Molecular characterization and expression analysis

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Molecular characterization and expression analysis
of the porcine caveolin-3gene
Zhengmao Zhu
a,b
,Yong Li a ,Delin Mo a ,Kui Li b ,Shuhong Zhao
a,*
a
Key Lab of Agricultural Animal Genetics,Breeding,and Reproduction of Ministry of Education,
Huazhong Agricultural University,Wuhan 430070,PR China
b
Department of Gene and Cell Engineering,Institute of Animal Science,Chine Academy of Agricultural Sciences,Beijing 100094,PR China
Received 21April 2006Available online 2May 2006
Abstract
Caveolin-3is the muscle-specific form of the caveolin protein family and plays an important role in modulating both the morpholog-ical appearance and function of caveolae.In this study,we cloned and characterized caveolin-3from porcine muscle.The promoter of porcine caveolin-3contained three connsus E box elements and one ROR a 2monomeric binding motif.The deduced amino acid quence of porcine caveolin-3contains a WW domain.This gene was mapped to SSC13q23–q24by the SCHP and the IMpRH panel.RT-PCR analys showed that caveolin-3was expresd specifically in skeletal muscle and heart.And we provide the first evidence that caveolin-3has a certain regulated expression pattern during the prenatal period of the porcine skeletal muscle development.This result suggests that the caveolin-3gene might be a candidate gene of meat production trait and provides some information for establishing of an animal model using pig to study human caveolinopathies.Ó2006Published by Elvier Inc.
Keywords:Pig;Caveolin-3;Skeletal muscle;Differential gene expression
The caveolin family currently has three members,which are caveolin-1(CAV1),caveolin-2(CAV2),and
caveolin-3(CAV3).They are important in the formation of caveolar membranes as caveolin expression levels correlate very well with the morphological appearance of caveolae [1].Caveo-lae are small flask-shaped plasma membrane invaginations that participate in membrane trafficking,sorting,transport,and signal transduction [2].CAV3is the muscle-specific form of the caveolin protein family [3].Unlike CAV1and CAV2’s ubiquitous expression pattern,CAV3expresd restricted in muscle cells [4].Recent studies using caveo-lin-deficient mou models showed that caveolae and caveolins play a prominent role in various human patho-bi-ological conditions [5].CAV3mutations and expression alterations can result in caveolinopathies including limb
girdle muscular dystrophy,rippling muscle dia,distal myopathy,and hyperCKemia [6].
The number and size of myofibers are critical for meat production and quality [7].In the pig,the total number of muscle fibers is determined at the prenatal stages,while the fiber size is determined in the postnatal development process [8].Vistin et al.[9]reported that CAV3expression in type-2fibers is significantly higher than that in type-1fibers in human.Therefore,the skeletal muscle fiber type may be influenced by a CAV3-dependent transcriptional pathway,and the knowledge of modulation of CAV3gene will con-tribute to the understanding of muscle development.
Although mou has been ud as human caveolinopa-thies models,the pig is considered as an important experi-mental animal model of human dia,becau pigs and humans share similar anatomical,physiological,and path-ological characteristics.The knowledge of porcine CAV3,therefore,will also contribute to the understanding and development of porcine models for human caveolinopa-thies prevention and treatment.
0006-291X/$-e front matter Ó2006Published by Elvier Inc.doi:10.1016/j.bbrc.2006.04.132
*
眼中Corresponding author.Fax:+862787280408.
E-mail address:shzhao@mail.hzau.edu (S.Zhao).
/locate/ybbrc
BBRC
In this study,we characterized the mRNA quences of porcine CAV3and cloned the porcine CAV3promoter, investigated its expression patterns and chromosome assignments.
