Go to Top Go to Bottom
Asian-Australas J Anim Sci > Volume 33(11); 2020 > Article
Cahyadi, Park, Seo, Jin, Choi, Heo, Kang, Jo, and Lee: Association of the thyroid hormone responsive spot 14 alpha gene with growth-related traits in Korean native chicken

Abstract

Objective

Thyroid hormone responsive spot 14 alpha (THRSP) has been used to investigate the regulation of de novo lipogenesis because the variation of THRSP mRNA content in the tissue affects directly the ability of that tissue to synthetize lipids. Also, this gene responds to thyroid hormone stimulation and high level of carbohydrate feeding or insulin-injection. This study was carried out to investigate variations within THRSP and their effects on body and carcass weights in Korean native chicken (KNC).

Methods

A total of 585 chickens which represent the five lines of KNC (Black, Gray-Brown, Red-Brown, White, and Yellow-Brown) were reared and body weight data were recorded every two weeks from hatch until 20 weeks of age. Polymerase chain reaction-restriction fragment length polymorphism, DNA chips for Agilent 2100 Bioanalyzer, and Fluidigm Genotyping Technology, were applied to genotype selected markers. A linear mixed-effect model was used to access association between these single nucleotide polymorphism (SNP) markers and growth-related traits.

Results

A total of 30 polymorphisms were investigated in THRSP. Of these, nine SNPs for loci were selected to perform association analyses. Significant associations were detected between g.-49G>T SNP with body weight at 20 weeks of age (BW20), g.451T>C SNP with growth at 10 to 12 weeks of age (GR10-12), and g.1432A>C SNP with growth at 14 to 16 weeks trait (GR14-16) and body weight at 18 weeks of age (BW18). Moreover, diplotype of the THRSP gene significantly affected body weight at 12 weeks of age (BW12) and GR10-12 traits. Diplotype of ht1/ht2 was favorable for BW12 and GR10-12 traits.

Conclusion

These results suggest that THRSP can be regarded as a candidate gene for growth traits in KNC.

INTRODUCTION

Korean native chicken (KNC) is an important genetic resource in Korea. The KNC conservation program was launched by the National Institute of Animal Science in 1994 [1,2]. After the two decades of the conservation program’s duration, the five lines of KNC, which are distinguished by plumage color, have been established and successfully released in the Korean market and are 2 to 3 times more expensive than commercial broilers [3,4]. The advantages of KNC in Korean market are that they have higher protein and lower fat contents compared to commercial broiler. However, like most indigenous chicken breeds, KNC have a relatively slower growth performance that implies a longer rearing time [1,3]. The heritability value (h2) of growth traits of KNC is high at an early age and becomes low to moderate as the chickens age [5]. Therefore, breeding strategies based on either quantitative or molecular genetics to improve growth performance of KNC are required.
Body weight in chickens can be easily observed and measured [6]. However, it is a direct indicator of growth which is a complex, and highly orchestrated phenomenon involving multiple genetic factors and environmental conditions with a host of different hormones and their receptors in various tissues [7]. Hence, it is not surprising that most of growth-related traits are regarded as complex quantitative traits.
Regulation of body weight has been an interesting topic to investigate since it is tightly related to obesity, it requires an understanding of lipid metabolism and energy balance processes [811]. Lipid synthesis involves multiple genes in highly complicated and integrated processes since they will maximize the storage of energy when food nutrients are abundant and repress energy expenditure when nutrients are restricted. Thyroid hormone responsive spot 14 (THRSP) gene, which is widely studied as a model of de novo lipogenesis regulation, encodes a small acidic THRSP protein. This protein acts as a transcription factor involved in control of lipogenic enzymes since it responds to thyroid hormone stimulation [1214]. It is located in the chromosomal region affecting obesity susceptibility, and its expression is increased by carbohydrate feeding or insulin-injection [15]. In humans, THRSP is associated with obesity, growth and differentiation of breast cancer cells [16,17]. In chickens, THRSP was firstly analyzed by Cogburn et al [18] using microarray analysis that showed a differential expression of the THRSP sequence tag in liver of chickens genetically selected for fast or slow growth rate. It was mapped at 1q41–44 [19], where the quantitative trait locus (QTL) region for abdominal fat yield, skin fatness, and abdominal fatness were found [2022]. The polymorphisms within this gene have been investigated using polymerase chain reaction-restriction fragment length polymorphism (PCR-RFLP) approach which revealed that they were associated with growth-related and fatness traits in chicken F2 intercross pedigrees [23,24]. Hence, the objective of this study was to investigate polymorphisms of THRSP and to evaluate their effects on the growth-related traits in KNC.

