CN114438102B - Strawberry ethylene response FaERF13 gene and application thereof in changing strawberry fruit maturity - Google Patents
Strawberry ethylene response FaERF13 gene and application thereof in changing strawberry fruit maturity Download PDFInfo
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- C12N15/00—Mutation or genetic engineering; DNA or RNA concerning genetic engineering, vectors, e.g. plasmids, or their isolation, preparation or purification; Use of hosts therefor
- C12N15/09—Recombinant DNA-technology
- C12N15/63—Introduction of foreign genetic material using vectors; Vectors; Use of hosts therefor; Regulation of expression
- C12N15/79—Vectors or expression systems specially adapted for eukaryotic hosts
- C12N15/82—Vectors or expression systems specially adapted for eukaryotic hosts for plant cells, e.g. plant artificial chromosomes (PACs)
- C12N15/8216—Methods for controlling, regulating or enhancing expression of transgenes in plant cells
- C12N15/8218—Antisense, co-suppression, viral induced gene silencing [VIGS], post-transcriptional induced gene silencing [PTGS]
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- C12N15/00—Mutation or genetic engineering; DNA or RNA concerning genetic engineering, vectors, e.g. plasmids, or their isolation, preparation or purification; Use of hosts therefor
- C12N15/09—Recombinant DNA-technology
- C12N15/63—Introduction of foreign genetic material using vectors; Vectors; Use of hosts therefor; Regulation of expression
- C12N15/79—Vectors or expression systems specially adapted for eukaryotic hosts
- C12N15/82—Vectors or expression systems specially adapted for eukaryotic hosts for plant cells, e.g. plant artificial chromosomes (PACs)
- C12N15/8241—Phenotypically and genetically modified plants via recombinant DNA technology
- C12N15/8242—Phenotypically and genetically modified plants via recombinant DNA technology with non-agronomic quality (output) traits, e.g. for industrial processing; Value added, non-agronomic traits
- C12N15/8243—Phenotypically and genetically modified plants via recombinant DNA technology with non-agronomic quality (output) traits, e.g. for industrial processing; Value added, non-agronomic traits involving biosynthetic or metabolic pathways, i.e. metabolic engineering, e.g. nicotine, caffeine
- C12N15/8249—Phenotypically and genetically modified plants via recombinant DNA technology with non-agronomic quality (output) traits, e.g. for industrial processing; Value added, non-agronomic traits involving biosynthetic or metabolic pathways, i.e. metabolic engineering, e.g. nicotine, caffeine involving ethylene biosynthesis, senescence or fruit development, e.g. modified tomato ripening, cut flower shelf-life
Abstract
The invention discloses strawberry ethylene regulation and controlFaERF13The gene and the application thereof,FaERF13the nucleotide sequence of the gene is shown as SEQ ID NO.1, and the amino acid sequence of the protein is shown as SEQ ID NO. 2. The invention is realized by putting strawberriesFaERF13After the gene is separated, the VIGS vector is constructed to transform strawberry fruits, the mature period of the gene silencing fruits is obviously earlier than that of negative control, which shows thatFaERF13The gene is a negative regulatory factor for strawberry fruit ripening,FaERF13the gene has important application value in cultivating new strawberry varieties with early or late ripening (storage resistance) of fruits.
Description
Technical Field
The invention belongs to the technical field of plant genetic engineering, and particularly relates to a strawberry fruit maturation regulatory gene FaERF13 and application thereof.
Background
Strawberry (Fragaria x ananasa Duch.) is a perennial herb of Rosaceae, and is favored by consumers because of its special fragrance, beautiful fruit, and rich vitamin C. The growth and development and the maturity of the facility strawberries are relatively consistent, and the strawberries are easy to form and come into the market in a centralized way. Therefore, the research on how to change the mature period of the strawberries, lead the strawberries to market or prolong the fruit period can obtain higher economic value. Fruits are classified into a jump-type fruit and a non-jump-type fruit according to whether or not a peak in respiration occurs before the fruit ripens. The respiratory transition of the fruit is accompanied by ethylene transition, and the fruit without transition has no ethylene transition before and after fruit ripening. Currently, studies on the regulation of fruit ripening by ethylene have focused mainly on fruits of the respiratory depression type, while the function and mechanism in regulating the ripening of fruits of the non-respiratory depression type, such as strawberry, are not yet clear. Strawberry fruit is capable of producing ethylene and expressing ethylene receptor genes. Exogenous ethephon can promote the coloration and softening of strawberry fruits and partially restore the coloration of RNAi strain fruits.
