EP0889904A2 - Gene codant une proteine se liant a un acide gras du coeur de porc et procede d'identification des caracteristiques du polymorphisme responsables du poids du corps - Google Patents

Gene codant une proteine se liant a un acide gras du coeur de porc et procede d'identification des caracteristiques du polymorphisme responsables du poids du corps

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Publication number
EP0889904A2
EP0889904A2 EP97914649A EP97914649A EP0889904A2 EP 0889904 A2 EP0889904 A2 EP 0889904A2 EP 97914649 A EP97914649 A EP 97914649A EP 97914649 A EP97914649 A EP 97914649A EP 0889904 A2 EP0889904 A2 EP 0889904A2
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EP
European Patent Office
Prior art keywords
alleles
pig
pigs
fabp
gene
Prior art date
Legal status (The legal status is an assumption and is not a legal conclusion. Google has not performed a legal analysis and makes no representation as to the accuracy of the status listed.)
Withdrawn
Application number
EP97914649A
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German (de)
English (en)
Inventor
Frans Gerbens
Current Assignee (The listed assignees may be inaccurate. Google has not performed a legal analysis and makes no representation or warranty as to the accuracy of the list.)
Noord Nederlands Varkensstamboek BV
Pig Genes Bv Io
ProVa BV
Stamboek Zuid BV
Stichting Instituut voor Dierhouderij en Diergezondheid (ID-DLO)
Original Assignee
Noord Nederlands Varkensstamboek BV
Pig Genes Bv Io
ProVa BV
Stamboek Zuid BV
Stichting Instituut voor Dierhouderij en Diergezondheid (ID-DLO)
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Application filed by Noord Nederlands Varkensstamboek BV, Pig Genes Bv Io, ProVa BV, Stamboek Zuid BV, Stichting Instituut voor Dierhouderij en Diergezondheid (ID-DLO) filed Critical Noord Nederlands Varkensstamboek BV
Priority to EP97914649A priority Critical patent/EP0889904A2/fr
Publication of EP0889904A2 publication Critical patent/EP0889904A2/fr
Withdrawn legal-status Critical Current

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Classifications

    • C—CHEMISTRY; METALLURGY
    • C07—ORGANIC CHEMISTRY
    • C07K—PEPTIDES
    • C07K14/00—Peptides having more than 20 amino acids; Gastrins; Somatostatins; Melanotropins; Derivatives thereof
    • C07K14/435—Peptides having more than 20 amino acids; Gastrins; Somatostatins; Melanotropins; Derivatives thereof from animals; from humans
    • C07K14/475—Growth factors; Growth regulators
    • C—CHEMISTRY; METALLURGY
    • C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
    • C12Q—MEASURING OR TESTING PROCESSES INVOLVING ENZYMES, NUCLEIC ACIDS OR MICROORGANISMS; COMPOSITIONS OR TEST PAPERS THEREFOR; PROCESSES OF PREPARING SUCH COMPOSITIONS; CONDITION-RESPONSIVE CONTROL IN MICROBIOLOGICAL OR ENZYMOLOGICAL PROCESSES
    • C12Q1/00—Measuring or testing processes involving enzymes, nucleic acids or microorganisms; Compositions therefor; Processes of preparing such compositions
    • C12Q1/68—Measuring or testing processes involving enzymes, nucleic acids or microorganisms; Compositions therefor; Processes of preparing such compositions involving nucleic acids
    • C12Q1/6809—Methods for determination or identification of nucleic acids involving differential detection

