct gctacctctg catcttcaag ctctctacag 7260

catccactgg atcacagtac gttgatttct ggagagacgc gtgaagctga tcatggttta 7320

tcagtgaaat tacaagtcca aaaacttatt cagcaagcaa tgtctcatga aaatctttgc 7380

caaaattatg ttgggtggtg tccattttgg tag 7413

<210> 2

<211> 2470

<212> PRT

<213> tomato

<400> 2

Met Ala Ala Thr Val Gln Ala Ile Arg Tyr Pro Val Ala Thr Thr Gly

1 5 10 15

Ala Gly Asn Ile Asp Ala Leu Asn Arg Val Leu Ala Asp Leu Cys Thr

20 25 30

Arg Gly Asn Pro Lys Asp Gly Ala Ala Leu Thr Leu Arg Arg Leu Val

35 40 45

Glu Glu Glu Ala Arg Asp Leu Ser Gly Glu Ala Phe Ala Arg Phe Met

50 55 60

Asp His Leu Tyr Glu Arg Val Thr Thr Ser Leu Asp Ser Asn Glu Val

65 70 75 80

Ser Glu Asn Leu Gly Ala Leu Arg Ala Ile Asp Glu Leu Ile Asp Val

85 90 95

Thr Ile Ser Glu Asn Ala Ser Lys Val Ala Lys Phe Ser Asn Tyr Met

100 105 110

Arg Val Ala Phe Glu Thr Lys Arg Asp Pro Glu Ile Leu Val Leu Ala

115 120 125

Ser Lys Val Leu Gly His Leu Ala Arg Ser Gly Gly Ala Met Thr Ala

130 135 140

Asp Glu Val Glu Arg Gln Val Lys Val Ala Leu Glu Trp Leu Arg Gly

145 150 155 160

Glu Arg Ile Glu Tyr Arg Arg Phe Ala Ala Val Leu Ile Leu Lys Glu

165 170 175

Met Ala Glu Asn Ala Ser Thr Val Phe Asn Val His Val Pro Glu Phe

180 185 190

Val Asp Ala Ile Trp Val Ala Leu Arg Asp Pro Thr Leu Ala Val Arg

195 200 205

Glu Lys Ala Val Glu Ala Leu Arg Ala Cys Leu Arg Val Ile Glu Lys

210 215 220

Arg Glu Thr Arg Trp Arg Val Gln Trp Tyr Tyr Arg Met Phe Glu Ala

225 230 235 240

Thr Gln Asp Gly Leu Gly Arg Asn Ala Pro Val His Ser Ile His Gly

245 250 255

Ser Leu Leu Ala Val Gly Glu Leu Leu Arg Asn Thr Gly Glu Phe Met

260 265 270

Met Ser Arg Tyr Arg Glu Val Ala Glu Ile Val Leu Arg Tyr Leu Glu

275 280 285

His Arg Asp Arg Leu Val Arg Leu Ser Ile Thr Ser Leu Leu Pro Arg

290 295 300

Ile Ala His Phe Leu Arg Asp Arg Phe Val Thr Asn Tyr Leu Thr Ile

305 310 315 320

Cys Met Asn His Ile Leu His Val Leu Lys Ile Pro Ala Glu Arg Ala

325 330 335

Ser Gly Phe Ile Ala Leu Gly Glu Met Ala Gly Ala Leu Asp Gly Glu

340 345 350

Leu Ile Asn Tyr Leu Pro Thr Ile Thr Ser His Leu Arg Asp Ala Ile

355 360 365

Ala Pro Arg Arg Gly Arg Pro Ser Leu Glu Ala Leu Ala Cys Val Gly

370 375 380

Asn Ile Ala Lys Ala Met Gly Pro Thr Met Glu Pro His Val Arg Gly

385 390 395 400

Leu Leu Asp Pro Met Phe Ser Ala Gly Leu Ser Val Thr Leu Val Asp

405 410 415

Ser Leu Asp Leu Leu Thr Glu Ser Ile Pro Pro Leu Leu Pro Thr Ile

420 425 430

Gln Asn Arg Leu Leu Glu Cys Ile Ser Ala Ile Leu Ser Arg Ser His

435 440 445

His Ala Met Ser Arg Gln Ser Ala Ala Leu Ser Arg Gly His Leu Ala

450 455 460

Thr Val Thr Pro Gln Val Pro Glu Leu Ser Gly Ser Ala Leu Val Gln

465 470 475 480

Leu Ser Leu Gln Thr Leu Ala Arg Phe Asn Phe Lys Gly His Asp Leu

485 490 495

Leu Glu Phe Ala Arg Glu Ser Val Val Val Tyr Leu Glu Asp Glu Asp