ERF transcription factors are ethylene response factors involved in the ethylene signal transduction pathway and are one of the AP2/ERF transcription factor superfamily. The proteins encoded by the AP2/ERF superfamily genes all contain a conserved AP2/ERF domain, having a DNA-binding domain AP2 domain of 60-70 amino acids, consisting of 3 beta-sheets and 1 alpha helix. The AP2 domain was originally identified in the APETAL2 protein that regulates flower growth in Arabidopsis. A protein module which is combined with the GCC box, namely an ERF domain, is found in tobacco ethylene response proteins ERF1, ERF2, ERF3 and ERF4, and the transcription level of ethylene response genes is regulated. The ERF gene family is the most important large family in the plant AP2/ERF transcription factor superfamily, and plays a very important role in the process of plant growth and development and stress response. Studies show that FaERF9 can combine with FaMYB98 to jointly induce the transcription level of a key synthase encoding gene FaQR of strawberry flavor substances 4-hydroxy-2, 5-dimethyl-3 (2H) furanone, and the AP2/ERF family can also regulate biosynthesis of plant hormones including ethylene, cytokinin, gibberellin and abscisic acid. AP2/ERF is also involved in the response of auxin, cytokinin, abscisic acid and jasmonic acid signals and is a key factor in the attachment of plant hormone signals.
At present, whether ethylene regulates the ripening of strawberry fruits and the mechanism problem in the ethylene are to be solved urgently, and the effect of an ERF gene in regulating the ripening of strawberry fruits is explored, so that a gene breeding tool can be provided for regulating the ripening period of the strawberry. On one hand, the elucidation and application of the regulation and maturation functions of the related genes have the effects of promoting early ripening of strawberries, achieving the purpose of increasing economic added value by marketing in advance, on the other hand, fresh fruits of the strawberries are not easy to deteriorate due to storage, and the application of the related genes can delay the maturation period, so that the effect of reducing logistics and storage cost due to peak shifting of the fruit period is achieved.
Disclosure of Invention
The invention aims to overcome the defects in the prior art, provides a strawberry ethylene response gene FaERF13 and application thereof in changing the maturity of strawberry fruits, and aims to provide the strawberry ethylene response gene FaERF13, meet the use requirements and also provide application of the strawberry ethylene response gene FaERF13.
In order to achieve the above object, the present invention adopts the following technical scheme:
a strawberry ethylene response gene FaERF13 has a nucleotide sequence shown in SEQ ID NO. 1.
The amino acid sequence of the expression protein of the strawberry ethylene response gene FaERF13 is shown as SEQ ID NO. 2.
The pNC-TRV2-VIGS vector containing the strawberry ethylene response gene FaERF13.
The pNC-Cam1304-35S overexpression vector containing the strawberry ethylene response gene FaERF13.
The pNC-Cam1304-RNAi vector containing the strawberry ethylene response gene FaERF13.
The effect of the strawberry ethylene response gene FaERF13 in changing the ripening of strawberry fruits.
The beneficial effects are that: the FaERF13 gene is cloned by taking strawberry variety 'peach fumigation' as a material. The real-time fluorescent quantitative detection technology is used for detecting the expression mode of the FaERF13 gene in different maturity stages, and meanwhile, the VIGS carrier is constructed to instantaneously transform strawberry fruits, so that the FAERF13 gene proves that the FAERF13 gene is a negative regulation factor for strawberry fruit maturity, and the carrier for stably over-expressing or silencing the FaERF13 gene is constructed, so that the FAERF13 gene has important application value in the breeding fields of improving strawberry yield, promoting strawberry maturity and the like.