Definitions

  • the invention relates to the field of molecular biology as well as to the field of breeding methods for farm animals, in particular pigs.
  • the invention relates to the use of diagnostic methods derived from the field of molecular biology to be applied in breeding programmes that select animals on production traits that improve their breeding value.
  • breeding programmes By selecting animals on their breeding value calculated mainly from phenotypic measurements of production traits, breeding has greatly improved the genotype for production traits of livestock animals.
  • breeding programmes have selected for phenotypic characteristics of animals.
  • more recently selection for genotypic characteristics that are associated with improved production traits have gained interest in the field. Selection for phenotypic characteristics entails mainly selection of the offspring or siblings or other relatives of the animals to be selected whereas selection of specific genotypic characteristics allows for earlier and specific detection of animals of interest.
  • One of the former methods is a marker assisted selection wherein polymorphisms in markers identified in a random manner are associated with production traits .
  • meat production is closely linked to embryonic muscle formation, and, consecutively, to the distribution of muscle cells and fat cells.
  • Biologically production is concentrated in defined tissues of the animal, e.g. muscle tissue for lean meat production.
  • various levels of selection pressure have been applied to different tissues (i.e. muscle, fat and bone) .
  • selecting for lean meat, and thus the abscence of fat one may lose certain traits that are wanted after all, i.e. traits that are associated with taste and thus with the consumers perception of the final product.
  • H-FABP heart fatty acid-binding protein
  • Fatty acid binding proteins are small intracellular proteins involved in fatty acid transport from the membrane to the sites of ⁇ oxidation and/or triacylglycerol or phospholipid synthesis (Veerkamp and Maatman, 1995). Furthermore, FABP's modulated the intracellular fatty acid concentration (Veerkamp et al.,1993) . Fatty acid metabolism has historically been linked to insuline resistance (Randle, 1963), and therefore mutations in FABP genes may be associated with changes in cellular insulin resistance or dependency, fatty acid oxydation and fatty acid binding.
  • FABP's are members of a family of intracellular Iipid binding proteins comprising at least eight structurally distinct types originating from: adipocytes, brain, epidermal cells, heart, intestinal cells, ileal cells, liver and myelin cells.
  • H-FABP heart type FABP
  • MDGI mammary derived growth inhibitor
  • H-FABP can induce cardiac myocyte hypertrophy in vitro, when added to the culture (Burton et al.,1994) and also promotes functional differentiation of mammary epithelial cells in vitro (Yang et al.,1994).
  • no secretion of H-FABP has been detected so far.
  • H-FABP H-FABP gene specific nucleic acid molucule or pig H-FABP gene specific fragments thereof comprising or hybridising to the nucleotide sequence as shown in figure 1, or its complementary sequence or the RNA equivalents thereof.
  • the locus of this gene is on porcine chromosome 6.
  • the pig H-FABP gene can be assigned functions in the regulation of intramuscular fat, thereby changing the ratio of fat deposited within the muscle versus fat deposited outside the muscles, i.e. in backfat depots. Since production and deposition of fat is energy consuming and takes away energy for other purposes, such as muscle growth, the regulation of intramuscular fat is correlated to the regulation of growth, and thus body weight and average daily gain and feed efficiency. Also, H-FABP can regulate myocyte (and thus muscle) hypertrophy and thus also muscle regeneration. Since FABP's are involved in fatty acid transport they can influence fatty acid oxidation rates, the metabolism of fatty acid derivatives in the tissue and the fatty acid composition of cells and thus of meat.
  • H-FABP may regulate cellular insulin dependency.