MATERIALS AND METHODS

Animal resources and DNA extraction

A total of 585 individuals of KNC (282 males and 303 females) comprising 68 F1 nuclear families ranging in size from 3 to 20 chickens were used in this study. They were divided into five lines on the basis of plumage colors (Black, 88; Gray-Brown, 110; Red-Brown, 135; White, 122; and Yellow-Brown, 130) [25]. They were reared under controlled feeding and environmental conditions in the National Institute of Animal Sciences (NIAS), Republic of Korea. This study was also performed to meet recommendations described in “The Guide for the Care and Use of Laboratory Animals” published by the Institutional Animal Care and Use Committee of the National Institute of Animal Science (2012-C-037) in Korea. Body weights were measured at every two weeks from hatch to 20 weeks of age. Descriptive statistics of body weight and growth-related traits in the KNC population can be found elsewhere [26].
For the molecular genetic analysis, blood samples were taken from wing veins using 3 mL tubes containing ethylenediaminetetraacetic acid. Moreover, these blood samples were processed to isolate genomic DNA according to the method described by Miller et al [27]. The DNA concentration was measured using NanoDrop 2000c UV-Vis Spectrophotometer (Thermo Fisher Scientific Inc., Waltham, MA, USA). Isolated DNA genome was stored in the refrigerator at −20°C to maintain DNA quality until used.

Polymerase chain reaction and genotyping

Three pairs of primer sets and restriction enzyme information used in this study are shown in Supplementary Table S1. Two primers were designed using primer3 version 0.4.0 (http://frodo.wi.mit.edu/primer3/) based on GenBank reference NC_006088.3, and a primer pair for 9-bp indel detection that was previously reported by Wang et al [15]. The PCR was carried out in 20 μL volume containing 25 ng per μL DNA genome, 0.8 μL each primer, 1.6 μL 10 mM dNTP, 2.0 μL 10× reaction buffer, 0.2 μL Hot Start Taq Polymerase (GenetBio, Daejeon, Korea), and 12.6 μL double distilled water. The PCR reactions were carried out in the following steps: pre-denaturation at 94°C for 10 minutes, 35 cycles of 94°C for 30 seconds, annealing temperature (°C) for each primer for 30 seconds (Supplementary Table S1), and 72°C for 30 seconds, and followed by final extension at 72°C for 10 minutes. Reaction was performed using either GeneAmp PCR system 2700 (Applied Biosystems, Foster City, CA, USA) or C1000TM Thermal Cycler (BioRad, Hercules, CA, USA). The PCR products were visualized by 2% standard agarose gels stained with ethidium bromide (GenetBio, Korea). Initially, DNA pools containing 3 individual samples were used for PCR and sequencing to screen polymorphisms of THRSP. The PCR-RFLP was applied to confirm 3 single nucleotide polymorphism (SNPs) and a 3-bp indel. Approximately, 15 μL of PCR product was digested with 2 units of each restriction enzyme (Supplementary Table S1) based on the protocol provided by company (New England Biolabs Inc., Ipswich, MA, USA). Furthermore, the digested PCR product was separated on 3% agarose gels to identify genotype variations (Supplementary Figure S1). The 9-bp indel was confirmed using DNA Chip for The Agilent 2100 Bioanalyzer (Agilent Technologies Inc., Santa Clara, CA, USA) by following company protocol (Supplementary Figure S2). In addition, eleven of SNPs were genotyped using standard protocol of Fluidigm 192.24 SNPtype Genotyping Technology (Fluidigm, South San Francisco, CA, USA).