Drawings
FIG. 1 is a diagram showing the spatiotemporal expression pattern of the FaERF13 gene;
FIG. 2 is a plant VIGS vector PNC-TRV2 plasmid map;
FIG. 3 is a graph of FaERF13 silenced versus negative group phenotype change controls;
FIG. 4 is a graph of FaERF13 gene silencing group versus negative group expression patterns;
FIG. 5 is a plasmid map of plant over-expression vector pNC-Cam 1304-35S;
FIG. 6 is a plasmid map of plant RNAi vector pNC-Cam 1304-RNA.
Detailed Description
The invention is further described below with reference to examples and figures.
Example 1
FaERF13 Gene cloning
And (3) taking cDNA of the strawberry fruits which are 'peach fumigation' as a template, amplifying the gene coding region fragments, connecting the cloning vectors and sequencing to obtain the gene nucleotide sequence.
(1) Primer design
Based on the octaloid strawberry genome, a gene FaERF13 amplification primer is designed
Primer F：5’-CTATCTAATCTTTGCCTTGG-3’
Primer R：5’-ATGGTTGTCTGTCCGTACAT-3’
(2) PCR amplification
RNA is extracted by taking the desclea receptacle of the 'peach fumigation' strawberry in the mature period as a template, a cDNA template is obtained through reverse transcription, and the CDS sequence of FaERF13 is amplified through PCR:
PCR amplification System (Total 50. Mu.l):
the reaction procedure:
(3) Ligation of purified fragments with cloning vector
Connection carrier pEASY-Blunt Cloning Kit (full gold, beijing)
Reaction system (5 μl):
PCR Product： 50ng
pEASY-Blunt Zero Cloning Vector 1μL
ddH 2 O up to 5μL
reaction conditions: 25 ℃ for 10min
E.coli transformation: thawing freshly prepared or frozen E.coli Trans-T1 competent cells at-80℃on ice; adding the ligation product into 50 mu L competent cells, and lightly mixing, and ice-bathing for about 20 min; heat-beating in water bath at 42 deg.C for 60sec, and rapidly placing on ice for 2min; adding 500 mu L of LB liquid medium, shaking at 37 ℃ and 200rpm for 1h; centrifuging at 4000rpm for 2min, discarding part of supernatant, retaining 100 μl of bacterial liquid, and mixing; the bacterial liquid was smeared on LB screening plates containing kan and cultured upside down at 37℃overnight.
(4) Positive clone screening and sequencing analysis
Selecting single colony from the screening culture plate, inoculating the single colony into LB liquid culture medium, and shaking at 37 ℃ and 250rmp for 6 hours; directly taking the bacterial liquid cultured for 6 hours as a template for bacterial liquid PCR detection
Reaction system (20 μl):
the reaction procedure:
clone positive for bacterial liquid PCR detection is sent to biological company (Nanjing) for sequencing
Example 2
FaERF13 Gene space-time expression Pattern analysis
The abundance of the FaERF13 gene in different development stages of fruits is quantified through a fluorescent quantitative PCR technology, a fluorescent quantitative PCR primer forward primer 5'-GGCGTATGACCGAGCTGCTT-3' and a fluorescent quantitative PCR primer reverse primer 5'-TACCGGCTCAACGTTGTTGGA-3' are designed based on the ORF region of the FaERF13 gene, and an inner reference primer forward primer 5'-TTCACGAGACCACCTATAACTC-3' and an inner reference primer 5'-GCTCATCCTATCAGCGATT-3' are designed based on the action gene. The reaction system refers to a Novain ChamQ SYBR Qpcr Master Mix kit, and the PCR reaction procedure is as follows: 95 ℃ for 30sec;95℃10sec,60℃30sec,40 cycles. Three biological replicates were performed for each trial. Using a fluorescence quantitative PCR instrument CFX96TM (Bio-RAD company), referring to the PCR system and program described by the manufacturer, performing fluorescence quantitative PCR to obtain the cycle number reaching the fluorescence threshold, and calculating the relative expression amount of the FaERF13 gene at different flowering phases and different parts of the flower organ.
The method comprises the steps of selecting four stages of fruits of peach fumigation strawberry G (green fruit), GW (green transformed gingko), W (gingko), R (red fruit), selecting three corbels with consistent colors as biological repetition in each stage, removing RNA (ribonucleic acid) extracted from the lean fruits, and carrying out expression analysis, wherein as the fruits develop, the expression level of FaERF13 genes is continuously reduced, and the fact that FaERF13 expression and strawberry fruit maturity are in negative correlation is proved by figure 2.