  • fat storage has an impact on embryo survival, and regulation of H-FABP will influence birth rates and littersize.
  • H-FABP regulates functional differentiation of mammary epithelial cells it may be involved in regulating the quantity and composition of the milk available, thus influencing the growth and survival of newborn animals.
  • the genetic variation within the pig H-FABP gene with respect to variation in regulation of expression can now be revealed and analysed for association with above production traits and physiological characteristics.
  • the present invention further provides a method to generate via recombinant DNA techniques an animal, such as small laboratory animals or farm animals, i.e. a pig, with additional genetic material originating from the pig H-FABP gene. Such animals may than encode wanted alleles of this gene and constitutively or transiently express allelic proteins or fragments thereof that enhance the production or physiological characteristics of those animals.
  • the invention further provides methods to generate proteins or (poly)peptides comprising various allelic proteins or fragments thereof derived from the pig H-FABP gene. Such peptides, or antibodies specifically directed against such peptides, may be used to influence production traits in the live animal, but may also be used in cell- culture systems in vitro . Such (poly)peptides or proteins, or antibodies specifically directed against these, may also be used in diagnostic test systems to select animals that express wanted forms of allelic proteins or fragments thereof encoded by the pig H-FABP gene.
  • the invention further provides methods localising, identifying or marking genes or alleles or quantitative trait loci, in particular those corresponding to the pig H- FABP gene, in samples, in particular biological samples, cells or tissues, such as but not limited to hair, skin or blood, of farm animals, in particular pigs, by allowing for specific amplification of genomic fragments of those genes or alleles or quantitative trait loci of pigs . Since marker assisted selection of animals is frequentally based upon genetic variation that exists within functional genes that influence a production trait directly, i.e.
  • genes such as the pig H-FABP that regulates fatty acid binding one of the methods that the invention provides is a method that identifies or marks loci or genes and that can distinguish between characteristics of alleles of those genes which characteristics serve as markers in selection programmes for animals with specific versions of those genes that are directly linked with improved production traits.
  • the invention further provides a method wherein polymorphic restriction sites within functional genes and thus different alleles of those genes are identified by allowing for specific amplification of genomic fragments of those genes, in particular by allowing for specific amplification of fragments of the H-FABP gene.
  • Amplification methods are well known in the art, the best known being PCR. A short description of the PCR used herein is given in the experimental part. Other primers, enzymes and conditions can of course be applied.
  • a suitable method of identifying wanted alleles is a restriction endonuclease treatment. Suitable restriction enzymes for pig H-FABP alleles are Mspl, Haelll or Hinfl, but others may also be used. By these methods large numbers of pigs can be rapidly genotyped for studies in which genotypic variation can be associated with growth characteristics and other production traits of pigs.
  • the methods according to the invention can be developed into diagnostic assays or kits by which selection of pigs with alleles of interest can be performed in routine screening protocols employed in breeding programmes. With such protocols better results of selection can be expected when genes responsible for regulation of commercially interesting body tissues can be rapidly identified and controlled.
  • such testing protocols can be used to identify, select and breed farm animals, such as pigs, which have better production traits, such as IMF% or backfat thickness or average daily weight gain or feed efficiency, than the average animal in the population.