Single marker association analysis

The phenotypic data of growth-related traits from the 585 F1 progeny were in normal distribution (Data not shown). Single marker association analyses between the selected DNA markers and the traits using the F1 progeny was conducted using the following linear mixed-model:
Yijklmn=μ+Gi+Sj+Bk+Ll+Pm+ɛijklmn
where, Yijklmn is the phenotype of the nth animal, μ is overall mean, Gi is the fixed effect of THRSP genotype i, Sj is the fixed effect of sex j, Bk is the fixed effect of batch k, Ll is the fixed effect of line l, and Pm is random additive polygenic effect of animal m, ɛijklmn is the random residual associated with the nth animal. The mean and variance for random additive polygenic effects can be defined as: P~N(0, Aσa2), where A is based on relationship matrix computed from the nuclear pedigree in this study and σa2 is the additive polygenic variance. The mean and variance for the residual random effect of individuals can be defined as: ɛ~N(0, Iσe2), where I is the identity matrix and σe2 is the residual variance.
The significance level of fixed effects in the mixed-effects model above was computed by the Wald test in ASReml program (VSN International, Hemel Hempstead, UK). Nominal p<0.05 was regarded as significant for all tests.

Haplotype-based association analysis

The F1 progeny were used to establish haplotypes at THRSP locus using the SNP markers showing nominal significance (p<0.05) for growth-related traits (Figure 1). The FImpute program based on pedigree information was applied to establish the haplotypes (REF. BMC Genomics 15, 478, 2014). Subsequently, the haplotype data of the F1 chickens were used to conduct haplotype-based association study. The effect of haplotypes on the traits of interests were evaluated by the following general linear model using the MINITAB program (MINITAB, State College, PA, USA):
Yijklmn=μ+Di+Sj+Bk+Ll+Dam(l,m)+ɛijklmn
Yijklmn is the phenotype of the nth animal, μ is overall mean, Di is the fixed effect of THRSP diplotype i (Table 1), Sj is the fixed effect of sex j, Bk is the fixed effect of batch k, Ll is the fixed effect of line l, and Dam(l,m) is fixed effect of mth dam within lth line, ɛ is the random residual term of the model.

RESULTS

Detection of polymorphisms in the THRSP gene

Five sequences of THRSP were deposited in the National Center for Biotechnology Information (NCBI) website (GenBank accession numbers: KF574271-KF574275). A total of 30 polymorphisms were identified by aligning the sequences using ClustalW2 online software (http://www.ebi.ac.uk/Tools/msa/clustalw2/). These variations were spread from upstream to downstream of the gene. Eleven SNPs and an 3-bp indel at 1,355 bp from ATG were found in the flanking regions and untranslated regions of the THRSP gene, thirteen SNPs and an 3-bp indel at 965 bp from ATG were identified in intron, and three SNPs were also observed in the exon 1 of the THRSP gene. In addition, an 9-bp indel at 236 bp from start codon in the exon region of the THRSP gene which is identified in previous studies to be associated with body weight and abdominal fat traits in chickens was detected in present study (Table 1).

The effects of the THRSP polymorphisms on growth-related traits

A total of 30 polymorphisms in the THRSP gene have been identified in this study. Of these, nine polymorphisms were investigated to be associated with the body weight traits using a mixed-effects model in 585 F1 KNC progeny. Among the SNP markers listed in the Table 1, a genotype call rate less than 90% was not used for further association studies. Three SNPs showed significant association with growth-related traits (Figure 1). The g.−49G>T SNP of the THRSP gene was significantly associated with body weight at 20 weeks of age (BW20). Homozygote GG population had a higher BW20 than other genotypes. A SNP, namely g.451T>C, was also significantly associated with growth at 10 to 12 weeks (GR10–12). Moreover, g.1432A>C was statistically associated with growth 14 to 16 weeks and body weight at 18 weeks. The CC genotype of the g.1432A>C SNP was favorable for those traits. Additionally, four SNPs, namely g.−49G>T, g.696G>C, g.1419A>C, and g.1432A>C, were suggestively associated with some growth-related traits in F1 KNC population (Figure 2).
We also constructed haplotypes of the THRSP gene to evaluate the effect of the THRSP haplotype on growth-related phenotypes in the F1 chicken population. Haplotype and diplotype frequencies of the THRSP gene in the F1 chickens are listed in the Table 2. The results of haplotype-based association analysis indicated that there were significant association between the diplotype and BW12. Ht1/ht2 diplotype was clearly favorable for BW12 (Table 3). Additionally, significant association of the diplotype with GR10–12 was detected in the F1 animals. Ht1/ht2 diplotype was also highest in growth at 10 to 12 weeks of age. Ht4/ht4, where the most KNC population included in, was relatively moderate to high BW12 and GR10–12 traits (Table 3).

