CN102382830B - Cloning and activity analysis of drought and salt damage induced promoter - Google Patents
Cloning and activity analysis of drought and salt damage induced promoter Download PDFInfo
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Abstract
The invention relates to cloning and activity analysis of a drought and salt damage induced promoter, and belongs to the technical field of plant gene engineering. The invention provides a corn drought and salt damage induced promoter sequence containing a DNA nucleotide sequence of -1bp to -1659p areas relative to an initial transcription site of SEQ ID NO: 1, and simultaneously provides a drought and salt damage induced plant expression vector for corn transformation, wherein the plant expression vector contains a corn drought induced promoter sequence and a 5' un-translated region of corn dehydrin genes; and polymerase chain reaction (PCR) primers of SEQ ID NO: 2 and SEQ ID NO: 3 are suitable for amplifying DNA fragments containing the sequence of SEQ ID NO: 1. The cloning and activity analysis can be used for promoting efficient expression of adversity related genes, are used in drought and salt tolerant transgenic plants, and have significance for solving the problems of food crisis, ecological deterioration and the like in drought and salt damage districts.
Description
Technical field
The invention belongs to the plant gene engineering technology field; Relate to the section of DNA sequence; It can be used as the promoter regulation expression of gene under arid and two kinds of environment stresses of salt damage; Specifically, the present invention relates to a kind ofly derive from corn dehydrin gene (Zmdhn2), and arid that in plant, efficiently expresses and salt damage inducible promoter sequence.
Background technology
In Global climate change aggravation, resource environment pressure increases, and the financial crisis influence increases the weight of, and under the background that the descending trend of grain-supply deepens constantly, ensures that China's grain security is a urgent and important strategic task all the time.Environment stress is to influence the most important factor of grain yield and one of challenge in the present agriculture prodn.The annual crop production reduction that is directly caused by environment stresses such as saline and alkaline, arid, low temperature is above about 20% of gross output.
Corn is one of China's three generalized grain kinds, and cultivated area and output are all at the forefront in the world, and to ensureing national food safety, it is extremely important to improve the main farm produce supply capacity.Corn belongs to adverse circumstance responsive type plant, very easily receives the interference and the injury of polynary environment stresses such as low temperature, arid, salt marsh in breeding time.Solve serious polynary environment stress problem in the Maize Production, improve corn yield, except traditional breeding method, the biological breeding technique that transforms based on gene genetic is an important selection.Though existing many gene engineering method are cultivated resistant variety, have obvious defects, that is: in genetically engineered, the promotor that is used for controlling destination gene expression is main with composition type expression promoters such as CaMV35S, corn Ubiquitin mainly.Though this promotor can make the goal gene overexpression, it is the promotor of the constitutive expression of no environment, tissue and temporal.Though utilize the overexpression of composition type expression promoter control resistance related gene can improve the anti-adversity ability of plant under adverse environmental factor; But also can the degeneration-resistant albumen of overexpression in the cell under the home condition; To plant is a kind of energy wastage; And the long-time excessive accumulation of intracellular degeneration-resistant albumen possibly hinder the normal growth metabolism of plant under the home, causes that the form of plant changes, and causes transgenic plant serious stagnate even dead of growing.Therefore, utilize the adverse circumstance inducible promoter to realize that the regulation and control adversity gene of efficient, controlled and special (fixed point, regularly, quantitatively) becomes the focus of current research in the resistance of improvement crop.
