CN110628808B - Arabidopsis AtTCP5 gene and application thereof in regulating plant height - Google Patents

Abstract

The invention discloses an arabidopsis AtTCP5 gene and application thereof in regulating and controlling plant height. The expression quantity and/or activity of the protein of the amino acid sequence shown in SEQ ID No.1 coded by the AtTCP5 gene can influence plant cell proliferation, wherein a plant over-expressing the AtTCP5 has a phenotype that the plant height is obviously shortened, so that the plant height is regulated. The AtTCP5 gene has wide application value in plant height regulation, and molecular breeding by using the gene is expected to breed lodging-resistant plants with practical production application value.

Description

Arabidopsis AtTCP5 gene and application thereof in regulating plant height

Technical Field

The invention belongs to the technical field of plant genetic engineering, and particularly relates to an arabidopsis gene AtTCP5 capable of reducing plant height and application of the gene.

Background

The plant height is an important measurement standard of the plant type of the crops, and proper reduction of the plant height of the crops plays an important role in improving the lodging resistance of the crops and changing the distribution of nutrient substances in the vegetative growth and reproductive growth processes, and finally influences the yield of the crops.

The arabidopsis thaliana is one of the most classical model plants, and has the advantages of short reproductive cycle, more offspring, simple growth conditions, self-flower closed-flower pollination, sequenced whole genome, abundant and accurate comments, small plant size and the like, so that the arabidopsis thaliana plays a great role in basic scientific research. The research of arabidopsis thaliana as a model plant is very convenient at present, and a plurality of results of gene research can be applied to the genetic modification of crops.

The traditional breeding method usually adopts natural mutation strains to hybridize with other varieties with better character performance, and selects strains with excellent and stable inheritance characters in the offspring to breed new varieties. A plurality of excellent varieties have been bred by adopting a cross breeding method. However, the traditional breeding method still has huge defects, wherein the breeding period is long, the floor space is large, the manpower consumption is high, and the character linkage caused by gene linkage is difficult to break, so that the development of the traditional breeding is restricted. However, in recent years, molecular biology and genetic engineering techniques are rapidly developed, and a target gene can be transferred into a variety to be improved through a transgenic technology, so that plant traits are more specifically and efficiently changed, and a new thought is provided for improving grain yield.

The TCP gene family is a plant-specific transcription factor family, and the transcription factors participate in each process of plant growth and development and are a very key plant development regulation and control element. TCP transcription factors are named for the first few genes found to belong to this class of transcription factors: TEOSINTE BRANCHED 1(TB1) in maize, CYCLOIDEA (CYC) in snapdragon, and PROLIFERATING CELL FACTORS 1 and 2(PCF1 and PCF2) in rice. Research shows that the TCP transcription factor family plays a role in hormone signal pathways such as signal pathways of auxin, cytokinin, jasmonic acid, strigolactone and the like, and mainly regulates the division and expansion processes of cells.

The TCP gene family is mainly divided into two major classes, TCP class i and TCP class ii. The first class of TCP is mainly expressed in meristems and is responsible for regulating cell cycle and further regulating processes such as cell division, and the like, which indicates that the first class of TCP transcription factors can be used as activating factors of cell division to promote the growth and proliferation of plants; the second class of TCP genes can be further divided into a class CIN TCP transcription factor and a class CYC/TB1 TCP transcription factor. Wherein, the CIN TCP transcription factor has the main function of inhibiting cell proliferation, and in Arabidopsis, the multi-mutant of the CIN TCP gene shows a rolling phenotype of excessive growth of leaf edges; the CYC/TB1 class of TCP transcription factors are reported to play a role in regulating floral organ symmetry and inhibiting the activity of lateral meristems.

TCP14 and TCP15 in a TCP transcription factor family belong to a first class of TCP transcription factors, and in Arabidopsis thaliana, double mutants of TCP14 and TCP15 show the phenotype of strain height change and internode shortening, which shows that the first class of TCP transcription factors in Arabidopsis thaliana have the function of promoting the plant height to be changed.