DISCUSSION

In this study, nine novel SNPs and a 9-bp indel reported previously were used for the association analyses. Interestingly, the genotypic value of the heterozygotes of the markers in Figure 1 and 2 showed dominance effects (i.e., deviation from the mean value of the two homozygotes) [28]. Therefore, no SNP with significant additive effect was detected in this study.
Significant associations were detected between g.−49G>T SNP, g.451T>C, and g.1432A>C with some growth-related traits in KNC. Those SNPs were located at 5′-flanking region, intron, and 3′-flanking region of THRSP gene. No exonic SNP showed significant association with growth related traits. Hence, expression levels might play an important role in the phenotypic variation of chicken growth. After transcription factor binding motif analysis using MEME suite [29] and JASPAR database [30], we revealed that the position of g.−49G>T SNP is co-localized with MEF2D (myocyte enhancer factor-2D) binding motif sequence in the 5′-flanking region of the THRSP gene (i.e., g.−52~g.−47). Therefore, further expression studies can be worthwhile to characterize g.−49G>T SNP. Association results of the three significant SNPs and diplotypes indicated their possible role in controlling of body weight and growth traits of mature KNC.
Previous works showed that polymorphisms of the THRSP were associated with body weight, carcass traits, fat deposition, cholesterol and lipoprotein [15,23,24]. Particularly, A213C SNP was associated with body weight at 5 to 12 weeks of age and carcass weight, and two SNPs in 5′-flanking region were associated with body weight at hatch and BW at 28 days of age [23,24]. Indeed, the stronger effect of 9-bp indel has previously been discovered to be affecting body weight at 8 to 12 weeks of age and also carcass weight in chickens [23]. However, they were not confirmed in KNC population, except for an 9-bp indel in the exon region. There are several possible explanations for this discrepancy. First, the previously identified association may not be a direct effect of the polymorphisms but rather the result of linkage disequilibrium. Second, the effect of the polymorphisms on the body weight traits may be too small to be detected in our F1 nuclear families. In the case of previously reported associations found in chicken F2 intercross pedigrees [23,24], they only used the general linear model procedure without including an additive relationship matrix to access the association between THRSP and body weight traits.
In conclusion, three novel SNPs in the THRSP gene were associated with body weight and growth-related traits in KNC. These results provided additional support for association between polymorphisms in the THRSP and growth-related traits. It could be useful to be developed as candidate gene for growth traits in KNC population.

Notes

CONFLICT OF INTEREST

We certify that there is no conflict of interest with any financial organization regarding the material discussed in the manuscript.

ACKNOWLEDGMENTS

This work was supported by the Korea Institute of Planning and Evaluation for Technology in Food, Agriculture, Forestry and Fisheries through the Golden Seed Project, Ministry of Agriculture, Food and Rural Affairs (213010-05-4-SB250) and the “Cooperative Research Program for Agriculture Science & Technology Development (PJ0128202020)” of the Rural Development Administration, Republic of Korea.

