CN102174486A - RDR701 protein and encoding gene as well as application thereof - Google Patents
RDR701 protein and encoding gene as well as application thereof Download PDFInfo
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- CN102174486A CN102174486A CN 201110030731 CN201110030731A CN102174486A CN 102174486 A CN102174486 A CN 102174486A CN 201110030731 CN201110030731 CN 201110030731 CN 201110030731 A CN201110030731 A CN 201110030731A CN 102174486 A CN102174486 A CN 102174486A
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Abstract
The invention relates to an RDR701 protein and an encoding gene as well as application thereof, in particular to a key functional site of an RNA (Ribonucleic Acid) dependent RNA polymerase RDR701 protein. A nucleotide sequence, corresponding to the site, for specifically expressing the RDR701 protein is SEQ ID NO:1; an amino acid sequence of the RDR701 protein is SEQ ID NO:2; rice of which the functional site of the RDR701 protein is subjected to missense mutation shows temperature-sensitive sterility; and by adopting the technology, a temperature-sensitive sterile rice mutant can be bred; and the protein is applied to rice cross breeding and has a broad prospect.
Description
Technical field
The present invention relates to RNA polymerase RDR701 albumen that a kind of RNA relies on and cultivating the application of paddy rice responsive to temperature type in sterile.
Background technology
RDR is the RNA polymerase that RNA relies on, and finds in virus that the earliest it comes the genome (Wang and Metzlaff, 2005) of replication-competent virus as the rna replicon enzyme of virus.In the eukaryote, the RDR that is proved to be the RNA composite reactive with RNA dependence first is the RDR1 in the tomato, and the QDE-1 albumen of following coarse arteries and veins born of the same parents bacterium (Neurospora crassa) also is found the activity of the RNA polymerase with RNA dependence.Nematode, find the RNA polymerase that exists RNA to rely in Arabidopis thaliana and the yeast immediately.Adopt the tomato RDR1 of biochemical means purifying and the RDR6 of Arabidopis thaliana can utilize long single stranded RNA to be synthetic its sequence complementary strand of masterplate, and this activity has disclosed their biochemical function having or not under the condition of primer and can both carrying out external.In vivo, the RNA of not all strand can be as the RDR masterplate, and " aberrant RNA " hypothesis has more reasonably been annotated the feature of Eukaryotic RDR target site at present.This hypothesis is thought in the most eukaryotes or external some unusual RNA (aberrant RNA), for example 3 ' hold the tail (polA-) or the 5 ' end that lack polyadenylic acid to lack cap (uncapped) structure, can be used as the target site of RDR, thereby be transformed into two strands.
In eukaryote, RDR synthetic double-stranded RNA is processed into all lengths by Dicer, and various types of siRNA have mediated various silencing process.Usually the effect mediation that depends on RDR produces secondary or multistage microRNA makes that the signal of RNA silence is increased, and is extensive adopted mode.Like this, even there is the little RNA of very a spot of elementary silence, also can mediate and keep lasting silence.This phenomenon is very general in plant and nematode.External source or endogenous dsRNA precursor form primary little RNA under the processing of Dicer, be assembled in the corresponding AGO albumen, after the transcript of target gene is cut, form secondary microRNA under the effect of RDR.In nematode, primary microRNA at first with after the special AGO-RDE-1 combination, mediates the cutting of target gene transcript, thereby RDR is recruited cleavage site.RDR is a masterplate with the transcript that cuts off, directly synthetic secondary microRNA.And how to determine each unitary length when directly synthetic for RDR, not clear at present.Because this microRNA be directly synthetic, is not the product of Dicer, so the secondary microRNA in the nematode is 5 ' end of pair phosphorus or three phosphorus, and this secondary microRNA all originates from the antisense strand of primary little RNA said target mrna.Different with nematode is that the secondary microRNA of plant is the product of DCL, so their 5 ' end is single phosphorus.The RDR of plant can utilize the cut product of the said target mrna of primary microRNA to be masterplate, synthetic long dsRNA, thus be processed to form secondary microRNA by DCL.