Materials and methods
Animal and tissue.Two pure breed,Duroc and Chine indigenous Tongcheng pigs,were ud in this study.One Duroc boar was mated with ven Duroc sows and one Tongcheng boar was mated with ven Ton-gcheng sows.The fetal skeletal muscles from ven ages(45,55,65,75,90, 105,and114days postconception(dpc))were harvested and then snap-frozen in liquid nitrogen.
cDNA library construction.Poly(A)+RNA for library construction was isolated from a55dpc porcine fetus skeletal muscle from Tongcheng pig by using the Oligotex direct mRNA midi kit and oligotex buffer t kit (Qiagen Inc.,Valencia,CA),a SMARTäcDNA library construction kit (Clontech,Palo Alto,USA)was ud for construction the full-length cDNA library.In order to check the inrt size of the clones of the cDNA library,plasmid DNA was extracted from200randomly lected clones with BioDev plasmid DNA purification kit(BioDev Inc.,China).The SfiIA and SfiIB restriction enzymes were ud to check the inrted fragments in the plasmids.The clones with the average inrted size of ab关于等待的作文
数字变变变out2kb were then quenced from the50-end with M13rever primer by ABI377automated DNA quencer.One quence showed high simi-larity with human CAV3gene.
Rever transcript-PCR analysis of CAV3gene expression.Total RNA isolated from the longissimus dorsi by using Trizol reagent(Gibco-BRL, Rockville,USA)following the supplier’s protocol,and treated with RNa-free DNa(Fermentas,Vilnius,Lithuania),was rever-tran-scribed into cDNA using Omniscript Rever Transcripta(Qiagen Inc., Valencia,CA)and the oligo(dT)primer.The rever transcription was carried out in afinal volume of20l L containing1l g of total RNA.The mixture was incubated for1.5h at37°C.Two microliters of the resulting single-stranded cDNA was amplified25cycles with CAV3-specific primers (Table1).A specific primer pair(Table1)that amplified the309-bp houkeeping gene GAPDH was ud as internal control.PCR
were parated by electrophoresis on2.0%agaro gels and visualized by ethidium bromide staining.The PCR fragments were purified and directly quenced to confirm the correct amplification of the porcine CAV3gene.
For temporal expression analysis of CAV3,total RNAs were isolated from longissimus dorsi muscle from various fetal developmental stages (45,55,65,75,90,105,and114dpc),three individual fetus we
re detected in each stage in order to do ANOVA analysis.For tissue-specific expression analysis,total RNAs were isolated from various tissues(heart, liver,skeletal muscle,lung,spleen,kidney,and bladder).
SYBR Green real-time RT-PCR analysis of gene expression patterns. The expression level of CAV3gene in three developmental stages was detected by SYBR Green I assay.Each real-time PCR(in20l L)reaction contained1·PCR buffer(TaKaRa),2.0mM MgCl2,500l M each dNTP, 0.4l M primers(Table1),0.3·SYBR Green I,and1U Taq DNA poly-mera(TaKaRa)plus2l L normalized template cDNA.The cycling conditions consisted of an initial,single cycle for3min at95°C followed by35cycles of cycling consisting of15s at94°C,20s at61°C,20s at 72°C,andfluorescence acquisition at82°C for1s.The specific PCR products were confirmed by melting curve analysis.cDNAs from three fetus muscle samples in each stage were ud to detect the expression changes of the target gene,and all PCRs were performed in triplicate and gene expression levels were quantified relatively to the expression of b-actin using Gene Expression Macro software(Bio-Rad,Richmond, CA,USA)by employing an optimized comparative C t(DD C t)value method.Expression levels were considered not detectable when the C t value of the targeted gene35in the sample tissue.The-test was conducted to identify genes differing in expression,p<0.05was considered significant.
Cloning and analysis of porcine CAV3promoter.To clone the50flanking region of the porcine CAV3gene,the human CAV3gene(GenBank Accession No.9930104)was aligned with Bos Taurus chromosome22 genomic contig(GenBank Accession No.NW_930087)to obtain homolo-gous quence for designing the primers.The primers spanned the50flanking region and the exon1(Table1).The amplified fragment was quenced and deposited in GenBank(GenBank Accession No.DQ415642).Promoter prediction analysis was performed using Neural Network Promoter Pre-diction(www.fruitfly/q_tools/promoter.html),CONSITE (b.ki./cgi-bin/CONSITE/consite),and Transcription Regulatory Element Search(bioportal.bic.nus.edu.sg/tres/).
