CN111978388B

CN111978388B - Rice UV-B light receptor gene OsUVR8a and application thereof

Info

Publication number: CN111978388B
Application number: CN202010910399.2A
Authority: CN
Inventors: 黄烯; 任慧; 胡珊
Original assignee: Xiamen University
Current assignee: Xiamen University
Priority date: 2020-09-02
Filing date: 2020-09-02
Publication date: 2022-02-25
Anticipated expiration: 2040-09-02
Also published as: CN111978388A

Abstract

A rice UV-B light receptor gene OsUVR8a and application thereof relate to the field of biotechnology and genetic engineering. Provides a rice UV-B light receptor gene OsUVR8 a. Provides a protein OsUVR8a coded by the rice UV-B light receptor gene. Provides a recombinant vector containing the rice UV-B light receptor gene OsUVR8 a. Provides a transgenic plant CFP-OsUVR8 a/Nip. Provides the application of the rice UV-B light receptor gene OsUVR8a and the light receptor gene coding protein OsUVR8a in the research and cultivation of anti-radiation plants. According to the ultraviolet intensity of different regions, the OsUVR8a gene is selectively over-expressed, so that rice varieties which are more tolerant to ultraviolet light with different intensities can be cultivated, and the yield of rice can be improved by utilizing an ultraviolet light energy source.

Description

Rice UV-B light receptor gene OsUVR8a and application thereof

Technical Field

The invention relates to the field of biotechnology and genetic engineering, in particular to a rice UV-B light receptor gene OsUVR8a and application thereof.

Background

Light is one of the most important environmental factors influencing the life activities of animals and plants in nature, and researches show that the light plays a vital role in regulating and controlling the whole growth and development process of plants from seedlings to seedlings. The process of completing cell differentiation and tissue and organ formation after the germination of plant seeds is called photomorphogenesis, which is the first step of ensuring the plant to benefit light energy for growth and development and basically determines the quality of subsequent growth and propagation of plant. The UV-B light with the wavelength of 280-315 nm is an important component of sunlight, and in previous researches, the UV-B light with long wavelength and low energy can promote the photomorphogenesis of plants, help the plants to resist high-intensity UV-B radiation and is beneficial to the growth and development of the plants, so that the UV-B light is used as an important environmental factor to participate in the regulation and control of the morphogenesis process of plant seedlings (Jenkins, G.I. (2009).

With the increasing intensity of ultraviolet rays on the ground surface, researches on how plants, especially food crops related to human survival, can deal with the irradiation of ultraviolet rays and the influence of ultraviolet rays on the growth of the crops are urgent. Studies have shown that in Arabidopsis thaliana, the UV-B photoreceptor AtUVR8 can receive UV-B light signals, convert from dimers to monomers, initiate transduction of UV-B light signals, and achieve adaptation and tolerance of Arabidopsis thaliana to UV-B (Rizzini, l., Favory, j.j., Cloix, c., Faggionato, d., O' Hara, a., Kaiserli, e., Baumeister, r., Schafer, e., Nagy, f., Jenkins, g.i., et al (2011) Perception of UV-B by the Arabidopsis UVR8 protein science 332:103 + 106). In recent years, homologous proteins of AtUVR8, which have high homology with AtUVR8, were cloned and identified in plants such as Populus diversifolia, apple, Chlamydomonas reinhardtii, Chrysanthemum morifolium and Betula platyphylla sequentially by different subject groups. After being transferred back into the mutant of Arabidopsis uvr8, the homologous proteins can complement the phenotype of the uvr8 mutant, show UV-B light-induced hypocotyl shortening and can up-regulate important genes in a UV-B light signal path. The above results indicate that the function of the AtUVR8 homologous protein in these species is very conserved.