The RDR albumen of plant can be divided into four families, i.e. RDR1, RDR2, RDR3 and RDR6.Present discovering in Arabidopis thaliana, they separately or participated in the route of synthesis of various microRNAs in the mode of partial function redundancy.Wherein RDR1 and RDR6 fellowship the microRNA in exogenous virus or the transgenosis process synthetic, thereby playing a role aspect the intrusion of opposing virus; RDR2 is responsible for the generation of some siRNA-hcsiRNAs relevant with the heterochromatinization of tumor-necrosis factor glycoproteins and transposon, thereby mediates their silence; Thereby RDR6 has also participated in the generation approach regulation and control growth and development of plant process of many endogenous siRNA, for example tasiRNA and natsiRNA in addition.
At present, mainly also rest on theoretical research stage, do not know the functional site that it is crucial, also be difficult to study the functional deficiency that this site mutation causes about the functional study of RDR protein family.Owing to lack clear and definite indication, therefore the research about RDR protein function critical sites needs a large amount of work.The inventor has found the critical sites of the homologous protein RDR701 of RDR6 in the paddy rice unexpectedly by long-term and arduous research, has obtained the sterile rice mutant of temperature sensitive by sudden change RDR701 critical sites, has broad application prospects.
Summary of the invention
An object of the present invention is to provide the proteic critical sites of RDR701, the disappearance of this critical sites or the alternative change that will cause the RDR701 protein function, the proteic sequence of RDR701 is the aminoacid sequence shown in the SEQ ID NO:2 in the sequence table, and its critical sites is the 172nd a tryptophane.RDR701 albumen is by the nucleotide coding shown in the SEQ ID NO:1 in the sequence table.
Another object of the present invention provides a kind of method of cultivating the temperature sensitive male sterile rice, specifically proteic the 172nd tryptophane of the RDR701 shown in the SEQ ID NO:2 in the sequence table in the rice cell is carried out missense mutation, the tryptophane that is preferably the 172nd is mutated into halfcystine, and more preferably the Nucleotide with the 516th of SEQ ID NO:1 in the sequence table becomes " T " by " G ".
" missense mutation " of the present invention is meant that certain amino acid whose codon of coding after base is replaced, becomes the another kind of amino acid whose codon of coding, thereby the amino acid kind and the sequence of polypeptide chain are changed.
Particularly, the invention provides the albumen of a kind of RDR701, this proteic aminoacid sequence is SEQ ID NO:2.
More specifically, the invention provides above-mentioned proteic missense mutation albumen, this Argine Monohydrochloride sequence is the 172nd the tryptophane generation missense mutation of SEQ ID NO:2.
Particularly, the 172nd tryptophane generation missense mutation is a halfcystine for the tryptophane missense mutation among the SEQ ID NO:2.
The present invention also provides a kind of coding above-mentioned proteic gene, and the nucleotides sequence of this gene is classified SEQ ID NO:1 as.
More specifically, the invention provides the base mutation gene of said gene, it is characterized in that the 516th " G " in the nucleotide sequence of SEQ ID NO:1 sports " T ".
The present invention also provides a kind of method of cultivating the temperature sensitive male sterile rice, it is characterized in that above-mentioned proteic the 172nd tryptophane in the rice cell is carried out missense mutation.Preferably, be halfcystine with the tryptophane missense mutation.
Particularly, it is to insert by tissue culture, chemical reagent mutagenesis or fixed point T-DNA that above-mentioned proteic 172 tryptophane in the rice cell is carried out missense mutation, is that proteic the 172nd tryptophane of SEQ ID NO:1 carries out missense mutation with sequence.Be that proteic the 172nd tryptophane of SEQ ID NO:1 carries out missense mutation with sequence preferably by tissue culture.
The present invention also provides proteic the 516th tryptophane of a kind of RDR701 of comprising to be mutated into the rice cell of non-tryptophane.