Example 3FaERF13-VIGS vector construction and transient transformation of "peach-stunted" fruits
(1) Vector construction
Designing a small fragment primer FaERF13.1_F:5'CAGCGCTAAAACCCGAGGA-3'; faERF13.1_R:5 '-CAAGCCACACTCTCGCTCCA') the FaERF13.1 sequence was amplified using FaERF13cDNA as template and the PCR procedure was the same as in example 1 using the Nimble cloning kit to insert the FaERF13 expression cassette into the VIGS vector PNC-TRV2.
The system is as follows
PCR Product 50ng
pNC-Cam1304-35S 20-120ng
Nimble Mix 5μL
ddH 2 O was added to 10. Mu.L and incubated at 25℃for 10min.
(2) Preparation of the liquor
The vector plasmid was transformed into Agrobacterium GV3101 competent by liquid nitrogen freeze thawing: removing competent cells from-80deg.C, thawing at 4deg.C, and placing on ice; adding the plasmid to be transformed into 100 mu l of competent cells, stirring by hands to dial the bottom of the tube, standing for 10min on ice, liquid nitrogen for 5min, water-bath at 37 ℃ for 5min, and ice-bath for 5min; adding 900 mu l of liquid medium without antibiotics to obtain YEB, and culturing for 2-3 hours at 28 ℃ in a shaking way; the cells were collected by centrifugation at 6000rpm for one minute, and about 100. Mu.l of the supernatant was left to gently blow the resuspended pellet onto a YEB plate containing Kan (50. Mu.g/m) and Rif (50. Mu.g/m), and inverted in a 28℃incubator for 2-3 days.
PNC-TRV2-FaERF13 Agrobacterium positive strain was identified in the same manner as in example 1. Inoculating into YEB liquid medium containing Kan (50 μg/m) and Rif (50 μg/m), culturing at 28deg.C for 16 hr with shaking table, removing supernatant at 4000rpm for 10min to collect thallus, adding heavy suspension (10 mM MgCl) gradually 2 10mM MES, 200. Mu.M acetosyringone), OD600 was measured using a spectrophotometer, the final bacterial mass was resuspended to a final concentration of 2.0, the resuspension was allowed to stand at room temperature in the dark for 3h, PNC-TRV1 resuspension was compared with PNC-Equal volume mixing of TRV2-FaERF13.1 heavy suspension ready for injection
(3) Instant transformation of strawberry fruits
And selecting GW and W-period strawberry peach fumigation fruits, injecting agrobacterium resuspension from the receptacle part, and carrying out infection. PNC-TRV1 resuspension was mixed with PNC-TRV2 empty suspension in equal volumes as a negative control. Phenotypic changes were observed by daily photographing after infection. The fruits began to change color the next day in the VIGS group, and the fruits were fully ripe the fourth day, while the development of the MOCK group fruits did not change much, and remained in the gingko stage, with a degree of ripeness significantly lower than in the VIGS group (fig. 3).
(4) qRT-PCR detection and phenotypic observation of VIGS silenced fruits
In order to verify whether the premature change of the strawberry fruits in the VIGS treatment group is related to the transfer of the FaERF13 gene, the invention adopts a common fluorescent quantitative PCR method to detect the expression quantity of the FaERF13 genes in the MOCK group CK group and the VIGS treatment group, and the fluorescent quantitative PCR method is the same as that in example 2.
The expression pattern of FaERF13 in different treatment groups is found (figure 4) by taking reference gene FaActin1 as a control, the expression quantity of FaERF13 in the VIGS treatment group is about 50% lower than that in the CK group, the silencing of FaERF13 transcription factor is proved to be effective, and the silencing of FaERF13 transcription factor leads to the great advance of the ripening of 'peach-fumigated' strawberry fruits, and the negative regulation effect of FaERF13 transcription factor on the ripening of strawberry fruits is proved.