Researches show that the chlamydomonas reinhardtii is more tolerant to UV-B light and high-intensity white light after being subjected to low-dose UV-B light adaptation treatment, and the important regulation and control effect of a UV-B light receptor on plant surface-adapted UV-B light in the early stage of biological evolution is realized. The existence of UV-B light receptors in Hangzhou white chrysanthemum and white birch in northeast indicates that garden flowers and plants can be cultivated and vegetation environment can be improved by using UV-B light, and the UV-B light receptors of populus euphratica suggest that the tolerance to the UV-B light is a great factor for enabling the populus euphratica to thrive in desert and improve ecological environment. Research on UV-B light receptors in apples and grapes also provides a good theoretical basis for applying UV-B light energy to fruit tree cultivation, and adaptation mechanisms of rice, corns, sorghum and cotton to UV-B light show great application value of UV-B light in agricultural production. Therefore, the UV-B light and the UV-B light receptor have wide application prospects in the actual production of forest, economic crops and food crops.

The rice is native to China and India, and people in Yangtze river basin of China have been planting rice as early as seven thousand years ago. Rice is an important grain crop in China, and the yield of the rice is directly related to the grain safety in China. At present, the environmental problem of ultraviolet light pollution is more and more prominent, and the yield of rice as a highlight crop is inevitably influenced by the increase of the intensity of ultraviolet light, so that if the UV-B light receptor gene in the rice can be cloned and analyzed, the method has considerable practical application value for helping the rice adapt to the ultraviolet light, if the method can be applied to the current breeding strategy, the rice variety which is more tolerant to the ultraviolet light is expected to be cultivated, even the yield of the rice is expected to be improved by utilizing an ultraviolet light energy source, and the method has remarkable significance for guaranteeing the global grain safety.

Disclosure of Invention

The first purpose of the invention is to provide a rice UV-B light receptor gene OsUVR8 a.

The second purpose of the invention is to provide a protein OsUVR8a coded by the rice UV-B light receptor gene.

The third purpose of the invention is to provide a recombinant vector containing the rice UV-B light receptor gene OsUVR8 a.

The fourth purpose of the invention is to provide a transgenic plant CFP-OsUVR8a/Nip containing the rice UV-B light receptor gene OsUVR8 a.

The fifth purpose of the invention is to provide application of the rice UV-B light receptor gene OsUVR8 a.

The sixth purpose of the invention is to provide the application of the rice UV-B light receptor gene coding protein OsUVR8 a.

The rice UV-B light receptor gene OsUVR8a has a cDNA sequence shown in SEQ ID No. 1.

SEQ ID NO.1：

The protein OsUVR8a coded by the rice UV-B light receptor gene is derived from Oryza sativa Japonica Group, and the amino acid sequence of the protein OsUVR8a is shown as SEQ ID NO. 2.

SEQ ID NO.2

The protein OsUVR8a coded by the rice UV-B light receptor gene consists of 453 amino acid residues, and is a homologous protein of Arabidopsis UV-B light receptor protein AtUVR 8.

On the basis of the scheme, the invention also provides an expression cassette, a recombinant expression vector, a transgenic plant and a recombinant strain containing the rice UV-B light receptor gene OsUVR8 a.

A plant expression vector can be used for constructing a recombinant expression vector containing the OsUVR8a gene. When the recombinant expression vector is constructed by using OsUVR8a, any constitutive promoter or enhanced promoter or other plant gene promoters can be added in front of the transcription initiation nucleotide.

The plant expression vector comprises a binary agrobacterium vector and the like.

The plant expression vector may also contain a 3' untranslated region of a foreign gene, i.e., a polyadenylation signal and any other DNA segment involved in mRNA processing or gene expression.

In order to facilitate the identification and selection of transgenic plant cells or plants, plant expression vectors therefor may be processed, for example, by adding genes encoding enzymes or luminescent compounds which produce a color change (GUS gene, luciferase gene, fluorescent protein tag, etc.) and antibiotic markers having resistance (ampicillin marker, kanamycin marker, etc.) which can be expressed in plants.

The transgenic plant of the rice UV-B light receptor gene OsUVR8a is CFP-OsUVR8 a/Nip.