In case being provided, the above-mentioned method that provides cultivates into the temperature sensitive male sterile rice; those skilled in the art utilizes hybridization at an easy rate, backcross etc., and the means proterties that the paddy rice temperature sensitive is sterile is transferred in other rice varieties or the wild-rice, and rice varieties that the temperature sensitive that obtains thus is sterile or wild-rice are also included protection scope of the present invention in.
The method for expressing that generally acknowledged in the field under the method for expressing of albumen used herein and amino acid, gene and Nucleotide was, amino acid as shown in table 1 or amino-acid residue symbol, those skilled in the art can instead release corresponding trinucleotide codon according to it.In the specific embodiment of the present invention, the proteic gene order of employed coding shown in SEQ ID NO:2 is shown in SEQ ID NO:1.
Table 1 amino acid abbreviations table
For the ease of understanding, below will the present invention be described in detail by concrete drawings and Examples.It needs to be noted that specific examples and accompanying drawing only are in order to illustrate, not constitute limitation of the scope of the invention.Obviously those of ordinary skill in the art can illustrate according to this paper, within the scope of the invention the present invention is made various corrections and change, and these corrections and change are also included in the scope of the present invention.In addition, the present invention has quoted open source literature, and these documents also are in order more clearly to describe the present invention, and their full text content is all included the present invention in and carried out reference, just look like they full text in specification sheets of the present invention repeated description the same excessively.
Description of drawings
Conservative site generation nonsense mutation on the RDR701 gene among Fig. 1 mutant rl: (A) RL gene synoptic diagram.The black square is represented gene extron; White square is represented gene 5 ' and 3 ' UTR; Black line is represented intron.Mutational site and near sequence in wild-type and the rl mutant have been listed in the rectangular frame in left side, below.Corresponding amino acid sequence is listed in the right frame.Wherein the site sequence that suddenlys change in the mutant marks with redness.(B) in several japonica rice and the rice variety, the comparison of the Nucleotide of RL gene." * " mark mutant mutating alkali yl position." In " represents rice variety; " Ja " represents japonica rice variety.The specifying information of each kind sees Table 2-1.(C) comparison of the RDR701 family member aminoacid sequence of several plant." * " mark mutant mutating acid position.Used protein sequence is numbered: RL (Q8LHH9), At RDR701 (NP_190519), Nt RDR701 (AAU21242), Hv RDR701 (AAR91036), Pp RDR701 (ARF82437).
Fig. 2 RDR701 gene complementation the phenotype of rl mutant: (A) function carrier of RDR701 gene makes up synoptic diagram.Initial and the termination site of restriction enzyme site and genes encoding marks in the top; Corresponding position information marks in the below.The coding initiation site is defined as 1.(B) phenotype of wild-type and transgenic progeny under the high temperature.WT: spend 11 in the wild-type; The transgenic progeny of Control:rl transfer empty carrier; PRL:RL is the have complementary functions transgenic progeny of carrier of rl transfer.
The phenotype analytical of Fig. 3 rl mutant Xiao Hua: A, B are respectively and spend 11 Xiao Hua outside and external morphology in the wild-type.C-K is the Xiao Hua of mutant rl.Le: coetonium; Pa: inner glume; St: stamen; Pi: gynoecium."＜" mark is coetonium.
The phenotype analytical of Fig. 4 rl mutant seed: A: spend 11 seed in the wild-type; B-J: the seed of mutant rl.
The form of wild-type and rl mutant Xiao Hua under Fig. 5 treatment of different temperature:
(A) circadian temperature curve.The different different Temperature Treatment of curve color representative.DAT represents mean daily temperature, in order to weigh treatment of different temperature.
(B) mean daily temperature is 28 ℃, the Xiao Hua of wild-type and mutant.W represents wild-type, and M represents mutant.Arrow shows coetonium position.