Disclosure of Invention

The invention aims to provide a gene for regulating the height of a strain and a protein coded by the gene. We found that the expression of the TCP5 gene of the CIN TCP family in Arabidopsis thaliana obviously shortens the plant height of the plant. This result further illustrates: the first class of TCP transcription factors may act as cell division activators to promote plant growth and proliferation; the main function of the CIN TCP transcription factor is to inhibit cell proliferation.

The invention provides the following technical scheme:

a protein for regulating plant height is derived from Arabidopsis thaliana (Arabidopsis thaliana), is named AtTCP5, and is a protein shown by the following amino acid sequence 1) or 2):

1) SEQ ID No: 1;

2) SEQ ID No:1, and the derived protein has the function of reducing the plant height.

SEQ ID No:1 consists of 360 amino acid residues and contains the TCP domain. Methods for substituting, deleting or adding amino acid residues are well known to those skilled in the art, and usually, genetic engineering means is used to mutate the encoding gene and then express the corresponding protein. The one to ten amino acid residues substituted, deleted or added may be amino acid residues in a non-conserved region, the alteration of which does not affect the function of the protein. By expressing the protein in the plant and observing the plant height condition of the obtained plant strain, whether the protein after the change has the function of regulating and controlling the plant height can be judged.

The expression quantity and/or activity of the protein AtTCP5 for regulating the plant height provided by the invention can influence the plant cell proliferation, wherein the plant over-expressing AtTCP5 has the phenotype that the plant height is obviously shortened, thereby realizing the regulation of the plant height.

The protein for coding the plant height regulation is an arabidopsis AtTCP5 gene, and the sequence of the protein can be a cDNA sequence of the gene, a genome DNA sequence of the gene, or a DNA sequence which has more than 90 percent of homology with the sequences and codes the same functional protein. Such as SEQ ID NO:2 and the genomic DNA sequence shown in SEQ ID NO:3, and (b) 3.

Expression vectors comprising the above nucleotide sequences and expression control sequences operatively linked to the nucleotide sequences are also within the scope of the present invention.

The protein for regulating the plant height of the plant or the application of the coding gene thereof provided by the invention comprises but is not limited to the following aspects:

1) controlling the plant height of the plant;

2) creating a new plant variety with improved lodging resistance;

3) changing the distribution of nutrients during vegetative and reproductive growth;

4) affecting the yield of the crop.

The invention also provides a method for reducing plant height, which comprises the steps of introducing the arabidopsis AtTCP5 gene or homologous gene thereof into plant cells, tissues or organs, and culturing the transformed plant cells, tissues or organs into plants to obtain transgenic plants with reduced plant height.

The arabidopsis AtTCP5 gene or a homologous gene thereof is typically introduced into a plant cell, tissue or organ by a plant expression vector. The plant expression vector comprises the arabidopsis AtTCP5 gene (or homologous gene thereof) sequence and an expression regulation sequence operatively connected with the gene sequence. In a preferred embodiment, the expression control sequence comprises a constitutively high expressing control sequence, such as the CaMV35S promoter.

In the method for reducing the plant height of the plant, the AtTCP5 gene may be a cDNA sequence of the gene, a genomic DNA sequence of the gene, or a DNA sequence having 90% or more homology with the cDNA sequence and encoding the same functional protein. The DNA sequence having 90% or more homology with the above sequence and encoding the same functional protein is obtained by isolating and/or modifying and/or designing the cDNA or genomic DNA sequence of the gene by a known method. It will be appreciated by those skilled in the art that minor changes in nucleotide identity in a gene sequence will in many cases not result in a reduction or enhancement in the efficiency of the gene. Methods for altering the sequence of genes, and methods for testing the effectiveness of such altered genes, are well known to those skilled in the art.

The AtTCP5 gene or its homologous sequence can be introduced into plant tissue, cell or organ via plant expression vector. The starting vector for constructing the plant expression vector can be any binary vector which can be used for transforming plants by agrobacterium tumefaciens or agrobacterium rhizogenes or a vector which can be used for plant microprojectile bombardment, such as a Gateway series vector (such as pB7FWG2 and the like), a pBin series vector (such as pBin 19 and the like), a pJim series vector (such as pJim 19 and the like), a pCAMBIA series vector (such as pCAMBIA 1301 and the like), per8, pX6 or other derivative plant expression vectors, and the starting vector can also be a vector which can be replicated in prokaryotes, such as a pENTR/D-TOPO vector, a pUC series vector or a pBluescript series vector and the like.