The rice UV-B light receptor gene OsUVR8a can be applied to research and cultivation of anti-radiation plants and can induce photomorphogenesis of plants under UV-B light.

The rice UV-B light receptor gene coding protein OsUVR8a can be applied to research and cultivation of anti-radiation plants and can be applied to UV-B light-induced photomorphogenesis.

The invention also obtains double mutant plant OsUVR8 of OsUVR8a and homologous gene OsUVR8b thereof by using CRISPR-Cas9 technology. The plant does not have the normal function a of OsUVR8, the height of a seedling is increased under UV-B light, the length between a first section and a second section is increased, the tolerance capability to the UV-B light is weakened, and the photo-configuration of the rice induced by the UV-B light is not facilitated.

The OsUVR8a is used as a UV-B light receptor gene of important grain crop rice, has considerable practical application value for stronger and stronger ultraviolet light in the environment adapted to the rice, can be applied to the current breeding strategy, selectively over-expresses the OsUVR8a gene according to the ultraviolet intensity of different areas in China, can cultivate rice varieties more tolerant to the ultraviolet light with different intensities, and can improve the yield of the rice by utilizing ultraviolet light energy; has very important significance for cultivating the rice variety which is more tolerant to ultraviolet light and can utilize the ultraviolet light energy source to realize higher yield, thereby further improving the rice yield.

Drawings

FIG. 1 shows the results of homology comparison of amino acid sequences of rice UV-B light receptor protein OsUVR8a and Arabidopsis UV-B light receptor protein AtUVR 8.

FIG. 2 is a table chart and statistical results of double mutant plant OsUVR8 of wild type rice and rice UV-B light receptor genes OsUVR8a and OsUVR8B under the illumination conditions of white light and white light plus UV-B light.

FIG. 3 shows the results of UV-B photodamage experiments on wild type rice and double mutant plants OsUVR8a and OsUVR8B of the UV-B photoreceptor genes OsUVR8 of rice.

FIG. 4 is a table chart and statistical results of wild type rice and rice UV-B photoreceptor gene OsUVR8a overexpression plant CFP-OsUVR8a OE under white light and white light plus UV-B illumination conditions.

FIG. 5 shows the results of UV-B photo-damage experiments on wild type rice and rice UV-B photo-receptor gene OsUVR8a overexpression plant CFP-OsUVR8a OE.

Detailed Description

The following examples will further illustrate the present invention with reference to the accompanying drawings.

As shown in FIG. 1, the present inventors found that a gene OsUVR8a homologous to the UV-B light receptor gene AtUVR8 in Arabidopsis thaliana is present in rice through sequence homology alignment, and subsequently cloned this gene from rice, confirming that the encoded protein can also respond to UV-B light signals and help plants to resist UV-B light damage. Meanwhile, rice plants with double mutations of OsUVR8a and homologous genes OsUVR8B thereof are found to be incapable of responding to UV-B light to promote the growth and development of rice, the tolerance capability of the rice plants to the UV-B light is obviously weakened, rice materials over-expressing OsUVR8a are compact ideal plant types under white light and UV-B light, the rice materials have strong tolerance capability to the UV-B light, and a good foundation is provided for the growth and development and fructification of the later stage of the rice materials.

Example 1: the cDNA clone of the rice UV-B light receptor gene OsUVR8 a.

1) Extraction of total RNA of rice:

wild type (Nipponbare) leaves of rice grown for 2 weeks under white light were taken, put into 1.5mL of an EP tube without RNase, snap frozen with liquid nitrogen, and either ground immediately or stored at-80 ℃.