(C-E) being respectively mean daily temperature is 30 ℃, when 32 ℃ and 34 ℃, and the Xiao Hua of wild-type and mutant.The per-cent of below is in the tassel, the ratio that different phenotype Xiao Hua are shared.
Embodiment
Method therefor is molecular biology commonly used, tissue culture technique and agronomy hand if no special instructions among the following embodiment
The method that volume is put down in writing.
The acquisition of embodiment 1.RDR701 albumen critical sites
We screen from tissue culture and obtain a mutant with glume development defective under the high temperature, called after rl.The phenotype of the F2 offspring plant that 136 strain kinds mutant rl and wild-type under hot conditions are backcrossed, we find that wherein 29 strains show as the unusual phenotype of glume development.Roughly meet normal plant and unusual 3: 1 segregation ratio of plant, illustrate that the phenotype of rl mutant is controlled by a recessive single-gene.The 368 strain F2 colonies that methods analyst rl by map based cloning and long-grained nonglutinous rice 93-11 hybridization are created, the Fine Mapping of Primary Location and STS marker mutational site the most at last is locked in the scope of No. 1 karyomit(e) 46kb between markers RH1-4 and RH1-8.RNA polymerase-RL (Os01g34350) gene that the analysis gene discovery that this zone comprised exists a RNA to rely on therebetween in common data base TIGR5.0.The biosynthetic process that studies show that all kinds of siRNA of this proteinoid wide participation in Arabidopis thaliana.Co-exist in six RDR in the Arabidopis thaliana, wherein three members' function is found, and RDR1 participates in the generation of siRNA in the virus infection process; Thereby RDR2 mainly is responsible for the generation of the 24nt siRNA relevant with heterochromatin to be kept genomic stable; RDR701 also is responsible for the generation that a class participates in the tasiRNA that regulation and control blade polarity grows in participating in the virus defense process siRNA synthetic.Carry out the homology search with the conservative RdRp structural domain of RDR albumen and find to have 5 RDR members in the paddy rice, be respectively RL-RDR705.Utilize the proteic full length sequence constructing system of RDR evolutionary tree in Arabidopis thaliana and the paddy rice, the RDR701 height homology of RL and Arabidopis thaliana as can be seen on the structure of evolutionary tree, thereby hint out that they may have similar function, promptly involved in plant lateral organ polar is set up.Therefore we determine that RL is the candidate gene of sudden change.
The design sequencing primer carries out full gene sequencing to RL, and the result shows that a nonsense mutation has taken place to hold 516 in first exon distance 5 ', and Nucleotide has become " T " by " G "; Corresponding amino acid has become halfcystine (C) (accompanying drawing 1-A) by tryptophane (W).In order to determine whether that this mutational site is essential for proteic function, we check order to this site in several typical japonica rice of paddy rice and rice variety, and the result shows that the nucleotides sequence in this mutational site is listed in (the accompanying drawing 1-B) for guarding between each kind.Utilize the protein sequence of RL in NCBI, to carry out the BLASTp search, the RDR protein sequence of several species that the comparison homology is higher, the amino acid that found that this mutational site also is (the accompanying drawing 1-C) that guards in several species.These presentation of results, guard on evolving in this mutational site of rl mutant, and this site is essential for proteic function.