Chromosomal assignment of the porcine CAV3gene.A somatic cell hybrid panel(SCHP)[10]was ud for chromosomal assignments and the radiation hybrid(IMpRH)was employed to improve the mapping reso-lution[11].Pig-specific primers(Table1)were designed in the30UTR of pig caveolin-3gene for chromosomal mapping.The PCRs were performed in a volume of20l L of1·PCR buffer(Promega),consisting of20ng of cell hybrid line DNA,10pmol of each primers,100l M of each dNTPs, 1.5mM MgCl2,and2.0U Taq DNA Polymera(Promega).The PCR profile was3min at94°C followed by35cycles of30s at94°C,30s at 56°C,30s at72°C,and afinal extension of5min at72°C.Analy
sis of the PCR results was performed with the software available on ulou.inra.fr/lgc/pig/hybrid.htm[12]and the IMpRH mapping ulou.inra.fr/)[13]for SCHP and RH mapping, respectively.Two-point RH analysis was ud for identification of linkage groups with LOD score threshold of3.0.
Table1
Primers ud in this study
Primer name Primer quence(50–30)Annealing temperature(°C) CAV3mi-RT-PCR PL CTCTGGGGCTTCCTGTTC52
CAV3mi-RT-PCR PR CATCACCTTGATGTTGCTG
GAPDH mi-RT-PCR PL CCTTCATTGACCTCCACTAC52
GAPDH mi-RT-PCR PR GTTGTCATACTTCTCATGGTTC
CAV3qPCR PL CGGCGTGTGGAAGGTGAG61
CAV3qPCR PR AAGGTGCGGATGCAGAGTG
b-Actin qPCR PL GGACTTCGAGCAGGAGATGG61
b-Actin qPCR PR GCACCGTGTTGGCGTAG GG
50flanking region PL AGAGCCTTACTGGTCTGTTT62
50flanking region PR CGATCTCCTTGAAGTGAATGT
Mapping PL GTGATGCTGCGGAAAGAA60
Mapping PR GGATGGGAACTGAGTGCTG
Note:The primer pairs CAV3mi-RT-PCR PL and PR,GAPDH mi-RT-PCR PL and PR were employed in mi-quantitative RT-PCR,the primer pairs CAV3qPCR PL and PR,b-actin qPCR PL and PR were ud in real-time RT-PCR.50flanking region PL and PR were ud for identifying porcine caveolin-3’sflanking region and the primer pair mapping PL and PR was employed for radiation hybrid mapping.
8Z.Zhu et al./Biochemical and Biophysical Rearch Communications346(2006)7–13
Results斜挎包男
Molecular cloning and quence analysis of porcine CAV3gene
The 950-bp full-length porcine CAV3cDNA (clone ID:sus_zm51;GenBank Accession No.AY731093)was identi-fied by randomly quencing the Chine Tongcheng pig 55-day fetal longissimus dorsi muscle full-length cDNA library [14].Sequence analysis showed that porcine CAV3cDNA contains an open-reading frame of 456nucleotides (nt),encoding a protein of 151amino acids with a calculat-ed molecular mass of 17.55kDa and an isoelectric point (p I )of 9.4.The 89nt 50-UTR and 405nt 30-UTR contain-ing a connsus polyadenylation signal (AATAAA)were also identified.The putative methionine initiation codon occurs in favorable quence context for the initiation of translation with an A in the À3position and a G in the +4position [15].To investigate the transcriptional regula-tion of porcine CAV3expression,we cloned and quenced the 50flanking region of the porcine CAV3gene (Fig.1).A connsus TATA box was identified at 29bp from the 50transcriptional start site.In a computer homology arch
for other cis -acting regulatory elements that have been implicated in skeletal muscle gene regulation,we identified three connsus E box elements that reprent putative binding sites for basic helix-loop-helix transcription factors and one ROR a 2element that functions as a constitutive trans -activator of gene expression (Fig.1).Alignment with the 50flanking region between porcine CA
V3and human CAV3gene showed that there were 419bp in the 50flank-ing region of porcine CAV3which had no homologous fragment in the human CAV3(Fig.1,underlined).