Abiotic stress; Like arid, high salt, low temperature and high temperature etc.; Related gene expression in can the inducing plant body; One of main mechanism of its transcriptional regulatory is crucial cis element specific combination in relevant transcription factor of environment stress (adjusting albumen) and the upstream region of gene promotor, strengthens the activity of promotor, improves the expression amount of goal gene.Some reports about the adverse circumstance promotor are arranged both at home and abroad, and the rd29A promotor is induced by adverse circumstances such as ABA, arid, high salt and low temperature, is (Li Xinling etc., 2007 of most widely used inducible promoter in the present adversity resistant plant genetically engineered; Yang Chunxia etc., 2008; Wu Meihua etc., 2005; Yamaguchi-Shinozaki and Shinozaki, 1993).Yamaguchi-Shinozaki etc. (2001) utilize CaMV 35S constitutive promoter and rd29A adverse circumstance inducible promoter to obtain the transfer-gen plant of DREB 1A respectively; The result shows the transfer-gen plant of the stress tolerance of rd29A::DREB 1A transfer-gen plant apparently higher than 35S::DREB1A, shows that the rd29A promotor can more effectively improve the flexibility of plant to environment stresses such as low temperature, arid and high salt.Brown etc. (2001) have found to receive the blt101.1 gene promoter of low temperature induction first in winter wheat; And identified the TCA like element (TCATCTTCTT) of a high conservative through gradual change disappearance, show this element and to induce goal gene under low temperature stress, to efficiently express closely related.Takumi etc. (2003) are separated to the promoter sequence of a low temperature response gene Wcor15 upper reaches 1.7kb at little wheat seeds; The result shows and includes 4 CRT/DRE like sequences in this section; Comprise CCG AC core motif allow, this promotor of experiment proof receives low temperature and photoinduction.
Utilize the adverse circumstance inducible promoter to drive efficiently expressing of adversity gene, thereby the improvement crop has become present research focus to the flexibility of environment stress.Yet the inducible promoter that can be applied to transgenic research at present still seldom.Therefore, for the interaction between the confirming of the promoter related clone of new adverse circumstance, the concrete sequence of cis-acting elements, each element, and the research of the transcription factor of doing mutually with these elements remains the emphasis of promotor research from now on.
Summary of the invention
An object of the present invention is to provide a kind of arid and salt damage inducible promoter sequence, this promoter sequence can be induced the target gene high level expression, and this promotor derives from corn dehydrin gene (Zmdhn2).
Second purpose of the present invention provides a kind of arid and salt damage inducible promoter carrier that is used for Plant Transformation, and this carrier instructs arid and salt damage inducible promoter sequence and the 5 ' non-translational region of high level expression target gene Zmdhn2.
The 3rd purpose of the present invention provides a kind of transient expression system that detects promoter activity, and this system can reflect the height of promotor induced activity.
Be to realize above-mentioned purpose, the present invention has cloned the promoter region of corn dehydrin gene (Zmdhn2), and has made up and be used for the plant transformed expression vector, comprises the above-mentioned promotor of the 5 ' non-translational region that has corn dehydrin gene (Zmdhn2) in the said carrier.Utilize said carrier in maize, to carry out abduction delivering subsequently, and it is active to observe this promotor high level relevant with the salt damage inducibility with arid.
The corn arid that the present invention provides a kind of acquisition and salt damage inducible promoter sequence comprise with respect to the transcription initiation site of SEQ ID NO:1-1bp is to-the dna nucleotide sequence of 1659bp district (seeing sequence table), arid and salt damage can be induced the high-level activity of promotor of the present invention in plant.
Corn arid that obtains and salt damage inducible promoter sequence source are from plain (dehydrin) gene (Zmdhn2) of corn dehydration.
5 ' non-translational region of the corn dehydrin gene (Zmdhn2) that a kind of clone obtains; Comprise with respect to the transcription initiation site of SEQ ID NO:1+1bp is to+the dna nucleotide sequence of 65bp district (seeing sequence table); Non-translational region of the present invention can be renderd a service through strengthening the translation that imports the target foreign gene in the plant, and induces the high level expression of target gene.
For realizing another purpose, the present invention provides a kind of plant expression vector that is used for the drought-inducible of corn conversion, and said plant expression vector comprises 5 ' non-translational region of corn drought-inducible promoter sequence and corn dehydrin gene (Zmdhn2).
The above-mentioned drought-inducible carrier that is used for the corn conversion is meant can be at the binary vector of the lasting expression alien gene of transgenic plant.Said binary vector can be the RB (right border sequence) and the LB (left margin sequence) of any T-DNA of comprising (transfer DNA), can in the presence of agrobacterium tumefaciens (Agrobaterium tumefaciens) Ti-plasmids, transform the binary vector of plant.This carrier can be through being usually used in the binary vector of association area, for example pCAMBIA1301 carrier, pBI121 carrier.
The present invention provides the PCR primer of SEQ ID NO:2 and SEQ ID NO:3, is suitable for increasing comprising the sequence DNA fragment of SEQ ID NO:1.