  • Better production traits such as BW or daily weight gain will increase the production per year expressed as amount of meat per animal raised.
  • a population of animals with a higher and less variable IMF% will result in a more homogenous product (meat) which is also better appreciated by putative customers because of a better taste.
  • selection for higher IMF% may be possible while at the same time selection against fat deposition in other depots, such as backfat, can be performed.
  • the porcine H-FABP gene has been isolated, characterized and chromosomally localized. Poylymorphisms in this gene have been identified. To test the association between bodyweight (BW) and percentage of intramuscular fat (IMF %), animals with different polymorphisms were selected, their bodyweight was measured and the amount of IMF after slaughter was measured.
  • BW bodyweight
  • IMF intramuscular fat
  • a porcine genomic DNA EMBL3/SP6/T7 library (Clontech Laboratories Inc. Palo Alto, CA) was screened using the plaque hybridization method (Sambrook et al. ,1989) .
  • plaques were transferred to replica nitrocellulose filters and incubated in denaturation buffer (1.5 M NaCl, 0.5 M NaOH) for 2 minutes, neutralisation buffer (1.5 M NaCl, 0.5 M Tris-HCl pH 8.0) for 5 minutes and fixation buffer (0.2 M Tris-HCl pH 7.5, 2X SSC(0.3 M NaCl, 0.03 M Sodium citrate)) for 30 s.
  • denaturation buffer 1.5 M NaCl, 0.5 M NaOH
  • neutralisation buffer 1.5 M NaCl, 0.5 M Tris-HCl pH 8.0
  • fixation buffer 0.2 M Tris-HCl pH 7.5, 2X SSC(0.3 M NaCl, 0.03 M Sodium citrate
  • the filters were prehybridized (6X SSC, 0.5% ( w/v) SDS, 5X Denhardt's and lOOug/ml NaOH treated salmon sperm DNA) for two hours at 67 ⁇ C and hybridized in identical buffer with the addition of the radioactively probe at 67°C overnight.
  • the filters were washed four times with 2X SSC, 0.1% ( w/v) SDS for 30 min at room temperature. Twenty plaques that showed positive signals on both replica filters were isolated and each subjected to two additional rounds of low density plaque purification. DNA of these clones was isolated using the plate lysate method (Sambrook et al.,1989).
  • PCR amplifications were performed on 1 ⁇ l of a 1:1000 dilution of phage DNA preparations or 50 ngr of genomic DNA in 50 ⁇ l containing 0.2 U Super Tth polymerase (SphaeroQ, Leiden, Nl) in 10 mM Tris-HCl pH 9.0, 50 mM KC1, 1.5 mM MgCI, 0.1% ( w/v) gelatin, 1% Triton X-100, 0.5 ⁇ M of each primer (Pharmacia Biotechnologies, Uppsala, Sweden) and 0.2 ⁇ M of each dNTP (Boehringer Mannheim, Mannheim, Germany) .
  • primer sequences as used for the poly-T microsatellite amplification are
  • PCR#1 (Table 1) was performed on DNA of all the phage clones to detect porcine H-FABP intron 1 specific fragments. Two of the three phage clones containing the H-FABP gene were used to subclone the various SacI and Kpnl (Boehringer Mannheim, Mannheim, Germany) restriction digestion fragments of the gene region in pBS. Unfortunately, neither of these phage clones contained exon 4 and the 3' untranslated region as detected by restriction analysis. The intron 3/exon 4 splice junction was cloned as PCR#2 (Table 1) product using porcine (Great England) genomic DNA as a template.
  • the products of two independent PCR reactions were cloned to identify errors by the Super Tth polymerase upon analysis.
  • the 3' untranslated region was isolated using the 5*/3' RACE-PCR kit (Boehringer Mannheim, Mannheim, Germany) with porcine (Meishan) muscle cDNA as the template and porcine H- FABP exon 1 or 3 (Table 1) specific primers in combination with the provided poly-A primer.
  • PCR products were cloned in the pT7Blue vector (Novagen Inc. ,Madison, U.S.A.).
  • H-FABP (sub)clones were transformed and the recombinant plasmid DNA was isolated and purified with the Wizard Maxiprep kit (Promega, Madison, WI, U.S.A.).
  • the nucleotide sequence was determined by dideoxy sequencing, partially by cycle sequencing (Perkin Elmer, ) or autoread sequencing (Pharmacia Biotechnologies, Uppsala, Sweden) and the analysis was performed on a ABI 373 (Applied Biosystems) or ALF DNA sequenator (Pharmacia Biotechnologies, Uppsala, Sweden) respectively.