First, the rice leaves were ground after drying in a mortar, pestle, spatula or the like at 180 ℃ for 4 hours and then cooling to room temperature, and then precooling the above-mentioned apparatus with liquid nitrogen. 100 μ L of the powder was dispensed, 500uL of RNA lysate was added and dissolved on ice until no flocculent precipitate was formed. Subsequently, 500. mu.L of RNA diluent was added and well mixed by aspiration, and after standing at room temperature for 5min, centrifugation was carried out at 12000rpm for 10min at room temperature, 800. mu.L of supernatant was taken out and put into a new RNase-free EP tube, and 400. mu.L of absolute ethanol was added thereto, and the mixture was inverted and well mixed. Then, the mixture was transferred to a 2mL spin column, centrifuged at 12000rpm for 1min, after discarding the filtrate, 600. mu.L of RNA wash was added, centrifuged at 12000rpm for 1min, and after discarding the filtrate, the mixture was idled at 12000rpm for 2 min. 50uL of DNase I was added to the center of the membrane and incubated for 30 min. After incubation, 600. mu.L of RNA wash was added, centrifuged at 12000rpm for 1min, the filtrate was discarded, the operation was repeated once more, and the solution was left to idle at 12000rpm for 2min after discarding the filtrate. Adding 50 μ L of nuclease-free water, centrifuging at 12000rpm for 2min, and repeating centrifugation to improve RNA yield.

2) Reverse transcription of RNA into cDNA

Mu.g of RNA and 0.5. mu.g of oligo (dT) were added, annealed at 70 ℃ for 5min, and then reverse transcribed using the following system:

the reverse transcription mixing system is shown in table 1:

TABLE 1

Reagent	Volume (μ L)
		Nuclease-free Water	6.9
Go Script^TM 5×Raection Buffer	4
		PCR Nuclease Mix	1
MgCl₂	1.6
		Go Script Reverase Transcriptase	1
Inhibitor	0.5

3) Primers P1 and P2 are designed according to the coding region sequence of the rice UV-B light receptor gene OsUVR8a, and the sequences of the primers are as follows:

P1:5’-ACGCGTCGACATGCATTCGACCGGCATGGAGATGG-3’

P2:5’-CGGGATCCTCAGACGCGCATCCTCTTCACATCT-3’

the PCR product is detected in 1% agarose gel by electrophoresis to obtain a band with the expected size, and then the band is cut and recovered. Connecting the target fragment with pCAMBIA2300-35S-CFP vector by T4 enzyme connection method, transferring the connection product into Escherichia coli DH5 alpha competent cell, screening positive clone according to Carna resistance marker on pCAMBIA2300-35S-CFP vector, and then confirming whether the recombinant plasmid contains complete and correct target gene nucleotide sequence by sequencing. Thus, a recombinant vector of pCAMBIA2300-35S-CFP-OsUVR8a was obtained.

Example 2: rice UV-B photoreceptor genes OsUVR8a and OsUVR8B are knocked out by using a CRISPR-Cas9 technology, and a double-mutant plant OsUVR8 is constructed.

The CRISPR knock-out of rice was carried out according to pYIISPR/Cas 9-mono vector System of monocotyledon plants proposed by Liu flare light teacher's laboratory of southern agriculture university (Ma, X., Zhang, Q., Zhu, Q., Liu, W., Chen, Y., Qiu, R., Wang, B., Yang, Z., Li, H., Lin, Y., et al. (2015). A Robust CRISPR/Cas9System for Convenient, High-Efficiency Multiplex Genome edition in monochip and dictionary plants.molecular Plant 1274-. The OsUVR8a and OsUVR8b genes respectively select Site1 and Site2 as targets, and the sequences are as follows:

Site1：5’-AGGAGCTGCAGCTCTACAG-3’

Site2：5’-GAGGACCGGCTGGTGCCGA-3’

vectors for knocking out the rice OsUVR8a and OsUVR8b genes were constructed according to the above-mentioned method. The constructed vector is transferred into agrobacterium, infection transformation is carried out on the callus of wild rice, transgenic crops are constructed (the transgenic crops can be constructed by a conventional method or are entrusted to a professional company for cultivation, such as a laboratory (Beijing) limited company at the front of the brand-name Xingwang systematic crop design), T0 generation plants are obtained through induced differentiation, and then the T0 generation plants are screened.

Example 3: screening of double mutant OsUVR8 of rice UV-B light receptor gene OsUVR8a and OsUVR 8B.