About the induction type carrier (pCAMBIA1301) that is used for plant of the present invention, the promotor of described corn corn dehydrin gene (Zmdhn2) and 5 ' non-translational region are positioned at the front of foreign gene in plant expression vector.The present invention provides the pCAMBIA1301 that makes up through in the promotor of inserting corn dehydrin gene of the present invention (Zmdhn2) and 5 ' non-translational region to the carrier that contains gus reporter gene.But said gus reporter gene is a foreign gene, and expection can replace with any other useful foreign gene.
The present invention provides a kind of and uses the arid and the transgenic plant of salt damage induction type binary vector, and a kind ofly uses the corn mature embryo that carries out transient expression according to the invention.
The present invention provides from the arid of corn dehydrin gene (Zmdhn2) and salt damage inducible promoter and 5 ' non-translational region, makes it possible in plant, carry out arid and salt damage inducible expression according to promotor of the present invention and 5 ' non-translational region.
Beneficial effect of the present invention is to can be used for starting efficiently expressing of adverse circumstance genes involved, is applied in the transgenic plant of drought-enduring and salt tolerant, for problems such as solving arid and salt damage area crisis in food, ecological degeneration positive effect is arranged all.
Description of drawings
Fig. 1 is corn (B73) genome electrophorogram
Fig. 2 is a Zmdhn2-pro PCR electrophorogram
Wherein, M is DL2000Marker, and size is followed successively by 2000bp, 1000bp, 750bp, 500bp, 250bp and 100bp from top to bottom, and 1-5 is the purpose band
Fig. 3 reclaims electrophorogram for Zmdhn2-pro
Fig. 4 connects bacterium liquid PCR figure behind the T carrier for Zmdhn2-pro
Wherein, M is DL2000Marker, and 1-5 is PCR result
Fig. 5 is Zmdhn2-pro restriction enzyme digestion and electrophoresis figure
Wherein, M is DL2000Marker, and 1-5 is that enzyme is cut the result
Fig. 6 is that electrophorogram is received in the switchback of Zmdhn2-pro enzyme
Fig. 7 is that pCAMBIA1301::Zmdhn2Pro bacterium liquid PCR identifies electrophorogram among the intestinal bacteria Top10
Wherein, M is DL2000Marker, and 1-6 is bacterium liquid PCR result
Fig. 8 for pCAMBIA1301::Zmdhn2Pro change Agrobacterium after bacterium liquid PCR detect electrophorogram
Wherein, M is DL2000Marker, and 1-6 is bacterium liquid PCR result
Fig. 9 is the plant expression vector construction synoptic diagram
Figure 10 is the GUS colored graph (front) of Zmdhn2 promotor
Figure 11 is the GUS colored graph (back side) of Zmdhn2 promotor
Wherein, among Figure 10 and Figure 11: be followed successively by from left to right: pCAMBIA1301 is untreated, Zmdhn2 is untreated, Zmdhn2 through PEG induce, Zmdhn2 induces through NaCl
Embodiment
Introduce specific embodiment below in conjunction with accompanying drawing: enforcement of the present invention can make those skilled in the art how more be expressly understood embodiment of the present invention.Invention has been described although combined preferred embodiment of the present invention, and following purpose of description is exemplary, rather than limits scope of the present invention.
Embodiment 1: the clone of the promotor of corn dehydrin gene (Zmdhn2)
The promotor of corn dehydrin gene (Zmdhn2) (promoter sequence comprise with respect to the transcription initiation site of SEQ ID NO:1-1bp is to the dna nucleotide sequence in-1659bp district), in 5 ' region sequence of corn Zmdhn2 gene, obtain evaluation.
(accession number: L35913), sequence table shows the corn arid of said gene of the present invention and the dna sequence dna of salt damage inducible promoter and 5 ' non-translational region at NCBI GenBank in the Zmdhn2 login.In SEQ ID NO.1, translation initiation codon ATG has underscore, utilizes
Http:// www.fruitfly.org/seq_tools/promoter.htmlPrediction transcription initiation site in website is with+1 mark.And with the promoter Analysis website (
Http:// bioinformatics.psb.ugent.be/webtools/plantcare/html/) analyzed the core parts of promotor.