Figure 1
The genotypic means and standard errors of growth-related traits represent significant associations between the genotypes of the THRSP gene and growth-related traits. p values were calculated using a mixed-effects model analysis. BW00 is body weight at birth; BW02 is body weight at 2 weeks of age; BW04 is body weight at 4 weeks of age; BW06 is body weight at 6 weeks of age; BW08 is body weight at 8 weeks of age; BW10 is body weight at 10 weeks of age; BW12 is body weight at 12 weeks of age; BW14 is body weight at 14 weeks of age; BW16 is body weight at 16 weeks of age; BW18 is body weight at 18 weeks of age; BW20 is body weight at 20 weeks of age; GR0–2 is body weight gain at 0 to 2 weeks of age; GR2–4 is body weight gain at 2 to 4 weeks of age; GR4–6 is body weight gain at 4 to 6 weeks of age; GR6–8 is body weight gain at 6 to 8 weeks of age; GR8–10 is body weight gain at 8 to 10 weeks of age; GR10–12 is body weight gain at 10 to 12 weeks of age; GR12–14 is body weight gain at 12 to 14 weeks of age; GR14–16 is body weight gain at 14 to 16 weeks of age; GR16–18 is body weight gain at 16 to 18 weeks of age; GR18–20 is body weight gain at 18 to 20 weeks of age.
ajas-19-0541f1.jpg
Figure 2
The genotypic means and standard errors of growth-related traits represent suggestive-significant associations between the genotypes of the THRSP gene and growth-related traits. p values were calculated using a mixed-effects model analysis. THRSP, thyroid hormone responsive spot 14 alpha gene.
ajas-19-0541f2.jpg
Table 1
Distribution of the THRSP gene polymorphism in Korean native chicken
No. SNP name SNP ID in chicken genome1) Location SNP information
1. g.−49G>T rs14937292 5′-flanking region Novel
2. g.−17G>A rs738325925 5′-flanking region Novel
3. g.−1T>C rs316023211 5′-flanking region Novel
4. g.11A>G rs740327674 5′-UTR Novel
5. g.17T>C rs14937290 5′-UTR Novel
6. g.128C>T rs735083064 Exon Novel
7. g.137A>G (c. 114A>G) rs314316303 Exon Novel
8. g.235A>C rs733948711 Exon Novel
9. 9 bp indel at 236 bp from ATG - Exon Wang et al [15]; Cao et al [23]; D’Andre Hirwa et al [24]
10. g.451T>C rs314618339 Intron Novel
11. g.489T>G rs1059169337 Intron Novel
12. g.696G>C rs316849944 Intron Novel
13. g.751C>T rs1057579406 Intron Novel
14. g.774G>A rs313687968 Intron Novel
15. g.785C>T rs734655240 Intron Novel
16. g.800A>C rs738084821 Intron Novel
17. g.814C>G rs315982665 Intron Novel
18. g.818G>C rs312478376 Intron Novel
19. g.868A>C rs314685615 Intron Novel
20. g.899G>A rs316757568 Intron Novel
21. g.911C>A rs1059055828 Intron Novel
22. 3 bp indel at 965 bp from ATG rs1058294719 Intron Novel
23. g.1072C>T rs1060381112 Intron Novel
24. g.1208C>T rs312815759 3′-UTR Novel
25. g.1232A>G rs1058578106 3′-UTR Novel
26. g.1357T>C rs317692975 3′-UTR Novel
27. 3 bp indel at 1355 bp from ATG rs1060155017 3′-UTR Novel
28. g.1393G>A rs315260792 3′-UTR Novel
29. g.1419A>C rs316843064 3′-UTR Novel
30. g.1432A>C rs313275625 3′-flanking region Novel

THRSP, thyroid hormone responsive spot 14 alpha; SNP, single nucleotide polymorphism.

1) SNP ID in the chicken genome was collected from Ensembl website (http://asia.ensembl.org/).

Table 2
Haplotype and diplotype frequencies of the THRSP gene in the F1 chickens
Haplotype1) Number Frequency2) Diplotype Number Frequency
G-C-C (ht1) 124 0.16 ht1/ht1 53 0.137
G-T-A (ht2) 95 0.122 ht1/ht2 18 0.046
T-T-C (ht3) 32 0.041 ht3/ht3 16 0.041
T-T-A (ht4) 447 0.577 ht4/ht4 196 0.505
G-T-C (ht5) 28 0.036 ht2/ht4 32 0.082
T-C-C (ht6) 50 0.064 ht5/ht6 28 0.072
ht4/ht2 23 0.059
ht6/ht2 22 0.057

THRSP, thyroid hormone responsive spot 14 alpha.

1) The 1st locus is THRSP g.−49G>T, the 2nd locus is THRSP g.451T>C, the 3rd locus is THRSP g.1432A>C.

2) The haplotypes with frequency less than 0.03 are dropped in this table.

Table 3
Effects of diplotype of the THRSP gene on BW12 and GR10–12 (LS means±SE)
Diplotype BW121) p-value GR10–122) p-value
ht1/ht1 877.4±39.19ab 0.028 142.9±27.59a 0.007
ht1/ht2 1,024.9±44.63b 278.3±31.41c
ht3/ht3 1,001.1±55.19ab 278.3±38.85abc
ht4/ht4 997.0±17.16ab 248.2±12.08bc
ht2/ht4 948.9±39.46ab 246.0±27.78abc
ht5/ht6 934.1±58.13ab 169.7±40.92abc
ht4/ht2 1,001.9±46.17ab 226.3±32.50abc
ht6/ht2 876.4±42.10a 155.8±29.64ab

THRSP, thyroid hormone responsive spot 14 alpha; SE, standard error.