1) Rice DNA is extracted by a CTAB method.

(1) Taking 2cm long rice young leaves, and grinding with liquid nitrogen;

(2) adding 550 μ L CTAB, incubating at 65 deg.C for 30min, and shaking for 3 times;

(3) adding 550 μ L of 24: 1 chloroform/isoamyl alcohol after incubation, mixing, standing at room temperature for 10min (the operation has peculiar smell, and is carried out in a fume hood);

(4) centrifuging at 12000rpm for 5min at room temperature;

(5) centrifuging, adding 400 μ L supernatant into an EP tube precooled at-20 deg.C and filled with 500 μ L anhydrous ethanol, standing at-20 deg.C for 15 min;

(6) centrifuging at 12000rpm for 5min at room temperature, and removing the supernatant;

(7) adding 300 μ L75% ethanol, mixing, centrifuging at 12000rpm for 5min at room temperature, and removing supernatant;

(8) air-drying at room temperature until no ethanol exists, and adding 100ul ddH₂And dissolving the O.

2) Designing PCR amplification primers P3 and P4 according to the target point position of OsUVR8a, designing PCR amplification primers P5 and P6 according to the target point position of OsUVR8b, and carrying out PCR amplification by using DNA of a plant to be screened as a template. The sequences of these primers are as follows:

P3:5’-CCTAGGCGGTGGTTAATTATCTG-3’

P4:5’-GGAGGGTGTCGTAATACATGAAA-3’

P5:5’-TTGCCGGCGCTTCTATC-3’

P6:5’-ATGGGCAGATGGCTGTTT-3’

sequencing the PCR product, comparing the obtained sequencing result with the original sequence, and confirming the mutation type.

Example 4: construction of rice UV-B light receptor gene OsUVR8a overexpression plant CFP-OsUVR8 a.

The CFP-OsUVR8a with constitutive expression is transferred into a wild rice variety by utilizing a recombinant vector pCAMBIA2300-35S-CFP-OsUVR8a to obtain a T0 generation plant and screen the T0 generation plant. Transplanting T0 generation plants into field for cultivation, and collecting T1 generation seeds. Screening T1 generation seeds with G418 resistant culture medium, transplanting the line meeting the separation ratio of 3: 1 into the field for cultivation, and collecting T2 generation seeds. And screening seeds of the T2 generation by using a G418 resistant culture medium, transplanting a fully alive line into a field for cultivating and breeding, wherein the line can be used for carrying out an over-expression plant of the UV-B light receptor gene OsUVR8a for experiment.

Example 5: and detecting the influence of the rice UV-B light receptor gene OsUVR8a on the growth and development of the rice under the UV-B light.

In order to detect the influence of the gene OsUVR8a of the rice UV-B light receptor gene on the growth and development of rice under UV-B light by using the transgenic materials (OsUVR8 and CFP-OsUVR8a) constructed above, the inventor lays rice seeds on a culture dish, carries out dark treatment at 37 ℃ for 3 days, then respectively carries out the dark treatment for 7 days under the illumination conditions of white light and white light plus UV-B light, and then photographs are taken to preserve the growth condition of each plant. The total length of the entire overground part of rice, the length of the first internode and the length of the second internode were measured by Image J software, and the measurement results are shown in fig. 2 and 4.

Example 6: experimental method of UV-B photodamage experiment.

The inventor spreads the rice on a culture dish, treats the rice in the dark at 37 ℃ for 3 days, respectively treats the rice for 7 days under the illumination conditions of white light and white light plus UV-B light, then transfers the plant into a culture solution for continuous culture, treats the rice seedlings growing under the two light conditions with the white light plus UV-B light for 2 days, and then recovers the rice seedlings for 1 day under the white light. Then, the tips of the two inverted blades are taken out for scanning, and clear pictures are obtained for observing the damage degree of the blades, as shown in fig. 3 and 5.