The CTAB method is extracted corn gene group DNA (Fig. 1), is that template amplification obtains the purpose fragment with the genomic dna, through 1% agarose gel electrophoresis analysis; Amplify the specific band that length is 1724bp (Fig. 2); And utilize glue to reclaim test kit and reclaim (Fig. 3), reclaim fragment and be connected with the pMD18-T carrier and obtain recombinant plasmid pMD18-T::Zmdhn2Pro, transformed into escherichia coli Top10; Shake bacterium behind the picking list bacterium colony and carry out bacterium liquid PCR detection (Fig. 4), the bacterium liquid that is positive is preserved and is also sent company's order-checking.
More particularly, the promotor of the corn dehydrin gene (Zmdhn2) through pcr amplification clone and the 5 ' non-translational region (seeing sequence table) of 65bp, above-mentioned PCR the primer is presented in the table 1 in detail.For pcr amplification, response procedures: 95 ℃ of 3min; 95 ℃ of 30s, 58 ℃ of 30s, 72 ℃ of 2min, 31 circulations; 72 ℃ of 10min; 16 ℃ stop.
Table 1: primer
| Upstream primer | ?5′CGGAATTCGGACCTTACCTCTTTATGATTC?3′ | SEQ?ID?NO：2 |
| Downstream primer | ?5′TGCCATGGAACGTACAGTACAGATGGCTTT?3′ | SEQ?ID?NO：3 |
Embodiment 2: the structure of corn dehydrin gene (Zmdhn2) promoter expression vector
5 ' the non-translational region Zmdhn2Pro (seeing sequence table) of corn dehydrin gene (Zmdhn2) promotor that in embodiment 1, be cloned and 65bp is inserted in the carrier, thereby makes up corn dehydrin gene (Zmdhn2) promoter expression vector.
More particularly; Cut recombinant plasmid pMD18-T::Zmdhn2Pro (Fig. 5) and plant expression vector pCAMBIA1301 with EcoRI and NcoI enzyme respectively; Reclaim purpose fragment (Fig. 6) and pCAMBIA1301 respectively; Connect with the T4 ligase enzyme, obtain pCAMBIA 1301::Zmdhn2Pro expression vector, be used to drive the expression of gus gene.Obtained promoter fragment (Fig. 7) through the PCR evaluation, identical with expected results.Utilize electric shocking method to transform Agrobacterium EHA105, and carry out bacterium liquid PCR and detect (Fig. 8).
In Fig. 9, with the coding β-glyconic acid aldehydase gene GUS be reporter gene, selective marker is a hygromycin gene.35S-pro represents the promotor of HPTII in addition, and 35S-ter represents its terminator, and Zmdhn2Pro represents the promotor of GUS, and Nos-ter represents its terminator.
Embodiment 3: the activity of identifying corn arid of the present invention and salt damage inducible promoter
Be to identify the adverse circumstance induced activity of promotor, utilize the method for Jefferson etc. (EMBO J, 1987) that the embryo arid of corn is handled the activity that detects GUS again.
3.1 the preparation of acceptor material
Earlier corn seed is soaked 1d in water, let it fully absorb water, put into 24 ℃ environment vernalization 1d, then that the seed rip cutting is subsequent use into two.
3.2 the preparation of Agrobacterium bacterium liquid
Agrobacterium EHA105 is inoculated on YEP (50mg/L rif, the 50mg/L kan) solid medium, and 28 ℃ of dark cultivations are to growing single bacterium colony.Picking list bacterium colony is seeded in 5mL and contains in the corresponding antibiotic YEP liquid nutrient medium, in 28 ℃, and the 200r/min shaking culture.Treat that bacterium liquid grows to OD
600, bacterium liquid is carried out re-activation in 1: 10 ratio at=0.4～0.6 o'clock.Work as OD
600=0.4～0.6 o'clock, draw bacterium liquid and place aseptic centrifuge tube, the centrifugal 10min of 5000rpm abandons supernatant, and is resuspended, subsequent use with isopyknic MS liquid nutrient medium.
3.3 Agrobacterium is infected
Get the part that radicle and bud are arranged in the seed and put into the 50ml centrifuge tube, add the good Agrobacterium bacterium liquid of an amount of activation, after suction filtration infects about 5min seed is put on the moistening filter paper, the dark place is cultivated 24h altogether under 24 ℃ of conditions.