Example 4
Construction of pNC-Cam1304-35S over-expression vector and RNA interference vector of FaERF13 and application prospect thereof
(1) Construction of pNC-Cam1304-35S FaERF13 over-expression vector and application prospect thereof
FaERF13-pNC-Cam1304-35S primers 5'-AGTGGTCTCTGTCCAGTCCTCTATCTAATCTTTGCCTTGG-3' and 5'-GGTCTCAGCAGACCACAAGTATGGTTGTCTGTCCGTACAT-3' with NC cloning linker were designed, and the FaERF13 obtained in example 1 was used as template cDNA, amplified to obtain the insert of FaERF13, and PCR amplification and product recovery were performed in the same manner as in example 1.
The insert was cloned rapidly in one step using a Nimble Cloning kit (NC Biotech) with the pNC-Cam1304-35S vector, the system was the same as in example 3, the escherichia coli transformation and positive clone screening method was the same as in example 1, and the positive bacterial fluid was sequenced by the biology company (nanjing) to obtain the pNC-Cam1304-35s:faerf13 overexpression vector.
The FaERF13-Cam1304-35S over-expression vector converts strawberries through an agrobacterium-mediated method, a new variety of strawberries with fruits matured later is expected to be obtained, and the fruit producing period (commodity period is prolonged) of the strawberries can be prolonged by combining the early-maturing variety with the genetically improved variety.
(2) pNC-Cam1304-RNAi: faERF13 vector construction and application prospect thereof
The primer design with NC joint, faERF13 insert acquisition, NC cloning process and the construction process of the pNC-Cam1304-35S FaERF13 vector are the same, the pNC-Cam1304-RNAi vector has two NC cloning frames in front and back, and the FaERF13 insert is assembled in front and back directions simultaneously through one-step reaction, so that the pNC-Cam1304-RNAi is obtained: ERF13 interferes with the vector. After plant cells such as strawberries are transformed by the vector, the vector is transcribed efficiently and can spontaneously form an RNAi hairpin structure, and the FaERF13 gene is silenced rapidly and efficiently.
pNC-Cam1304-RNAi: the ERF13 interference vector can be used for transforming strawberry tissue culture seedlings by using an agrobacterium-mediated method or a gene gun method, is expected to obtain a new variety with early mature fruits, and has great potential for application in the aspects of early marketing time and commodity competitiveness enhancement of late maturing strawberry varieties.
Sequence listing
<110> university of Yangzhou
<120> strawberry ethylene response FaERF13 gene and application thereof in changing strawberry fruit maturity
<160> 12
<170> SIPOSequenceListing 1.0
<210> 1
<211> 639
<212> DNA
<213> strawberry (Fragaria x ananasa Duch.)
<400> 1
atgtacggac agacaaccat agactcagta gattccgact tggctcttct tgaatccatt 60
cgccaaaacc ttctaggtga tcatatggcg tctcaggccg ccctcttcgg cggcgacgtc 120
gacgacatca tccccttgtt tttccccagc tctgctactg cttcaacctg catgagtggg 180
ttttttacca cagagaactg gtgggaatca gcgctaaaac ccgaggactc tgaaaccgaa 240
gctgtaagtg ccccagggga cgacgtcgag gttaaactga gggagcaagc tcgtgttgcg 300
catgaggtgc acgcgccaag ggtagcgagg catttcaggg gagtgaggag gaggccatgg 360
gggaagtacg cggcggagat aagagatccg gcgaagaatg gagcgagagt gtggcttggg 420
acttacgaga ctcctgagga cgccgggctg gcgtatgacc gagctgcttt caagctgcgc 480
ggacgtaaag ctaagctcaa ctttcctcac ttgattggct ccaacaacgt tgagccggtt 540
agggttaccc cgaagcggcg gcgctcgccg gagcagtact ctacttcgac ctcgtcggca 600
tcggaggaca gcgaatctcc caaggcaaag attagatag 639
<210> 2
<211> 212
<212> PRT
<213> strawberry (Fragaria x ananasa Duch.)