A GenBank databa arch using BLAST revealed that the predicted porcine CAV3amino acid quence has high similarity with other mammalian CAV3protein quences,with a 92%identity to human,an 88%identity to rat and mou,respectively.The phylogenic relationship among all the hitherto characterized members of the caveolin fam-ily was illustrated according to the phylogenetic distance calculated by the CLUSTALW1.83program [16](Fig.2).CAV2first diverged into a cluster,then CAV1and CAV3form two parate clusters.It is generally accepted that CAV2is expresd in a variety of cell types [17].
helix in CAV3predicted by www.cbs.dtu.dk/rvices/TMHMM-2.0)showed that 1–78amino acid residues are inside,79–101amino acid residues are transmembrane helix,while 102–151amino acid residues are outside [18].This is in consis-tent with the Kyte–Doolittle plot of the hydrophilicity,with 74-hydrophilic N-terminal domain,a 33-amino acid membrane-spanning gment,and a 44-amino acid hydro-philic C-terminal domain.We arched for putative WW domains within the caveolin-3protein quence.WW
domain is characterized as containing the conrved aro-matic residues Tryptophan(W,Trp),Phenylalanine(F, Phe),or Tyrosine(Y,Tyr),and Proline(P,Pro).The two W residues
parated by29amino acids are highly con-rved among caveolin family members.There is also a highly conrved residue followed with the cond trypto-phan.The conrved amino acid residues are indicated with an asterisk in Fig.3.租赁合同印花税税率
Chromosomal location of the porcine CAV3gene By PCR analysis of DNA from a somatic cell hybrid panel and a INRA-University of Mininesota porcine radi-ation hybrid(IMpRH)panel,the porcine CAV3was mapped to SSC13q23–(1/2)q24with an error risk less than0.5%.clost marker is SW960with LOD score of3.47.The conrved syntenic region in human is on chro-mosome3p25,where the human CAV3is located.
Temporal expression of CAV3gene during skeletal muscle development
To detect the expression pattern of CAV3gene in por-cine skeletal muscle,we performed the mi-quantitative RT-PCR analys of ven developmental time points, and samples from three individual fetus were
detected,
respectively,in each stage.The analys showed that CAV3expression level was incread from the 45dpc to the 75dpc fetal muscle,whereas the expression level was down-regulated from the 75dpc to the 114dpc fetal muscle of both Duroc and Tongcheng pig (Fig.4a).Moreover,pig CAV3was significantly differentially expresd among the stages in both Tongcheng pig (p =9.47e À05<0.01)and Duroc (p =5.84e À05<0.01).In Tongcheng pig fetal muscle,the mRNA abundance of CAV3in the 75dpc was 1.75-fold of that in the 114dpc,and in Duroc fetal muscle,the mRNA abundance of CAV3in the 75dpc was 1.96-fold of that in the 114dpc.The real-time RT-PCR results showed similar expression trend with the mi-quantitative RT-PCR analys at the corresponding three stages (Fig.4a and b).And pig CAV3was also signif-icantly differentially expresd among thes
e stages in both Tongcheng pig (p =3.67e À014<0.01)and Duroc (p =5.84e À012<0.01).At the 90dpc,the real-time RT-PCR results showed that the expression level of CAV3gene of Tongcheng pig was significantly lower than that in the Duroc (p =0.0016<0.01)(Fig.4b).
Tissue-specific expression of porcine CAV3gene
Tissue expression patterns of porcine CAV3gene were also analyzed using RT-PCR.Strong expression of CAV3was obrved in skeletal muscle and heart,and expression of porcine CAV3was undetectable in liver,kidney,lung,spleen,and bladder (Fig.5).Our result indicated that por-cine CAV3strictly expresd in striated muscle tissues.Discussion
CAV3gene has been so far characterized in human [19],mou [20],chicken [20],and zebrafish [21].In this study,we have obtained the full-length porcine CAV3cDNA from the pig 55dpc fetal longissimus dorsi muscle full-length cDNA library and cloned the porcine caveolin-3promoter.We also mapped this gene to SSC13q23–(1/2)q24.When browsing QTLs in the latest relead Pig QTL Databa (www.animalgenome/cgi-bin/QTLdb/brow )[22],veral QTLs,such as longissimus muscle area,shear force,backfat weight,feed intake,body weight at birth,and average daily gain,were mapped to this small chromosomal region.