  • Porcine genomic DNA was isolated as described (Sambrook et al.,1989) from EDTA treated blood stored at -80°C. One hundred ng of genomic DNA was used for PCR amplification in 50 ⁇ l reaction as described before. The primer sequences and its corresponding product size and annealing temperature for each combination are given in table 2. Fifteen ⁇ l of the PCR reaction was used for restriction digestion with 2 units of Haelll, Hinfl or Mspl (Boehringer Mannheim) in a total volume of 20 ul . For Haelll and Hinfl the recommended buffer conditions were additionally used whereas Mspl was added directly to the PCR buffer.
  • Restriction digestion fragments were loaded on a 2% ( ⁇ fspl) or 3% (Haelll and Hinfl ) agarose (Sigma, St Louis, MO, U.S.A.) gel and after electrophoresis the RFLP patterns were scored by two persons, independently.
  • FABP gene to a specific chromosome by PCR.
  • Duroc pigs were selected on the basis of the amount of IMF in their slaughtered relatives.
  • Performance traits recorded for each pig were live weight at 180 days of age (BW) , backfat thickness (BFT) and for each dam the number of piglets produced alive in first (FPP) or second parity (SPP), respectively.
  • meat quality traits i.e. cooking loss, drip loss, intamuscular fat percentage, minolta colour, pH and shear force were measured in a subset of the slaughtered animals.
  • H-FABP PCR-RFLP genotyping Blood or hairroots were collected from each animal to isolate genomic DNA for H-FABP PCR-RFLP genotyping.
  • the final dataset comprises information from in total 2345 pigs including pedigree.
  • H-PABP genotype information is available for at least one of the PCR-RFLPS.
  • P(XX), (Xx), (xx) are the estimated chance for each genotype for each animal .
  • exon-intron splice junctions were located in comparison with the porcine H-FABP cDNA and the murine H-FABP/MDGI (Treuner et al.,1994) gene sequence.
  • a potential TATA-box was located 92 bp upstream the ATG start codon and in the (3'UTR) a consensus poly-A signal sequence was identified (see Fig.l).
  • the coding sequences showed 92%, 91%, 87% and 85% identity to the bovine, human, mouse and rat H-FABP sequences at the nucleotide level and the deduced amino acid sequence were 92%, 90%, 87% and 86% identical, respectively (Billich et al. , 1988; Peeters et al. , 1991; Binas et al. , 1992; Claffey et al . , 1987).
  • a panel comprising genomic DNA of 7 pig breeds each represented by unrelated animals was used to detect genetic variation in the 5' upstream region, intron 2 and intron 3 of the porcine H-FABP gene. Therefore, part of the 5 ' upstream region was amplified on DNA of this panel using PCR (Table 2) and digested with the restriction enzyme Hinfl . The Hinfl digestion showed two alleles a single fragment of 256 bp (allele h) or two fragments of 197 and 59 bp (allele H) . Similarly intron 3 (PCR#3,Table 1) and intron 2 (Table 2) were tested for genetic variation with the enzymes ⁇ fspl and Haelll respectively, and both showed genetic variation in intron 2.
  • Haelll showed one fragment of 850 bp (allele D) and/or fragments of 400 and 450 bp (allele d) . Accurate size determination revealed that these three fragments were 684 bp, 278 bp and 406 bp. Afspl showed a fragment of 850 bp (allele a) and/or fragments of 750 and 50 bp (A) . Accurate size determination revealed that these fragments were 814 bp, 703 bp and 111 bp. Both sites of genetic variation are approximately 300 (285) bp apart.
  • Table 3 represents the allele frequencies of the different PCR-RFLPs in the different pig breeds tested.
  • the mendelian inheritance pattern of the three PCR- RFLPs was analysed in a porcine family comprising 3 generations of a Great Yorkshire breed. The genotypes of the individual pigs show consistent patterns of inheritance in this family.
  • the porcine H-FABP gene was chromosomally localized using a porcine H-FABP gene intron 3 specific PCR which amplified no rodent homologous. Amplification on DNA of two independently established pig/rodent cell hybrid panels and comparison with the cytogenetically (panel A and B) and reference loci data (panel A) revealed a single significant association of the H-FABP gene with chromosome 6 (Table 4) for both cell hybrid panels.