500 505 510

Gly Ala Thr Arg Lys Asp Ala Ala Leu Cys Cys Cys Lys Leu Ile Ala

515 520 525

Asn Ser Phe Leu Ala Met Ser Ser Thr Gln Phe Ser Pro Ser Arg Ile

530 535 540

Asn Arg Ala Ser Gly Lys Arg Arg Arg Leu Val Glu Glu Ile Val Gln

545 550 555 560

Lys Leu Leu Ile Ala Ala Val Ala Asp Ala Asp Val Thr Val Arg His

565 570 575

Ser Ile Phe Ser Ser Leu Tyr Ala Asp Gly Gly Phe Asp Glu Phe Leu

580 585 590

Ala Gln Ala Asp Ser Leu Thr Ala Ile Phe Ala Thr Leu Asn Asp Glu

595 600 605

Asp Phe Glu Val Arg Glu Tyr Ala Ile Ser Leu Ala Gly Arg Leu Ser

610 615 620

Glu Lys Asn Pro Ala Tyr Val Leu Pro Ala Leu Arg Arg His Leu Ile

625 630 635 640

Gln Leu Leu Thr Tyr Leu Glu Gln Ser Ser Ala Asp Asn Lys Cys Lys

645 650 655

Glu Glu Ser Ala Lys Leu Leu Gly Cys Leu Ile Arg Asn Cys Glu Arg

660 665 670

Leu Val Leu Pro Tyr Val Ser Pro Ile His Lys Ala Leu Val Ala Lys

675 680 685

Leu Cys Glu Gly Thr Gly Val Asn Ala Asn Ser Gly Ile Ile Ser Gly

690 695 700

Val Leu Val Thr Val Gly Asp Leu Ala Arg Val Gly Gly Phe Ala Met

705 710 715 720

Arg Gln Tyr Ile Ser Glu Leu Met Pro Leu Ile Val Glu Ala Leu Leu

725 730 735

Asp Gly Ala Ala Val Thr Lys Arg Glu Val Ala Val Ser Thr Leu Gly

740 745 750

Gln Val Val Gln Ser Thr Gly Tyr Val Ile Thr Pro Tyr Asn Glu Tyr

755 760 765

Pro Gln Leu Leu Gly Leu Leu Leu Lys Leu Leu Asn Gly Glu Leu Ala

770 775 780

Trp Ser Thr Arg Arg Glu Val Leu Lys Val Leu Gly Ile Met Gly Ala

785 790 795 800

Leu Asp Pro His Val His Lys Arg Asn Gln Gln Ser Leu Pro Gly Ser

805 810 815

His Gly Glu Val Thr Arg Val Thr Gly Asp Pro Gly Gln His Ile Arg

820 825 830

Ser Met Asp Glu Leu Pro Thr Asp Leu Trp Pro Ser Phe Ala Thr Ser

835 840 845

Glu Asp Tyr Tyr Ser Thr Val Ala Ile Asn Ser Leu Met Arg Ile Leu

850 855 860

Arg Asp Pro Ser Leu Ser Ser Tyr His Gln Lys Val Val Gly Ser Leu

865 870 875 880

Met Phe Ile Phe Lys Ser Met Gly Leu Gly Cys Val Pro Tyr Leu Pro

885 890 895

Lys Val Leu Pro Asp Leu Phe His Ile Val Arg Ile Cys Glu Asp Gly

900 905 910

Leu Lys Glu Phe Ile Thr Trp Lys Leu Gly Thr Leu Val Ser Ile Ala

915 920 925

Arg Gln His Ile Arg Lys Tyr Leu Pro Glu Leu Leu Ser Leu Ile Ser

930 935 940

Glu Leu Trp Ser Ser Phe Ser Leu Pro Ala Ala Asn Arg Pro Val His

945 950 955 960

Ile Ala Pro Ile Leu His Leu Val Glu Gln Leu Cys Leu Ala Leu Asn

965 970 975

Asp Glu Phe Arg Lys Tyr Leu Pro Asp Ile Leu Pro Cys Cys Ile Gln

980 985 990

Val Leu Thr Asp Ala Glu Arg Phe Asn Asp Tyr Thr Tyr Val Ile Pro

995 1000 1005

Ile Leu His Thr Leu Glu Val Phe Gly Gly Thr Leu Asp Glu His Met

1010 1015 1020

His Leu Leu Phe Pro Ala Leu Ile Arg Leu Phe Lys Val Asp Ala Ser

1025 1030 1035 1040

Val Glu Val Arg Arg Gly Ala Ile Arg Thr Leu Thr Arg Leu Ile Pro

1045 1050 1055

Cys Val Gln Val Thr Gly His Ile Ser Ser Leu Val His His Leu Lys