<400> 2
Met Tyr Gly Gln Thr Thr Ile Asp Ser Val Asp Ser Asp Leu Ala Leu
1 5 10 15
Leu Glu Ser Ile Arg Gln Asn Leu Leu Gly Asp His Met Ala Ser Gln
20 25 30
Ala Ala Leu Phe Gly Gly Asp Val Asp Asp Ile Ile Pro Leu Phe Phe
35 40 45
Pro Ser Ser Ala Thr Ala Ser Thr Cys Met Ser Gly Phe Phe Thr Thr
50 55 60
Glu Asn Trp Trp Glu Ser Ala Leu Lys Pro Glu Asp Ser Glu Thr Glu
65 70 75 80
Ala Val Ser Ala Pro Gly Asp Asp Val Glu Val Lys Leu Arg Glu Gln
85 90 95
Ala Arg Val Ala His Glu Val His Ala Pro Arg Val Ala Arg His Phe
100 105 110
Arg Gly Val Arg Arg Arg Pro Trp Gly Lys Tyr Ala Ala Glu Ile Arg
115 120 125
Asp Pro Ala Lys Asn Gly Ala Arg Val Trp Leu Gly Thr Tyr Glu Thr
130 135 140
Pro Glu Asp Ala Gly Leu Ala Tyr Asp Arg Ala Ala Phe Lys Leu Arg
145 150 155 160
Gly Arg Lys Ala Lys Leu Asn Phe Pro His Leu Ile Gly Ser Asn Asn
165 170 175
Val Glu Pro Val Arg Val Thr Pro Lys Arg Arg Arg Ser Pro Glu Gln
180 185 190
Tyr Ser Thr Ser Thr Ser Ser Ala Ser Glu Asp Ser Glu Ser Pro Lys
195 200 205
Ala Lys Ile Arg
210
<210> 3
<211> 20
<212> DNA
<213> Artificial sequence (Artificial Sequence)
<400> 3
ctatctaatc tttgccttgg 20
<210> 4
<211> 20
<212> DNA
<213> Artificial sequence (Artificial Sequence)
<400> 4
atggttgtct gtccgtacat 20
<210> 5
<211> 20
<212> DNA
<213> Artificial sequence (Artificial Sequence)
<400> 5
ggcgtatgac cgagctgctt 20
<210> 6
<211> 21
<212> DNA
<213> Artificial sequence (Artificial Sequence)
<400> 6
taccggctca acgttgttgg a 21
<210> 7
<211> 22
<212> DNA
<213> Artificial sequence (Artificial Sequence)
<400> 7
ttcacgagac cacctataac tc 22
<210> 8
<211> 19
<212> DNA
<213> Artificial sequence (Artificial Sequence)
<400> 8
gctcatccta tcagcgatt 19
<210> 9
<211> 19
<212> DNA
<213> Artificial sequence (Artificial Sequence)
<400> 9
cagcgctaaa acccgagga 19
<210> 10
<211> 20
<212> DNA
<213> Artificial sequence (Artificial Sequence)
<400> 10
caagccacac tctcgctcca 20
<210> 11
<211> 40
<212> DNA
<213> Artificial sequence (Artificial Sequence)
<400> 11
agtggtctct gtccagtcct ctatctaatc tttgccttgg 40
<210> 12
<211> 40
<212> DNA
<213> Artificial sequence (Artificial Sequence)
<400> 12
ggtctcagca gaccacaagt atggttgtct gtccgtacat 40
Claims (3)
1. Strawberry ethylene response geneFaERF13Application of strawberry ethylene response gene in negative regulation of strawberry fruit ripeningFaERF13The nucleotide sequence is shown as SEQ ID NO. 1.
2. Comprises strawberry ethylene response geneFaERF13Is a plant interference vector of (a), the strawberry ethylene response geneFaERF13The nucleotide sequence is shown as SEQ ID NO. 1.
3. Use of the plant interference vector of claim 2 for promoting fruit ripening in strawberry.
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| CN110894221A (en) * | 2019-12-12 | 2020-03-20 | 中国科学院植物研究所 | Strawberry maturation associated transcription factor gene FaNAC2 and application thereof |
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| CN109576288A (en) * | 2018-12-05 | 2019-04-05 | 浙江五合生物科技有限公司 | A kind of strawberry ABA degradation pathway key enzyme FveCYP707A4a gene and its application |
| CN110894221A (en) * | 2019-12-12 | 2020-03-20 | 中国科学院植物研究所 | Strawberry maturation associated transcription factor gene FaNAC2 and application thereof |
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