3.4GUS dyeing
With inducing culture 6h among the NaCl of 20%PEG or 200mM, water treatment is used in contrast respectively, then corn seed is put in GUS and detects in the liquid, and suction filtration 3min spends the night in 37 ℃.GUS detects liquid: 3mg/ml X-gluc (5-bromo-4-chloro-3-indoles-β-D-glucuronide), 50mM buffer solution of sodium phosphate (PH=7.0), 10mM EDTA, the 0.5mM Tripotassium iron hexacyanide, 0.5mM yellow prussiate of potash, 0.1%TritonX-100,10% methyl alcohol.As shown in the figure (Figure 10, Figure 11) active through demonstrating high-caliber GUS in PEG and the NaCl inductive radicle, and it is active to demonstrate low-level GUS without the inductive tissue.
Industrial applicibility
As stated, the present invention provides corn dehydrin gene (Zmdhn2) arid and salt damage inducible promoter and 5 ' non-translational region.
The present invention provides respectively corn dehydrin gene (Zmdhn2) arid and salt damage inducible promoter and 5 ' non-translational region is inserted into pCAMBIAl301 and the plant expression vector that makes.
Identify that through using described plant expression vector the promotor and the 5 ' non-translational region of corn dehydrin gene of the present invention (Zmdhn2) make it possible to carry out the expression of drought-inducible.Therefore, it is active that the present invention can be used for improving drought-enduring startup with resistant gene of salt, the food crop or the vegetation of the drought-enduring and salt tolerant that final acquisition can be used to produce.
The sequence of SEQ ID NO.1 (band functional element mark)
(i) sequence signature: (A) length :-1bp is to-1659bp; + 1bp is to+65bp; (B) type: Nucleotide; (C) chain property: strand.
(ii) molecule type: Nucleotide
(iii) sequence description: SEQ ID NO.1
-1659?
GGACCTTACC?TCTTTATGAT?TCATCCTATC?GTCTGACTAG?GCAACTAGTC?GACTGCCACT
CAT-box
-1599?AGCCACTTGA?CTTGCATCTG?CCTTAGCCCC?ACAACCAAAG?TCCAAATAAG?GTGATCACGC
CAT-box CAAT-box
-1539?GAATCGGCAG?AATGACCG
CC?AATAACTTAT?ATACATACTA?CATTAGATCT?TGTGGATGAT
MBS CAAT-box
-1479?AAATTCTACA?GATTTCAACT?AAAGTTATCA?CCTGATAGCT?GGTGTGTA
CA?CGACAAATTT
G-box?CAAT-box
-1419?GACATGTACC?GACGTAGGTT?GACGAAGTTT?CATGGATCCT?CGAGGCTAGT?TAGATTTTTT
CGTCA-motif
-1359?ATTTCTCTCA?AAAATTTGTT?AGCTGCCTTA?TAAATACTAG?AGTCGTTCTA?TAAAAGTTAA
-1299?ATCTCATTAT?TCGGATATTC?ATCGTTATGT?CGTCTTTATT?ATTTTGGTAT?TGTTCTTGCG
-1239?TCACAACTTT?TAATTCAATA?CATCAATTAG?CTAGAAAATA?GACTCATGAA?TTAAGATAAT
CGTCA-motif CAAT-box?CAAT-box
-1179?TTATAAAGAT?ATAGATTTCT?TATTCCATAA?CGAGTGCTTA?AACTTATAAA?TTAGCTAGAA
-1119?AATAGACTCA?TGAATTAAGA?TAATATATCA?ATTAGCTAGA?AAATAGACTC?ATGAATTAAG
CAAT-box
-1059?ATAATTTATA?AAAAATATTT?TTTATCGTAT?AGTATTGATC?AATTGATTTG?ATTGGCATTC
CAAT-box
-999 CTTATATCGT?AATCTGGCTA?TTGTCGGGCC?AAATCCCCCC?TCACACCCCC?ACCGGATATG