Embodiment 2:RDR701 function complementation experiment
Cut BAC by enzyme, we have made up and have comprised RDR701 gene 5 ' end 2093bp, the carrier that has complementary functions of whole gene regions 4890bp district and 3 ' end 528bp.Employed BAC (b005E01) is from country of Chinese Academy of Sciences cara gene.Adopt Sal I and Xba I enzyme to cut BAC, reclaim the band that length is 7499bp.To reclaim product is masterplate, utilizes primer CX3107 (SEQ ID NO:3:5 '-ACCGAATCCGATTGATCTTC-3 ') and the purpose band of CX3109 (SEQ ID NO:4:5 '-CCAATGCAGGACGTCTTCTT-3 ') amplification to determine that the recovery band is wanted for us on the RL gene.This fragment has comprised 5 ' 2086bp of RDR701 gene, gene regions 4890bp and 3 ' end 523bp.Simultaneously, expression vector pCAMBIA2300-Actin I-ocs being carried out identical enzyme cuts.To be connected on the expression vector pCAMBIA2300 from the purpose fragment that BAC downcuts.PCR and enzyme are cut the correct cloned plasmids of checking and are transformed Agrobacterium Agl I, be used to transform the callus of rl mutant, under hot conditions, commentaries on classics has the transfer-gen plant of this carrier all to recover the phenotype of clever shell anormogenesis, and it then still is the phenotype (accompanying drawing 2) of coetonium aristiformization that commentaries on classics has the plant of empty carrier.This proves absolutely that it is this proteic key function site that this point sudden change on the RDR701 gene has caused the phenotype of glume development defective under the high temperature, the tryptophane that RDR701 albumen is the 172nd.
The phenotype of embodiment 3:rl mutant has temperature dependency
The coetonium developmental defect (accompanying drawing 3) that shows in various degree of Xiao Hua on the same fringe of the mutant of rl, corresponding fertility also have nothing in common with each other (accompanying drawing 4).The coetonium part aristiform that presents top or sidepiece of some Xiao Hua is interior coetonium separately not closed.This part Xiao Hua can be solid, but seed is not by clever shell bag quilt; The coetonium needle-like that becomes fully of most of Xiao Hua, anther development is not full, and their major parts are acarpous; Coetonium or the interior coetonium completely dissolve of indivedual Xiao Hua, floral organ is exposed, and stamen presents shrinkage, the phenomenon that minority has the floral organ number to increase.This part Xiao Hua is fully sterile.The phenotype of rl is subjected to such environmental effects very big, the dysplastic phenotype of performance under some condition, but under some condition, mutant is grown again normally.In the plantation of the region of different light time, for example the summer in Hangzhou (long day) and Hainan winter (short day), the phenotype of rl presents similar variation, we have got rid of the influence of photoperiod for mutant.Rice tillering regenerated characteristic has been given us very big enlightenment, sows in Hangzhou when the May, and the August, rl presented typical coetonium dysplastic phenotype when blooming, and most of Xiao Hua is barren.But its existing tassel sliced off allow its long again tillering during regeneration, along with the commentaries on classics of Hangzhou nine, October temperature is low, the Xiao Hua on new long tillering grows normal and solid again.This points out us may temperature for the phenotype of rl very big influence to be arranged.
In order further to determine the relation between temperature and the rl phenotype, we have carried out the Temperature Treatment experiment in the growth cabinet of accurate controlled temperature.14.5 hours same photoperiods, and under the condition of same relative humidity 75%, adopt four different mean daily temperatures (DAT), respectively to spending 11 to handle in rl and the wild-type.Mean daily temperature is respectively 28 ℃, and 30 ℃, 32 ℃ and 34 ℃ (accompanying drawing 5-A).When DAT is 28 ℃, all the same grow normally (the accompanying drawing 5-B) of the Xiao Hua of all rl with wild-type; When DAT is 30 ℃, although coetonium phenomenon of separating in about Xiao Hua of about 10% occurs in the same tassel of rl, most of little flower development normal (accompanying drawing 5-C); When temperature is upgraded to DAT=32 ℃, the Xiao Hua of all wild-types all grows normally, but has 15% Xiao Hua the phenotype of coetonium part aristiformization to occur in the same tassel of rl approximately; 70% the coetonium aristiform that becomes fully of Xiao Hua is arranged approximately, the stamen shrinkage; 15% the Xiao Hua of also having an appointment shows coetonium disappearance or the interior coetonium aristiform (accompanying drawing 5-D) that all becomes.When mean daily temperature reaches 34 ℃, although the Xiao Hua of rl presents more serious dysplastic phenotype, show as coetonium or interior coetonium completely dissolve, the flower pesticide atresia, the filigree number increases, can not educate (accompanying drawing 5-E) fully, but it should be noted that the little flower development of wild-type also has been subjected to certain influence, shows as the Xiao Hua that degeneration is arranged on the tassel.Therefore, the Temperature Treatment experiment shows that rl is a temperature sensitive mutant, is proportionate between the seriousness of phenotype and the temperature.Analyze under the differing temps phenotype of wild-type and rl simultaneously and determined that also the high and low temperature condition of this mutant is respectively 32 ℃ and 28 ℃.Under four temperature condition, do not have obvious phenotypes difference at vegetative growth phase between wild-type and the rl in addition, illustrate that the phenotype of this temperature-sensitive mutant is mainly reflected in the growth of floral organ.