When the AtTCP5 gene or its homologous sequence is used to construct plant expression vector, any one of reinforced, constitutive or inducible promoter may be added before its transcription initiation nucleotide. The constitutive expression promoter can be a cauliflower mosaic virus (CAMV)35S promoter, a maize Ubiquitin promoter or a rice actin1 promoter and the like; the inducible promoter can be a promoter induced by low temperature, drought, ABA, ethylene, saline alkali or chemistry and the like. The above promoters may be used alone or in combination with other plant promoters. In addition, when the gene of the present invention is used to construct plant expression vectors, enhancers, including translational or transcriptional enhancers, may be used, and these enhancer regions may be ATG initiation codon or initiation codon of adjacent regions, etc., but must be in the same reading frame as the coding sequence to ensure proper translation of the entire sequence. The translational control signals and initiation codons are widely derived, either naturally or synthetically. The translation initiation region may be derived from a transcription initiation region or a structural gene.

In order to facilitate the identification and screening of transgenic plant cells or plants, plant expression vectors to be used may be processed, for example, by adding a gene encoding an enzyme or a luminescent compound which can produce a color change (GUS gene, GFP gene, luciferase gene, etc.), an antibiotic marker having resistance (neomycin phosphotransferase (NPTII) gene, Hygromycin phosphotransferase (Hygromycin phosphotransferase) gene, gentamycin marker, kanamycin marker, etc.), or a chemical agent resistance marker gene (e.g., herbicide resistance gene), etc., which can be expressed in plants. After the screening by the method, molecular detection means such as Southern, PCR or dot hybridization can be adopted to detect the transgenic plant so as to determine whether the transgenic plant is transformed with a target gene.

The plant expression vector carrying the AtTCP5 gene or its homologous sequence of the present invention can be obtained by using protoplast-chemical ligation (Ca)²⁺PEG), Ti plasmid, Ri plasmid, plant virus vector, direct DNA transformation, pollen tube introduction, micro injection, electric excitation, gene gun, agrobacterium mediation and other conventional biological methods, and culturing the transformed plant cell, tissue or organ into a plant; the tissues and organs may include pods, callus, stem tips, leaves, seeds, etc. of the host plant.

The invention also provides a method for obtaining a highly dwarf plant line, comprising:

1) introducing the expression vector of the invention into the plant body;

2) and (3) expressing the expression vector in a host cell to obtain the plant line with the strain height dwarfing character.

In a preferred embodiment, the plant line is from arabidopsis thaliana.

The invention over-expresses the AtTCP5 gene in Arabidopsis thaliana, and the over-expressed plant can be observed to have a phenotype that the plant height is obviously shortened. Therefore, the AtTCP5 gene has wide application value in plant height regulation, and molecular breeding by using the gene is expected to breed lodging-resistant plants with practical production application value.

Drawings

FIG. 1 shows the expression levels of the TCP5 gene in wild-type Arabidopsis thaliana (WT) and TCP5 overexpressing the strain No. 3 (TCP5OX-3) and strain No. 9 (TCP5OX-9) in transgenic plants.

FIG. 2 shows the phenotype of wild type Arabidopsis thaliana (WT) and strain No. 3 (TCP5OX-3) and strain No. 9 (TCP5OX-9) in TCP5 overexpressing transgenic plants.

Detailed Description

The methods used in the following examples are conventional unless otherwise specified, and specific procedures can be found in: molecular Cloning: A Laboratory Manual (Sambrook, J., Russell, David W., Molecular Cloning: A Laboratory Manual, 3)^rd edition，2001，NY，Cold Spring Harbor)。

Example 1 cloning of genes

Cloning of genomic DNA sequence of AtTCP5 gene:

the genomic DNA sequence (SEQ ID No:2), CDS sequence (SEQ ID No:3) and amino acid sequence (SEQ ID No:1) of the AtTCP5 gene were obtained from the Arabidopsis thaliana genomic database (http:// www.arabidopsis.org /).