-939 GCTAGGTTCA?GCCGCCCAGC?GACGGATCAG?CGGGTCCCGC?AGAGCTAGCG?TCCAGACCCA
GCC?box
-879 CATGTCAACG?TCGAAGCCTC?TAGGCCTCTC?CCGTCGTCCG?AGTCATCCAA?CCCCCGTACG
ABRE
-819 TGGAGTGGCG?GAATGCAATA?ATCACTCTCG?TGCCCCGGCC?GCGTCGCGTA?GGCAACTTGC
ABRE GC-motif?ABRE
-759 AAACTTCAAC?AACGACAAGT?TCATTCGACT?AAACTTCCTT?AGATTTAACA?GAACCTGACA
TGA-element
-699 TTATTAGTAA?CATTTGTACT?GTTTCTGAAC?GAGCAATTGG?AAGATCGATC?TAAAACAACA
-639 GTTCATTCGA?AACGGAGGCA?GTAGACACAT?CTCTCCTGTA?AAAAAGGAGG?GGTACCGTCA
CGTCA-motif
-579 AGCTGAAAGT?GAAACTGACG?ACGCTAACGC?TTGATTGTTG?GGTTGTCCAG?TTGAAAGTGA
TGACG-motif MBS
-519 AACTGACATT?CGAAACTGAA?GCGAGCCCGA?CGAGTCCAAA?GCCAACAACA?ACAAA
AAAAC
MBSI
-459
GCTTACGTCT?TCAACCAAAG?TACTTGAATG?AAACTTACGA?GTAGAAAACT?GATGAACGGA
ACE
-399?AGACAAGCGT?CGCGCGGGCA?GCGGAGCAGG?TGAGAGAGTT?AAGGGGACCG?ACATCGGGCC
CE3
-339?GTCCACGCTT?TCTCGCCCGC?GCGCGTCGCC?GACCTCTCCC?GTCCCGACGC?CGTCCACAGC
CCGTCC-box CCGTCC-box?CCGTCC-box
-279?AGCAGCAGTA?CCGAACCTCG?AGGCACCAGC?CACGTCGGAA?CCAGGCCGCC?CCACGCGAGG
G-box
-219?GGCGCGACAC?GCGTCGTGGC?GCTACTGGCC?GTGGCCCTGC?CCACCCGCCC?GCCACCAGGC
CE3
-159?CATGTCCCCG?TCGGCCGCGG?GCTGATCTGT?CTGTCCTTCG?CGTCGCGTAG?CCTACGCGTC
ABRE
-99 GCACGCATCA?ACGCGGCTGG?AACGTTACAC?TGCGTACACC?TTTCGGTTCC?TTTCCTTTAG
LTR
+1
-39 CTTGCTCTAT?ATAAGGACGC?CTCCGGCCTA?GCCACCTCCA?CCATCCGAAA?GCCCAAGACA
LTR
+22 CTGAGCTGGT?CTGTTTAGTT?GGAAAGCCAT?CTGTACTGTA?CGTTTTTCGC?CGATCATG?。
Claims (3)
1. the corn of acquisition arid and salt damage inducible promoter sequence; It is characterized in that said corn arid and salt damage inducible promoter sequence are shown in SEQ ID NO:1; Wherein SEQ ID NO:1-1bp to-1659p district is the dna sequence dna of corn dehydrin gene Zmdhn2 promotor, SEQ ID NO:1+1bp extremely+the 65bp district is the dna sequence dna of the 5 ' non-translational region of corn dehydrin gene Zmdhn2.
2. one kind is used for the arid of corn conversion and the plant expression vector of salt damage induction type, it is characterized in that said plant expression vector comprises said corn arid of claim 1 and salt damage inducible promoter sequence.
3. a PCR primer is right; Its sequence is shown in SEQ ID NO:2 and 3; Said primer is to being suitable for increasing corn arid and salt damage inducible promoter sequence according to claim 1; Wherein: the sequence of SEQ ID NO:2 is 5 ' CGGAATTCGGACCTTACCTCTTTATGATTC3 ', and the sequence of SEQ ID NO:3 is 5 ' TGCCATGGAACGTACAGTACAGATGGCTTT 3 '.
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| 刘晓敏等.玉米逆境诱导型启动子克隆及其植物表达载体构建.《生物技术通报》.2011,(第3期),第86-90页. * |
| 植物非生物胁迫诱导启动子顺式元件及转录因子研究进展;郭晋艳等;《生物技术通报》;20110430(第4期);第16-21页 * |
| 王志新等.植物诱导型启动子的研究进展.《大豆科技》.2011,(第3期),第5-9页. * |
| 郭晋艳等.植物非生物胁迫诱导启动子顺式元件及转录因子研究进展.《生物技术通报》.2011,(第4期),第16-21页. |
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Granted publication date: 20121205 Termination date: 20131110 |