In our study,porcine CAV3was found strictly expresd in striated muscle tissues.The result is generally in agree-ment with the report that CAV3is the muscle-specific form of the caveolin protein family [4].We identified three con-nsus E box elements and one ROR a 2element in the 50flanking region of the porcine CAV3gene,and we found that there were 419bp in the 50flanking region of porcine CAV3which had no homologous fragment in the human CAV3promoter region.Biederer et al.[23]reported that basic helix-loop-helix transcription factors mediated specific induction of CAV3gene expression during embry-onic development.And Lau et al.[24]uncovered that ROR a regulated the expression of CAV3in skeletal muscle cells.The difference of 50flanking region of CAV3in por-cine may affect its expression and requires further investigation.
Our study also showed that during the porcine muscle fiber hyperplasia period,the expression of CAV3was up-regulated,while in the period of the muscle
fiber
Fig.4.The temporal distribution of porcine caveolin-3mRNA in the fetal skeletal muscle from veral different stages.(a)Semi-quantitative RT-PCR results.Error bars indicate the SD (n =3)of relative mRNA expression levels of CAV3to GAPDH.The values were normalized to the GAPDH (houkeep
ing gene)expression.Pig CAV3was significantly differentially expresd among the stages in both Tongcheng pig (p =9.47e À05<0.01)and Duroc (p =5.84e À05<0.01).d45,45dpc;d55,55dpc;d65,65dpc;d75,75dpc;d90,90dpc;d105,105dpc;d114,114dpc.(b)Real-time RT-PCR results.Error bars indicate the SD (n =3)of relative mRNA expression levels of CAV3to b -actin.The values were normalized to endogenous b -actin expression.And pig CAV3was also significantly differentially expresd among the stages in both Tongcheng pig (p =3.67e À014<0.01)and Duroc (p =5.84e À012<0.01).d33,33dpc;d65,65dpc;d90,90
dpc.
Fig.5.Porcine CAV3gene expression in different tissues by RT-PCR.
Z.Zhu et al./Biochemical and Biophysical Rearch Communications 346(2006)7–1311
hypertrophy stages,the expression of CAV3was down-reg-ulated.It was generally agreed that pig skeletal musclefiber number stopped increasing from65dpc to85dpc,and after that the pig skeletal musclefiber began to hypertro-phy[7].This result indicates that porcine CAV3may have a relationship with musclefiber development.
We also showed that the expression of CAV3in Duroc was higher than that in Tongcheng pig.CAV3expression is significantly higher in type-2fibers than that in type-1fibers in human[9].There are more type-1fibers in Chine indigenous pigs than in the introduced pigs such as Duroc and Large White[25].Gueguen et al.[26]showed that the number of primaryfibers did not differ significantly between breeds,whereas the condary/primary ratio was dramatically reduced in Meishan pig.Duroc is well known for its high growth rate and lean meat percentage,whereas Tongcheng pig has low growth rate and low lean meat per-centage,but it has good meatflavor.Thus,the expression profile of porcine CAV3indicated that CAV3gene could have some relationship with meat production which is affected by musclefiber type.
In summary,we have isolated and characterized the porcine CAV3gene.Data prented in our study
pro-vide an expression and structural basis for future studies on porcine CAV3function and will potentially lead to a better understanding of the mechanism of caveolae sig-naling in skeletal muscle.The expression differences in pig breeds with different musclefiber types indicate a possible relationship to meat production.Our data also contribute to the understanding and development of por-cine models for human caveolinopathies prevention and treatment.
Acknowledgments
We thank Dr.Martine Yerle of INRA,France,for pro-viding the RH panel,and Huang Zhi,Cao Jianghua,and Professor Yu Chuanzhou for help with sample collection. The rearch was supported by National Natural Science Foundation of China(30571328,30400311),China Postdoc-toral Science Foundation(2005037086),National High Science and Technology Foundation of China (2004AA222170),and Key Project of National Basic Re-arch and Developmental Plan(G2000016103)of China. References
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