1) BW12, body weight at 12 weeks of age.

2) GR10–12, growth at 10 to 12 weeks of age.

a–c LS means with different superscripts within same column are significantly different at p<0.05.

REFERENCES

1. Cahyadi M, Seo DW, Jin S, et al. Association of SNPs in ODC and PRDM16 with body weight traits in Korean native chicken. Korean J Poult Sci 2013; 40:157–62. https://doi.org/10.5536/KJPS.2013.40.2.157
crossref pdf
2. Seo DW, Hoque MR, Choi NR, et al. Discrimination of Korean native chicken lines using fifteen selected microsatellite markers. Asian-Australas J Anim Sci 2013; 26:316–22. https://doi.org/10.5713/ajas.2012.12469
crossref pmid pmc pdf
3. Sang BD, Hong SK, Kim HK, et al. Estimation of genetic parameters for economic traits in Korean native chickens. Asian-Australas J Anim Sci 2006; 19:319–23. https://doi.org/10.5713/ajas.2006.319
crossref pdf
4. Hoque MR, Lee SH, Jung KC, et al. Discrimination of Korean native chicken populations using SNPs from mtDNA and MHC polymorphisms. Asian-Australas J Anim Sci 2011; 24:1637–43. https://doi.org/10.5713/ajas.2011.11144
crossref pdf
5. Manjula P, Park HB, Seo D, et al. Estimation of heritability and genetic correlation of body weight gain and growth curve parameters in Korean native chicken. Asian-Australas J Anim Sci 2018; 31:26–31. https://doi.org/10.5713/ajas.17.0179
crossref pmid pdf
6. Cahyadi M, Jo C, Lee JH. Quantitative trait loci and candidate genes for the economic traits in meat-type chicken. Worlds Poult Sci J 2014; 70:329–42. https://doi.org/10.1017/S0043933914000348
crossref
7. Spencer GSG. Hormonal systems regulating growth. A review. Livest Prod Sci 1985; 12:31–46. https://doi.org/10.1016/0301-6226(85)90038-7
crossref
8. Friedman JM, Halaas JL. Leptin and the regulation of body weight in mammals. Nature 1998; 395:763–70. https://doi.org/10.1038/27376
crossref pmid pdf
9. Barsh GS, Farooqi IS, O’Rahilly S. Genetics of body-weight regulation. Nature 2000; 404:644–51. https://doi.org/10.1038/35007519 https://doi.org/10.1038/35007519
crossref pdf
10. Schutz Y. Dietary fat, lipogenesis and energy balance. Physiol Behav 2004; 83:557–64. https://doi.org/10.1016/j.physbeh.2004.09.015
crossref pmid
11. Fall T, Ingelsson E. Genome-wide association studies of obesity and metabolic syndrome. Mol Cell Endocrinol 2014; 382:740–57. https://doi.org/10.1016/j.mce.2012.08.018
crossref pmid
12. Seelig S, Liaw C, Towle HC, Oppenheimer JH. Thyroid hormone attenuates and augments hepatic gene expression at a pretranslational level. Proc Natl Acad Sci USA 1981; 78:4733–7. https://doi.org/10.1073/pnas.78.8.4733
crossref pmid
13. Kinlaw WB, Church JL, Harmon J, Mariash CN. Direct evidence for a role of the “spot14” protein in the regulation of lipid synthesis. J Biol Chem 1995; 270:16615–8. https://doi.org/10.1074/jbc.270.28.16615
crossref pmid
14. Jump DB, Narayan P, Towle H, Oppenheimer JH. Rapid effects of triiodothyronine on hepatic gene expression: Hybridization analysis of tissue-specific triiodothyronine regulation of mRNAS14. J Biol Chem 1984; 259:2789–97.
pmid
15. Wang X, Carre W, Zhou H, Lamont SJ, Cogburn LA. Duplicated Spot 14 genes in the chicken: characterization and identification of polymorphisms associated with abdominal fat traits. Gene 2004; 332:79–88. https://doi.org/10.1016/j.gene.2004.02.021
crossref pmid
16. Chagnon YC, Perusse L, Bouchard C. The human obesity gene map: the 1997 update. Obes Res 1998; 6:76–92. https://doi.org/10.1002/j.1550-8528.1998.tb00318.x
crossref pmid