1060 1065 1070

Leu Val Leu Asp Gly Asn Lys Glu Glu Leu Arg Lys Asp Ala Ile Asp

1075 1080 1085

Ala Leu Cys Cys Leu Ala His Ala Leu Gly Glu Asp Phe Thr Ile Phe

1090 1095 1100

Ile Pro Ser Ile His Lys Leu Met Val Lys His Arg Leu Gln His Lys

1105 1110 1115 1120

Glu Phe Glu Glu Ile Gln Gly Arg Val Glu Lys Arg Glu Pro Leu Ile

1125 1130 1135

Phe Gly Ser Thr Thr Ala Gln Arg Leu Asn Arg Arg Leu Pro Val Glu

1140 1145 1150

Val Ile Ser Asp Pro Leu Ser Asp Gly Glu Ser Asp Leu Tyr Glu Val

1155 1160 1165

Gly Thr Asp Met Gln Lys Gln Leu Arg Asn His Gln Val Asn Asp Gly

1170 1175 1180

Arg Leu Arg Thr Ala Gly Glu Ala Ser Gln Arg Ser Thr Lys Glu Asp

1185 1190 1195 1200

Trp Ala Glu Trp Met Arg His Phe Ser Ile Glu Leu Leu Lys Glu Ser

1205 1210 1215

Pro Ser Pro Ala Leu Arg Thr Cys Ala Arg Leu Ala Gln Leu Gln Pro

1220 1225 1230

Phe Val Gly Arg Glu Leu Phe Ala Ala Gly Phe Val Ser Cys Trp Ser

1235 1240 1245

Gln Leu Asn Glu Ala Ser Gln Arg Gln Leu Val Arg Ser Leu Glu Met

1250 1255 1260

Ala Phe Ser Ser Pro Asn Ile Pro Pro Glu Ile Leu Ala Thr Leu Leu

1265 1270 1275 1280

Asn Leu Ala Glu Phe Met Glu His Asp Glu Arg Pro Leu Pro Ile Asp

1285 1290 1295

Ile Arg Leu Leu Gly Ala Leu Ala Glu Lys Cys Arg Ala Phe Ala Lys

1300 1305 1310

Ala Leu His Tyr Lys Glu Met Glu Phe Glu Gly Ala Leu Ser Asn Arg

1315 1320 1325

Arg Asp Ala Asn Pro Val Ala Val Val Glu Ala Leu Ile His Ile Asn

1330 1335 1340

Asn Gln Leu His Gln Tyr Glu Ala Ala Val Gly Ile Leu Thr Tyr Ala

1345 1350 1355 1360

Gln Gln His Leu Gly Val Gln Leu Lys Glu Ser Trp Tyr Glu Lys Leu

1365 1370 1375

Gln Arg Trp Asp Asp Ala Leu Lys Ala Tyr Thr Ala Lys Ala Ser Gln

1380 1385 1390

Ala Ser Ser Pro His Leu Cys Leu Asp Ala Thr Leu Gly Arg Met Arg

1395 1400 1405

Cys Leu Ala Ala Leu Ala Arg Trp Glu Glu Leu Asn Asn Leu Cys Lys

1410 1415 1420

Glu Tyr Trp Thr Pro Ala Glu Pro Ala Ala Arg Leu Glu Met Ala Pro

1425 1430 1435 1440

Met Ala Ala Asn Ala Ala Trp Asn Met Gly Glu Trp Asp Gln Met Ala

1445 1450 1455

Glu Tyr Val Ser Arg Leu Asp Asp Gly Asp Glu Thr Lys Leu Arg Val

1460 1465 1470

Leu Gly Asn Thr Ala Ser Ser Gly Asp Gly Ser Ser Asn Gly Thr Phe

1475 1480 1485

Tyr Arg Ala Val Leu Leu Val Arg Arg Gly Lys Tyr Asp Glu Ala Arg

1490 1495 1500

Glu Tyr Val Glu Arg Ala Arg Lys Cys Leu Ala Thr Glu Leu Ala Ala

1505 1510 1515 1520

Leu Val Leu Glu Ser Tyr Glu Arg Ala Tyr Ser Asn Met Val Arg Val

1525 1530 1535

Gln Gln Leu Ser Glu Leu Glu Glu Val Ile Glu Tyr Cys Thr Leu Pro

1540 1545 1550

Pro Thr Gly Asn Pro Val Ala Glu Gly Arg Arg Ala Leu Val Arg Asn

1555 1560 1565

Met Trp Asn Glu Arg Ile Lys Gly Ala Lys Arg Asn Val Glu Val Trp

1570 1575 1580

Gln Ala Leu Leu Ala Val Arg Ala Leu Val Leu Pro Pro Thr Glu Asp

1585 1590 1595 1600

Ile Glu Thr Trp Ile Lys Phe Ala Ser Leu Cys Arg Lys Asn Gly Arg