  • Table 5 shows the result of mean values and their standard deviations of IMF and bodyweight for different fixed effects which were taken into account in this analyses.
  • the PEST program was used to be able to use family information in the analyses of the different fixed effects.
  • the used model contained the same fixed effect as the model with SAS but also contains a random animal effect.
  • a pedigree file was used containing family relations up to two generations back. Table 7, 10, 11, 12 and 13 show predicted values for the different fixed effects and their standard errors .
  • the value tl-l/2 ⁇ is taken from a confidence table and has a value of 1.96 for a 95% confidence interval.
  • Table 8 shows the difference between different genotypes of each polymorphism and its 90% and 95% critical values.
  • Bodyweight is significantly different for the different genotypes in all three polymorphisms.
  • the genotypes of the three polymorphisms tested show a significant (95%) difference in bodyweight (BW) . All three polymorphisms can be used in selection for bodyweight.
  • the genotypes of the three polymorphisms show a distinct, albeit non-significant difference in IMF percentage. If there is a difference between different genotypes of 0.20, 50 animals of the least frequent genotype (AADDhh) would be needed to reach a significant (95%) difference of 0.2.
  • Tables 9, 10 and 11 show that when more animals are tested, statistically significant differences among the three polymorphisms can indeed be found, for instance for IMF, backfat thickness and BW. Also, tables 12 and 13 show that the effect on IMF, as measured by RFLP testing, can stil be found when the effects are corrected for backfat thickness and/or growth.
  • FIG. 1 The porcine H-FABP gene sequence including 1632 bp of the 5 'upstream region and 200 bp of the 3' untranslated region. Exons are represented by bold capital letters and the deduced amino acid sequence is shown directly beneath it. Standard one letter amino acid symbols are used.
  • the putative TATA-box, the polyadenylation signal in the 3'UTR and the 13 nucleotide element are underlined. The size of the nondepicted intron sequences is shown between arrowheads.
  • the polymorphic Haelll (GGCC), Hinfl (GATTC), Mspl (CAGG) sites and the polymorphic microsatellite sequence (poly-T) are depicted bold and underlined.
  • Heart fatty acid binding protein is a novel regulator of cardiac myocyte hypertrophy. Biochem. Biophys. Res. Comm. vol 205 no 3:1822-1828. Chevalet, C. and Corpet, F. (1986) Statistical decision rules concerning synteny or independence between markers . Cytogenet. Cell Genet. 43:132-139.
  • Table 1 Primer sequences and the corresponding size and annealing temperatures for PCR reactions of different regions of the porcine H-FABP gene.
  • Table 2 The primer sequences and combinations used for PCR- RFLP detection with the corresponding annealing temperature
  • PI Pietrain
  • WP Wild Pig.
  • the second line indicates the nuaber of unrelated animals tested.
  • Table 4 Chromosomal localization of the porcine hFABP gene by two independent cell hybrid panels.
  • the percentage concordance aa determined by the total o£ equal abaence/preaenc ⁇ in both data aeta divided by 20.
  • the ⁇ value r ⁇ preaenta the correlation between both dataaets.
  • a gene can be aaa gnad to a chromosome when the concordance s high and the ⁇ ⁇ a more than 0.74 (ayntanic) . Values far ⁇ between 0.59 and 0.74 g ve no validation of the assignment.
  • Panel A Rettenberger et a . , 1994a
  • Panel B Zijlstra et al., 1994a
  • N Number of animals (N) , their mean and standard deviations (std) for intramuscular fat (IMF) and bodyweight (BW) difference from 110 kg at 180 days) for different fixed effects (effect) .
  • IMF intramuscular fat
  • BW bodyweight
  • Table 6 Values of significance for different fixed effects influencing intramuscular fat (IMF) and bodyweight (BW) using a large model (lrg) and using seperate models per fixed effect (ind) (correcting for fixed effects test farm and sexe) .