Gene-specific PCR primers were designed based on the CDS sequence of the AtTCP5 gene:

F1：5’-CACCATGAGATCAGGAGAATGTGA-3’(SEQ ID NO:4)；

R1：5’-TCAAGAATCTGATTCATTAT-3’(SEQ ID NO:5)。

the genomic sequence of the AtTCP5 gene (not containing 3 'untranslated region and 5' untranslated region) was cloned from the genomic DNA of Arabidopsis thaliana (Arabidopsis thaliana) Columbia ecotype by Pfu high fidelity enzyme amplification cloning using the pair of primers. The PCR product is connected into pENTR-D-TOPO vector by TOPO reaction, vector with forward insert (ATG of gene near M13F sequencing primer) is screened and sequenced, the plasmid with correct sequencing is named TOPO-TCP5sc, and is used for constructing plant expression vector of AtTCP5 gene.

(II) construction of the AtTCP5 gene plant expression vector:

in order to construct a plant expression vector of AtTCP5 gene, the TOPO-TCP5sc vector is connected into a pK2GW7 vector (purchased from university of Gente) through LR reaction, after positive clone is screened and sequenced correctly, the positive clone is named as 35S-TCP5, and the positive clone is used as a final vector to carry out the next experiment.

Example 2 obtaining of transgenic Arabidopsis thaliana constitutively expressing AtTCP5 Gene

(I) agrobacterium transformation:

agrobacterium GV3101 competent cells were transformed with 35S-TCP5 vector by electric excitation, plated with solid LB medium containing 50 mg/l gentamicin, 50 mg/l rifampicin and 50 mg/l spectinomycin, and after 2 days of dark culture at 28 ℃ positive clones were selected with primers F1 and R1. The obtained positive clone was inoculated into a liquid LB medium containing 50 mg/l gentamicin, 50 mg/l rifampicin and 50 mg/l spectinomycin, shake-cultured at 28 ℃ for 24 hours, 0.5 ml of bacterial liquid was taken, and the strain was preserved for Arabidopsis transformation.

(II) Arabidopsis thaliana transformation:

1 ml of the strain was added to 10 ml of liquid LB medium containing 50 mg/l gentamicin, 50 mg/l rifampicin and 50 mg/l spectinomycin, shake-cultured at 28 ℃ for 24 hours, and the whole strain was transferred to 300 ml of liquid LB medium containing 50 mg/l gentamicin, 50 mg/l rifampicin and 50 mg/l spectinomycin, and shake-cultured at 28 ℃ for 12 hours. The obtained 300 ml of bacterial liquid is centrifuged at 3000 rpm for 15 minutes, the supernatant is discarded, the bacteria are resuspended in 100 ml of infection buffer, and the formula of the infection buffer is as follows (1 liter): 50 g of cane sugar, 2.2 g of MS powder and 200 microliter of silwet, and water is added to one liter.

Selecting an arabidopsis thaliana plant which shoots for one to two weeks and has good growth condition, cutting off siliques and opened flowers, leaving apical meristems and buds, soaking the treated arabidopsis thaliana inflorescences in 100 ml of resuspended bacterial liquid for 1-2 minutes, treating the soaked plant in the dark at 22 ℃ for 24 hours, and then placing the plant in the light for growth.

(III) screening transformed plants:

after the transformed Arabidopsis plants continued to grow for about one month, the mature seeds were collected, divided into 1.5 ml centrifuge tubes, 1 ml 75% ethanol (containing 0.5% triton x-100) was added, sterilized by shaking for 15 minutes, then washed 1 time with absolute ethanol, and then blown dry in a clean bench. The sterilized seeds were uniformly placed on 1/2MS solid medium containing 50 mg/L kanamycin and 100 mg/L, dark-treated at 4 ℃ for 48 hours, then placed under 22 ℃ for light culture for 6-7 days, and screened for normal-growing positive seedlings to be transplanted into soil. When the plant grows to about 21 days, cutting true leaves of about 0.5 square centimeter to extract DNA, carrying out PCR reaction by using primers on the vector and primers on the genes, and screening out positive plants.