1605 1610 1615

Ile Ser Gln Ala Arg Ser Thr Leu Ile Lys Leu Leu Gln Phe Asp Pro

1620 1625 1630

Glu Thr Thr Pro Ala Thr Val Arg Tyr His Gly Pro Pro Gln Val Met

1635 1640 1645

Leu Ala Tyr Leu Lys Tyr Gln Trp Ser Leu Gly Glu Asp His Lys Arg

1650 1655 1660

Lys Glu Ala Phe Ala Arg Leu Gln Asp Leu Ala Met Asp Leu Ser Arg

1665 1670 1675 1680

Thr Ala Thr Leu Gln Pro Val Met Gln Asn Ala Leu Val Ala Ser Ser

1685 1690 1695

Gly Ala Pro Leu Val Ala Arg Ile Tyr Leu Arg Leu Gly Thr Trp Lys

1700 1705 1710

Trp Ala Leu Ser Pro Gly Leu Asp Asp Asp Ser Ile Gln Glu Ile Leu

1715 1720 1725

Ser Ala Phe Arg Asn Ala Thr His Cys Ala Thr Lys Trp Gly Lys Ala

1730 1735 1740

Trp His Thr Trp Ala Leu Phe Asn Thr Ala Val Met Ser His Tyr Thr

1745 1750 1755 1760

Leu Arg Gly Phe Ala Asn Ile Ala Ala Gln Phe Val Val Ala Ala Val

1765 1770 1775

Thr Gly Tyr Phe His Ser Ile Ala Cys Gly Ala His Ala Lys Gly Val

1780 1785 1790

Asp Asp Ser Leu Gln Asp Ile Leu Arg Leu Leu Thr Leu Trp Phe Asn

1795 1800 1805

His Gly Ala Thr Ser Glu Val Gln Met Ala Leu Gln Lys Gly Phe Thr

1810 1815 1820

His Val Asn Ile Asn Thr Trp Leu Val Val Leu Pro Gln Ile Ile Ala

1825 1830 1835 1840

Arg Ile His Ser Asn Asn His Ala Val Arg Glu Leu Ile Gln Ser Leu

1845 1850 1855

Leu Val Arg Ile Gly Gln Ser His Pro Gln Ala Leu Met Tyr Pro Leu

1860 1865 1870

Leu Val Ala Cys Lys Ser Ile Ser Asn Leu Arg Arg Ala Ala Ala Gln

1875 1880 1885

Glu Val Val Asp Lys Val Arg Gln His Ser Gly Val Leu Val Asp Gln

1890 1895 1900

Ala Gln Leu Val Ser Lys Glu Leu Ile Arg Val Ala Ile Leu Trp His

1905 1910 1915 1920

Glu Met Trp His Glu Ala Leu Glu Glu Ala Ser Arg Leu Tyr Phe Gly

1925 1930 1935

Glu His Asn Ile Glu Gly Met Leu Lys Val Leu Glu Pro Leu His Glu

1940 1945 1950

Met Leu Glu Glu Gly Ala Met Arg Asn Asn Thr Thr Met Lys Glu Lys

1955 1960 1965

Ala Phe Ile Gln Ala Tyr Arg Ile Glu Leu Leu Glu Ala Tyr Glu Cys

1970 1975 1980

Cys Met Lys Tyr Arg Arg Thr Gly Lys Asp Ala Glu Leu Ile Gln Ala

1985 1990 1995 2000

Trp Asp Leu Tyr Tyr His Val Phe Arg Arg Ile Asp Lys Gln Leu Gln

2005 2010 2015

Thr Leu Thr Thr Leu Asp Leu Gln Ser Val Ser Pro Glu Leu Leu Glu

2020 2025 2030

Cys Arg Asp Leu Glu Leu Ala Val Pro Gly Thr Tyr Arg Ala Asp Thr

2035 2040 2045

Pro Val Val Thr Ile Ala Ser Phe Ala Pro Gln Leu Val Val Ile Thr

2050 2055 2060

Ser Lys Gln Arg Pro Arg Lys Leu Thr Ile His Gly Ser Asp Gly Lys

2065 2070 2075 2080

Asp Tyr Ala Phe Leu Leu Lys Gly His Glu Asp Leu Arg Gln Asp Glu

2085 2090 2095

Arg Val Met Gln Leu Phe Gly Leu Val Asn Thr Leu Leu Glu Asn Ser

2100 2105 2110

Arg Lys Thr Ala Glu Lys Asp Leu Ser Ile Gln Arg Tyr Asp Val Ile

2115 2120 2125

Pro Leu Ser Pro Asn Ser Gly Leu Ile Glu Trp Val Pro Asn Cys Asp

2130 2135 2140