17. Donnelly C, Olsen AM, Lewis LD, Eisenberg BL, Eastman A, Kinlaw WB. Conjugated linoleic acid (CLA) inhibits expression of the Spot 14 (THRSP) and fatty acid synthase genes and impairs the growth of human breast cancer and liposarcoma cells. Nutr Cancer 2008; 61:114–22. http://dx.doi.org/10.1080/01635580802348666
crossref
18. Cogburn LA, Wang X, Carre W, et al. Systems–wide chicken DNA microarrays, gene expression profiling, and discovery of functional genes. Poult Sci 2003; 82:939–51. https://doi.org/10.1093/ps/82.6.939
crossref pmid pdf
19. Carre W, Diot C, Fillon V, et al. Development of 112 unique expressed sequence tags from chicken liver using an arbitrarily primed reverse transcriptase–polymerase chain reaction and single strand conformation gel purification method. Anim Genet 2001; 32:289–97. https://doi.org/10.1046/j.1365-2052.2001.00792.x
crossref
20. Ikeobi CON, Woolliams JA, Morrice DR, et al. Quantitative trait loci affecting fatness in the chicken. Anim Genet 2002; 33:428–35. https://doi.org/10.1046/j.1365-2052.2002.00911.x
crossref pmid
21. Lagarrigue S, Pitel F, Carre W, et al. Mapping quantitative trait loci affecting fatness and breast muscle weight in meat–type chicken lines divergently selected on abdominal fatness. Genet Sel Evol 2006; 38:85 https://doi.org/10.1186/1297-9686-38-1-85
crossref
22. Ankra-Badu GA, Shriner D, Le Bihan-Duval E, et al. Mapping main, epistatic and sex–specific QTL for body composition in a chicken population divergently selected for low or high growth rate. BMC Genomics 2010; 11:107 https://doi.org/10.1186/1471-2164-11-107
crossref
23. Cao ZP, Wang SZ, Wang QG, Wang YX, Li H. Association of spot14 alpha gene polymorphisms with body weight in the chicken. Poult Sci 2007; 86:1873–80. https://doi.org/10.1093/ps/86.9.1873
crossref pmid pdf
24. d’Andre Hirwa C, Yan W, Wallace P, et al. Effects of the thyroid hormone responsive spot14α gene on chicken growth and fat traits. Poult Sci 2010; 89:1981–91. https://doi.org/10.3382/ps.2009-00582
crossref pdf
25. Hoque MR, Lee SW, Lee JH. DNA markers in chicken for breed discrimination. CNU J Agric Sci 2012; 39:211–7. https://doi.org/10.7744/cnujas.2012.39.2.211
crossref pdf
26. Cahyadi M, Park HB, Seo DW, et al. Variance component quantitative trait locus analysis for body weight traits in purebred Korean native chicken. Asian-Australas J Anim Sci 2016; 29:43–50. https://doi.org/10.5713/ajas.15.0193
crossref pmid pdf
27. Miller SA, Dykes DD, Polesky HF. A simple salting out procedure for extracting DNA from human nucleated cells. Nucleic Acids Res 1988; 16:1215 https://doi.org/10.1093/nar/16.3.1215
crossref pmid pmc pdf
28. Falconer DS, Mackay TFC. Introduction to quantitative genetics. 4th edHarlow, UK: Addison-Wesley Longman; 1996.

29. Bailey TL, Boden M, Buske FA, et al. MEME SUITE: tools for motif discovery and searching. Nucleic Acids Res 2009; 37:W202–8. https://doi.org/10.1093/nar/gkp335
crossref pmid pmc pdf
30. Khan A, Fornes O, Stigliani A, et al. JASPAR 2018: update of the open-access database of transcription factor binding profiles and its web framework. Nucleic Acids Res 2018; 46:D260–6. https://doi.org/10.1093/nar/gkx1126
crossref pmid pdf


Editorial Office
Asian-Australasian Association of Animal Production Societies(AAAP)
Room 708 Sammo Sporex, 23, Sillim-ro 59-gil, Gwanak-gu, Seoul 08776, Korea   
TEL : +82-2-888-6558    FAX : +82-2-888-6559   
E-mail : jongkha@hotmail.com               

Copyright © 2020 by Asian-Australasian Journal of Animal Sciences. All rights reserved.

Close layer
prev next