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Abstract

La présente invention, qui concerne une séquence nouvellement découverte du gène H-FABP du porc, concerne également des procédés d'utilisation du gène considéré et de ses produits. L'invention concerne plus particulièrement des procédés de détection d'allèles différenciés du gène H-FABP du porc, lesquels allèles différenciés sont associés à des différences au niveau des caractéristiques du génotype et/ou du phénotype des porcs possédant ces allèles. L'invention concerne notamment des procédés permettant de faire la distinction entre allèles aboutissant à des phénotypes différents, en l'occurrence, grâce à des techniques d'amplification sélective de matériel génétique dérivé du gène H-FABP du porc. Ces techniques conviennent particulièrement à la sélection d'animaux dans le cadre de programmes de sélection. L'invention concerne enfin des programmes de sélection utilisant de telles techniques.
EP97914649A 1996-03-28 1997-03-27 Gene codant une proteine se liant a un acide gras du coeur de porc et procede d'identification des caracteristiques du polymorphisme responsables du poids du corps Withdrawn EP0889904A2 (fr)

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Application Number Priority Date Filing Date Title
EP97914649A EP0889904A2 (fr) 1996-03-28 1997-03-27 Gene codant une proteine se liant a un acide gras du coeur de porc et procede d'identification des caracteristiques du polymorphisme responsables du poids du corps

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Application Number Priority Date Filing Date Title
EP96200855 1996-03-28
EP96200855 1996-03-28
PCT/NL1997/000157 WO1997035878A2 (fr) 1996-03-28 1997-03-27 Gene codant une proteine se liant a un acide gras du coeur de porc et procede d'identification des caracteristiques du polymorphisme responsables du poids du corps
EP97914649A EP0889904A2 (fr) 1996-03-28 1997-03-27 Gene codant une proteine se liant a un acide gras du coeur de porc et procede d'identification des caracteristiques du polymorphisme responsables du poids du corps

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EP0889904A2 true EP0889904A2 (fr) 1999-01-13

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EP (1) EP0889904A2 (fr)
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CA (1) CA2256983A1 (fr)
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Cited By (1)

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CN108103207A (zh) * 2017-12-28 2018-06-01 中国农业科学院北京畜牧兽医研究所 Brca1、jaml及其调控基因在品种选育中的应用

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WO1998015837A1 (fr) * 1996-10-07 1998-04-16 Meat And Livestock Commission Methode de selection de fibres musculaires de type duroc
GB2333154B (en) * 1996-10-07 2001-05-09 Meat And Livestock Commission Assay for duroc muscle fibre tyre
EP1015634A1 (fr) * 1997-09-18 2000-07-05 Pig Genes B.V. I.O. Gene codant la proteine se liant a l'acide gras des adipocytes du porc, et procedes de localisation, d'identification ou marquage des genes ou alleles ou loci de caractere quantitatif des animaux de ferme
GB9929140D0 (en) 1999-12-10 2000-02-02 Univ Geneve Diagnostic assay for stroke
AU8898001A (en) 2000-09-08 2002-03-22 Univ Iowa State Res Found Inc Novel prkag3 alleles and use of the same as genetic markers for reproductive andmeat quality traits
CN101935706B (zh) * 2010-09-02 2012-05-30 中国农业科学院北京畜牧兽医研究所 一种检测猪肉品质性状的方法及专用引物对
CN113913536B (zh) * 2021-11-30 2022-08-16 湖北省农业科学院畜牧兽医研究所 猪eepd1基因第一外显子变异作为肌内脂肪含量性状的遗传标记及应用

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EP0580767B1 (fr) * 1991-04-19 2000-12-27 Iowa State University Research Foundation, Inc. Marqueurs genetiques de la taille de portees de porcs
DE4338817A1 (de) * 1993-11-13 1995-06-14 Max Delbrueck Centrum Gensequenzen von Cellular X Binding Proteinen

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Cited By (1)

* Cited by examiner, † Cited by third party
Publication number Priority date Publication date Assignee Title
CN108103207A (zh) * 2017-12-28 2018-06-01 中国农业科学院北京畜牧兽医研究所 Brca1、jaml及其调控基因在品种选育中的应用

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WO1997035878A2 (fr) 1997-10-02
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CA2256983A1 (fr) 1997-10-02
AU2180397A (en) 1997-10-17

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