Thr Leu His Gln Leu Ile Arg Glu Tyr Arg Asp Ala Arg Lys Ile Thr

2145 2150 2155 2160

Leu Asn Gln Glu His Lys Leu Met Leu Ser Phe Ala Pro Asp Tyr Asp

2165 2170 2175

Asn Leu Pro Leu Ile Ala Lys Val Glu Val Phe Glu Tyr Ala Leu Gln

2180 2185 2190

Asn Thr Glu Gly Asn Asp Leu Ser Arg Val Leu Trp Leu Lys Ser Arg

2195 2200 2205

Thr Ser Glu Val Trp Leu Asp Arg Arg Thr Asn Tyr Thr Arg Ser Leu

2210 2215 2220

Ala Val Met Ser Met Val Gly Tyr Leu Leu Gly Leu Gly Asp Arg His

2225 2230 2235 2240

Pro Ser Asn Leu Met Leu His Arg Phe Ser Gly Lys Ile Leu His Ile

2245 2250 2255

Asp Phe Gly Asp Cys Phe Glu Ala Ser Met Asn Arg Glu Lys Phe Pro

2260 2265 2270

Glu Lys Val Pro Phe Arg Leu Thr Arg Met Leu Val Lys Ala Met Glu

2275 2280 2285

Val Ser Gly Ile Glu Gly Asn Phe Arg Ser Thr Cys Glu Asn Val Met

2290 2295 2300

Gln Val Leu Arg Leu His Lys Asp Ser Val Met Ala Met Met Glu Ala

2305 2310 2315 2320

Phe Val His Asp Pro Leu Ile Asn Trp Arg Leu Phe Asn Phe Asn Glu

2325 2330 2335

Val Pro Gln Met Ser Thr Leu Ala Ser Ala His Val Pro Pro Val Val

2340 2345 2350

Asn Ser Glu Asp Ser Ser Ser Asp Arg Glu Leu Leu Gln Pro Gln Arg

2355 2360 2365

Gly Ala Arg Glu Arg Glu Leu Leu Gln Ala Val Asn Gln Leu Gly Asp

2370 2375 2380

Ala Asn Glu Val Leu Asn Glu Arg Ala Val Ala Val Met Ala Arg Met

2385 2390 2395 2400

Ser Asn Lys Leu Thr Gly Arg Asp Phe Ala Ala Thr Ser Ala Ser Ser

2405 2410 2415

Ser Ser Leu Gln His Pro Leu Asp His Ser Thr Leu Ile Ser Gly Glu

2420 2425 2430

Thr Arg Glu Ala Asp His Gly Leu Ser Val Lys Leu Gln Val Gln Lys

2435 2440 2445

Leu Ile Gln Gln Ala Met Ser His Glu Asn Leu Cys Gln Asn Tyr Val

2450 2455 2460

Gly Trp Cys Pro Phe Trp

2465 2470

<210> 3

<211> 38

<212> DNA

<213> Artificial sequence

<400> 3

cctccatggg gatccgaccg gaaatttccc tctatacc 38

<210> 4

<211> 37

<212> DNA

<213> Artificial sequence

<400> 4

cgtgagctcg gtaccgcacc ggattatgat aatttgc 37

Claims

1. The application of nucleic acid molecules for destroying SlTOR gene expression in improving tomato glandular hair density is characterized in that the nucleotide sequence of the SlTOR gene is shown as SEQ ID No. 1.

2. The use of a substance that reduces the content of a protein encoded by the SlTOR gene of claim 1 in increasing the density of tomato glandular hairs, wherein the amino acid sequence of the protein encoded by the SlTOR gene is shown in SEQ id No.2, and the substance is AZD8055.

3. A method for improving tomato glandular hair density is characterized in that SlTOR genes in target tomato plants are silenced through a gene editing technology, or AZD8055 is applied to the outside of the target tomato plants, so that the obtained tomato plant glandular hair density is improved, and the nucleotide sequence of the SlTOR genes is shown in SEQ ID No. 1.