Claims (9)
1. the albumen of a RDR701, this proteic aminoacid sequence is SEQ ID NO:2.
2. the described proteic missense mutation albumen of claim 1, this Argine Monohydrochloride sequence is the 172nd the tryptophane generation missense mutation of SEQ ID NO:2.
3. according to the albumen under the claim 2, wherein the 172nd tryptophane generation missense mutation is a halfcystine for the tryptophane missense mutation.
4. coding claim 1 described proteic gene, the nucleotides sequence of this gene is classified SEQ ID NO:1 as.
5. the base mutation gene of the described gene of claim 4 is characterized in that the 516th " G " in the nucleotide sequence of SEQ ID NO:1 sports " T ".
6. a method of cultivating the temperature sensitive male sterile rice is characterized in that described proteic the 172nd tryptophane of claim 1 in the rice cell is carried out missense mutation.
7. method according to claim 6 is characterized in that with described proteic the 172nd the tryptophane missense mutation of claim 1 in the rice cell be halfcystine.
8. the method for the described cultivation temperature sensitive of claim 6 male sterile rice, it is characterized in that it is to insert by tissue culture, chemical reagent mutagenesis or fixed point T-DNA that described proteic 172 tryptophane of claim 1 in the rice cell is carried out missense mutation, is that proteic the 172nd tryptophane of SEQ ID NO:1 carries out missense mutation with sequence.
9. method according to claim 8 is characterized in that it is to be that proteic the 172nd tryptophane of SEQ ID NO:1 carries out missense mutation by tissue culture with sequence that described proteic 172 tryptophane of claim 1 in the rice cell is carried out missense mutation.
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Cited By (2)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| CN113930444A (en) * | 2020-06-28 | 2022-01-14 | 中国科学院遗传与发育生物学研究所 | Rice OsRDR6 protein and application of encoding gene thereof in regulation and control of plant male fertility |
| CN114073760A (en) * | 2020-08-21 | 2022-02-22 | 北京大学 | Application of RDR protein in tumor treatment |
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Non-Patent Citations (4)
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| 《GENBANK》 20080215 Nagasaki H et al GENBANK：AB353923.1 全文 1，4 , * |
| 《Trends Plant Sci》 20061231 Wassenegger M et al Nomenclature and functions of RNA-directed RNA polymerase 全文 1-9 第11卷, 第3期 * |
| 《中国农业科学院博士学位论文》 20060601 汪得凯 三个与水稻株型相关突变体的鉴定与基因克隆 全文 1-9 , * |
| 《中国农业科学院研究生院博士学位论文》 20091231 卫波 普通小麦和短柄草微RNA的发现及蕃茄小分子RNA合成相关基因RDR1的功能研究 全文 1-9 , * |
Cited By (3)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| CN113930444A (en) * | 2020-06-28 | 2022-01-14 | 中国科学院遗传与发育生物学研究所 | Rice OsRDR6 protein and application of encoding gene thereof in regulation and control of plant male fertility |
| CN114073760A (en) * | 2020-08-21 | 2022-02-22 | 北京大学 | Application of RDR protein in tumor treatment |
| CN114073760B (en) * | 2020-08-21 | 2024-02-13 | 北京大学 | Application of RDR protein in tumor treatment |
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