Sequence listing

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Glu Ala Val Pro Glu Ala Pro Glu Arg Ser Val Val Leu Ile Ser Ala

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Gly Ala Ser His Ser Val Ala Leu Leu Ser Gly Gly Val Val Cys Ser

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Trp Gly Arg Gly Glu Asp Gly Gln Leu Gly His Gly Asp Ala Glu Asp

50 55 60

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

65 70 75 80

Gly Val Ala Ser Val Val Ile Cys Gly Ala Asp His Thr Thr Ala Tyr

85 90 95

Ser Asp Glu Glu Leu Gln Leu Tyr Ser Trp Gly Trp Gly Asp Phe Gly

100 105 110

Arg Leu Gly His Gly Asn Ser Ser Asp Val Phe Asn Pro Gln Pro Ile

115 120 125

Gln Ala Leu Gln Gly Val Arg Ile Thr Gln Ile Ala Cys Gly Asp Ser

130 135 140

His Cys Leu Ala Val Thr Val Ala Gly His Val His Ser Trp Gly Arg

145 150 155 160

Asn Gln Asn Gly Gln Leu Gly Leu Gly Asn Thr Glu Asp Ser Leu Leu

165 170 175

Pro Gln Lys Ile Gln Ala Phe Glu Gly Val Arg Val Lys Met Ile Ala

180 185 190

Ala Gly Ala Glu His Thr Ala Ala Val Thr Glu Asp Gly Asp Leu Tyr

195 200 205

Gly Trp Gly Trp Gly Arg Tyr Gly Asn Leu Gly Leu Gly Asp Arg Asp

210 215 220

Asp Arg Leu Ile Pro Glu Lys Val Ser Ser Val Asn Gly Gln Lys Met

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Val Leu Val Ala Cys Gly Trp Arg His Thr Ile Thr Val Ser Ser Ser

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Gly Ser Ile Tyr Thr Tyr Gly Trp Ser Lys Tyr Gly Gln Leu Gly His

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Gly Asp Phe Glu Asp His Leu Val Pro His Lys Leu Glu Ala Leu Lys

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Asp Thr Thr Ile Ser Gln Ile Ser Gly Gly Trp Arg His Thr Met Ala

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Leu Ala Ala Asp Gly Lys Leu Tyr Gly Trp Gly Trp Asn Lys Phe Gly

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Gln Val Gly Val Gly Asp Asn Glu Asp His Cys Ser Pro Val Gln Val

325 330 335

Asn Phe Pro Asn Glu Gln Lys Val Val Gln Val Ala Cys Gly Trp Arg

340 345 350

His Thr Leu Ala Leu Thr Glu Ala Lys Asn Val Phe Ser Trp Gly Arg

355 360 365

Gly Thr Ser Gly Gln Leu Gly His Gly Glu Ile Val Asp Arg Asn Ile

370 375 380

Pro Lys Met Ile Asp Ala Leu Ser Ser Asp Gly Ser Ala Cys Lys Gln

385 390 395 400

Leu Glu Ser Ser Lys Ala Val Pro Met Ser Ala Lys Val Trp Val Ser

405 410 415

Pro Ser Glu Arg Tyr Ala Ile Val Pro Asp Glu Lys Ala Gly Lys Gly

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Ile Pro Ala Gly Asn Gly Thr Glu Thr His Val Pro Gln Gly Asp Val

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Lys Arg Met Arg Val

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Claims

1. Rice UV-B light receptor geneOsUVR8aThe application in cultivating rice with better tolerance to ultraviolet light is characterized in that the rice UV-B light receptor geneOsUVR8The cDNA sequence of (A) is shown in SEQ ID NO. 1.

2. The application of the protein OsUVR8a coded by the rice UV-B light receptor gene in cultivating rice more tolerant to ultraviolet light is characterized in that the amino acid sequence of the protein OsUVR8a coded by the rice UV-B light receptor gene is shown in SEQ ID NO.2, consists of 453 amino acid residues and is a homologous protein of the Arabidopsis UV-B light receptor protein AtUVR 8.