Example 3 detection of AtTCP5 Gene expression level in transgenic Arabidopsis thaliana of AtTCP5 Gene

Leaves of positive transgenic Arabidopsis thaliana adults were cut, total plant RNA was extracted with TRIzol reagent (Invitrogen), DNA was digested with DNase I (Takara), and then reverse transcription was performed using a reverse transcription kit of Invitrogen to obtain cDNA of a transgenic plant. The real-time quantitative PCR primers realF and realR with gene specificity are designed according to the cDNA sequence of the AtTCP5 gene, the expression level of the AtTCP5 gene in a transgenic plant is detected through the real-time quantitative PCR, and the Arabidopsis thaliana ACTIN8 gene (the primers are realF2 and realR2) is selected as an internal reference gene. The sequences of the primers are as follows:

realF：5'-TCTCAAGAACATTCGGTGGC-3'(SEQ ID NO:6)

realR：5'-CTACGTCATCTTTTGCTGCT-3'(SEQ ID NO:7)

realF2：5'-TCCAGGCATTGTCCACAGAA-3'(SEQ ID NO:8)

realR2：5'-ACCTGCTCCTCCTTAGACAT-3'(SEQ ID NO:9)

the relative expression conditions of the ATCP5 gene in wild type and transgenic rice plants through real-time quantitative PCR detection are shown in figure 1, compared with the wild type, the expression quantity of TCP5 in a No. 3 strain of an overexpression transgenic plant is increased by about 170 times, and the expression quantity of TCP5 in a No. 9 strain is increased by about 100 times.

Example 4 transgenic Arabidopsis phenotype Observation of AtTCP5 Gene

After sterilization and cleaning of wild type and TCP5 overexpression transgenic positive plants by using 75% ethanol, placing the plants on a 1/2MS culture medium for growth, culturing the plants for 6-7 days at 22 ℃ under normal illumination until cotyledons are opened normally for greening, transplanting the plants into nutrient soil when the roots are about 0.5 cm long, growing the plants for about 35 days at 22 ℃ under normal illumination, observing the difference of the overall phenotype of the plants compared with the wild type, and observing that the height of the TCP5 overexpression plants is obviously shortened compared with the wild type (figure 2).

The invention obtains the overexpression transgenic arabidopsis of the TCP5 gene by transforming arabidopsis, and discovers that the overexpression transgenic arabidopsis strain of the TCP5 gene is short in height by observation. This is of great significance for obtaining high yield crops by transgenic methods.

SEQUENCE LISTING

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Claims (7)

1. The application of AtTCP5 protein derived from arabidopsis thaliana or coding gene thereof in regulating and controlling plant height, wherein the AtTCP5 protein is SEQ ID NO:1 amino acid sequence of a protein; the application is to improve arabidopsis in arabidopsisAtTCP5And obtaining the transgenic plant with reduced plant height by the expression amount of the gene.

2. Use according to claim 1, wherein said Arabidopsis thaliana isAtTCP5The nucleotide sequence of the gene is shown as SEQ ID NO:2 or SEQ ID NO:3, respectively.

3. Use according to claim 1 or 2, wherein said arabidopsis thaliana is usedAtTCP5The gene is introduced into plant cell, tissue or organ, and the transformed plant cell, tissue or organ is cultured into plant to obtain transgenic plant with reduced plant height.

4. Use according to claim 3, wherein said Arabidopsis thaliana isAtTCP5The gene is introduced into a plant cell, tissue or organ by a plant expression vector.

5. The use of claim 4, wherein said plant expression vector comprises said Arabidopsis thalianaAtTCP5The sequence of the gene and expression regulatory sequences operatively linked to the gene sequence.

6. The use of claim 5, wherein said expression control sequence is a control sequence that drives constitutive high expression of said genome.

7. The use of claim 6, wherein the plant expression vector is primed with CaMV35SThe rotor drives the Arabidopsis thalianaAtTCP5Expression of the gene.

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